Zoogeography of Arachnida

Monographiae Biologicae 94
Petar Beron
Zoogeography
of Arachnida
Monographiae Biologicae
Volume 94
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Petar Beron
Zoogeography of Arachnida
Petar Beron
National Museum of Natural History,
Bulgarian Academy of Sciences
Sofia, Bulgaria
ISSN 0077-0639 ISSN 2215-1729 (electronic)

Monographiae Biologicae
ISBN 978-3-319-74417-9 ISBN 978-3-319-74418-6 (eBook)
https://doi.org/10.1007/978-3-319-74418-6
Library of Congress Control Number: 2018932984
© Springer International Publishing AG, part of Springer Nature 2018

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E. Haeckel, Kunstformen der Natur, 1904
“One of the primary suppositions of sciences
is that world is orderly, that there are
patterns, and that patterns need causal
explanation. The same can be said of the
science of historical biogeography.”
J. Cracraft (1988)
To the memory of Vassil B. Guéorguiev
Bulgarian entomologist, biospeleologist, and
zoogeographer, for our friendship and his
ideas
Contents
1 Introduction�� 1
2 Builders of Arachnology�� 7
Bibliography�� 33
3 Systems of Arachnida�� 35
4 The Fossil Arachnida �� 45
5 Factors Determining the Distribution of Arachnida�� 59
5.1 Some Basic Notions�� 59
5.1.1 Zoogeography and Biogeography:
The New Approach�� 59
5.1.2 Centers of Origin and Dispersal�� 60
5.1.3 Development of Atlantic Ocean�� 61
5.1.4 Development of the Indian Ocean �� 64
5.2 Paleogeography and the Past Distribution of Arachnida�� 65
5.2.1 Age of the Faunas�� 65
5.3 Passive Dispersal and Phoresy of Arachnida: Ballooning,
Rafting, and Continental Drift�� 72
5.3.1 Dispersal (Natural and by Man Activities)�� 72
5.4 Ecological Factors�� 76
5.4.1 Climatic and Other Barriers and Bridges�� 76
5.4.2 Real and “Fake” Disjunctions�� 77
5.4.3 Old World-New World Disjunctions�� 79
5.4.4 West African-Indomalayan Disjunction�� 81
5.4.5 Strange (Old?) Disjunctions�� 81
5.4.6 Disjunctions in the Southern End of the World
(New Zealand, Australia, South Africa, Patagonia)�� 81
5.4.7 Boreomontane and Arctic-Alpine Disjunctions�� 81
5.4.8 Transpacific Disjunctions�� 82
xi
xii Contents
5.5 Endemics and Relicts in Arachnida�� 83

5.5.1 Endemism�� 83
5.5.2 Relicts and Relictness�� 94
5.6 The Northern Limits of Arachnida�� 97
6 Review of the Classification of the Orders of Arachnida �� 135
6.1 Palpigradi�� 135
6.2 Schizomida�� 136
6.3 Thelyphonida (Uropygi)�� 138
6.4 Amblypygi�� 140
6.5 Ricinulei�� 143
6.6 Scorpiones �� 144
6.7 Pseudoscorpiones�� 146
6.8 Opiliones �� 149
6.9 Solifugae (Solpugida, Solifugi)�� 159
6.10 Araneae �� 170
6.11 Opilioacarida �� 185
6.12 Trombidiformes�� 189
6.13 Sarcoptiformes�� 193
6.14 Holothyrida �� 194
6.15 Ixodida�� 198
6.16 Mesostigmata�� 199
7 Regional Arachnogeography�� 319
7.1 Holarctic�� 319
7.1.1 Zoogeography and Arachnozoogeography�� 319
7.2 Europe (General)�� 320
7.2.1 Geography, General Zoogeography,
and Paleogeography�� 320
7.2.2 Arachnogeography�� 323
7.3 Balkan Peninsula and Aegaeis�� 328
7.4 Apennine Peninsula�� 342
7.5 Iberian Peninsula and the Pyrenees�� 344
7.6 Mediterranean Sea and Its Islands�� 348
Contents xiii
7.7 Balearic Islands �� 350

7.8 Sicily, Sardinia, Corsica, and Elba�� 352
7.9 Malta �� 357
7.10 Central and Northern Europe, Great Britain, Ireland,
Island, and Faroe Islands�� 359
7.11 Arachnida of the Atlantic Islands
(Macaronesia, St. Helena, Ascension)�� 363
7.11.1 Canary Islands, Azores, and Madeira�� 363
7.12 Cabo Verde�� 367
7.13 St. Helena and Ascension�� 368
7.13.1 Geography and Paleogeography (General)�� 368
7.14 Russia (North of Caucasus), Belarus, Ukraine,
Siberia, Altai, China (North of Yangtze), and Mongolia�� 370
7.15 Sakhalin and Kuril Islands�� 374
7.16 Caucasus and Transcaucasia �� 375
7.17 Turkey, Cyprus, Sinai, the Dodecanese Islands, Iran,
Afghanistan, the Near East, Middle Asia,
and the Arabian Peninsula�� 379
7.17.1 Geography, General Zoology, and Paleogeography�� 379
xiv Contents
7.18 Socotra�� 391

7.19 Karakorum, Hindu Kush, Pamir, Tian Shan, Himalaya,
and Tibet�� 393
7.20 Indian Peninsula�� 397
7.21 Ceylon (Sri Lanka)�� 400
7.22 Andaman and Nicobar Islands�� 403
7.23 Southeast Asia (Burma, Thailand, Indochina,
Peninsular Malaysia, China South of Yangtze, and Hainan)�� 404
7.24 Taiwan �� 421
7.25 The Malay Archipelago�� 423
7.26 The Philippines�� 431
7.27 Japan (the Ryukyus Excluded)�� 435
7.28 Ryukyu Islands�� 438
Contents xv
7.29 Korea (North and South) �� 441

7.30 Africa, North of 20oN�� 444
7.31 Africa Between 20oN and Zambezi-Kunene
(Tropical or Intertropical Africa)�� 453
7.32 Differences Between the Arachnofaunas
of East and West Africa�� 475
7.33 Tropical South America�� 476
7.34 Galapagos�� 502
7.35 Juan Fernandez Islands �� 505
7.36 North America, North of Rio Grande (incl. Greenland) �� 506
7.37 Polynesia and Micronesia �� 514
7.38 Hawaii �� 519
7.39 Antarctic and Subantarctic Arachnofauna�� 522
xvi Contents
8 Some Peculiarities of the Distribution of Arachnida�� 625

8.1 Mediterranean and Central European Arachnofauna�� 625
8.2 Arachnofauna of the Ancient Mediterranean�� 625
8.3 The Boundary Neotropica-Nearctic�� 626
8.3.1 Palpigradi�� 626
8.3.2 Ricinulei�� 627
8.3.3 Pseudoscorpiones�� 627
8.3.4 Opiliones�� 627
8.3.5 Amblypygi �� 628
8.3.6 Uropygi (Thelyphonida)�� 628
8.3.7 Schizomida�� 628
8.3.8 Araneae�� 628
8.3.9 Opilioacarida �� 629
8.4 Mexico: The Great American Interchange of Fauna �� 629
8.5 Arachnida of the Antilleans and Mesoamerica �� 639
8.5.1 Central America�� 639
8.6 Antillean (Caribbean) Islands �� 650
8.6.2 Comparison Between the Faunas of Central
America and the Antilleans �� 669
8.6.3 Conclusion�� 673
8.6.4 Opiliones�� 673
8.7 Patagonia: South Chile, Argentina, Uruguay, and Tierra
del Fuego�� 674
8.7.2 Arachnogeographical Character of South Chile
and Argentina�� 675
8.8 Falkland Islands (Malvinas)�� 679
8.9 The Boundary Between Palearctic and Paleotropical �� 680
8.9.1 The Boundary Between Western Palearctic
and Afrotropical Arachnofaunas�� 680
8.10 Arachnida in the Afrotropical and Palearctic Regions�� 681
8.10.1 Analysis and Comments�� 681
8.11 The Boundary Across China �� 688
8.11.1 The Transitional Zone�� 688
8.11.2 Arachnogeography of Himalaya�� 689
8.11.3 East Asiatic (Himalayan-Chinese) Region
of Lopatin (1980)�� 690
8.12 The Cape Arachnofauna�� 690
8.12.1 Africa South of Zambezi-Kunene�� 690
Contents xvii
8.13 Arachnofauna of Madagascar and the Adjacent Islands�� 701

8.14 Madagascar �� 701
8.15 Mascarene Islands �� 711
8.16 Seychelles (incl. Aldabra), Comores�� 713
8.16.2 Remarkable Distributions and Absences�� 726
8.17 Evaluation of Wallace’s Line and the Other Lines
in Southeast Asia According to the Arachnida�� 727
8.17.1 Wallace’s Line �� 728
8.17.2 Lydekker’s Line �� 728
8.17.3 Weber’s Line�� 728
8.17.4 Development of Wallacea and the Surrounding
Lands of Southeast Asia and the Malayan
Archipelago �� 730
8.17.5 The “Lines” and Some Other Groups�� 731
8.17.6 Arachnida and the Lines�� 732
8.17.7 Conclusions �� 743
8.18 The Arachnida of the Papuan Subregion�� 746
8.18.1 New Guinea and Melanesia�� 746
8.19 Arachnogeography of Australia and Tasmania �� 753
8.19.1 Australia�� 753
8.20 Tasmania�� 764
8.21 Parasitiformes (Some Ixodida and Mesostigmata):
Australia and New Zealand �� 767
8.22 New Zealand �� 768
8.23 New Caledonia�� 774
8.24 Lord Howe Island�� 777
8.25 Conclusion�� 778
8.25.1 Conclusion�� 782
9 Cave Arachnida�� 833
9.1 Cave Life in Different Orders �� 833
9.2 Distribution of Cave Arachnida in Different
Geographical Areas �� 842
xviii Contents
9.3 Conclusions �� 846

10 High-Altitude Arachnida (Partly from Beron 2008,
Updated in Beron 2016)�� 853
10.1 Introduction �� 853
10.2 Major Groups of Arachnida in the Old World Living
At or Above 2200 m�� 860
10.3 Level of Knowledge on Opiliones in the High Mountains
and Credibility of Conclusions�� 864
10.4 Maximum Altitudes of Spiders Living Above 2200 m
in the Old World�� 864
10.5 High-Altitude Arachnida in the Oreal of the Himalayas,
Compared with the Alps�� 870
10.6 High-Altitude Arachnida in the Mountains of Central Asia �� 880
10.7 Afrotropical High-Altitude Arachnida�� 882
10.8 Conclusions �� 884
10.9 Champions in Altitude (Above 4500 m) in the Old World �� 884
10.10 Some Personal Notes on the High-Altitude Arachnofauna
of the Americas and Elsewhere�� 886
11 Zoogeographical Notes on Different Orders of Arachnida�� 893
11.1 Palpigradi�� 893
11.2 Schizomida�� 894
11.3 Amblypygi�� 894
11.4 Thelyphonida (Uropygi)�� 895
11.5 Ricinulei�� 896
11.6 Solifugae �� 896
11.7 Scorpiones �� 897
11.8 Pseudoscorpiones�� 897
11.9 Opiliones �� 898
11.10 Araneae �� 900
11.11 Opilioacarida �� 902
12 Zoogeographical Subdivisions of the World�� 905
13 General Conclusions�� 931
Index�� 939
Chapter 1
Introduction
Abstract The classical zoogeographical subdivision of the land fauna, done by

Sclater and Wallace in the nineteenth century, is still “official” in our time. This
subdivision has been based on vertebrates, mostly mammals and birds. The ocean
of tiny creatures (insects, arachnids, etc.) has been (and still is) largely neglected.
Some attempts have been done to restructure the old scheme (Lopatin, Krizhanovskiy,
Morrone, etc.), but nothing has been done for the total of Arachnida – old groups of
non-flying invertebrates, quite important for zoogeography and already relatively
well known.
The classical zoogeographical subdivision of the land fauna, done by Sclater and
Wallace in the nineteenth century, is still “official” in our time. This subdivision has
been based on vertebrates, mostly mammals and birds. The ocean of tiny creatures
(insects, arachnids, etc.) has been (and still is) largely neglected. Some attempts
have been done to restructure the old scheme (Lopatin, Krizhanovskiy, Morrone,
etc.), but nothing has been done for the total of Arachnida – old groups of non-flying
invertebrates, quite important for zoogeography and already relatively well known.
By venturing into this mammoth task, to put together the information about all
orders in Arachnida and to elaborate on their distribution could be explained (besides
with my long experience as a lecturer of zoogeography) with the words of one of the
prominent arachnologists of our time, the Brazilian Adriano Kury “In a moment of
deprivation of common sense” (Kury, on line). Quot potui – feci!
Some 60 years ago, such analysis would have been much more incomplete, much
more difficult, and on completely different bases. Since the middle of the twentieth
century, so many new descriptions have appeared, revisions on the whole cladistics
taxonomy and zoogeography, so we have now a picture much closer to reality. At
the same time, the huge amount of information became very hard to digest, especially
for a single analyzer. It is fortunate that some competent specialists provided
catalogues, bibliographies, and checklists of whole groups of Arachnida and of
particular countries or regions. We must mention some (not all) of these very helpful
arachnological manuals, published in the twentieth and twenty-first centuries:
The “smaller orders”: Harvey (2003)
Palpigradi: Kraepelin (1901), Rowland and Sissom (1980), Condé (1996)
© Springer International Publishing AG, part of Springer Nature 2018 1

P. Beron, Zoogeography of Arachnida, Monographiae Biologicae 94,
https://doi.org/10.1007/978-3-319-74418-6_1
2 1 Introduction
Schizomida: Harvey (1992), Reddell and Cokendolpher (1995)

Amblypygi: Mullinex (1975), Quintero (1981 1986)
Thelyphonida (Uropygi): Rowland and Cooke (1973)
Ricinulei: Tuxen (1974)
Solifugae: Kraepelin (1901), Gromov and Savary (Bibliography on line)
Scorpiones: Fet, Sissom, Lowe and Braunwalder (2000)
Pseudoscorpiones: Chamberlin (1931), Beier (1932a 1932b 1963), Harvey (1990
1992), Schawaller (1980, bibliography)
Opiliones: Roewer (1923), Rosas Costa (1950 – Cyphophthalmi), Hallan (2005,
checklist), Cokendolpher (1997, bibliography), Giribet (2000, Cyphophthalmi),
Kury (2003, catalogue New World Laniatores; 2012, description of catalogues
and checklists), Pinto-da-Rocha, Machado and Giribet (Eds.) (2007), Schönhofer
(2013, Dyspnoi)
Araneae: Roewer (1942), Brignoli (1983), Platnick (The World Spider Catalog)
Opilioacarida: Hammen van der (1966), Beron (2014)
Holothyrida: Beron (2014)
Prostigmata: Beron (2008b, 2016)
Mesostigmata: Beron (2016)
Regional lists and bibliographies:
Old World: Beron (2008a, high altitude Arachnida)
New World: Kury (2003, Opiliones Laniatores)
Africa: Dupré (2013, Scorpiones), Lamoral and Reynders (1975, Scorpions,
Ethiopian Region), Scharff (1990, Linyphiidae), Starega (1984, Phalangiidae;
1992, African Opiliones, except Phalangiidae)
East Africa: Probst (1973, Scorpiones)
South Africa: Lawrence (1931, Opiliones; 1955, Solifugae, Scorpiones and
Pedipalpi, Schizomida, 1969), Kauri (1961, Opiliones), Lotz (2009 – Opiliones)
Madagascar: Lawrence (1959, Opiliones), Lourenço (1996, Scorpiones)
Australia: Forster (1955 and further), Glauert (1963, Scorpiones, W. Australia),
Koch (1977, Scorpiones, Australia and Papua), Harvey (1992)
Europe: Beier (1963, Pseudoscorpiones), Stol (1993, 2007, Opiliones, North
Europe), Deltshev and Blagoev (2001, Araneae, Bulgaria)
North America: Hoff (1958, Pseudoscorpiones), Cokendolpher and Lee (1993,
Opiliones)
Texas: Rowland and Reddell (1976, Texas)
South America: Benavides and Giribet (2007, Opiliones Neogoveidae), Mello-
Leitão (1945, Scorpiones), Kury (2003, Laniatores), Caporiacco (1948, Guianas,
Opiliones), Ringuelet (1959, Argentina, Opiliones)
West Indies: Cokendolpher and Camilo-Rivera (1989, Opiliones, bibliography)
Mexico: Díaz Nájera (1975, Araneae), Kury and Cokendolpher (2000, Opiliones),
Vazquez Rojas (1981 – Solifugae)
Madagascar: Emerit (1974, Araneae, Gasteracanthinae), Lawrence (1959,
Opiliones), Lourenço (1996, Scorpiones), Araneae (Goodman and Benstead
2005)
1 Introduction 3
New Zealand: Forster (1967–1973, Araneae; 1978, Opiliones), Dumbleton, 1953,

Heath, 1977 (Ixodida)
Oceania: Chamberlin (1934)
Chile: Cekalovic (1975, Solifugae; 1976, Arachnida; 1983, Scorpiones; 1984,
Pseudoscorpiones and Palpigradi; 1986, Opiliones), Beier (1964,
Pseudoscorpiones)
Northern Asia: Eskov (1994, Linyphiidae); Fet (1988, Scorpiones, the former
USSR); Marusik, Eskov, and Kim (1992, Araneae); Mikhailov (1998 1999 2000
2002 2013, Araneae, the former USSR); Staręga (1978 – Opiliones of the former
USSR)
Panama: Fairchild et al. (1966 – Ixodida)
Neotropical Region: Guglielmone et al. (2003, Ixodida)
Georgia: Djaparidze (1960 – Ixodida)
India: Murthy and Ananthakrishnan (1977, Pseudoscorpiones), Pocock (1900,
Arachnida), Siliwal, Molur and Biswas (2005, Araneae), Tikader (1987, Araneae)
Nepal: Clifford et al. (1975, Ixodida)
Iran: Abassian-Lintzen (1960, Ixodida), Mozaffarian and Marusik (2001, Araneae)
Arachnida: Blick, Hänggi, and Thaler (2002, Germany, Switzerland, Austria,
Belgium, and the Netherlands), Maes et al. (1989, Nicaragua), Zaragoza (2007 –
Iberian Peninsula)
Palpigradi: Kraepelin (1901), Rowland and Sissom (1980), Condé (1996)
Solifugae: Aliev and Gadzhiev (1983, Azerbaydjan), Armas and Teruel (2005,
Cuba), Gromov and Savary (bibliography), Levy and Shulov (1964, Israel)
Schizomida: Armas 2004 (Cuba, Dominican Rep.), Harvey (1992, 2000, Australia;
2001, Seychelle Isl.); Reddell and Cokendolpher (1995)
Amblypygi: Armas (2004, 2009b, Antilleans), Mello-Leitão (1931, Brazil)
Thelyphonida (Uropygi): Mello-Leitão (1931, Brazil)
Ricinulei: Tuxen (1974, Africa)
Scorpiones: Acosta and Maury (1998, Argentina); Acosta and Ochoa (2002,
Bolivia); De Armas (2009a, Antilleans); Fet (1988, former USSR; 1994,
Tukmenistan; 2010, Europe); Koch (1977, Australia); Lamoral (1979, Namibia);
Levy and Amitai (1980, Palestina); Lourenço and Méndez (1984, Panama);
Lourenço (1995a, Ecuador; 1996, Madagascar; 1997, Colombia); Maury (1984,
Paraguay); Mello-Leitão (1932, Brazil); Zhu, Qi, and Song (2004, China);
Ojanguren-Affilastro (2005 – Argentina)
Pseudoscorpiones: Agnarsson (1998, Iceland); Beier (1932, 1965a, New Guinea;
1966, Philippines); Chamberlin (1934, Oceania); Ćurčić (1974, Yugoslavia);
Ćurčić, Dimitriević, and Legakis (2004, Serbia, Montenegro, Macedonia); de
Lessert (1911, Switzerland); El-Hennawy (1988, Egypt); Gardini (2000, Italia);
Harvey (1990); Kunt (2008, Turkey); Legg and O’Connor (1997, Ireland);
Mahnert (1981–1988, Kenya; 1975, Malta; 2004, Austria); Petrov (1997,
Bulgaria); Schawaller (1980, bibliography; 1994b, Thailand; 1995, China);
Telnovs (2002a, Latvia); Zaragoza (2000, bibliography, Iberian Peninsula,
Baleares, Macaronesia); Legg (1988, Great Britain); Murthi and Anasnthakrishnan
(1977, India)
4 1 Introduction
Opiliones: Agnarsson (1998, Iceland); Babalean (1992, Romania); Bayram et al.
(2010, Turkey); Bezdecka (2008, Czech Republic); Blick and Komposh (2004,
Central and Northern Europe); Canals (1936, Chile); Cawley (2002, Ireland);
Chevrizov (1980, URSS); Cokendolpher (1990, Egypt); Cokendolpher and
Camilo-Rivera (1989, West Indies); Cokendolpher and Lee (1993, Greenland,
Canada, USA, Mexico); Gritzenko (1978, Asian part of USSR); Forster (1954,
New Zealand); Hillyard and Sankey (1989, British Fauna); Kim, D.-H., J.-W. Lee,
and J.-P. Kim (2006, Korea); Klimeş (2000, Czech and Slovak Rep.); Komposch
(2004, Hungary); Komposch and Gruber (2005, Austria); Lawrence (1959,
Madagascar); Lotz (2009, Southern Africa); Martens (1978, Germany; many
papers, Nepal); Mheidze (1964, Georgia); Novak (2004, Croatia; 2005, Bosnia
and Herzegowina); Novak et al. (2006, Slovenia); Prendini (2010, Seychelles);
Rafalski (1960, 1961, Poland); Rafalski and Staręga (1997, Poland); Rambla
(1967, Portugal); Redikorzev (1936 – Soviet Union); Ringuelet (1959, Argentina;
1963, Uruguay); Roewer (1923); Shavanova (2004, Belarus); Šilhavý (1956,
Czechoslovakia); Staręga (1976, Bulgaria; 1978, Soviet Union; 1992, Afrotropical
Region; 2000, Poland); Suzuki (1985a, 1985b, Thailand); Telnovs (2002b,
Latvia); Tsurusaki (1993, Japan)
Araneae: Aakra and Hauge (2000, 2003, Norway, Svalbard, Jan Mayen); Agnarsson
(1996, Iceland); Bosmans (2009, Belgium); Bosmans and Chadzaki (2005,
Greece); Bosmans and de Keer (1985, Pyrenees); Blagoev (2002, Macedonia);
Bonnet (1945–1961, bibliography); Brignoli (1983); Cardoso and Morano
(2010, Iberian Peninsula); Deltshev (2005, Bulgaria); Deltshev and Blagoev
(2001, Bulgaria); Deltshev, Curčić, and Blagoev (2001, Serbia); Drensky (1936,
Balkan Peninsula); Evenhuis (2006, Fiji); Forster et al. (1967–1973, New
Zealand); Forster and Forster (1973, New Zealand), Gajdos, Svaton, and Sloboda
(1999, Slovakia); Kim (1990, Korea); Kostanjšek and Kuntner (2015, Slovenia);
Larsen and Scharff (2003, Greenland); Le Peru (2007, France); Marinu and
Verneau (2002, Corsica); Marusik, Eskov, Logunov, and Basarukin (1993,
Sakhalin and Kuril Isls); Marusik, Eskov, Koponen, and Vinokurov (1993,
Yakutia); Kronestedt (2001, Sweden); Marusik, Logunov, and Koponen (2000,
Tuva); Merrett, Locket, and Millidge (1985, Great Britain); Mikhailov and Fet
(1994, Turkmenistan); Mikhailov (1997, 1998, 1999, 2000, 2002, 2013, the
former Soviet Union); Milosević (2002, Croatia); Nikolić and Polenec (1981, f.
Yugoslavija); Platnick (2000–2011. The World Spider Catalog, Version 11.5.);
Proszynski and Staręga (1971, Poland); Relys and Spungis (Latvia); Roewer
(1942); Song, Zhang, and Daigin (2002, Singapore); Starega (2000, Poland);
Suman (1964, Hawaii), Tikader (1970, Sikkim); Topcu, Demir, and Seyyar
(2005, Turkey); Varol (2003, Turkey); Vilkas (1992, Lithuania); Weiss and Urak
(2000, Romania)
Opilioacarida: Beron (2014), Vasquez and Klompen (2002, North and Central
America; 2010, Madagascar)
Holothyrida: Beron (2014), Lehtinen (1995)
Ixodida: Drensky (1955, Bulgaria), Feider (1965, Romania), Filippova (1966,
Argasidae of the World), Starkoff (1958, Ixodida of Italy), Georgieva and
1 Introduction 5
Gecheva (2013, Ixodidae of Bulgaria), Gregson (1956, Ixodida of Canada),

Anastos (1950, Ixodida of Indonesia), Roberts (1964, Ixodida of Tasmania;
1970, Ixodida of Australia), Dumbleton 1953, Heath 1977 (Ixodida of New
Zealand), Hoogstraal (1953, Ixodida of Madagascar)
Prostigmata: Beron (2008a – Calyptostomatoidea and Erythraeoidea of the world)
Oribatida: Shtanchaeva 2003 (Caucasus), Collof and Haliday 1998 (Australia),
Bayartogtokh 2010 (Mongolia), Vu Quang Manh (Vietnam)
This book would have been impossible without the generous help of many arach-
nologists and biogeographers since the 1960s, when I started gathering material for
it. Some of these people I knew personally, I received regularly their publications
and advice, and also I benefited from their hospitality in many institutes and muse-
ums; I had also access to their private book and reprint collections. I cannot enumer-
ate all of them, but I will mention some, without their titles.
Bulgaria – Ch. Deltshev, B. Gruev, V.B. Guéorguiev, P. Mitov, B. Petrov, S. Lazarov
Serbia – B. Ćurčić, I. Karaman, S. Matvejev
Slovenia - I. Hadži
Croatia – T. Novak
Romania – M. Dumitresco, D. Dumitrescu, Z. Feider, S. Avram
Hungary – Y. Kontchán, S. Mahunka, Z. Varga
Czech Rep. – J. Buchar, M. Daniel, F. Kovařik, J. Kratochvil, M. Krumpál, V.
Šilhavý
Poland – J. Rafalski, W. Staręga, R. Haitlinger, J. Mąkol, G. Gabryś, W. Weselowska
Italy – P.M. Brignoli, G. Callaini, L. Delle Cave, G. Gardini, G. Lazzeroni,
I. Marcellino, M. La Greca, A. Vigna Taglianti
France – L. Berland, B. Condé, J. Heurtault, C. Juberthie, M. Judson, P. Leclerc,
R. Legendre, W.R. Lourenço, M. Vachon, A. Vandel, I. Coineau
Great Britain – G. Legg, R. Hall, P.J.A. Pugh
Belgium – P. Benoit, R. Bosmans, A. Fain, R. Jocqué
Netherlands – L. van der Hammen
Austria – M. Beier, E. Christian, J. Gruber, H. Franz, C. Komposch, H. Schatz,
K. Schmölzer, K. Thaler, H. Janetschek
Switzerland – V. Mahnert
Denmark – H. Enghoff, S.L. Tuxen
Finland – J. Kaisila, P. Lehtinen
Sweden – H. Lohmander
Germany – J.A. Dunlop, J. Haupt, J. Martens, P. Müller, W. Schawaller, P. Weygoldt,
J. Wunderlich
Spain – M. Rambla, J.A. Zaragoza
Russia – S. Golovatch, Yu. Marusik, A. Tanasevitch, O.L. Krijanovskij,
U. Shtanchaeva, K.G. Mikhailov
Kazakhstan – A.V. Gromov
Georgia – D. Kobakhidze
Iran – A. Saboori
Turkey – K.B. Kunt
6 1 Introduction
United States – E. Benedict, J.C. Chamberlin, J.C. Cokendolpher, J. Cracraft, V. Fet,

O.F. Francke, G. Giribet, J.L. Gressitt (Hawaii), C. C. Hoff, C. and M. Goodnight,
F. Howarth (Hawaii), H. Hoogstraal, E. Mayr, R.W. Mitchell, W.B. Muchmore,
M. Muma, N. Platnick, L. Prendini, J.R. Reddell, J. M. Rowland, P.A. Selden,
W. Shear
Canada – S. Peck
Cuba – L. de Armas, R. Teruel, M.A. Ituralde-Vinent, G. Alayón Garcia
Panama – D. Quintero, Jr.
Australia – B. Haloway, M. Harvey, V.V. Hickman, C.J. Humphries, L.E. Koch,
D. Lohman, B. Main, P.V. Taylor, R. Southcott
New Zealand – R.R. Forster
Mexico – A. Hoffmann, G. Halffter, J.J. Morrone
Brazil – L.J. de Araújo Feio, A.B. Kury, the family H. and B. Soares
Venezuela – M.A. González - Sponga
Argentina – V. Vitali-di-Castri
Japan – K. Morikawa, M. Shimojana, S. Suzuki, N. Tsurusaki
China – J.X. Zhang, Shuqiang Li
Korea – Kap Yong Paik
Philippines – L. Corpuz Raros
India – M.S. Mani
South Africa – R.F. Lawrence, H. Martin
Israel – F.D. Por
My sincere gratitude goes also to the colleagues who helped me a lot in preparing
the maps and with other important “technicalities” – Mr. Assen Ignatov, Mrs. Sylvia
Tosheva, and others.
Chapter 2
Builders of Arachnology
Abstract In many older authors (Aristotle and others), we can find information
about spiders, scorpions, and other Arachnida. The scientific studies however started
with the paper of Clerck (1757), the only one recognized as valid before the system
of Linnaeus (1758).
In many older authors (Aristotle and others), we can find information about spiders,
scorpions, and other Arachnida. The scientific studies however started with the
paper of Clerck (1757), the only one recognized as valid before the system of
Linnaeus (1758). Before trying to discuss the zoogeographical problems of
Arachnida, we owe a tribute to the builders of the arachnology by saying a few
words about the life of some (by no means all) founding fathers. Some of them
(Beier, Vachon, Brignoli, Kratochvil, Hadži) were known personally to the present
author. To the correspondence with others (Chamberlin, Lawrence, Šilhavý, Suzuki,
all other “living classics”), he owes much of his devotion to arachnology.
Clerck, Carl (1709–1765) – the names in his paper of 1757 Svenska Spindlar were
recognized as valid, and it is the oldest recognized paper in the zoological nomen-
clature. It started the scientific study of Arachnida. Presently, 53 spider species carry
names given by Clerck.

https://doi.org/10.1007/978-3-319-74418-6_2
8 2 Builders of Arachnology
Linnaeus, Carolus (Carl von Linné) (1707–1778) – professor of medicine and

botany at Uppsala (Sweden) and founder of the system of plants and animals
(Systema Naturae, Ed. X, 1758)
C. Linnaeus (Portrait by A. Roslin, 1775)

2 Builders of Arachnology 9
Thorell, Tord Tamerlan Theodor (1830–1901) – born in Sweden but lived long-
time in Genoa and author of numerous important contributions to the system of
Arachnida (On European spiders, 1869; Synonym European spiders, 1870–1873,
many papers with descriptions of new taxa from Malaysian and Papuan collections,
Arctica, Cameroon, etc.). Thorell described 12 families of spiders (Ctenizidae,
Theraphosidae, Palpimanidae, Hersiliidae, Uloboridae, Oxyopidae, Amaurobiidae,
Zodariidae, Philodromidae, etc.). Important papers are devoted also to Opiliones
(13 publications on Opiliones from SE Asia, Indonesia, Burma, Argentina, USA,
Europe, and West Asia).
T.T.T. Thorell
Sørensen, William Emil (1848–1916) – Danish arachnologist. Described 157

valid species of Opiliones. Some important papers are as follows: 1898, Arachnida
Groenlandica (Acaris exceptis); 191, Opiliones (in I. Sjostedt’s Kilimanjaro – Meru
volumes); and 1932, Descriptiones Laniatorum.
William Emil Sørensen (1848–1916)

Hansen, H.J. (1855–1936) – Danish arachnologist. Some important papers:

Hansen & Sorensen (1904) – established the modern system of Opiliones; Described
many new taxa in Arachnida.
Hans Jakob Hansen

Mello-Leitão, Cândido Firmino de (1886–1948) – Brazilian arachnologist, pro-

fessor, and explorer of Opiliones (198 publications, describing 347 valid species),
Uropygi, Amblypygi, Scorpiones, Araneae, and some smaller orders in Brazil.
Some important papers were Pedipalpos do Brasil e algumas notas sobre a ordem
(1931), Opiliões do Brasil (1932, 511 pp.), Escorpiães Sul-Americanos (1945, 468
pp.), and others.
Cândido Firmino de Mello-Leitão

Pocock, Reginald Innes (1863–1947) – English arachnologist and curator of the

collections of Arachnida and Myriapoda from 1885 to 1904. Author of many papers
on various groups of Arachnida.
R.I. Pocock (From P. Bonnet, 1945)

Kraepelin, Karl Matthias Friedrich Magnus (1848–1915) – German arachnolo-

gist, worked in the Zoology Museum in Hamburg, and author of many descriptions
of various groups of Arachnida. Among the publications of Kraepelin are a volume
on Palpigradi and Solifugae in Das Tierreich and many others.
Karl Kraepelin
Roewer, Carl-Friedrich Roewer (1881–1963) – German arachnologist, profes-

sor, and director of the Overseas Museum in Bremen (1933–1945). His huge works
laid the base of the system of Opiliones and of Solifugae. Since his first paper
(Revision der Opiliones Plagiostethi, 1910, 294 pp.), he indicated a very mighty
arrival in the field of taxonomy of Opiliones. In 1923 he published the biggest
monograph on this group ever written – “Die Weberknechte der Erde” – followed by
series of voluminous supplements (Weitere Weberknechte I–XX, 1927–1957) and
by numerous publications, describing most of the known genera of Opiliones from
all over the world and 2260 valid species. Another big achievement of Roewer was
the system of Solifugae (1934, 1941). In his papers he described almost all genera
of this order. Roewer wrote (without computer) also a Catalogue of Araneae from
1758 to 1940 (1942) and articles on other Arachnida (mostly Araneae). Some of the
localities indicated by him have been questioned (Martens and v. Helversen 1972).
His huge collection is preserved in Senckenberg Museum.
C.-F. Roewer (From O. Kraus)

Beier, Max (1903–1979) – his main fields have been the pseudoscorpions and the
Mantodea. After several revisions, Beier’s two volumes of the series appeared Das
Tierreich (Pseudoscorpionidea I and II, 1932). Together with the monograph of
J.C. Chamberlin, published in the previous year, they laid the foundation of
Pseudoscorpionology. In his papers on Pseudoscorpiones from all over the world,
Beier published many new taxa and contributed greatly to the study of faraway
countries. Many of his papers are devoted to Mantodea and Orthoptera. Max Beier
has been an associate and director of Naturhistorisches Museum in Vienna.
Max Beier
Chamberlin, Joseph Conrad (1898–1962) – one of the founders of the system of

Pseudoscorpiones and nephew of R.V. Chamberlin. The second part of his thesis,
entitled The Arachnid Order Chelonetida (1931), is the most important for the clas-
sification of the order. He described many new genera and the 12 of the 26 existent
families: Lechytiidae, Bochicidae, Ideoroncidae, Neobisiidae, Menthidae,
Atemnidae, Geogarypidae, Pseudochiridiidae, Pseudogarypidae, Hyidae,
Syarinidae, and Withiidae. From 1921 to 1962, J.C. Chamberlin published 41
papers on Pseudoscorpiones of many countries and also other papers on Crustacea,
systematic and economic entomology, etc.
Joseph Conrad Chamberlin

Brignoli, Paolo Marcelo (1942–1986) – Italian arachnologist, professor in Rome

and Aquila, and specialist mainly in world spiders. From 1967 to 1985, he wrote
205 publications, describing 25 new genera and 370 new species. He has many
contributions to the fauna of Greece, Mexico, Ethiopia, Nepal, Italy, and other coun-
tries, partly as a result of his own expeditions in different continents. Also he has
publications in other Arachnida (Opiliones, Schizomida, etc.). He expressed inter-
esting theoretical views on cave spiders. He wrote a catalogue of world spiders
(1940–1981) of 755 pages.
Paolo Marcelo Brignoli

Simon, Eugène (1848–1924) – French zoologist, working mainly on spiders and

birds (Trochilidae). His main arachnological works are, 1864–1884, Histoire
Naturelle des Araignées; 1875, Les Arachnides de France; and 1879, Arachnides de
France (VII volume, analyze); also he had many descriptions of spiders collected by
expeditions in Indochina, the Philippines, Greece, Syria, Yemen, Algeria, Egypt,
and other countries. Simon described 40 families (out of the total of 117 accepted
families in the world), many new genera, and 4650 species of spiders and other
Arachnids (about 3790 species are still considered valid) more than any other
arachnologist.
E. Simon
Koch Carl Ludwig (1778–1857) – German arachnologist. His principal works,

Übersicht des Arachnidensystems, Nürnberg, 1837–1850; Deutschlands Crustaceen,
Myriapoden und Arachniden, Regensburg, 1835–1844; and Die Arachniden (1831–
1848) (16 volumes), are the base of European arachnology. He is the father of
another prominent arachnologist – Ludwig Carl Christian Koch (1825–1908).
C.-L. Koch
Banks, Nathan (1868–1953) – American arachnologist and entomologist. Author

of many descriptions of new genera and species in all orders of Arachnida, together
with studies on Trichoptera and other insects.
Nathan Banks
Šilhavý, Vladimir (1913–1984) – Czech physician and prominent specialist in

Opiliones. Working mostly on the fauna of the Caribbean, Mexico, the Balkans, and
Central Asia, he described many new genera and species and wrote the book on
Opiliones of Czechoslovakia (1956).
Vladimir Šilhavý (Contributed by P. Bezdĕčka)

Hadži, Jovan (1884–1972) – Slovenian arachnologist and academician. He stud-

ied the scorpions, opilions, and pseudoscorpions of former Yugoslavia, describing
many new species. His main arachnological works were Opilionidea. In: Catalogus
Faunae Jugoslaviae, III/4 (1973b) and Neue Taxa der Weberknechte (Opilionidea)
aus Jugoslawien (1973a), both published posthumously.
Jovan Hadži (Slovenia Wikipedia)

Petrunkevitch, Alexander (1875–1964) – American arachnologist of Russian ori-

gin (Ukrainian-born). From 1910 to 1939, he described over 130 spider species. He
was a professor (since 1917) in Yale University, where he taught from 1910 to 1944.
Many important papers on fossil Arachnida: A Monograph of the Terrestrial
Paleozoic Arachnida of North America (1913); Tertiary Spiders and Opilionids of
North America (1922); Inquiry into the Natural Classification of Spiders, Based on
a Study of Their Internal Anatomy (1933); Palaeozoic and Mesozoic Arachnida of
Europe (1953); Arachnida. In; Treatise on Invertebrate Paleontology (1955); Amber
Spiders in European Collections (1958), papers on the spiders of Puerto Rico, of
Mexico, etc. He described also many new families and higher taxa, including the
valid fossil families Protosolpugidae, Kustarachnidae, Eotrogulidae,
Nemastomoididae, Dolichophoninae, Archaeoctonidae, Centromachidae,
Mazoniidae, Isobuthidae, and Calcitronidae.
A. Petrunkevitch (Courtesy, Yale University)

Kratochvil, Josef (1909–1992) – Czech Zoologist and Academician, worked on

Araneae, Opiliones, and Mammalia. He wrote 209 papers, including many on
Arachnida (from 1930 to 1970), and several books. Kratochvil described many new
taxa, especially from the caves of the Balkan Peninsula, including the genera
Paralola and Tranteeva from Bulgarian caves (1958). He was a director of many
scientific institutions.
Josef Kratochvil
Lawrence Reginald Frederick (1897–1987) – the most prominent researcher of

South African Arachnida and Myriapoda. R. Lawrence published 23 papers on
Opiliones (describing 224 valid species) and many new genera and species on all
groups of Arachnida and Myriapoda.
Reginald Frederick Lawrence

Vachon Max (1908–1991) – French specialist in Scorpiones and Pseudoscorpiones.

Prof. Vachon was Director of the Laboratory of Invertebrates (Paris Museum of
Natural History).
Max Vachon
Suzuki, Seisho (1914–2011) – During the period from 1938 to 1991, Dr. Suzuki
wrote important series of 135 papers on Opiliones (Japan, Taiwan, Nepal, Malaysia,
Philippines, Thailand, Sakhalin, etc.), with many discoveries and descriptions of
valid species and several other taxa, also with zoogeographic analysis. He wrote
also some papers of spiders.
Suzuki Seisho
Gonzalez-Sponga Manuel-Angel (1929–2009) – Venezuelan arachnologist and

professor. Described 219 valid species and many other taxa of Opiliones, many
scorpions, and other Arachnids.
Manuel-Angel Gonzalez-Sponga (Courtesy of R. Pinto-da-Rocha)

Berlese Antonio (1863–1927) – Italian Acarologist who described thousands of

new genera and species of mites.
Antonio Berlese
Oudemans Anthonie Cornelis (1858–1943) – Dutch acarologist and one of the

founders of the actual system of Acari.
Anthonie Cornelis Oudemans

Staręga Wojciech (1939–2015) – Polish arachnologist and specialist in Opiliones

and spiders. He published many new taxa, the important revision of Bulgarian
Opiliones (1976), catalogues, and articles on African Opiliones and also on the
Opiliones of Russia, Central and East Asia, and other countries.
W. Starega (Courtesy of J. Cokendolpher)

Bibliography 33
Of course, the pioneers were (and still are) followed by many others, on a new level.
Now data on Arachnida is snowballing, and the systems change overnight. With the
new means of communication, we can now be on course to rapidly exchanging
information, thanks also to some of our colleagues, who took care to facilitate our
life by preparing catalogues of some orders and/or pages online. Such catalogues
are listed in the Bibliography, but here we enumerated some of their authors:
Brignoli P.M., Harvey M., Kury A., Giribet G., Cockendolfer J.C., Platnick N., Fet
V., Deltshev Hr., Blagoev G., Ćurčić B., Martens J., Morano E., Michailov K.,
Juberthie Ch., Rambla M., Weigoldt P., Prendini L., Lourenço W., Drensky P.,
Gardini G., Saaristo M., Raven R.J., Lehtinen P., Rowland M., Dunlop J., Selden
A.A., Thaler K., Wunderlich J.W., Shear W., and Muchmore W.B.
Bibliography
Birabén M (1949) Necrológio – Cândido F. de Mello-Leitão. Revista de la Socedad Entomologica

Argentina. Buenos Aires 14(4):238–240
Buchar J (1984) Memory of MUDr. Vladimír Šilhavý, CSc. Acta entomologica bohemosl
81:157–158
Carpenter FM, Darlington PJ Jr (1954) Nathan Banks, a biographic sketch and list of publications.
Psyche 61:81–110
Judson M, Chamberlin DC (1998) Joseph C. Chamberlin 1898 – 1962. J Arachnol 26:411–418
Kaltenbach A (1980) Hofrat Professor Dr. Max Beier zum Gedenken. Annalen des Naturhistorischen
Museums Wien 83:763–781
Kury AB, Baptista RLC (2004) Arachnological papers published by Cândido Firmino de Mello-
Leitão (Arachnida). Publicações Avulsas do Museu Nacional, Rio de Janeiro 105:1–17
Muster C, Dunlop JA (2006) Le temps marche si vite – in memory of Konrad Thaler. J Arachnol
34:281–284
Osella G (1987) Professor Dr Paolo Marcello Brignoli (1942–1986). Bull Br Arachnol Soc 7(6):186
Prószynski J (1997) In memoriam Prof. Dr. Jan Rafalski (1909-1995). Proceedings of the 16th
European Colloquium of Arachnology, Marek Zabka, editor. Siedlce, 8 – 9
Roesler R (1997/98) Karl Ludwig Koch (1778–1857). In Oberpfälzer Heimat. Auflage 42, Weiden
Vigna Taglianti A (1986) Paolo Marcello Brignoli (1942 – 1986). Fragmenta Entomologica Roma
19(1):267–271
Chapter 3
Systems of Arachnida
“Traditions die hard: it is difficult to understand why fishes or

birds should be divided into dozens of orders, whereas, for most
zoologists, a single order appears sufficient for all spiders or all
mites. From this it could be inferred that either the
arachnologists are more conservative or, in the eyes of most
zoologists, small differences between (nice) birds are more
significant than great differences between (ugly) spiders.”
P.M. Brignoli (1980, The Evolution of Arachnida)
Abstract Heymons (1901), the author of the term Chelicerata, united Merostomata
(Eurypterids and Xiphosurans) with Arachnida in one monophyletic group. Dunlop
and Selden (1997) restricted this term to Lemoneites, Xiphosura, Chasmataspida,
Eurypterida, and Arachnida. There were many attempts to create a “natural” system
of the monophyletic (Wheeler et al. 1993) class of Arachnida. Presently, it seems
well established that the class Arachnida (Lamarck 1801) is composed of 10 recent
orders plus 2–6 orders of “Acari” and variable number of fossil taxa. The seemingly
undisputed orders are Palpigradi, Solifugae, Amblypygi, Uropygi, Schizomida,
Ricinulei, Scorpiones, Pseudoscorpiones, Opiliones, and Araneae. The Acari have
been (and still are) subject of various systems (see Beron 2008, 2011), more often
considered as two (Acariformes and Parasitiformes) or three (+ Opilioacarida)
orders. In the recent Manual of Acarology, Krantz and Walter and eds. (2009) agree
on six orders of “Acari,” Mesostigmata, Holothyrida, Ixodida, Opilioacarida,
Sarcoptiformes, and Trombidiformes, grouped in two superorders: Acariformes and
Parasitiformes. Some authors (Stockwell 1989, Lourenço 2000) still raise some
orders into subclasses or even classes (Opilionida, Scorpionida).
Heymons (1901), the author of the term Chelicerata, united Merostomata

(Eurypterids and Xiphosurans) with Arachnida in one monophyletic group. Dunlop
and Selden (1997) restricted this term to Lemoneites, Xiphosura, Chasmataspida,

https://doi.org/10.1007/978-3-319-74418-6_3
36 3 Systems of Arachnida
Eurypterida, and Arachnida. There were many attempts to create a “natural” system
of the monophyletic (Wheeler et al. 1993) class of Arachnida. Presently, it seems
well established that the class Arachnida (Lamarck 1801) is composed of 10 recent
orders plus 2–6 orders of “Acari” and variable number of fossil taxa. The seemingly
undisputed orders are Palpigradi, Solifugae, Amblypygi, Uropygi, Schizomida,
Ricinulei, Scorpiones, Pseudoscorpiones, Opiliones, and Araneae. The Acari have
been (and still are) subject of various systems (see Beron 2008, 2011), more often
considered as two (Acariformes and Parasitiformes) or three (+ Opilioacarida)
orders. In the recent Manual of Acarology, Krantz and Walter and eds. (2009) agree
on six orders of “Acari,” Mesostigmata, Holothyrida, Ixodida, Opilioacarida,
Sarcoptiformes, and Trombidiformes, grouped in two superorders: Acariformes and
Parasitiformes. Some authors (Stockwell 1989, Lourenço 2000) still raise some
orders into subclasses or even classes (Opilionida, Scorpionida).
The Higher and Highest Arrangements.
The arrangement of Mello-Leitão (1931) in Portuguese:
Arachnideos
Subclass Ctenophoros
Order Escorpioes
Subclass Lipoctenos
Sectio Patelados
Susectio Megoperculados
Order Pedipalpos
Order Aranhas
Dubinin (1959, announced first in 1957) divided the “superclass” Chelicerata
into five classes: Merostomata, Scorpionomorpha, Solifugomorpha (with subclasses
Holactinochitinosi and Pedipalpides), Arachnida (subclasses Opiliomorphae,
Soluta, and Araneae), and Acaromorpha.
Van der Hammen (1977) proposed to subdivide his new phylum Chelicerata into
seven classes: Epimerata, Cryptognomae, Opilionidea, Apatellata, Arachnidea,
Merostomata, and Scorpionidea.
There is also a proposal by Savory (1977) to promote the suborder Cyphophthalmi
into full order. This was not accepted.
The Systems of Some Orders
“Pedipalpi” – under this name have been united three orders of Arachnida:
Amblypygi, Uropygi, and Schizomida.
Amblypygi
Millot (1949) – two families: Charontidae and Tarantulidae
Quintero (1986) – created the suborders Apulvillata and Pulvillata and the family
Charinidae
3 Systems of Arachnida 37
Suborder Apulvillata – families Phrynidae and Phrynichidae

Suborder Pulvillata – families Charinidae and Charontidae
Thelyphonida (Uropygi). Rowland and Cooke (1973) elevated the ancient
Pocock’s (1899) subfamilies into families, thus three genera (Hypoctonus,
Labochirus, and Thelyphonellus) in Hypoctonidae and the remaining 13 genera in
Thelyphonidae. Presently, only one family is accepted, Thelyphonidae, which is
divided into three subfamilies (Thelyphoninae, Hypoctoninae, and Typopeltinae).
Scorpiones
Stockwell (1989) – until this paper traditionally up to 8–9 families have been recog-
nized. He raised the group to class with three orders (Protoscorpiones,
Palaeoscorpiones, and Scorpiones).
Lourenço (2000) – based on the classification of Stockwell (1989), he kept the
scorpions as a class.
Class Scorpionida
Order Protoscorpiones Petrunkevich, 1949
Order Palaeoscorpiones Stockwell, 1989
Order Scorpiones C.L. Koch, 1837
Suborder Mesoscorpionina Stockwell, 1989
Suborder Neoscorpionina Thorell et Lindström, 1885
Infraorder Palaeosterni Stockwell, 1989
Infraorder Orthosterni Pocock, 1911 – includes the actual scorpions (20
recent families of the superfamilies Bothriuroidea, Buthoidea,
Chaeriloidea, Chactoidea, Vaejovoidea, and Scorpionoidea)
Fet et al. (2000) – their world catalogue includes 16 recent families and one fos-
sil family in the suborder Neoscorpionina, infraorder Orthosternina, superfamily
Scorpionidea, and the fossil suborder Branchioscorpionina.
Prendini and Wheeler (2005) – 18 families.
Prendini (2011a, b, c, d, e, f, g) – Order Scorpiones, one recent suborder
Neoscorpionina with 23 families (four fossil, but there is a chance Akravidae to be
found alive), so 19–20 recent families.
Pseudoscorpiones (Chelonethi) – see the table in Harvey (1992, p.1376).
Chamberlin (1931) – three suborders: Heterosphyronida, Monosphyronida, and
Diplosphyronyda
Beier (1932a, b) – three suborders: Chthonioidea, Neobisioidea, and Cheliferinea
Harvey (1992, 2011, 2013a, b) – two suborders Epiocheirata and Iocheirata, with
24 families in 7 superfamilies, in total 26 families
Suborder Epiocheirata
Superfamily Chthonioidea (four families – Chthoniidae, Lechytiidae,
Pseudotyrannochthoniidae, Tridenchthoniidae)
Superfamily Feaelloidea (two families – Feaellidae, Pseudogarypidae)

Suborder Iocheirata
Infraorder Hemictenata
Superfamily Neobisioidea (seven families – Neobisiidae, Bochicidae,
Gymnobisiidae, Hyidae, Ideoroncidae, Parahyidae, Syarinidae)
Infraorder Panctenata
Superfamily Garypoidea (six families – Garypidae, Garypinidae,
Geogarypidae, Larcidae, Menthidae, Olpiidae)
Superfamily Cheiridioidea (two families – Cheiridiidae, Pseudochiridiidae)
Superfamily Sternophoroidea (one familes – Sternophoridae)
Superfamily Cheliferoidea (four families – Cheliferidae, Atemnidae,
Chernetidae, Withiidae)
Opiliones
Thorell (1876a, b) – divided Opiliones into Palpatores and Laniatores
Loman (1902) – proposed the term Insidiatores (new suborder including
Triaenonychidae)
Pocock (1902) – introduced the name Plagiostethi (= Palpatores), divided into
Apagosterni and Eupagosterni. This was followed by Roewer (1910).
Hansen and Sørensen (1904) – established a system of three suborders:
Cyphophthalmi, Palpatores, and Laniatores. Palpatores were divided into the
tribes Eupnoi and Dyspnoi.
Mello-Leitão (1944) – proposed a phylogenetic tree of the Opiliones, noticing that
there was a sister-group relationship between Cyphophthalmi and Palpatores
Šilhavý (1961) – proposed subordinal rank of Eupnoi and Dyspnoi plus two more
suborders Gonyleptomorphi and Oncopodomorphi
Martens (1976) – The Palpatores are replaced by four superfamilies and the
Laniatores by three:
Palpatores – Troguloidea, Ischyropsalidoidea, Phalangoidea, and Caddoidea
Laniatores – Travunioidea, Oncopodoidea, Gonyleptoidea
Martens (1980 and further) – introduced the term Cyphopalpatores
(Cyphophthalmi plus Palpatores). It was followed by Cokendolpher and Lee
(1993) and other authors.
Shear (1980, 1993) – divided suborder Cyphophthalmi into two infraorders with
three superfamilies and six families:
Suborder Cyphophthalmi
Infraorder Temperophthalmi
Superfamily Sironoidea
Family Troglosironidae
3 Systems of Arachnida 39
Family Sironidae
Family Pettalidae
Infraorder Tropicophthalmi
Superfamily Stylocelloidea
Family Stylocellidae
Superfamily Ogoveoidea
Family Ogoveidae
Family Neogoveidae
This arrangement was accepted by Giribet (2000) and used since.
Cokendolpher and Lee (1993) – used the term Cyphopalpatores but in their cata-
logue retained the names Cyphophthalmi and Palpatores
Giribet et al. (2002) – introduced the term Dyspnolaniatores (Dyspnoi and its sister
group Laniatores)
Giribet and Kury (2007) – reviewed the phylogeny and biogeography of Opiliones
and agreed to maintain the classical and generally accepted subdivision of the
monophylic order Opiliones into Cyphophthalmi, Eupnoi, Dyspnoi, and
Laniatores. We shall follow this arrangement in the present book. They used the
old term Phalangida as a name of the clade formed by Eupnoi, Dyspnoi, and
Laniatores. Their summary tree:
Sironidae
Pettalidae
Troglosironidae Cyphophthalmi
Ogoveidae
Neogoveidae
Stylocellidae
Caddoidea
Phalangioidea Eupnoi
Ischyropsalidoidea
Troguloidea Dyspnoi
Travunioidea
Triaenonychoidea
Zalmoxoidea
Samooidea Laniatores
Phalangodoidea
Epedanoidea.
Gonyleptoidea.
System of Opiliones in Kury (2013):

Order Opiliones Sundevall, 1833 (4 suborders, 46 extant families, 3 fossil families,
1649 extant genera, 18 fossil genera, 6534 extant species, 37 fossil species)
Suborder Cyphophthalmi (six families)
Suborder Eupnoi
Superfamily Caddoidea (one family, Caddidae)
Superfamily Phalangioidea (four extant families – Monoscutidae,
Neopilionidae, Phalangiidae, Sclerosomatidae)
Suborder Dyspnoi
Superfamily Ischyropsalidoidea (three families – Ischyropsalididae,
Sabaconidae, Taracidae)
Superfamily Troguloidea (four extant families – Dicranolasmatidae,
Nemastomatidae, Nipponopsalididae, Trogulidae)
Suborder Laniatores
Infraorder Insidiatores
Superfamily Travunioidea (three families – Nippononychidae,
Paranonychidae, Travuniidae)
Superfamily Triaenonychoidea (two families – Synthetonychiidae,
Tryaenonychidae)
Infraorder Grassatores
Superfamily Assamioidea (two families – Assamiidae, Pyramidopidae)
Superfamily Epedanoidea (five families – Epedanidae, Petrobunidae,
Podoctidae, Sandokanidae, Tithaeidae)
Superfamily Gonyleptoidea (seven families – Agoristenidae, Cosmetidae,
Cranaidae, Gonyleptidae, Manaosbiidae, Stygnidae, Stygnopsidae)
Superfamily Phalangodoidea (one family – Phalangodidae)
Superfamily Samooidea (five families – Biantidae, Escadabiidae,
Kimulidae, Samoidae, Stygnommatidae)
Superfamily Zalmoxoidea (four families – Fissiphalliidae, Guasiniidae,
Icaleptidae, Zalmoxidae)
Araneae
Pocock (1892) – basic terms in the classification of spiders
Petrunkevitch (1939) – classification of Araneae
Dubinin (1959) – subclass Araneae with three orders: Liphistiomorphae,
Mygalomorphae, and Araneomorphae
Platnick and Gertsch (1976) – order Araneae; their system is now followed.
Suborder Mesothelae Pocock
Suborder Opisthothelae Pocock
Infraorder Mygalomorphae Pocock
Infraorder Araneomorphae Smith
Bibliography 41
Lehtinen (1978) – three groups without ranking: Theraphosomorpha, Araneomorpha,

and Amaurobiomorpha
Eskov and Zonshtein (1990) – order Araneida (Araneae)
1. Suborder Theraphosina (Orthognatha)
1 .1. Infraorder Liphistiomorphae (family Liphistiidae)
1.2. Infraorder Theraphosomorphae (Mygalomorphae)
1.3. Infraorder Filistatomorphae (family Filistatidae)
2. Suborder Araneina (Labidognatha)
2.1.
Infraorder Geralycosomorphae (labidognathous carboniferous
“liphistiomorphs”)
2.2. Infraorder Dysderomorphae
2.3. Infraorder Hypochilomorphae (Hypochiloidea and Austrochiloidea)
2.4. Infraorder Araneomorphae
Eskov and Zonshtein (1990) also group the seven infraorders into three grades dis-
tinguished by symplesiomorphies: Arthrabdominata, Sigillata, and Asigillata.
Other sources: Bristowe (1938), Dubinin (1957 1959), Giribet et al. (2002),
Giribet and Kury (2007), Hansen and Sørensen (1904), Harvey (1990, 1992), Kury
(2011a, b, 2013), Lehtinen (1967, 1975, 1978), Petrunkevitch (1939), Platnick and
Gertsch (1976), and Quintero (1983).
There are some historical reviews about the development of the research on dif-
ferent orders: Lourenço (2014, Scorpiones), Bonnet (1945, Araneae), and Savory
(1961, Arachnida).
The number of non-acarine Arachnida in the world is at least 58,900 spp.
(Mikhailov 2013). After the same source, the Acari would be 54,312 spp. or in total
113,212 spp. of Arachnida.
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Chapter 4
The Fossil Arachnida
Abstract Concerning the fossil orders, Dunlop (1996a, b) presents a revised con-
cept of the formerly accepted (Weygoldt and Paulus 1979; Shultz 1990) five fossil
orders of Arachnida. The subclasses of Petrunkevitch (1949) have been rejected,
Anthracomartida – synonymized with Trigonotarbida. It was concluded that the
position of Phalangiotarbida and Haptopoda remains obscure. Kustarachnida,
according to Dunlop (1996a, b), represents misidentified opilionids.
Concerning the fossil orders, Dunlop (1996a, b) presents a revised concept of the
formerly accepted (Weygoldt and Paulus 1979; Shultz 1990) five fossil orders of
Arachnida. The subclasses of Petrunkevitch (1949) have been rejected,
Anthracomartida – synonymized with Trigonotarbida. It was concluded that the
position of Phalangiotarbida and Haptopoda remains obscure. Kustarachnida,
according to Dunlop (1996a, b), represents misidentified opilionids.
According to the table in the paper of Dunlop et al. (2008), by March 2008, there
were 1593 spp. of fossil Arachnida, including 111 Scorpiones, 25 Opiliones, 38
Pseudoscorpiones, 5 Solifugae, 294 “Acari,” 1 Palpigradi, 15 Ricinulei, 979
Araneae, 9 Amblypygi, 7 Uropygi, and 4 Schizomida (actually the Schizomida are
6). Extremly rich fossil fauna has been found in the Burmese amber. “Fossils from
the mid-Cretaceous (c.99 Ma) Myanmar (Burma) amber include all extant orders of
Arachnida, including the earliest representatives of Schizomida, Parasitiformes, and
Palpigradi…The most abundant and diverse arachnid order is the Araneae, with 38
families, 93 genera, and 165 species recorded tom date” (Selden and Dong Ren
2017).
Going through the recent analytical work on the position of different (recent)
orders in the system of Arachnida, the following summary appears:
Palpigradi Palaeokoenenia mordax Rowland et Sissom, 1980, from the late
Tertiary (?Pliocene) of Arizona and the Electrokoenenia yaksha Engel et Huang
from the Cenomanian Burmese amber remain the only known fossil Palpigrades.

https://doi.org/10.1007/978-3-319-74418-6_4
46 4 The Fossil Arachnida
Solifugae Only five fossil species of these large arachnids, living mostly in dry
areas, are known (from Poland, USA, Brazil, and two from the Baltic and Dominican
amber, respectively) (Poinar and Santiago-Blay 1989; Dunlop et al. 2004).
Ricinulei Selden (1992) revised the fossil ricinuleids, describing 15 valid fossil
species in 4 genera and 2 families from the Pennsylvanian Coal Measures of Europe
and North America.
Amblypygi Eight valid species of fossil whip spider are accepted by Dunlop et al.
(2008): four from Europe and North America; two from Mexican and Dominican
amber, respectively; another one from Mexican amber (Poinar and Brown 2004);
and one from the Crato Formation in Brazil (Dunlop and Martill 2002). According
to Dunlop and Mrugalla (2015), there are already 11 spp. The Late Carboniferous
whip spider Graeophonus anglicus Pocock 1911, was redescribed by Dunlop et al.
(2007a, b). The species Electrophrynus mirus Petrunkevitch 1971, was considered
representing new genus and family, but both have been strongly contested by Armas
(2006) and Dunlop and Mrugalla (2015). According to Selden et al. (1991),
Amblypygi originated at least by the mid-Devonian (ca. 380 Ma). Presently, there
are no Amblypygi living in Europe, and very few are known from North America,
north of Rio Grande.
Thelyphonida (Uropygi) According to Harvey (2003), there are nine species of
fossil Uropygi, but after the recalculation of Tetlie and Dunlop (2008) and Dunlop
et al. (2008), their number has been reduced to six to seven. Cai and Huang (2017)
described from Upper Cretaceous Burmese amber the new genus and species of
Thelyphonida Mesothelyphonus parvus - the earliest Thelyphoninae known so far.
Schizomida Five fossil Schizomida genera have been described so far:
Antillostenochrus Armas et Teruel and Rowlandius Reddell et Cokendolpher from
the Miocene amber of the Dominican Republic; Calcitro Petrunkevitch from China,
Oligocene, and the USA, Pliocene (Calcitronidae); and Onychothelyphonus Pierce
and Calcoschizomus Pierce from the USA, Pliocene (Hubbardiidae) (Petrunkevitch
1945; Pierce 1951). From them, only Rowlandius contains extant species.
Scorpiones The phylogenetic position of the scorpions remains a key question to
resolve in studies of Chelicerate evolution (Weygoldt 1998; Dunlop and Braddy
2001). Scorpions are considered since longtime as the oldest known arachnids
(Selden 1993a, b). Pocock (1893) divided the Arachnida into Ctenophora (scorpions)
and Lipoptena (the remaining Arachnida). Dubinin (1962) unites Scorpiones and
Eurypterida in a class (now clade) Scorpionomorpha. As scorpions are known as (sea
?) forms from the Silurian and the Devonian, very different from the extant forms, the
dispute is between paleontologists and neontologists. The familial and suprafamilial
classification of the scorpions is very controversial (see Soleglad and Fet 2003;
Prendini and Wheeler 2005). The extant scorpions are assigned to the Orthosterni – a
group or “parvorder” (Soleglad and Fet 2003) – known from the Carboniferous to the
present. The number of superfamilies is not definitely established.
4 The Fossil Arachnida 47
Despite their hard cover, the fossil scorpions are relatively few in numbers. Fet
et al. (2000), based on the monograph of Kjellesvig-Waering (1986), quote 78 spp.
The further transformations and new descriptions from Baltic amber (Lourenco
2012), Burmese amber (Santiago-Blay et al. 2004), etc. increased this number to
111 by 2008 (Dunlop et al. 2008). The table in the last mentioned paper indicates
one curious (and unique among Arachnida) particularity of fossil scorpions: 79 spp.
are from Paleozoic age, 16 from Mesozoic, and 16 from Cenozoic.
The older concept that scorpions are very primitive (partly because are very
ancient) and are the ancestors of the other arachnids is strongly denied by cladists,
which find (Shultz 1990) that they are derived arachnids forming a Dromopoda
clade together with Opiliones, Solifugae, and Pseudoscorpiones. Some authors
(Uchida 1966) still support the primitive character of the scorpions. The cladistic
analysis of Dunlop and Braddy (2001) did not provide an undisputable end of the
contest between the three main hypotheses: (1) Scorpions are the sister group of all
other arachnids; (2) scorpions are derived group of arachnids as part of Dromopoda
clade (and perhaps sister group of opilionids); and (3) scorpions are sister group of
eurypterids. Support exists for any of the alternative hypotheses, based on
morphological and molecular data and their interpretation. We can only admire the
perception of the older authors (Pocock, Dubinin), which had in hand much less
instruments for comparative analysis.
Gess (2013) published “the earliest record of terrestrial animals in Gondwana”
from South Africa. This was the new genus and species Gondwanascorpio
emzantsiensis, a scorpion from a Famennian (Late Devonian) formation.
Pseudoscorpiones In the catalogue of Harvey (1990) were listed 32 valid fossil
pseudoscorpions from amber (Burmese, Chinese, Baltic, and Dominican). Schwaller
et al. (1991) published the oldest pseudoscorpion from the mid-Devonian of Gilboa,
New York State (the first Paleozoic pseudoscorpion). More details could be found
in Harvey (1990) and Spahr (1993). Dunlop et al. (2008). After the new data of
Hendericks (2005) and Judson (2007), respectively, from Baltic and Dominican
amber, the total number of fossil pseudoscorpions amounts on 38 species (Dunlop
et al. 2008). According to Harms and Dunlop (2017), overall 16 of the 26 [27?] fam-
ilies of living pseudoscorpions have been documented from fossils and 49 currently
valid species are recognized. “The present-day distributions of some group
(Faellidae and Pseudogarypidae) is relictual and highlights past extinction events”
(Harms and Dunlop 2017). Harvey et al. (2017) described from the Burmese amber
(mid-Cretaceous, ca. 90 Mya) the oldest member of Chthoniidae Weygoldtiella
plausus Harvey et al.
Opiliones The fossil record of this important group has been summarized by Dunlop
(2007a, b). As we can see from the summary table in Dunlop’s paper, listing all
known by this time fossil Opiliones, the fossil record of this group, containing now
more than 6500 described species, consists of only 31 entries, part of them unnamed.
Eight of them are from the Paleozoic (from Lower Devonian in Scotland to Upper
Carboniferous in France). Only two taxa are recognized as genuine Mesozoic har-
vestmen (from the Lower Cretaceous in Australia and from Upper Cretaceous
Myanmar amber). None of them belongs to the modern families. The remaining 21
entries are Cenozoic and belong to the present-day families Trogulidae, Phalangiidae,
Sclerosomatidae, Nemastomatidae, Caddidae, Kimulidae, Samoidae, and
Cladonychiidae and are even assigned to the modern genera Trogulus, Kimula,
Caddo, Dicranopalpus, Leiobunum, Nemastoma, Sabacon, etc. Sometimes these
findings cast new light over the present-day distribution of families like Caddidae or
Kimulidae and help in restoring the paleodistribution of the Opiliones. Actually, only
25 species are recognized as valid (Dunlop et al. 2008), with only one sp. from the
Mesozoic. Ten of the findings come from the Baltic amber, considered to be of
Eocene age (some 38–54 million years ago), mostly old information, summarized by
Koch and Berendt (1854) and Menge (1854). The modern research in the Dominican
amber by Cokendolpher (1986) and Cokendolpher and Poinar (1982, 1998) brought
another four species. They belong to the family of Phalangodidae and Kimulidae,
still living in the Antillean area. The age of this fauna is not very different from the
age of Baltic amber fauna, but the research in the Dominican amber has much shorter
history (for the harvestmen since 1978), and there is a lot more to expect.
Only 14 fossil Opiliones species have been recorded before 1955, so the task of
Petrunkevitch (1955), who summarized the existing scant information concerning
fossil Opiliones, seems to be relatively simple. But it is amazing that these
widespread animals are so rare in the fossil record (the reasons are explained by
Dunlop, op. cit.). The oldest known Devonian harvestmen look very much like the
modern forms, and it is a well-founded guess that there are also Silurian or other
pre-Devonian Opiliones-like creatures, which are to be discovered. By Devonian
(the Opiliones are considered to be at least 400 million years old), most of the
known arachnid orders already existed as terrestrial, air-breathing animals. However,
Garwood et al. (2014) described a new fossil suborder and concluded that “The
discovery of Tetrophthalmi alters molecular divergence time estimates, supporting
Carboniferous rather then Devonian diversification for extant suborders and directly
impacting inference of terrestrialization history and biogeography.”
The peculiar suborder Cyphophthalmi is represented by Siro platypedibus
Dunlop et Giribet, 2003, a single specimen, described from Tertiary Bitterfeld
amber, and by a new genus and species Palaeosiro burmanicum Poinar, 2008, from
the Early Cretaceous Burmese amber (Upper Albian (100 to 105 m.y.B.P.)
Araneae The bulk of Arachnida (the Acari excluded), both living over 40 000 spp.
and fossil (979 spp.), belong to the order of spiders. A general review of the spider
fossil record is provided by Penney and Selden (2006a, b). The oldest record is from
Gilboa, New York (mid-Devonian, 380 Ma) (Selden et al. 1991). According to
Dunlop et al. (2008), the vast majority of fossil spiders (c. 820 spp.) originate from
amber: 540 spp. from the Baltic amber and 170 spp. from the Dominican amber.
There are also findings in the younger (Cenozoic) amber and copal of France,
Germany, Ukraine, China, etc. Spiders are described recently also from Cretaceous
ambers in Siberia, Canada, Burma, the USA, and Spain (see Dunlop et al. 2008).
Significant findings from the Cretaceous occurred also in Siberia and Mongolia
(Eskov and Zonshtein 1990) and Spain (Selden and Penney 2003) and from the
Crato Formation in Brazil (Selden et al. 2006).
4 The Fossil Arachnida 49
Acari As tiny creatures, mites are not many in the fossil record. After the publica-
tion of Krivolutsky (1973) about finding fossil Oribatida (including some recent
genera (Cultroribula, Achipteria) from the Jurassic and Cretaceous deposits in
Russia, it was accepted (Hammer and Wallwork 1979) that “families of oribatids
that we can recognize today were present during the Jurassic and undoubtedly ear-
lier.” It was proved some time later by Norton et al. (1988) with the finding of oriba-
tids in the terrestrial Devonian deposit near Gilboa, New York.
Describing the second fossil Opilioacarid, Dunlop et al. (2010a) analyzed the
status of the fossil Acari by this time. According to them and to Dunlop et al.
(2010b), 292 fossil actinotrichid species (including recent forms found as subfossils)
have been recorded. Actinotrichids were certainly present by the Early Devonian
(ca.410 Ma) and potentially much earlier. There are only 14 fossil species of
anactinotrichids, and their record goes back to Cretaceous (Poinar and Buckley
2008) – two opilioacarids, four gamasids, no holothyrids, and eight ticks. The small
number of fossil gamasids is really “puzzling.” Most genera of the fossil Gamasida
and Ixodida are the same as in the extant fauna. Only in the Burmese amber have
been found two new genera of Ixodidae (Cornupalpatum Poinar et Brown, 2003,
and Compluriscutula Poinar et Buckley, 2008).
Many years since this time, the fossil record of Oribatida is still poor (Dunlop
et al. 2008). According to the table of Dunlop et al. (2008), by March 2008, there
were known 294 “Acari.” Obviously, Oribatida are privileged as being preserved in
the older rock layers (not amber), because of their hard armor. Such armor, even
harder, has also the caeculids, but they are much smaller group, and only two species
have been described recently from Baltic and Dominican amber (Eocene). From the
Canadian amber has been described as new genus and species Proterythraeus south-
cotti Vercammen-Grandjean, 1972. From the Baltic amber have been recorded
another new genus (Arythaena Menge, 1854) and eight other Erythraeidae species,
assumed to belong to the present-day widespread genera Erythraeus, Balaustium,
and Leptus. Provided that the material still exists, these identifications should be
revised (Zacharda and Krivolutsky 1985).
The fossil Mesostigmata are rare. From the Baltic amber (Eocene, ca. 44–49 Ma),
Dunlop, Kontschán, and Zwanzig (2013) recorded Uropodina and Dunlop et al.
(2014) Laelapidae of the extant genus Myrmozercon.
Opilioacarida The first fossil opilioacarid to be described is Paracarus pristinus
Dunlop, Wunderlich et Poinar, 2004, from Eocene Baltic amber. It has been attached
to the genus Paracarus Chamberlin et Mulaik, 1942, represented by a modern spe-
cies from Central Asia, and represents the northernmost species of Opilioacarida.
The second fossil opilioacarid (? Opilioacarus aenigmus Dunlop, Sempf et
Wunderlich, 2010a) was described again from Baltic amber (Eocene, 45–50 Ma).
However, it has been assigned tentatively to the now Mediterranean genus
Opilioacarus With, 1903.
Paleozoogeography of Arachnida Important information concerning the past dis-
tribution of the Arachnida comes from the amber – Baltic, Dominican, Burmese,
Lebanese, Chinese, etc. The amber is of different geological age. Other findings of
fossil Arachnida are from the Crato Member of the Santana Formation in Brazil
(Martill et al., Eds, 2007), Kazakhstan, etc.
Here are some data showing the differences between the past and present
distribution of Arachnida:
Palpigradi Remaining are the findings of Palaeokoenenia mordax Rowland et
Sissom from the late Tertiary (?Pliocene) of the USA (Arizona) and of the
Electrokoenenia yaksha Engel et Huang from the Cenomanian Burmese amber
(Myanmar).
Ricinulei The findings of the ancient group in the Pennsylvanian Coal Measures of
Europe and North America (Nam.- Steph., 327–290 Ma) (Selden 1992) show a
much broader distribution in the distant past. Actually, no ricinuleid lives in the
Palearctic and (practically) in the Nearctic (only one sp. Pseudocellus dorotheae
from Texas, on the border of Mexico). There are no extant Ricinulei also in Asia, but
Wunderlich (2012) described? Poliocera cretacea from Burmese Cretaceous amber.
Scorpiones There are at least 45 fossil families of scorpions (almost entirely from
the suborder Branchioscorpionina).
Solifugae Fossil species have been described not only from countries, where there
are Solifugae even now (USA, Brazil), but also from Poland and the Baltic amber.
Amblypygi Most of the known fossil Amblypygi come from countries where the
order is represented now (Brazil, Mexico, Dominican Republic). However, four
species are recorded from Europe and North America (Nam.-Steph., 327–290 Ma).
Thelyphonida (Uropygi) Different genera from the now existing have been
described from the Carboniferous of North America and Europe. Presently, Uropygi
are not known from Europe (even closely) and from most of North America.
Schizomida One representative of the order was found from the Oligocene of
China. Presently, the only Schizomida known from this country is Bamazomus sia-
mensis from Hong Kong, within the present range of the order.
Opiliones There are fossil families of Opiliones, belonging to four recent subor-
ders: Cyphophthalmi (Sironidae), Eupnoi (Phalangiidae), Dyspnoi (Caddidae,
Nemastomatidae), and Laniatores. The material from Baltic amber (Eocene, ca.
44–49 Ma) has been presented by Koch and Berendt (1854) and supplemented or
corrected by Roewer (1939), Dunlop and Barov (2005), and Dunlop and Mitov
(2009). From the Bitterfeld amber (Germany, Oligocene–Chattian, 25.3–25.8 Ma)
have been described the first fossil cyphophthalmid Siro platypodibus Dunlop et
Giribet, 2003, and many other Opiliones, belonging to extant genera (Lacinius
Thorell, Amilenus Martens, Mitostoma Roewer, ?Histricostoma Kratochvil, Caddo
Banks, Dicranopalpus Doleschall, and even species like Lacinius erinaceus Staręga,
known from Caucasus) (Dunlop and Mitov 2009). It is supposed that Opiliones of
the extant genera and even species existed some tens of millions year ago (in the
Paleogene). However, the Opiliones are much older and, as specialists say, can be
Bibliography 51
traced back to at least the mid-Paleozoic. With the discovery of Caddo in North
Europe, it was supposed even that the now discontinuous species of this genus
(Japan and North America) have been spread in the Tertiary much broader. The fact
that extant species have been recorded in the Oligocene, more than 20 Ma ago,
raises the problem of the pace of evolution. Cokendolpher and Cokendolpher (1982)
reexamined the tertiary harvestmen from Colorado.
Araneae As it was said, the age of the fossil spiders is mostly from the Mesozoic,
the oldest known being at 380 Ma (mid-Devonian of Gilboa). The Russian arach-
nologist Eskov (1984) used the finding of a member of “Gondwanan” family
Archaeidae (Jurarchaea zherikhini) in the Upper Jurassic of Kazakhstan, as an
argument against the attempts to explain many distributions with Gondwana. The
actual distribution of the family (South Africa, Madagascar, Australia, Tasmania,
New Zealand, and Patagonia) was considered by Legendre (1977) for “typically
Gondwanan.” These spiders are known also from Baltic amber (seven species) and
probably their present kind Gondwanan distribution is a relict one from a much
more cosmopolitic distribution in the past. About the same time was published also
a revision by Forster and Platnick (1984), leaving within Archaeidae only four gen-
era from the Baltic amber, Australia, South Africa, and Madagascar. This does not
change very much the conclusions of Eskov. Also, as Wunderlich (1995) pointed,
Cyatholipidae are not “representing former parts of the Gondwanaland” (Griswold
1987) but rather a relict distribution of a very wide distribution in the Lower Tertiary
period.
Opilioacarida Paracarus pristinus from Baltic amber has been ascribed by
Dunlop, Wunderlich, and Poinar (2004) to the genus Paracarus, represented today
only by one species in Central Asia. The other fossil Opilioacarid (Opilioacarus (?)
aenigmus Dunlop, Sempf et Wunderlich, 2010a) is conditionally included in the
genus Opilioacarus, living today in the south of Europe (Italy, Greece) and in North
Africa.
Trombidiformes From Canadian amber has been described as new genus and spe-
cies Proterythraeus southcotti Vercammen-Grandjean, 1972. From the Baltic amber
have been recorded another new genus (Arythaena Menge, 1854) and eight other
Erythraeidae species. From the Early Cretaceous (Aptian) Crato Formation, Brazil,
has been described a new large, fossil mite (Arachnida: Acari, Erythraeoidea),
Pararainbowia martilli Dunlop 2007a. Bernini (1991) made general review of the
fossil Acarida, known by this time.
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Chapter 5
Factors Determining the Distribution
of Arachnida
Abstract As factors of distribution of Arachnida are outlined paleogeography and

paleodistribution, age of groups, barriers, bridges, ability to overcome them,
phoresy, dispersal, climate, orography and many other fundamental concepts.
Zoogeography appears to be one of the most amusing and Stimulation of the natural sci-
ences: every few years its fundamental concepts change and one can begin anew.
(P.M. Brignoli 1983)
5.1 Some Basic Notions
5.1.1 Zoogeography and Biogeography: The New Approach
In the article of Posadas et al. (2006), a review is made of the development of the
historical biogeography and the basic concepts of this science. The Argentinian
authors never mention the word “zoogeography.” This is also the approach of most
Western biogeographers. They also neglect entirely Russian and any other publica-
tion in non-Western languages. Most contemporary authors use the term “biogeog-
raphy”; while the notions are generally the same in zoogeography and
phytogeography, I think that when we have to deal with animals, it is entirely justi-
fied to use the word “zoogeography.”
Basic review of the main concepts and ideas would be useful to the readers of
this text. Most information derives from the analysis of Posadas et al. (2006).

https://doi.org/10.1007/978-3-319-74418-6_5
60 5 Factors Determining the Distribution of Arachnida
5.1.2 Centers of Origin and Dispersal
Species have center of origin, from which some individuals disperse by chance and
then evolve through natural selection (Darwin-Wallace traditional approach)
(Matthew 1915).
x
x x
Trying to explain the distribution of various animals, the older zoogeographers

“discovered” many land bridges between continents and islands, even entire conti-
nents. With the development of modern explanations of these distributions (conti-
nental drift, etc.), most of these bridges became unnecessary and now have mostly
historic value. They are explained in details by Müller (1974a, b).
Paleantarctic This is a term created by Jeannel (1942) and means a continent,
appearing in the Jurassic during the first breakup of Gondwana and the beginning of
the “opening up” of Indian Ocean.
Schuchert Land After Schuchert and Ihering, a North American-Pacific mountain
range, connecting North with South America during the Cretaceous (from West to
East from the Pacific Coast to the Missouri Basin).
Lemuria Imaginary land bridge across the Indian Ocean, used by the older authors
(Sclater, Ihering) to explain the connection of the fauna of Madagascar and India in
the Cretaceous or early Tertiary.
Beringia A land bridge (actually the notion is used to indicate a much larger
dry land), connecting several times during the Tertiary and during the cold peri-
ods of the Pleistocene (Müller 1974a, b) Eurasia and North America. Through it
massif exchange occurred of faunas and, finally, the penetration of Man in the
Americas.
Archiplata According to Ihering (1927), this was a land bridge in the place of
today’s Andean South America. In Cretaceous time it connected the Archinotis with
Schuchert land. This bridge was an attempt to explain the interchange of fauna
between North and South America in the Tertiary.
Archiguiana Supposed Cretaceous island in the region of Venezuela and the
Guianas.
5.1 Some Basic Notions 61
Mesosoic Archhelenis Land bridge, connecting South America with South West
Africa by way of Tristan da Cunha.
Archatlantis Ihering (1927) coined this land bridge to connect the Antilles and
Florida with North Africa and southern Spain and including the Azores, Canaries,
and Cabo Verde islands.
Archinotis Land bridge constructed by the older biogeographers to connect New
Zealand and Australia with southern South America through Antarctica and the
South Pacific islands (Cretaceous and early Tertiary).
Canary Bridge Connection of the Eastern Canary Islands (Lanzarote) with the
African mainland. Some biogeographers support this idea, under variations. After
Sauer and Rothe (1972), “Separation of the eastern Canaries from Africa might have
been by rifting and a land connection might still have existed in the lower Pliocene.”
The shells found on the islands indicate that until about 12 million years ago, they
were inhabited by large flightless birds.
Tyrrhenian Bridge Supposed Riss-Glacial land bridge between Tuscany and
Corsica and Sardinia. This bridge was invented in the efforts to explain mainly the
distribution of herpetofauna.
Galita Bridge Connecting Sardinia and Corsica with Africa.
Balearic Bridge Connecting Corsica, Sardinia, and Balearic Islands with Spain.
Provence Bridge Connecting Sardinia and Corsica with Provence.
Buffon’s Law of Endemism Different areas, no matter how similar their physical
conditions, generally house different species.
5.1.3 Development of Atlantic Ocean
According to Sclater et al. (1977), the development of Atlantic Ocean is as

follows:
165 Ma (Middle Jurassic) – The Atlantic Ocean is closed. The Baffin Bay is com-
pletely closed, and the shelf of Greenland fits neatly into the shelf of Labrador.
Iberia is overlapping slightly onto the Atlas Mountains. To the South there is
almost a perfect fit of the outline of South America and Africa.
Map 5.1 The reconstructed position of the Atlantic continent 165 Ma (After Sclater et al. 1977,
modified)
140 Ma (Late Jurassic–Early Cretaceous) – The main event between 165 and
140 Ma is the opening up of North Atlantic.
125 Ma (Early Cretaceous) – Between 165 and 140 Ma, Africa is still drifting away
from North America along the Newfoundland fracture zone. By 125 Ma Iberia
and Africa separate, and the North Atlantic is connected with Tethys. The proto-
Caribbean arc system has been as high and active as it is today.
110 Ma (boundary between Lower and Middle Cretaceous) – South America and
Africa start to separate. The North and South Atlantic are still separated by part
of Africa and South America. The elevated Walvis and Rio Grande Rise split the
South Atlantic into two separate basins.
95 Ma (Middle Cretaceous) – Africa and Iberia continue to separate from North
America. The rise between both parts of South Atlantic starts to sink, and the
water will be able to flow from the South to the North through the Vema Gap.
Europe starts separating from North America.
5.1 Some Basic Notions 63
80 Ma (Late Cretaceous–Senonian) – Greenland starts to move away both from

Europe and North America. Iberia is well separated from Africa and is far from
North America.
65 Ma (boundary between Cretaceous and Tertiary) – Europe is already far from
North America, and Iberia is fixed to Europe.
53 Ma (boundary Paleocene–Eocene) – Baffin Bay is starting to open; Europe is
separating much faster from Greenland and North America. The South Atlantic
is already wide and deep.
36 Ma (boundary Eocene–Oligocene) – The Atlantic Ocean acquired almost its
present shape.
21 Ma (Early Miocene), 10 Ma (Late Miocene) – Between the Oligocene and the
Miocene, the Cabo Verde Plateau is uplifted, and the Azores-Gibraltar Ridge
starts to form.
3.5 Ma – Uplift of the Isthmus of Panama and the waters of Pacific and Atlantic
Oceans are separated.
Map 5.2 The present Atlantic Ocean (After Sclater et al. 1977, modified)
Conclusions
1. Since the Middle Jurassic (165 Ma) till the Lower Cretaceous (125 Ma), Northern
Atlantic was completely closed basin.
2. Since the Lower Cretaceous (125 Ma) till Upper Cretaceous (110–95 Ma),
Southern Atlantic was also completely closed basin.
5.1.4 Development of the Indian Ocean
With a surface of 73,556,000 km², the Indian Ocean covers 20% of the total water
surface of the planet. There are many scientific and semi-scientific attempts to ana-
lyze the India-Madagascar paleoconnections, the history of the massifs, and the
remains of them as archipelagoes with most interesting fauna (Seychelles, Mascarene
and others), the speculations concerning the hypothetic continent Lemuria, and
many others.
The arbitrary boundary between the Indian and Atlantic Oceans is following the
20°E meridian; along the meridian 146°55′ E runs the boundary with the Pacific.
The average depth of Indian Ocean is 3890 m and the maximal 8047 m (Diamantina
Trench).
Speaking of “Sudamadie,” Jeannel (1965) is basing his conclusions on the two
landmasses – the bloc South Africa-Madagascar (the continental part) and the
remains of the so-called Subantarctic Plateau (the islands Crozet, Prince Edward,
Kerguelen, and McDonald). Jeannel is sticking to the term Lemuria as a firm land,
remains of which are the Comoro, Seychelles, and Mascarene Islands. He also
accepts unconditionally the idea of a former bridge (land connection) between
South Africa and South America as such connection is needed to explain the distri-
bution of some groups of insects between the two continents. But the riddle still
stays, as the analysis of Jeannel is based on the paleogeography of Furon (1959).
Since this time many new data was obtained, modifying the zoogeographical con-
clusions. Now it is accepted that the African, Indian, and Antarctic plates converge
in the Indian Ocean in the so-called Triple Point Rodriges.
Lemuria of Jeannel is called now Mascarene Plateau (2000 km long underwater
plateau East of Madagascar). Concerning the mid-ocean underwater ridges, there
are several such active ridges, keeping in mind that Indian Ocean is the youngest of
all oceans.
In his works Jeannel pays special attention to Kerguelen Islands, where deep lay-
ers of fossil plants are found from Eocene and Oligocene. According to Jeannel, this
area has been an extensive land with rivers and forests, similar to those existing now
in Tierra del Fuego. Now is considered that Kerguelen Archipelago (7215 km2,
almost the size of Corsica) is situated on a sunken microcontinent having a surface
of 2.2 million of km2.
The development of Indian Ocean has been outlined in details according to mod-
ern geophysical data by Ali and Aitchison (2008).
5.2 Paleogeography and the Past Distribution of Arachnida 65
5.2 Paleogeography and the Past Distribution of Arachnida
5.2.1 Age of the Faunas
Here seems useful to remind the reader the time span of the different periods (BP).
The oldest and the predominant record of the different groups of arachnids is as
follows:
Palpigradi Palaeokoenenia mordax Rowland et Sissom is from the late Tertiary
(?Pliocene). The Pliocene extended from 5.332 million to 2.588 million years before
present. Engel et al. (2016) published from Cenomanian (mid-Cretaceous) Burmese
amber a much older (94–97 My old) member of Palpigradi (Electrokoenenia yaksha
Engel et Huang), the first Mesozoic fossil of this order.
Solifugae Five fossil species known (from Poland, USA, Brazil and from the Baltic
and Dominican amber, respectively). The age of Baltic amber is 44 million years
ago (Eocene), of Dominican amber – 25–15 Ma.
Ricinulei Known from the Pennsylvanian Coal Measures of Europe and North
America (Nam.- Steph., 327–290 Ma). Wunderlich (2012) described a new fossil
species from the Cretaceous of Burma (Myanmar).
Amblypygi There are nine valid species of fossil whip spider:
–– Four from Europe and North America (Nam.-Steph., 327–290 Ma)
–– Three from Mexican amber (30 million years)
–– One from Dominican amber (15–45 million years)
–– One from the Crato Formation of Brazil (Early Cretaceous, mostly during the
early Albian age, about 108 million years ago)
Thelyphonida (Uropygi) There are 7–8 fossil Uropygi from Brazil (Mesoproctus
rowlandi Dunlop, 1998, Lower Cretaceous (Aptian) of the Crato Member of the
Santana Formation) and from Carboniferous of North America and Europe
(Geralinura, Proschizomus, Parageralinura, Protelyphonus). A new genus and spe-
cies (Mesothelyphonus parvus Cai et Huang, 2017) has been described from
Burmese amber (Cretaceous).
Schizomida Three fossil Schizomida have been described from Arizona, USA,
one from the Oligocene of China (the Oligocene epoch is from 34 to 23 million
years BP) and two from Dominican amber (Krüger and Dunlop 2010).
Scorpiones Known from the Silurian and the Devonian (sea forms ?), scorpions
are considered since longtime as the oldest known arachnids. The extant scorpions
are assigned to the Orthosterni, known from the Carboniferous to present.
By 2008 there were known 111 spp. of fossil scorpions (79 spp. are of Paleozoic
age, 16 of Mesozoic, and 16 of Cenozoic). According to Prendini (2011), fossil
scorpions include 45 families, 86 genera, and 120 species (Akravidae being consid-
ered by the present author as recent). They belong to the extinct suborder
Branchioscorpionina Kjellesvig-Waering, 1986 (42 families), and to suborder

Neoscorpionina Thorell et Lindström, 1885 (three families).
Pseudoscorpiones Most of the 38 fossil species of pseudoscorpions come from
amber (Burmese, Chinese, Baltic, and Dominican). Schawaller et al. (1991) pub-
lished the oldest pseudoscorpion from the mid-Devonian (Paleozoic) of Gilboa,
New York State (380 Ma), the only fossil family of Pseudoscorpiones (Dracochelidae;
Schawaller, Shear et Bonamo, 1991).
Opiliones Eight entries in the table of Dunlop (2007) are from the Paleozoic (from
Lower Devonian in Scotland to Upper Carboniferous in France). Two taxa are
Mesozoic (from the Lower Cretaceous in Australia and from Upper Cretaceous
Myanmar amber).
The remaining 21 entries are Cenozoic and belong to the present-day families.
Ten of the findings come from the Baltic amber of Eocene age (some 38–54 Ma
years ago), mostly old information. The different beds of Dominican amber have
been dated as Miocene (15–20 million years) to Upper Eocene (ca. 30–45 million
years). So, the age of this fauna is not very different from the age of Baltic amber
fauna, but the research in the Dominican amber has much shorter history (for the
harvestmen since 1978), and there is a lot more to expect.
It is of note the description of a new genus and species of Cyphophthalmi
(Palaeosiro burmanicum) by Poinar (2008) from the Burmese amber of Upper
Albian (100–105 my BP).
The oldest known Devonian harvestmen, considered to be at least 400 million
years old, look very much like the modern forms, and it is a well-founded guess that
there were also Silurian or other pre-Devonian Opiliones-like creatures, which are
to be discovered.
Garwood et al. (2014) described a new fossil suborder Tetrophthalmi, based on
two fossil species – the new Carboniferous harvestman Hastocularis argus and the
Devonian Eophalangium sheari. This finding supports Carboniferous rather than
Devonian diversification for extant suborders of Opiliones.
Araneae The vast majority of fossil spiders (c. 820 spp.) originate from amber:
540 spp. from the Baltic amber and Dominican amber c.170 spp. (Miocene to Upper
Eocene (15–20 million years to ca. 30–45 million years). There are also findings in
the younger (Cenozoic) amber and copal of France, Germany, Ukraine, China, etc.
Spiders have been described recently also from Cretaceous ambers in Siberia,
Canada, Burma, the USA, Spain, and Mongolia and from the Crato Formation of
Brazil. It means that the age of the fossil spiders is mostly from the Mesozoic, the
oldest known being at 380 Ma (mid-Devonian of Gilboa).
Acari Most of the Acari (except of Opilioacarida and Holothyrida) are not so
important zoogeographically, many of them being parasitic, and the others are not
adequately studied and are not treated in details in this publication.
Opilioacarida Two fossil spp. have been described from Baltic amber (44 million
years ago, Eocene): Opilioacarus (?) aenigmus Dunlop, Sempf et Wunderlich,
2010, and Paracarus pristinus Dunlop, Wunderlich et Poinar, 2004.
Acariformes
Trombidiformes Among the other Arthropoda from Baltic amber by Koch and
Berendt (1854) have been recorded eight spp. of Erythraeidae (ascribed to
“Rhyncholophus,” now Erythraeus, Balaustium, and Leptus). Menge (1854)
described from Baltic amber the new genus and species Arythaena troguloides;
Vercammen-Grandjean (1972) found in Canadian amber (Cretaceous) another new
genus and species: Proterythraeus southcotti. Zacharda (in Zacharda and Krivolutsky
1985) described Mesoanystis taymirensis from Taymyr Peninsula, Upper Cretaceous
amber (the oldest fossil anystid mite known), and Palaeoerythracarus sachalinensis
from Sachalin (Paleogene amber). Bartel et al. (2015) described three new spp. of
Fessonia (Smarididae) from Baltic amber, and Calyptostoma katyae Koniktyiewicz,
Wohltmann et Makol, 2016, was the first fossil Calyptostoma (described also from
Baltic amber).
Dunlop (2007) described from the Early Cretaceous (Aptian) Crato Formation
from Ceará State, Brazil, a new large, fossil mite (Arachnida: Acari, Erythraeoidea),
Pararainbowia martilli.
Sarcoptiformes
Oribatida From many places have been recorded fossil oribatid mites, because of
the hard tegument and their massif presence (up to 75%) in the soil fauna (Krivolutsky
and Druck 1986). Norton et al. (1988) recorded 14 fossil Oribatid spp. (including
the new genera Protochthonius and Devonacarus and the new family Devonacaridae)
from terrestrial Devonian deposit near Gilboa, New York. An acarid belonging to
the still-living genus Hydrozetes Berlese, 1902, was found in the Lower Sinemurian
(Jurassic) in a core drilling from Sweden (Sivhed and Wallwork 1978).
Parasitiformes
Mesostigmata Inclusions assignable to the tortoise mites (Mesostigmata,

Uropodina) have been described by Dunlop et al. (2013) from Baltic amber (Eocene,
ca. 44–49 Ma). This is the oldest record of Uropodina. “Fossil mesostigmatid mites
(Acari: Parasitiformes: Mesostigmata) are extremely rare, and specimens from only
nine families, including four named species, have been described so far. A new
record of Myrmozercon sp. described here from Eocene (ca 44–49 Myr) Baltic
amber represents the first – and so far only – fossil example of the derived, extant
family Laelapidae”.
Ixodida In the amber from Cretaceous has been recorded a larval argasid tick
(Carios jerseyi Klompen et Grimaldi, 2001). Another Mesozoic tick (Amblyomma,
Ixodidae) has been described from Dominical amber (Lane and Poinar Jr. 2009), the
first in the New World. Ixodes tertiarius Scudder, 1885, was described from
Oligocene amber (Wyoming, USA). Another tick (Ixodes succineus Weidner, 1964)
is known from the Baltic amber.
We borrowed from the paper of Giribet, Sharma et al. (2012) this chronology:
Geological intervals:
–– 0–35 Ma (separation of Mediterranean plate from Western Europe, separation of
Japan from Eurasia, connection of Iberia to Eurasia)
–– 35–45 Ma (separation of Borneo and Indomalay Archipelago from Eurasia)
–– 45–60 Ma (Balkans connected to Western Europe, Iberia connected to
Mediterranean plate, Balkans, and Japan).
–– 60–75 Ma (Iberia separated from Mediterranean plate, Balkans, and Japan, North
America separated from Western Europe, emergence of Indomalay Archipelago)
–– 75–110 Ma (Mediterranean plate separated from North America, Iberia con-
nected to Western Laurasia, Balkans separated from North America and Western
Europe)
–– 110–120 Ma (Iberia disconnected from other landmasses; Western Europe,
Mediterranean plate, and North America separated from Eastern Laurasia; emer-
gence of Borneo; Indomalay Archipelago nonexistent)
Mediterranean plate, and North America separated from Eastern Laurasia;
Borneo and Indomalay Archipelago nonexistent)
–– 180–250 Ma (Thai-Malay Peninsula disconnected from other landmasses;
Eastern Himalaya disconnected from North America, Western Europe, and
Iberia; Borneo and Indomalay Archipelago nonexistent)
–– 250–296 Ma (Borneo and Indomalay Archipelago nonexistent, other landmasses
connected)
Here seems useful to remind the reader the time span of the different periods (BP).
Silurian – 444–416 Ma (roughly, other figures exist in different sources). Known
scorpions (sea forms?)
Devonian – 416–359 Ma. Known scorpions, spiders, and pseudoscorpions (mid-
Devonian, 380 Mya)
Carboniferous – 359–299 Ma. Known scorpions Orthosterni (the presently existing
scorpions)
Permian – 299–251 Ma
Triassic – 251–200 Ma
Jurassic – 200–146 Ma
Cretaceous – 146–66 Ma
Tertiary – 66–2.6 Ma
Paleogene (Paleocene, Oligocene, Eocene) – 66–23 Ma
Neogene (Miocene, Pliocene) – 23 – 2.6 Ma
Quaternary – 2.6 Ma to the present
Pleistocene – 2.6–0.012 Ma
Holocene – 0.012 Ma to the present
Palaeokoenenia mordax Rowland et Sissom is from the late Tertiary (?Pliocene).
The Pliocene extended from 5.332 million to 2.588 million years before present.
Engel et al. (2016) published from Cenomanian (mid-Cretaceous) Burmese
amber a much older (94–97 My old) member of Palpigradi (Electrokoenenia yaksha
Engel et Huang), the first Mesozoic fossil of this order.
Solifugae Five fossil species known (from Poland, USA, and Brazil and from the
Baltic and Dominican amber, respectively). The age of Baltic amber is 44 million
years ago (Eocene), of Dominican amber – 25–15 Ma.
Ricinulei Known from the Pennsylvanian Coal Measures of Europe and North
America (Nam.- Steph., 327–290 Ma). Wunderlich (2012) described a new fossil
species from the Cretaceous of Burma (Myanmar).
Amblypygi There are nine valid species of fossil whip spider:
–– Four from Europe and North America (Nam.-Steph., 327–290 Ma)

–– Three from Mexican amber (30 million years)
–– One from Dominican amber (15–45 million years)
–– One from the Crato Formation of Brazil (Early Cretaceous, mostly during the
early Albian age, about 108 million years ago)
(Uropygi) Known from the Silurian and the Devonian (sea forms ?), scorpions are
considered since longtime as the oldest known arachnids. The extant scorpions are
assigned to the Orthosterni, known from the Carboniferous to present.
By 2008 there were known 111 spp. of fossil scorpions (79 spp. are of Paleozoic
age, 16 of Mesozoic, and 16 of Cenozoic). According to Prendini (2011), fossil
scorpions include 45 families, 86 genera, and 120 species (Akravidae being consid-
ered by the present author as recent). They belong to the extinct suborder
Branchioscorpionina Kjellesvig-Waering, 1986 (42 families) and to suborder
Neoscorpionina Thorell et Lindström, 1885 (three families).
Eight entries in the table of Dunlop (2007) are from the Paleozoic (from Lower
Devonian in Scotland to Upper Carboniferous in France). Two taxa are Mesozoic
(from the Lower Cretaceous in Australia and from Upper Cretaceous Myanmar
amber).
The remaining 21 entries are Cenozoic and belong to the present-day families.
Ten of the findings come from the Baltic amber of Eocene age (some 38–54 Ma
years ago), mostly old information. The different beds of Dominican amber have
been dated as Miocene (15–20 million years) to Upper Eocene (ca. 30–45 million
years). So, the age of this fauna is not very different from the age of Baltic amber
fauna, but the research in the Dominican amber has much shorter history (for the
harvestmen since 1978), and there is a lot more to expect.
Amber of Upper Albian (100 to 105 my BP). The oldest known Devonian har-
vestmen, considered to be at least 400 million years old, look very much like the
modern forms, and it is a well-founded guess that there were also Silurian or other
pre-Devonian Opiliones-like creatures, which are to be discovered. Garwood et al.
(2014) described a new fossil suborder Tetrophthalmi, based on two fossil species –
the new Carboniferous harvestman Hastocularis argus and the Devonian
Eophalangium sheari.This finding supports Carboniferous rather than Devonian
diversification for extant suborders of Opiliones.
Araneae The vast majority of fossil spiders (c. 820 spp.) originate from amber:
540 spp. from the Baltic amber and Dominican amber c.170 spp. (Miocene to Upper
Eocene, 15–20 million years to ca. 30–45 million years). There are also findings in
the younger (Cenozoic) amber and copal of France, Germany, Ukraine, China, etc.
Spiders are described recently also from Cretaceous ambers in Siberia, Canada,
Burma, the USA, Spain, and Mongolia and from the Crato Formation in Brazil. It
means that the age of the fossil spiders is mostly from the Mesozoic, the oldest
known being at 380 Ma (mid-Devonian of Gilboa).
Acari Most of the Acari (except of Opilioacarida and Holothyrida) are not so
important zoogeographically, many of them being parasitic, and the others are not
adequately studied and are not treated in details in this publication.
Opilioacarida Two fossil spp. have been described from Baltic amber (44 million
years ago, Eocene): Opilioacarus (?) aenigmus Dunlop, Sempf et Wunderlich,
2010, and Paracarus pristinus Dunlop, Wunderlich et Poinar, 2004.
Trombidiformes Among the other Arthropoda from Baltic amber by Koch and
Berendt (1854) have been recorded eight spp. of Erythraeidae (ascribed to
“Rhyncholophus,” now Erythraeus, Balaustium, and Leptus). Menge (1854)
described from Baltic amber the new genus and species Arythaena troguloides;
Vercammen-Grandjean (1972) found in Canadian amber (Cretaceous) another new
genus and species: Proterythraeus southcotti. Zacharda (in Zacharda and Krivolutsky
1985) described Mesoanystis taymirensis from Taymyr Peninsula, Upper Cretaceous
amber (the oldest fossil anystid mite known), and Palaeoerythracarus sachalinensis
from Sachalin (Paleogene amber). Bartel et al. (2015) described three new spp. of
Fessonia (Smarididae) from Baltic Amber, and Calyptostoma katyae Koniktyiewicz,
Wohltmann et Makol, 2016, was the first fossil Calyptostoma (described also from
Baltic amber). Dunlop (2007) described from the Early Cretaceous (Aptian) Crato
Formation from Ceará State, Brazil, a new large, fossil mite (Arachnida: Acari,
Erythraeoidea), Pararainbowia martilli.
Sarcoptiformes
Oribatida From many places have been recorded fossil oribatid mites, because of
the hard tegument and their massif presence (up to 75%) in the soil fauna (Krivolutsky
and Druck 1986).
Norton et al. (1988) recorded 14 fossil oribatid spp. (including the new genera
Protochthonius and Devonacarus and the new family Devonacaridae) from terres-
trial Devonian deposit near Gilboa, New York. An acarid belonging to the still-
living genus Hydrozetes Berlese, 1902, was found in the Lower Sinemurian
(Jurassic) in a core drilling from Sweden (Sivhed and Wallwork 1978).
Parasitiformes
Mesostigmata Inclusions assignable to the tortoise mites (Mesostigmata,

Uropodina) have been described by Dunlop et al. (2013) from Baltic amber (Eocene,
ca. 44–49 Ma). This is the oldest record of Uropodina. “Fossil mesostigmatid mites
(Acari: Parasitiformes: Mesostigmata) are extremely rare, and specimens from only
nine families, including four named species, have been described so far. A new
record of Myrmozercon sp. described here from Eocene (ca 44–49 Myr) Baltic
amber represents the first - and so far only - fossil example of the derived, extant
family Laelapidae”.
Ixodida In the amber from Cretaceous has been recorded a larval argasid tick
(Carios jerseyi Klompen et Grimaldi, 2001). Another Mesozoic tick (Amblyomma,
Ixodidae) has been described from Dominical amber (Lane and Poinar Jr. 2009), the
first in the New World. Ixodes tertiarius Scudder, 1885 was described from
Oligocene amber (Wyoming, USA). Another tick (Ixodes succineus Weidner, 1964)
is known from the Baltic amber.
We borrowed from the paper of Giribet, Sharma et al. (2012) this chronology:
Geological intervals:
–– 0–35 Ma (separation of Mediterranean plate from Western Europe, separation of
Japan from Eurasia, connection of Iberia to Eurasia)
–– 35–45 Ma (separation of Borneo and Indomalay Archipelago from Eurasia)
–– 45–60 Ma (Balkans connected to Western Europe, Iberia connected to
Mediterranean plate, Balkans, and Japan)
–– 60–75 Ma (Iberia separated from Mediterranean plate, Balkans, and Japan, North
America separated from Western Europe, emergence of Indomalay Archipelago)
–– 75–110 Ma (Mediterranean plate separated from North America, Iberia con-

nected to Western Laurasia, Balkans separated from North America and Western
Europe)
Mediterranean plate, and North America separated from Eastern Laurasia; emer-
gence of Borneo; Indomalay Archipelago nonexistent)
Mediterranean plate, and North America separated from Eastern Laurasia;
Borneo and Indomalay Archipelago nonexistent)
–– Dunlop et al. (2008), Dunlop and Martill (2002), Dunlop et al. (2004), Eskov and
Zonshtein (1990), Harvey (1990, 2003), Judson (2007), Kjellesvig-Waering
(1986), Krivolutsky and Krasilov (1977), Penney and Selden (2007), Poinar and
Brown (2004), Poinar and Santiago-Blay (1989), Rowland and Sissom (1980),
Selden (1993), Selden et al. (1991), Soleglad and Fet (2003), Tetlie and Dunlop
(2008)
5.3 P
assive Dispersal and Phoresy of Arachnida: Ballooning,
Rafting, and Continental Drift
5.3.1 Dispersal (Natural and by Man Activities)
Dispersal and Dispersion The difference between these two notions, sometimes
used indiscriminately, was explained by Platnick (1976): “Dispersion would be
defined as a property of individuals, the process by which an organism is able to
spread from its place of origin to another locality. Dispersal would be defined as a
property of taxa, the process by which a species or other group is able to spread
from its previously established range to a different range.”
Dispersal was and remains one of the major problems of zoogeography and a
rather controversial one. From the extreme fixists (stabilists) to the extreme mobil-
ists, one can observe the whole range of approach of the zoogeographer. Despite the
numerous achievements of earth science, the extreme fixists still believe that the
patterns of the land-sea distribution over the globe remain unchanged since the third
day of Creation. Their opponents, it is true, sometime deal freely with entire conti-
nents, displacing them to fit some particular distribution. The fixists like to refer to
the saying of L. Vialleton (1929, L’Origine des êtres vivants): “On a joué des dislo-
cations géologiques ou de l’existence de continents supposés au point de pouvoire
faire dire que certains biogéographes n’hesiteraient pas à inventer un continent pour
expliquer un genre de Coléoptère.” Many and many cases of distribution, however,
can be explained only by admitting direct land connections. To say that the
Galapagos and Aldabra tortoises or the moa of New Zealand have reached these
islands riding logs is, to put it mildly, not serious. The “stepping stones” could have
had significance with some groups, but the present author is firmly convinced that
5.3 Passive Dispersal and Phoresy of Arachnida: Ballooning, Rafting, and Continental… 73
even a gap of 10 meters of salt water is enough to prevent the dispersal of many
animal species and even groups.
The land invertebrates are often thought to disperse easier than the land verte-
brates. This is probably true (to some extend) for some groups of insects. Again,
how to explain the high endemism of insect fauna of Madagascar? There are many
groups not dispersing by floating logs. Yes, tropical rivers do carry sometimes logs
and even “floating islands” far into the ocean, but none of these “rafts” has brought
Uropygi (abundant in Southeast Asia) or many Papuan groups to Australia. Isopods
use to live in rotten logs, but nevertheless the fauna of Cuba is strictly neotropical
(the proximity of North America hasn’t affected it) (Vandel 1972a, b), and only two
widespread genera of woodlice are common to both Australia and Melanesia. As
Vandel (1973b) emphasizes: “Encore que l’Australie et la Nouvelle-Guinée soient,
au temps present, fort proche l’une de l’autre, leurs faunes isopodiques ne presen-
tent à peu près aucun point commun.” The remaining 20 genera and all the 53 spe-
cies of Melanesian Oniscidea have preferred to not undertake the risky rafting
across the 150 km-wide Torres Strait (the same is true for the numerous Australian
genera of Isopoda not represented in Melanesia). According to Szymkowiak et al.
(2007), “Despite similar geological history and close vicinity of both areas, the crab
spiders show great distinction between New Guinea and Australia. 36 genera on
both areas have been recorded, of which only 13 are common (Amyciaea, Cymbacha,
Diaea, Hedana, Loxoporetes, Mastira, Misumena, Porropis, Runcinia, Stephanopis,
Tharrhalea, Tmarus, and Xysticus), and 167 species, of which 6 are common. Thus,
similarity of the spider fauna of the family Thomisidae at genera level is near 37%,
while at species level it is 3.7%.”
Even if “to account for the endemic elements of the Hawaiian fauna would
require only about one successful establishment every 20,000 years or more”(Gressitt
1971a, b), it means that some of the groups of land animals do not disperse by air or
floating logs at all or meet with considerable difficulties in establishing themselves
on the new place. These “conservative” groups are of special interest for the zooge-
ographer, as they reflect the ancient state of the fauna of the particular area. On the
list of such “chosen” groups, the Arachnida stay very high, especially some of them.
A thorough review of the passive dispersal of Arachnida is due to Szymkowiak et al.
(2007).
In the nineteenth and twentieth centuries, voluntary or involuntary transport by
Man became a major factor in the dispersal of plants and mammals. Again the num-
ber of species of Arachnida, using ships or aircraft, is rather limited, and establish-
ment of Man-carried individuals is seldom successful. There are differences from
order to order.
As Wallwork (1979) points out, it is generally recognized as a principle of zoo-
geography that random dispersal by wind, water, birds, or Man results in dishar-
monic faunas.
Other sources: Berland (1934), Bristowe (1930), Cowrie and Holland (2006),
Darlington (1938), Duffey (1998), Gressit and Yoshimoto (1963), Schatz (1991),
Udvardy (1981)
Opiliones Studying the phalangids of the Antilles, Šilhavý (1973) indicates that
these animals are “from the phylogenetical point of view a very old order with small
ecological valence, conforming with difficulty to changing life conditions and being
very sensitive to the shortage of water and not very mobile. Phalangids cannot move
by “ballooning” as some spiders can and their possibility of migration among the
islands has been minimal even if we admit the possibility of their passive transfer by
hurricanes (either as living specimens or as eggs).” This opinion of the leading spe-
cialist in Opiliones confirms our view about the limited role of passive dispersal in
this group.
One very discussed case is the presence in New Caledonia of endemic, relic, and
very old (> 200 Ma) family of cave Opiliones of Cyphophthalmi (Troglosironidae,
13 spp.). They are close not to the nearest families (Pettalidae, Sironidae,
Stylocellidae) but to the far away Neogoveidae, living in the Americas. Sharma and
Giribet (2012) discussed the possible explanation of this relation. Cyphophthalmi
are poor dispensers, absent in the oceanic islands, and they certainly cannot over-
come marine water expanses.
Solifugae Among the 13 000 ship-trapped specimens of insects and arachnids,
Holzapfel and Harrell (1968) mention 275 specimens of spiders (Araneae), and
among them one specimen of “Solpugida” is probably a mistake. The easternmost
localities of Solifugae are in Vietnam (Dinorhax rostrumpsittaci Simon, fam.
Melanoblossiidae). They don’t live in apparently suitable Australia and are not
found east of Wallace’s Line. The “sun spiders” don’t seem to be very successful
candidates for “rafting,” let alone “ballooning.” Good example is their total absence
in Madagascar, despite the many species living in Southern Africa.
Pseudoscorpiones The tiny false scorpions, living under the bark or in the soil and
plant debris, should be transported easily by floating logs, by phoresy, or by human
activities. However, apart from the several cosmopolitic species and some store and
manure-inhabiting species, there is rather limited number of pseudoscorpions really
carried to faraway places. Even with the intense Europe-North America traffic, such
examples are rare (Muchmore 1969, 1972).
Pseudoscorpions sometime travel attached to birds or mammals (phoretic)
(Martens 1975). Seabirds have carried them to the subantarctic of very isolated
islands like St. Helena, Ascension, or Tristan-da-Kunha. If we compare two rela-
tively close and well-known faunae like those of Australia and New Zealand (58 and
25 local genera, respectively), we shall see that they have only ten genera and no
species in common.
Amblypygi Sometime people write amazing things. To quote Besch (1969), “The
Charontidae, an Indo-pacific group, are represented in South America only by the
genus Charinus, which except for its occurrence in the Galapagos Islands, is found
only in the islands of the Indian and Pacific Oceans. It is, therefore, considered to
have come from these islands to Galapagos Archipelago.” The author of this highly
5.3 Passive Dispersal and Phoresy of Arachnida: Ballooning, Rafting, and Continental… 75
improbable hypothesis evidently has overlooked the publication of Goodnight and

Goodnight (1946), describing Charinus Gertschi from Guiana and Suriname. The
subsequent founds of Charinus in Brazil (eight spp.), Venezuela, Peru, and many
other countries rather confirm the second alternative of the Goodnights that the
discovery of Charinus in Guiana “indicates our lack of knowledge of the neotropi-
cal arachnid fauna” (p.327). Actually, the genus Charinus Simon (by the way now
in the Charinidae family) is the most widespread genus from all Amblypygi, with 54
tiny species, limited mostly by the climate.
Thelyphonida (Uropygi) Another order missing in Australia, New Caledonia, and

New Zealand but occurring in South Asia, Papuan area, and the Philippines. The
finding of Mastigoproctus (American genus) with one specimen in Hong Kong is,
for us, a case of Man’s transport. However, there is another much bigger puzzle with
the description of the amblypygid Phrynus exsul Harvey, 2002, from a cave on
Flores (Indonesia), the first and only representative of the American family
Phrynidae and genus Phrynus. A relict like the Fiji iguanas?
There was a discussion about the Hypoctonus species found in West Africa (no
other Uropygi in or near Africa!) – whether a relict or a Man’s transport from East
Asia, where the other Hypoctonus live (Cooke and Shadab 1973). Finally, from the
species from West Africa was described a new genus Etienneus Heurtault, 1984
(another case of “fake disjunction”). The West African uropygids were declared
autochthonous and relict by Huff and Prendini 2009 (but why they didn’t spread
further in Africa?).
Schizomida Despite the statement of Besch (1969) that “they are distinguished by
the ease with which they are distributed into new areas by means of Man’s activi-
ties” (p. 730), very few of these tiny arachnids are actually being reported or sup-
posed to be transported by Man. Some are known from European greenhouses
(Zomus bagnallii Jackson, 1908 in Kew). Others have reached Hawaii or some other
Pacific islands (Marianas, Fiji, Marshall Islands, Samoa, New Caledonia). We agree
with Rowland (1972) that “the low agility and inability of Schizomida to disperse
through suboptimal areas may account for their geographical distribution. They lack
any secondary means of travel, such as phoresy, ballooning, resistant eggs, parasit-
ism or other means employed by arachnids” (p.153).
Scorpiones According to Vachon (1972) “Isometrus maculatus est, d’ailleurs, le

seul scorpion cosmopolite et son extension due, vraisemblablement, à l’homme”.
Similar observations made Berland (1932): “…cette espèce, frequentant les habita-
tions humaines et penetrant dans les navires, s’est repandue dans toute la région
chaude ou on la trouve parfois dans les îles les plus isolées, comme l’île de Paques,
par exemple.”
Araneae There are many articles describing the relatively active dispersal of spi-
ders, called “ballooning” (Braendegaard 1937; Brignoli 1983; Szymkowiak et al.
2007 and others). Brignoli (1983) says, “I would like to note that “dispersal” is a
rather poorly chosen term, not only because the distinction against “dispersion” is
possible only in English, but also because it is used for only one kind of change of
the range, more exactly only for the increase of it. But, as we all know, there are also
decreases of the ranges, which also involve movements of the species.”
Acari Wallwork (1979) insists that “It is very probably true that short-range over-
seas dispersal of Oribatids does occur, but the long-range dispersal over inhospita-
ble oceans which would be required to produce the global patterns that we can
observe today is unlikely.” For this author about 30% of the Oribatei Inferiores have
a cosmopolitan distribution, compared with only 15% of the Superiores. Oribatids
are an ancient group, and old genera have existed on the supercontinents and have
been carried away by the continent’s products of the breakup of Gondwana and
Laurasia.
5.4 Ecological Factors
5.4.1 Climatic and Other Barriers and Bridges
En somme, si nous laissons de coté l’Hologenèse qui est du domaine de la Génétique et la

Dérive des continents qui ne s’appuie sur rien de solide, il ne nous reste plus que les rela-
tions intercontinentales des géographies anciennes. Nous considerons comme la cause
essentielle des repartitions actuelles les variations du climat au cours des temps géologiques
et singulièrement pendant le Quaternaire. R. Furon (1953a, b)
The ecological factors (mostly temperature and humidity) of the territories are
determining the present-day distribution. In the past the distribution of some arach-
nids has been different but always reflecting the thermos and hygropreferendum of
the animals. Now we can outline several thermophilous groups with sharp limitation
in their distribution: Solifugae, Amblypygi, Uropygi, Schizomida, Ricinulei,
Holothyrida, and some lower-ranking groups like most Cyphophthalmi. Also the
scorpions have limitations – from the 111 spp. in the USA, only 1 species reaches
the extreme South of Canada.
Another temperature limitation is observed in the high mountains. The present-
day climatic zones of the Earth, so important for the distribution of animals, have
been classified by Russian-German climatologist W. Köppen first in 1884 and then
modified by himself, by Geiger, and other specialists (McKnight and Hess 2000).
Some isotherms clearly are limits of the distribution, mostly of the thermophilous
groups. The paleoclimates are also important to explain some distributions, mostly
the relictual (Elouard 1984).
Other factors are dryness/rainfall, length of the “moderate” seasons, etc. Some of
the limitations are not so evident. Why there are no Solifugae in Madagascar – only
419 km far from the richest area in the world what concerns the Solifugae? This
group has not reached Australia with its dry lands and deserts, maybe because of the
vast areas of rain forest and other unsuitable habitats for Solifugae. Why many
5.4 Ecological Factors 77
ancient orders are lacking from the ancient land New Zealand? The way of dispersal
and of reaching new territories by Arachnida are already known, but very few have
profited by “rafts” to reach Australia from New Guinea and vice versa.
There are many factors determining the ability of an arachnid to reach a new ter-
ritory, to survive on it and even to start further speciation. Important factors are also
time, glaciation, saturation of the new land with competitors, winds, etc. Some
groups are much more plastic and ubiquitous than others. Such are spiders, many
groups of mites, and to some extent Opiliones.
Interesting case of development in complete isolation are some caves like Movile
in Romania and Ayalon in Israel. In Movile have been found blind spiders (and
other arthropods) of families usually not entering caves (Clubionidae, Theridiidae,
Hahniidae). In Ayalon dead scorpions of a new and very strange family Akravidae
have been found, along with a new genus of pseudoscorpions (Ayalonia).
Time of speciation is also an important factor in the distribution of arachnids.
Some are like “living fossils.” Others, like the arachnids in the Hawaiian lava tubes,
have evolved in these tubes for a few thousand years in real troglobites.
The physical barriers are of different types – sea, high mountains, large rivers,
rain forest, deserts, and many others. They are not existing forever – the past distri-
bution of many groups proves that the paleogeography is determining quite different
paleozoogeography.
Except of some mites, Arachnida do not live in sea water and have little resis-
tance to it. All disjunctions, ubiquitous or pantropical species or genera remain from
the former situation of the territories or may be explained by dispersal via the three
W (wind, wings, waves) and more recently by the human transport (unlike some
other invasive animals, the arachnids are transported involuntary). In the deserts
prevail scorpions and solpugids, some other orders (spiders, mites) are barely repre-
sented, and some others (Ricinulei, Uropygi, Holothyrida) are completely missing.
In the high mountains, on the contrary, spiders are prevailing; there are many mites,
pseudoscorpions, and Opiliones; and the other orders are either missing or repre-
sented by low number of species (Beron 2008b).
The grassland is without Amblypygi and other arachnids needing rock shelters
and caves to hide, but spiders and mites are plentiful. The Holarctic broadleaf forest
and even more the taiga are already outside the termopreferendum of the “southern”
orders, and spiders, mites, pseudoscorpions, and Opiliones are masters of these
biomes. Even more this is true for the orophytic areas. Humidity is important for
most arachnids, and very few (scorpions, Solifugae, some mites) avoid areas with
high humidity.
5.4.2 Real and “Fake” Disjunctions
When you have a case of species of an American genus (and family!) found in a
cave of Flores (Indonesia), as in the case of Phrynus exsul (Harvey, 2002), you may
think of at least six options:
1. The species was wrongly identified. This applies usually for very old unchecked
identifications, which need reassessment.
2. It goes here for an animal, transported somehow (usually by Man) to a faraway
place.
3. To relict locality.
4. To lack of enough research in the area in between – the “gaps” might be in our
knowledge.
5. To the fact that this animal is different from the “congeners” in the main area and
should be given another name.
6. Mistake in the label. The case of the new Phrynus is really puzzling, as it has
been described by a very qualified researcher, found repeatedly; the distance is
so big that we cannot presume lack of collecting in between. Here are some cases
of real and “fake” disjunctions.
One of the classical groups to study disjunctions is the amphibians. Examples are
the family Proteidae (Proteus, Necturus), the Cryptobranchidae (one extant species
of Cryptobranchus in North America, two spp. of Andrias in China and Japan, and
one fossil in Europe), and the New Zealand frogs (four spp. of Leiopelma) with the
other two members of Leiopelmatidae of genus Ascaphus in North America. A good
example we were teaching in the University were the newts of the genus
Hydromantes, found in France, Italy, Sardinia, and California. Among the reptiles
long-standing example was the turtle “Podocnemis” madagascariensis in
Madagascar, considered member to a South American genus. However:
1. Some researchers created for the North American frogs a special family

Ascaphidae. Still, they are very close with Leiopelma within the suborder
Archaeobatrachia, and the problems of the origin of New Zealand frogs remain
open.
2. The status of the only European member of the American family Plethodontidae
(the lungless salamanders) is still a controversy. The three American species
remain in the genus Hydromantes; the seven species from southern France, Italy,
and Sardinia are given either generic or subgeneric status as Speleomantes and
Atylodes. Still, a problem remains how they occurred in so distant regions. In
2005 in South Korea was found a new lungless salamander (Karsenia Min et al.),
which indicates that the plethodontids were widespread once in the Holarctis.
So, we see that such sensations occur with Amphibia, let alone the
invertebrates!
3. The aberrant “Podocnemis” in Madagascar is no more a Podocnemis but is called
Erymnochelys madagascariensis. Together with other Neotropical elements, it
remains the only Madagascan member of the South American family
Podocnemididae.
4. The enigmatic West African whip scorpions (an order which is otherwise not
represented in or near Africa) have been longtime considered a zoogeographical
puzzle. In Senegal, Guinea-Bisau, Guinea, and Gambia, uropygids were
described as members of the Southeast Asian genus Hypoctonus (H. africanus
Henschel, 1899). At first it was presumed that they have been transported by
Man from Southeast Asia (Cooke and Shadab 1973), but finally they received a
separate generic status Etienneus (Heurtault, 1984) and were declared Gondwana
relicts by Huff and Prendini (2009).
Some mistakes are due to mislabeling or incorrect reading of old labels. One
Mediterranean scorpion has been published by Roewer from “Samoa,” but the
inquiry revealed the real locality – island Samos in Greece! In the materials pub-
lished from the collection (Roewer), there are several such “zoogeographical puz-
zles” (Helversen and Martens 1972).
5.4.3 Old World-New World Disjunctions
Amblypygi The genus Trichodamon Mello-Leitão, 1935 (Brazil, two spp.), is the
only member of the Old World family Phrynichidae in the Americas (tropical and
Southern Africa, Algeria, Morocco, Yemen, Saudi Arabia, Oman, Mauritius,
Seychelles, Zanzibar, Sao Tome, Madagascar, Sri Lanka, Brazil, 32 spp.). The find-
ing of an Old World species of Phrynus in Indonesia has been discussed above.
Thelyphonida (Uropygi) The genus Thelyphonellus Pocock, 1894 (Brazil,

Guyana, Venezuela) and Ravilops Viquez et Armas, 2005 (Dominican Republic),
are the only American representatives of the subfamily Hypoctoninae (Burma,
South China, Malaysia, West Africa, India, Sri Lanka). The Old World Mastigoproctus
transoceanicus Lazell, 2000 from Hong Kong is, for this author, a case of transport
by Man.
Scorpiones One of the most enigmatic disjunctions is the case of the eyeless
European scorpion Belisarius xambeui Simon, 1879 (France and Spain), placed in
Chactidae (Brotheinae) by Soleglad and Fet (2003) and in the new family
Troglotayosicidae (known from a cave in Ecuador) by Lourenço (1998), as base of
a new subfamily Belisariinae. In both cases is presumed a South American connec-
tion – a striking case of New World – Europe disjunction. Such relations are pre-
sumed in the recently described from Israel cave scorpion Akrav and in other groups
of animals (Isopoda terrestria).
Another classical case of disjunction is the occurrence of genus Heteronebo
Pocock on the islands Abd-El-Kuri (Yemen). Later (Francke 1978) some Heteronebo
have been found in the Caribbean. However, it was proved that the species described
by Pocock (Heteronebo granti and H. forbesii) do live on Abd-el-Kuri islands.
Lourenço (1998) tries to explain this peculiar distribution with the “generalized
tracks” of Croizat (1958). Similar explanation was proposed by Francke and
Soleglad (1981) for the disjunction Caraboctonidae – Iuridea.
Opiliones There are several interesting cases, analyzed (mostly in Japanese) by
Suzuki (1972, 1977). Interesting case is the family Caddidae (in the traditional
sense) – Caddella Hirst (South Africa), Hesperopilio Shear (Australia and Chile ),
Caddo Banks (Japan, North America, fossil in Europe), Acropsopilio Silvestri
(Japan, N. America, S. America, Australia, New Zealand), and Austropsopilio
Forster (Australia) (= Tasmanopilio Hickman, Tasmania). According to Groh and
Giribet (2015), Acropsopilionidae is no more a subfamily of Caddidae but a sepa-
rate family in Caddoidea and, together with Caddidae, is transferred in Dyspnoi.
Hesperopilio Shear is referred to Phalangioidea without indication of family. These
changes the genera enumerated are still remarkable as connection between South
America and Notogea. One genus (Jarmilana) of the Afrotropical family
Pyramidopidae has been described from a cave in Belize.
Americovibone Hunt et Cokendolpher is member of another interesting family of
Eupnoi – Neopilionidae. One species of this genus lives in Chile, and recently
another species has been described from New Zealand (Americovibone remota
Taylor, 2016).
The case with the genus Ceratomontia Roewer, 1915, with 22 spp., 18 known
from South Africa and 4 from South America (Argentina, Uruguay, S. Brazil), was
tested cladistically by Cruz Mendes and Kury (2008), who concluded that “The
consensus hypothesis did not recover a monophyletic Ceratomontia, instead, the
South African species constitute a clade with the Austromontia and Monomontia
species. The Ceratomontia species from South America form a clade that is sister-
group of the clade formed by the South African genera of the ingroup. The result
shows that Ceratomontia is involved in a “Gondwanan relationship” but also
includes other genera, suggesting that it is not a monophyletic group.”
Pseudoscorpiones Recently has been described from Spain and Portugal one
remarkable new genus of cave Pseudoscorpions of the South American and Antillean
family Bochicidae – Titanobochica Zaragoza et Reboleira, 2010 – and the genus
Troglobisium Beier, 1939, from Spain was transferred to the same family. Both were
declared relict (Reboleira et al. 2010).
Araneae There are many cases like the genus Caloctenus Keyserling – one species
in Ethiopia (Caloctenus abyssinicus Strand, 1917) and the other four in South
America. Do they really belong to one genus? Similar is the case with genus Vulsor
Simon – seven spp. in Madagascar and the Comoro Islands one (Vulsor occidentalis
Mello-Leitão, 1922) in Brazil.
Acari A good example is Trichosmaris papuana Beron, 2002 (Smarididae),

described by us from the center of New Guinea. The other members of the genus are
known from Guatemala and the USA.
5.4.4 West African-Indomalayan Disjunction
One of the classic cases of disjunctions is that between the animals of the tropical
rain forest of West and Central Africa and the wet forested areas of Southeast Asia,
separated by steppes, deserts, and dry mountains in between. Examples are apes,
peacocks, Tragulidae, Lorisidae, and many others. Are there such disjunctions
among Arachnida?
5.4.5 Strange (Old?) Disjunctions
This is found in many groups of Arachnida. Good example is the family Travuniidae,
found (mainly in caves) in South Europe, North America, and (Yuria Susuki, inc.
sedis) Japan. Another example from Opiliones – the family Sironidae (again: South
Europe, North America and the relict genus Suzukielus Juberthie in Japan!).
5.4.6 D
isjunctions in the Southern End of the World (New
Zealand, Australia, South Africa, Patagonia)
They are among the most interesting in the zoogeography and are explained by the
former continuity of these lands, the trans-Antarctic road and similar. Good exam-
ples are found among the Opiliones and the spiders (outlined further). Such disjunc-
tions have been detected also among other groups of animals and have provoked
some authors (Lopatin 1980) to speak about a greater Notogea, comprising also
Patagonia. Eskov and Golovatch (1986), analyzing the origin of the trans-Pacific
disjunctions, conclude that the “southern Pacific semicircle” have preserved a con-
siderable number of relicts extinct elsewhere…the fact is that patterns like South
America, New Zealand, Australia, as well as the other types of circum-Pacific dis-
junctions, are nothing but particular cases of the reduction of a formerly single
bipolar distribution. These authors insist also that “…the biogeographical signifi-
cance of recent relicts is limited unless supported by paleontology.”
5.4.7 Boreomontane and Arctic-Alpine Disjunctions
Holdhaus (1954) analyzed in details the distribution of some animals in the far
north of the Holarctic and, after a wide gap, in the mountains like Alps (boreo-
alpine or arctic-alpine distribution) or some other mountains. This type of distribu-
tion is usually attributed to the consequences of the ice age in Eurasia and North
America. Unlike Insecta, there are relatively few examples of boreomontane
disjunctions in groups like Arachnida, Myriapoda, or Isopoda (Buresch and Arndt

1926; Beron 1969a, 2008b). Among Bulgarian animals as such are considered the
mites Laelaps clethrionomydis and Hirstionyssus gudauricus and the spider
Meioneta gulosa.
A difference should be made between the notions “arctic-alpine” and “boreo-
montane” distribution. The animals with arctic-alpine distribution occur in the
Arctic and Subarctic tundra and high in the mountains of Middle or South Europe.
We speak about boreomontane distribution when animals inhabit the northern conif-
erous forests of Europe and North America and the relevant zones in the mountains
of Europe, Caucasus, and Central Asia. The term “boreo-alpine” is less accurate,
merging both above described and is better to avoid its use.
Araneae Many studies are due to Thaler (1976), Thaler and Knoflach (2002),
Relys (2000), and others for the spiders of the Alps.
Relys (2000) registered 13 arctic-alpine species in the Eastern Alps, the lowest
point being at 1618 spp. – Meioneta gulosa (L. Koch) – but most species have been
detected over 2100 m. At this altitude more than 10% of all spider specimens in a
community usually belong to arctic-alpine species. The 14 boreomontane species
were located at altitudes above 1900 m on the timberline ecotone. According to
Relys (2000), “despite high species richness neither boreomontane nor arctic-alpine
spiders, nor both groups together, can make up such a significant part of subalpine
spider communities as do alpine-endemic species (e.g., Pardosa oreophila Simon).”
According to Bosmans and De Keer (1987), 14 of the spider species of the
Pyrenees are arctic-alpine elements, 11 of them belonging to Linyphiidae, 1 to
Araneidae, 1 to Lycosidae, and 1 to Gnaphosidae.
5.4.8 Transpacific Disjunctions
Good examples are the Opiliones found in Japan and in North America. One (doubt-
ful) explanation is the Beringian bridge. This bridge is essential for the “Northern
Semicircle Eurasia – North America.” The “Southern Semicircle” (New Zealand,
Australia, Antarctica, Cape area, Patagonia) also provoqued many explanations,
Paleantarctic and others. Details with many examples (mostly insects) could be
found in the article of Eskov and Golovatch (1986). The conclusion in this article is
that the “southern Pacific semicircle” has preserved a considerable number of relicts
extinct elsewhere. And also “…the fact is that patterns like South America – New
Zealand – Australia, as well as the other types of circum-Pacific disjunctions, are
nothing but particular cases of the reduction of a formerly single bipolar
distribution.”
5.5 Endemics and Relicts in Arachnida 83
5.5 Endemics and Relicts in Arachnida
5.5.1 Endemism
Here the analyses are based on endemism to the level of family/subfamily and
genus/subgenus.
What is really endemism? Endemics are taxa which are known only from one
area (geographical unit or, more narrow, cave, or mountain). In many cases they are
considered endemic as long as they are not found elsewhere (especially the newly
described taxa).
For convenience, we use the name of countries to mark the endemic taxa. In use
are the terms “Bulgarian endemic,” or “Balkan endemic.” The political borders,
which are not natural and not permanent, do not make too much sense. If we take
our country, Bulgaria, we see that most Bulgarian mountains are on the borders with
the neighboring countries. An endemic species described from one part of the
mountain almost surely will be found in the other parts. Exceptions are very narrow
endemics like the cave animals (the Opiliones Paralola buresi from four caves in
Stara Planina and nowhere else).
So, the names of countries make sense in the case of islands (Malta, New
Guinea – the whole island, Madagascar, Mauritius, New Caledonia, New Zealand –
even there we find differences between the northern and the southern islands).
Islands (even big) are good examples for endemism (Sardinia, Tasmania, Mallorca).
We speak about endemics for Cuba and Jamaica but also about Antillean endemics.
To call “endemic” taxon living in “Russia” or “the USA” does not make sense – is
it Baltic shore or Sakhalin? Is it Florida or Oregon? Some countries like Brazil, DR
Congo, or Canada are very big but uniformed in nature. So, sometimes we have to
mark endemics only in a very broad sense (African, Indian, Australian). There is
also a notion of “subendemic” to indicate the main areal and some smaller penetra-
tions. We also have to distinguish between paleo- and neoendemics.
Other sources: Brignoli (1979), Gromov (1998), Komposch (2011), Lazzeroni
(1969), Lourenço (2007), Lourenço and Đình Sắc Pham (2010), Thaler (1976),
Prendini et al. (2006).
Some Endemic Genera and Families in Different Orders
Palpigradi
Allokoenenia Silvestri, 1913 – Guinea
Triadokoenenia Condé, 1991 – Madagascar
Solifugae
Galeodumus Roewer, 1960 – Afghanistan
Gluviema Caporiacco, 1937 – Somalia
Paragaleodiscus Birula, 1941 – Yemen
Roeweriscus Birula, 1937 – Iran

Zombis Simon, 1882 – Israel
Barrus Simon, 1880 – Egypt
Prosolpuga Roewer, 1934 – Namibia
Solpugista Roewer, 1934 – Namibia
Solpugisticella Turk, 1960 – Kenya
Solpuguna Roewer, 1933 – Namibia, South Africa, and Botswana
Toreus Purcell, 1903 – South Africa
Subfam. Melanoblossiinae – Southern Africa
Daesiella Hewitt – Namibia
Lawrencega Roewer, 1933 – Namibia, South Africa
Melanoblossia Purcell, 1903 – South Africa
Microblossia Roewer, 1941 – Namibia
Unguiblossia Roewer, 1941 – Namibia
Cordobulgida Mello-Leitão, 1938 – Argentina
Gaucha Mello-Leitão, 1924 – Chile
Gauchella Mello-Leitão, 1937 – Bolivia
Metacleobis Roewer, 1934 – Brazil
Mummucina Roewer, 1934 – Chile
Mummuciona Roewer, 1934 – Venezuela
Mummucipes Roewer, 1934 – Paraguay
Sedna Muma, 1971 – Chile
Uspallata Mello-Leitão, 1938 – Argentina, Chile
Acanthogylippus Birula, 1934 – Israel
Bdellophaga Wharton, 1981 – Namibia
Lipophaga Purcell, 1903 – Namibia, Southern Africa
Trichotoma Lawrence, 1968 – Namibia
Ammotrechesta Roewer, 1934 – Central America
Ammotrechinus Roewer, 1934 – Haiti, Jamaica
Ammotrechona Roewer, 1934 – Cuba
Antillotrecha Armas, 1994 – Dominican Republic, Leeward Islands, and Cuba
Campostrecha Mello-Leitão, 1937 – Ecuador
Dasycleobis Mello-Leitão, 1940 – Argentina
Neocleobis Roewer, 1934 – Galapagos
Scorpiones
Belisarius Simon, 1879 – French and Spanish Pyrenees
Alayotityus Armas, 1973 – Cuba
Ananteris Borelli, 1910 – Trinidad y Tobago
Tityopsis Armas, 1974 – Cuba
Rhopalurus Thorell, 1876 – Cuba and Hispaniola
Cazierius Francke, 1978 – Greater Antilles
Oiclus Simon, 1880 – Lesser Antilles
Didymocentrus Kraepelin, 1905 – Lesser Antilles and Central Cuba
Pseudochactas Gromov, 1998 – Uzbekistan, Tajikistan
Troglokhammouanus Lourenço, 2007 – Laos (cave)

Vietbocap Lourenço et Đình Sắc Pham, 2010 – Vietnam, Laos (caves)
Lanzatus Kovařik, 2001 – Iran
Saharobuthus Lourenço et Duhem, 2009 – Western Sahara
Somalibuthus Kovarik, 1998 – Somalia
Somalicharmus Kovarík, 1998 – Somalia
Amblypygi
Musicodamon Fage, 1929 – South Morocco, Algeria
Suborder Palaeoamblypygi – endemic suborder
Fam. Paracharontidae – endemic family
Paracharon Hansen, 1921 – Guinea-Bissau
Thelyphonida (Uropygi)
Glyptogluteus Rowland, 1973 – Philippines (Panay)
Chajnus Speijer, 1936 – Sunda Islands
Uroproctus Pocock, 1894 – India (Assam)
Mimoscorpius Pocock, 1894 – Philippines
Amauromastigon Mello-Leitão, 1931 – Brazil
Ravilops Viquez et Armas, 2005 – Dominican Republic
Etienneus Heurtault, 1984 – West Africa
Schizomida
Subfamily Megaschizominae
Megaschizomus Lawrence, 1969 – Mozambique, Republic of South Africa
Afrozomus Reddell et Cokendolpher, 1995 – Angola, DR Congo
Adisomus Cokendolpher et Reddell, 2000 – Brazil
Anepsiozomus Harvey, 2001 – Seychelles
Artacarus Cook, 1898 – Liberia, Ivory Coast
Attenuizomus Harvey, 2000 – North Australia
Brignolizomus Harvey, 2000 – Australia
Cubazomus Reddell et Cokendolpher, 1995 – Cuba
Draculoides Harvey, 1992 – Australia
Javazomus Reddell et Cokendolpher, 1995 – Java
Julattenius Harvey, 1992 – Australia (Queensland)
Kenyazomus Armas, 2014 – Kenya
Luisarmasius Reddell et Cokendolpher, 1995 – Cuba, Pinos, Puerto Rico
Mahezomus Harvey, 2001 – Seychelles
Mayazomus Reddell et Cokendolpher, 1995 – Mexico (Tabasco, Chiapas)
Neozomus Reddell et Cokendolpher, 1995 – India (Maharashtra)
Notozomus Harvey, 1992 – Australia (Queensland)
Oculozomus Reddell et Cokendolpher, 1995 – Sumatra
Ovozomus similis (Hirst, 1913) – Seychelles
Pacal Reddell et Cokendolpher, 1995 – Mexico
Schizomus Cook, 1899 – Sri Lanka
Secozomus latipes (Hansen, 1905) – Seychelles
Sotanostenochrus Reddell et Cokendolpher, 1991 – Mexico

Stewardpeckius Reddell et Cokendolpher, 1995 – Jamaica
Tayos Reddell et Cokendolpher, 1995 – Ecuador
Trithyreus Kraepelin, 1899 – Burma (= Myanmar)
Pseudoscorpiones
Neobalkanella Ćurčić, 2013 – Serbia
Chiliochthonius Vitali-di-Castri, 1975 – Chile
Congochthonius Beier, 1959 – DR Congo
Drepanochthonius Beier, 1964 – Chile
Francochthonius Vitali-di-Cstri, 1975 – Chile
Malcolmochthonius Benedict, 1978 – USA (California, Oregon)
Microchthonius Hadži, 1933 – Croatia
Neochthonius Chamberlin, 1929 – USA (California)
Selachochthonius Chamberlin, 1929 – South Africa, Lesotho
Arcanobisium Zaragoza, 2010 – Spain
Troglochthonius Beier, 1939 – Croatia, ? Sardinia
Anisoditha J.C. Chamberlin et R.V. Chamberlin, 1945 – Paraguay
Cryptoditha J.C. Chamberlin et R.V. Chamberlin, 1945 – Brazil
Haploditha Caporiacco, 1951 – Venezuela
Neoditha Feio, 1945 – Brazil
Sororoditha J.C. Chamberlin et R.V. Chamberlin, 1945 – Brazil
Verrucaditha J.C. Chamberlin, 1929 – USA
Afroroncus Mahnert, 1981 – Kenya
Ideoroncus Balzan, 1887 – Brazil, Paraguay
Nannoroncus Beier, 1955 – Kenya, Uganda
Shravana Chamberlin, 1930 – Thailand
Indohya Beier, 1974 – India
Pseudogarypus Ellingsen, 1909 – Canada, USA
Neopseudogarypus Morris, 1948 – Tasmania
Antillobisium Dumitresco et Orghidan, 1977 – Cuba
Apohya Muchmore, 1973 – Mexico
Bochica J.C. Chamberlin, 1930 – Grenada, Trinidad
Leucohya J.C. Chamberlin, 1946 – USA (Texas), Mexico
Paravachonium Beier, 1956 – Mexico
Troglobochica Muchmore, 1984 – Jamaica
Titanobochica Zaragoza et Reboleira, 2010 – Portugal
Troglohya Beier, 1956 – Mexico
Vachonium J.C. Chamberlin, 1947 – Mexico, Belize
Arcanobisium Zaragoza, 2010 – Spain
Titanobochica Zaragoza et Reboleira, 2010 – Portugal
Beierobisium Vitali-di-Castri, 1970 – Falkland Islands
Gymnobisium Beier, 1931 – South Africa, Lesotho
Vachonobisium Vitali-di-Castri, 1963 – Chile
Ammogarypus Beier, 1962 – Namibia, South Africa
Eremogarypus Beier, 1955 – Namibia

Meiogarypus Beier, 1955 – Namibia
Neogarypus Vachon, 1937 – Zimbabwe
Paragarypus Vachon, 1937 – Madagascar
Paramenthus Beier, 1963 – Israel
Hemisolinus Beier, 1977 – St. Helena
Alabamocreagris Ćurčić, 1984 – USA (Alabama)
Americocreagris Ćurčić, 1982 – USA (Oregon, Washington)
Chinacreagris Ćurčić, 1983 – China
Fissilicreagris Ćurčić, 1984 – USA (California)
Globocreagris Ćurčić, 1984 – USA (California)
Insulocreagris Ćurčić, 1987 – Croatia (Vis), Herzegovina
Lissocreagris Ćurčić, 1981 – USA
Minicreagris Ćurčić, 1981 – USA
Nepalobisium Beier, 1974 – Nepal
Novobisium Muchmore, 1967 – USA
Occitanobisium Heurtault, 1977 – France
Pedalocreagris Ćurčić, 1985 – Far East of Russia
Paedobisium Beier, 1939 – Romania, Spain
Pararoncus J.C. Chamberlin, 1938 – Japan
Protoneobisium Ćurčić, 1988 – Croatia
Roncobisium Vachon, 1967 – France
Saetigerocreagris Ćurčić, 1984 – USA
Simonobisium Heurtault, 1974 – France, Italy
Tartarocreagris Ćurčić, 1984 – USA (Texas)
Trisetobisium Ćurčić, 1982 – USA
Tuberocreagris Ćurčić, 1978 – USA
Afrowithius J.C. Chamberlin, 1931 – South Africa
Aisthetowithius Beier, 1967 – Kenya, Tanzania
Cryptowithius Beier, 1967 – Kenya
Cyrtowithius Beier, 1955 – Namibia, South Africa
Hyperwithius Beier, 1951 – Vietnam
Paragoniochernes Beier, 1932 – South Africa
Parallowithius Beier, 1955 – Namibia, South Africa
Plesiowithius Vachon, 1954 – Mauretania
Pogonowithius Beier, 1979 – DR Congo
Protowithius Beier, 1955 – Juan Fernandez Islands
Pseudatemnus Beier, 1947 – South Africa
Scotowithius Beier, 1977 – St. Helena
Sphallowithius Beier, 1977 – St. Helena
Thaumatowithius Beier, 1940 – Reunion, Mauritius
Trichotowithius Beier, 1944 – Ethiopia, Kenya
Tropidowithius Beier, 1955 – Peru
Amaurochelifer Beier, 1951 – Vietnam
Ancistrochelifer Beier, 1951 – Laos, Cambodia, Vietnam

Aperittochelifer Beier, 1955 – South Africa
Aporochelifer Beier, 1953 – Indonesia (Flores)
Aspurochelifer Benedict et Malcolm, 1979 – USA
Canarichelifer Beier, 1965 – Canary Islands, Ilhas Selvagens
Chamberlinarius Heurtault, 1983 – Ivory Coast
Cubachelifer Hoff, 1946 – Cuba, Dominican Republic
Ectoceras Stecker, 1875 – India
Florichelifer Hoff, 1964 – USA (Florida)
Haplochelifer J.C. Chamberlin, 1932 – USA
Hygrochelifer Murthy et Ananthakrishnan, 1977 – India
Idiochelifer J.C. Chamberlin, 1932 – USA
Juxtachelifer Hoff, 1956 – USA (New Mexico)
Kashimachelifer Morikawa, 1957 – Japan
Levichelifer Hoff, 1946 – Mexico, USA
Litochelifer Beier, 1948 – South Africa
Lophodactylus J.C. Chamberlin, 1932 – Brazil
Macrochelifer Vachon, 1940 – China
Mexichelifer Muchmore, 1973 – Mexico
Nannocheliferoides Beier, 1974 – India
Pachychelifer Beier, 1962 – Georgia
Paisochelifer Hoff, 1946 – USA
Papuchelifer Beier, 1965 – Indonesia (Papua), Papua New Guinea
Phorochelifer Hoff, 1956 – USA
Pilochelifer Beier, 1935 – Mauritius, Reunion
Pseudorhacochelifer Beier, 1976 – Madeira, Canary Islands
Pugnochelifer Hoff, 1964 – USA
Rhopalochelifer Beier, 1964 – South Africa
Sinochelifer Beier, 1967 – China
Stenochelifer Beier, 1967 – India
Telechelifer J.C. Chamberlin, 1949 – ? Sri Lanka
Tetrachelifer Beier, 1967 – Vietnam
Xenochelifer J.C. Chamberlin, 1949 – USA (California)
Scotowithius Beier, 1977 – St. Helena
Sphallowithius Beier, 1977 – St. Helena
Athleticatemnus Beier, 1979 – DR Congo
Brazilatemnus Muchmore, 1975 – Brazil
Caecatemnus Mahnert, 1985 – Brazil
Mesatemnus Beier et Turk, 1952 – Cyprus
Synatemnus Beier, 1944 – Tanzania
Acanthicochernes Beier, 1964 – Papua New Guinea, Solomon Islands
Acuminochernes Hoff, 1949 – USA
Adelphochernes Beier, 1937 – Philippines (Mindanao, Mindoro)
Ancalochernes Beier, 1932 – Mexico
Atherochernes Beier, 1954 – Venezuela
Cacoxylus Beier, 1965 – Solomon Islands (Guadalcanal), Papua New Guinea

Calidiochernes Beier, 1954 – Venezuela
Calymmachernes Beier, 1954 – Australia (W. Australia)
Caribochernes Beier, 1976 – Dominican Republic
Chiridiochernes Muchmore, 1972 – Indonesia (Sulawesi)
Chrysochernes Hoff, 1956 – USA (New Mexico)
Cocinachernes Hentschel et Muchmore, 1989 – Mexico
Conicochernes Beier, 1948 – Australia
Coprochernes Beier, 1976 – Costa Rica
Corosoma Karsch, 1879 – Brazil
Cyclochernes Beier, 1970 – Solomon Islands (Guadalcanal)
Dasychernes J.C. Chamberlin, 1929 – Colombia, Panama
Diplothrixochernes Beier, 1962 – Argentina
Epichernes Muchmore, 1982 – Mexico
Eumecochernes Beier, 1932 – Hawaii
Gelachernes Beier, 1940 – Solomon Islands, Papua New Guinea
Gobichernes Krumpál et Kiefer, 1982 – Mongolia
Hexachernes Beier, 1953 – Guatemala
Marachernes Harvey, 1992 – Australia
Petterchernes Heurtault, 1986 – Brazil
Selachochernes Mahnert, 2011 – Juan Fernandez Islands
Illinichernes Hoff, 1949 – USA
Maorichthonius Chamberlin, 1925 – New Zealand
Sathrochthoniella Beier, 1967 – New Zealand
Tyrannochthoniella Beier, 1966 – New Zealand
Heterochernes Beier, 1966 – New Zealand
Hemisolinus Beier, 1977 – Saint Helena
Antillobisium Dumitresco et Orghidan, 1977 – Cuba
Caribochernes Beier, 1976 – Dominican Republic
Cubachelifer Hoff, 1946 – Cuba, Dominican Republic
Mexichthonius Muchmore, 1975 – Mexico (Campeche)
Apohya Muchmore, 1973 – Mexico
Vachonium J.C. Chamberlin, 1947 – Mexico, Belize
Mexichelifer Muchmore, 1973 – Mexico
Ancalochernes Beier, 1932 – Mexico
Epichernes Muchmore, 1982 – Mexico
Papuchelifer Beier, 1965 – Indonesia (Papua), Papua New Guinea
Selachochthonius Chamberlin, 1929 – South Africa, Lesotho
Gymnobisium Beier, 1931 – South Africa, Lesotho
Afrowithius J.C. Chamberlin, 1931 – South Africa
Ammogarypus Beier, 1962 – Namibia, South Africa
Eremogarypus Beier, 1955 – Namibia
Meiogarypus Beier, 1955 – Namibia
Cyrtowithius Beier, 1955 – Namibia, South Africa
Paragoniochernes Beier, 1932 – South Africa
Parallowithius Beier, 1955 – Namibia, South Africa

Pseudatemnus Beier, 1947 – South Africa
Aperittochelifer Beier, 1955 – South Africa
Litochelifer Beier, 1948 – South Africa
Rhopalochelifer Beier, 1964 – South Africa
Cryptoditha J.C. Chamberlin et R.V. Chamberlin, 1945 – Brazil
Haploditha Caporiacco, 1951 – Venezuela
Neoditha Feio, 1945 – Brazil
Sororoditha J.C. Chamberlin et R.V. Chamberlin, 1945 – Brazil
Ideoroncus Balzan, 1887 – Brazil, Paraguay
Tropidowithius Beier, 1955 – Peru
Lophodactylus J.C. Chamberlin, 1932 – Brazil
Brazilatemnus Muchmore, 1975 – Brazil
Caecatemnus Mahnert, 1985 – Brazil
Calidiochernes Beier, 1954 – Venezuela
Corosoma Karsch, 1879 – Brazil
Dasychernes J.C. Chamberlin, 1929 – Colombia, Panama
Aporochelifer Beier, 1953 – Indonesia (Flores)
Chiridiochernes Muchmore, 1972 – Indonesia (Sulawesi)
Ectoceras Stecker, 1875 – India
Hygrochelifer Murthy et Ananthakrishnan, 1977 – India
Paramenthus Beier, 1963 – Israel
Athleticatemnus Beier, 1979 – DR Congo
Chiliochthonius Vitali-di-Castri, 1975 – Chile
Drepanochthonius Beier, 1964 – Chile
Francochthonius Vitali-di-Castri, 1975 – Chile

Vachonobisium Vitali-di-Castri, 1963 – Chile
Diplothrixochernes Beier, 1962 – Argentina
Australochelifer Beier, 1975 – Australia
Conicochernes Beier, 1948 – Australia
Pararoncus J.C. Chamberlin, 1938 – Japan
Kashimachelifer Morikawa, 1957 – Japan
Opiliones
Fam. Troglosironidae – New Caledonia
Troglosiro Juberthie, 1979 – New Caledonia
Pettalus Thorell, 1878 – Sri Lanka
Rakaia Hirst, 1925 – New Zealand
Aoraki Boyer et Giribet, 2007 – New Zealand
Neopurcellia Forster, 1948 – New Zealand – South Island
Purcellia Hansen et Soerensen, 1904 – Western South Africa
Parapurcellia Rosas Costa, 1950 – Eastern South Africa
Austropurcellia Shear, 1980 – Australia (Queensland)
Manangotria Shear et Gruber, 1996 – Madagascar
Karripurcellia Giribet, 2003 – Western Australia
Speleosiro Lawrence, 1931 – South Africa
Chileogovea Roewer, 1961 – Chile
Fam. Ogoveidae – equatorial Africa
Ogovea Hansen et Soerensen, 1914 – Bioko, Cameroon, Congo
Huitaka Shear, 1979 – Colombia
Shearogovea Giribet, 2011 – Mexico
Meghalaya Giribet, Sharma et Bastawade, 2007
Fangensis Rambla, 1994 – Thailand
Metasiro Juberthie, 1960 – Florida, Georgia, S. Carolina
Odontosiro Juberthie, 1961 – Portugal
Iberosiro de Bivort, 2004 – Portugal
Paramiopsalis Juberthie, 1962 – Portugal, Spain
Suzukielus Juberthie, 1970 – Japan
Marwe Shear, 1985 – Kenya
¿Tranteeva Kratochvil, 1958 – Bulgaria
? Ankaratra Shear et Gruber, 1996 – Madagascar (one sp., inc. sedis)
Caddella Hirst, 1925 – South Africa
Hesperopilio Shear, 1996 – Western Australia
Austropsopilio Forster, 1955 – Australia (New South Wales)
Tasmanopilio Hickman, 1957 – Tasmania
Yuria Suzuki, 1964 – Japan
Buemarinoa Roewer, 1956 – Sardinia
Fam. Synthetonychiidae – New Zealand (14 spp.)
Synthetonychia Forster, 1954 – New Zealand (14 spp.)
Fam. Escadabiidae – Brazil
Biantoncopus Martens et Schwendinger, 1998 – Philippines (Leyte)

Caenoncopus Martens et Schwendinger, 1998 – Sumatra
Palaeoncopus Martens et Schwendinger, 1998 – Sumatra
Konfiniotis Roewer, 1940 – Crete
Acuclavella Shear, 1986 – Idaho, etc.
Ceratolasma Goodnight et Goodnight, 1942 – Oregon
Crosbycus Roewer, 1914 – British Columbia
Ausobskya Martens, 1972 – Greece, southern Bulgaria
Banksula Roewer, 1949 – California (10 spp.)
Bishopella Roewer, 1927 – southeastern USA
Calicina Ubick et Briggs, 1989 – California
Crosbyella Roewer, 1927 – southeastern USA
Haasus Roewer, 1949 – Israel
Lola Kratochvil, 1937 – Dalmatia
Maiorerus Rambla, 1993 – Canary Islands
Microcina Briggs et Ubick, 1989 – California
Paralola Kratochvil, 1958 – Bulgaria
Proscotolemon Roewer, 1916 – Japan
Sitalcina Banks, 1911 – California
Tolus Goodnight et Goodnight, 1942 – Tennessee
Trojanella Karaman, 2005 – Serbia
Undulus Goodnight et Goodnight, 1942 – Alabama
Wespus Goodnight et Goodnight, 1942 – Arkansas
Aburista Roewer, 1935 – Ghana
Sokodea Roewer, 1935 – Togo
Typhloburista Lawrence, 1947 – Ivory Coast
Afroassamia Caporiacco, 1940 – Ethiopia
Assamiella marginata (Roewer, 1912) – Burma
Gudalura Roewer, 1927 – India
Micrassamula Martens, 1977 – Nepal
Neassamia Roewer, 1935 – Thailand
Nepalsia Martens, 1977 – Nepal
Nepalsioides Martens, 1977 – Nepal
Pechota Roewer, 1935 – Malacca
Popassamia Roewer, 1940 – Burma
Puria Roewer, 1923 – India (Maharashtra)
Tavoybia quadrispina Roewer, 1935 – Malacca
Umtaliella Lawrence, 1934 – Zimbabwe
Irumua Roewer, 1961 – DR Congo
Machadoessa Lawrence, 1951 – Angola
Mutadia Kauri, 1985 – DR Congo
Numipedia Kauri, 1985 – DR Congo
Typhlobunus Roewer, 1915 – Kenya
Araneae
Atrophothele Pocock, 1903 – Socotra
Aurecocrypta Raven, 1994 – Australia
Barycheloides Raven, 1994 – New Caledonia
Barychelus Simon, 1889 – New Caledonia
Cyrtogrammomma Pocock, 1895 – Guyana
Encyocrypta Simon, 1889 – New Caledonia
Fijocrypta Raven, 1994 – Fiji
Idiommata Ausserer, 1871 – Australia
Moruga Raven, 1994 – Australia
Natgeogia Raven, 1994 – New Caledonia
Nihoa Raven et Churchill, 1992 – Oceania
Orstom Raven, 1994 – New Caledonia
Ozicrypta Raven, 1994 – Australia
Plagiobothrus Karsch, 1891 – Sri Lanka
Questocrypta Raven, 1994 – New Caledonia
Seqocrypta Raven, 1994 – Australia
Synothele Simon, 1908 – Australia
Trittame L. Koch, 1874 – Australia (Queensland)
Tungari Raven, 1994 – Australia (Queensland)
Zophorame Raven, 1990 – Australia (Queensland)
Zophoryctes Simon, 1902 – Madagascar
Neodiplothele Mello-Leitão, 1917 – Brazil
Paracenobiopelma Feio, 1952 – Brazil
Ammonius Thorell, 1899 – Cameroon
Eubrachycercus Pocock, 1897 – Somalia
Reichlingia Rudloff, 2001 – Belize
Troglothele Fage, 1929 – Cuba
Fam. Halidae Jocqué, 1994 – Madagascar
Hala Jocqué, 1994 – Madagascar
Tolma Jocqué, 1994 – Madagascar
Teloleptoneta Ribera, 1988 – Portugal
Masirana Kishida, 1942 – Japan
Falcileptoneta Komatsu, 1970 – Japan
Rhyssoleptoneta Tong et Li, 2007 – China
Opilioacarida
Adenacarus Van der Hammen, 1966 – Yemen
Amazonacarus Vasquez, Araújo et Feres, 2014 – Brazil
Brasilacarus Vasquez, Araújo et Feres, 2014 – Brazil
Indiacarus Das et Bastawade, 2007 – India
Phalangiacarus Coineau et Van der Hammen, 1979 – Gabon
Siamacarus Leclerc, 1989 – Thailand
Vanderhammenacarus Leclerc, 1989 – Thailand
Parasitiformes
Holothyrida
Australothyrus van der Hammen, 1972 – Australia
Dicrognathus Gerlach, Lehtinen et Mandl, 2010 – Seychelles
Hammenius Lehtinen, 1981 – New Guinea
Haplothyrus Lehtinen, 1995 – New Caledonia
Holothyrus Gervais, 1842 – Mauritius
Indothyrus Lehtinen, 1995 – Sri Lanka
Leiothyrus van der Hammen, 1983 – New Guinea
Michaelothyrus Gerlach, Lehtinen et Mandl, 2010 – Seychelles
Sternothyrus Lehtinen, 1995 – Seychelles
Thonius Lehtinen, 1981 – New Guinea
Caribothyrus Kontschán et Mahunka, 2004 – Dominican Republic
Diplothyrus Lehtinen, 1999 – Brazil, French Guyana
Ixodida
Fam. Nuttalliellidae – Southern Africa
Nuttalliella Bedford, 1931 – Southern Africa – from Tanzania to Namibia
5.5.2 Relicts and Relictness
Are There Relicts at All? The Issue of Relicts
Types of Relicts The notion of “relict” is widely used by biogeographers, despite

the warning of some of them to their colleagues to not allow themselves to be car-
ried away too much (Elenevskiy and Radigina 2002).
This notion was used widely by biospeleologists, influenced by the book of
Jeannel “Les fossiles vivants des cavernes” (1943). Some researchers strongly
oppose this very notion, as does the prominent Italian Arachnologist Brignoli
(1979): “Le terme de “relicte” (ou même de “fossile vivant”) si souvent employé
pour les troglobies, n’a pour moi aucun sense.”
Checking in the dictionaries, we can find several interpretations of “relict.”
Merriam-Webster Dictionary: “A surviving species of an otherwise extinct group
of organisms; also: a remnant of a formerly widespread species that persists in an
isolated area.”
The Free Dictionary: “An organism or species of an earlier time surviving in an
environment that has undergone considerable change.”
Botanists have three concepts to classify relicts: geographic, taxonomic, and lin-
eage relicts.
Birstein (1947), confirmed by Vandel (1964): “Les rélicts comme des types ani-
maux (ou végétaux) dont l’évolution est arrêtée ou du moins fort ralentie, et qui ont
conservé le facies de leurs lointains ancêtres.”
In the recent paper of Grandcolas et al. (2014), relicts (geographical or philoge-

netic) are identified as “a species or a group of species remaining from a large group
that is mainly extinct.”
We shall not discuss here this problem on the scale of all animals and plants. For
us the statement of Brignoli is far too strong and cannot be supported. Of course,
relicts and even “living fossils” do exist – classical examples are the two fish species
of Latimeria, the two species of the reptile tuatara (Sphenodon), the okapi, and
many, many others. Worth mentioning are the marine relicts in the cave waters (the
Sphaeromides and the other Cirolanidae). High in the mountains of Papua New
Guinea, far from any sea, we found in the caves polychaete worms (Namanereis),
Isopoda Anthuridea (Cyathura), and snails of marine origin at almost 3000 m
(Selmistomia). There is no doubt that they are relicts. What, if not relict, is the
recently discovered in Laos rodent Laonastes, the only living member of a family
Diatomyidae, extinct 11 million years ago? Or the only member of the family
Microbiotheriidae, also known (at latest) from the Miocene – the small marsupial
from Chile Dromiciops gliroides (“monito del monte”)?
Here we shall limit our discussion using examples among Arachnida.
Amblypygi The species Paracharon caecus Hansen, 1921, in the family
Paracharontidae and even the suborder Palaeoamblypygi from Guinea-Bissau is
declared “living fossil” by various authors.
Scorpiones Among scorpions one of the most striking examples or relictness is the
recently described family Pseudochactidae Gromov, 1998. It contains for the
moment three genera, all considered relicts: Pseudochactas Gromov in Central
Asia, Troglokhammouanus Lourenço (cave in Laos), and Vietbocap Lourenço et
Đình Sắc Pham (caves in Vietnam and Laos) (Lourenço and Pham 2012, 2014). It is
a very primitive family (Prendini et al. 2006).
A relict scorpion of Buthidae family (new genus) was described from the Great
Rift Valley in Kenya (Riftobuthus inexpectatus Lourenço, Duhem et Cloudsley
-Thompson, 2010).
Another case of relict scorpion is Euscorpius oglasae Caporiacco, 1950, on the
island of Montecristo. This is a case of paleoendemism, according to Vignoli et al.
(2007).
Pseudoscorpiones Among the pseudoscorpions, especially the troglobites, there
are many which could qualify for the category of relics.
Many detailed studies on the pseudoscorpions, termed as relicts, are due to B.
Ćurčić and his team (Ćurčić, 1983, 1985, Ćurčić and Dimitriević 1984 and others).
Schawaller (1987) recorded a second relict locality of the northern species
Syarinus strandi (Ellingsen) in Central Europe (South Germany), after the finding
in Austria of this species, known from Norway (Mahnert 1976).
There are also other examples of relicts in European Syarinidae. One of the most
striking is Hadoblothrus aegaeus Beron, 1985, found by us in caves on Santorin and
Iraklia islands.
For us there is no doubt that the family Syarinidae contains some striking cases
of relictness. According to Beier (1963), “Syarinidae: Wohl bei keiner anderen
Familie ist der Reliktcharakter so ausgeprägt.”
Gardini (1982): “Hadoblothrus, monotipico, è di incerta affinità e representa uno
dei più singolari relitti della regione pugliese.”
It is curious that not a single representative of the family Syarinidae has been
found so far in the numerous (and partly well investigated) caves on the mainland of
Balkan Peninsula. Hadoblothrus aegaeus has been found in the islands from where
not a single cave Neobisiidae has been recorded. The closest Syarinidae to the
Balkan Peninsula are Pseudoblothrus Beier (one sp. in France, two species in
Crimea, two in Italy, and two in Switzerland, all living in caves, except for P. regalini
Inzeghi, 1983), Hadoblothrus gigas from South Italy (two caves), and Syarinus
strandi (Austria, Germany, Finland, and Norway; all other five Syarinus species
living in North America, outside caves).
Another striking example of relict distribution are two genera of the South
American family Bochicidae in caves of Spain (Troglobisium Beier, 1939) and
south Portugal (Titanobochica Zaragoza et Reboleira, 2010).
Opiliones Roewer (1935) declared the few Laniatores in Europe being all relicts.
Martens (1972), describing the first non-troglobite Laniatores on Balkan Peninsula
(Ausobskya athos from Greece), came to the conclusion that “The European
Laniatores should no longer be regarded as tertiary relics, for they are nowadays
widely distributed inhabitants of soil litter in areas not covered by ice during the
glacious periods.” At least, for genera like Lola in Dalmatia, Paralola in Bulgaria,
and Trojanella in Serbia (also in Stara Planina, both very restricted monotypic tro-
globites), for us, there is no doubt that they represent fauna and environment of the
past. Concerning the relicts in the other countries, we must mention the North
American cave harvestmen (Briggs 1971, 1974) and the whole family Troglosironidae
in New Caledonia (Giribet et al. 2012).
In the fam. Pyramidopidae, Meiorerus randoi Rambla from a cave in Fuerteventura
and Conomma orientale (Roewer) from Moshi (Tanzania) are considered relictual
(Sharma et al. 2011).
Thelyphonida (Uropygi) Huff and Prendini (2009) consider the only African
genus Etienneus Heurtault in West Africa as Gondwanan relict.
Araneae Interesting is the case of the relict family Liphistiidae, sole member of
the suborder Mesothelae. This family is found in Japan, China, and Southeast Asia.
According to Xu X et al. (2015), these spiders are “living fossils,” and the suborder
Mesothelae is an ancient clade sister of all modern spiders. Again according these
authors, Liphistiidae genera originate in Asia in the Paleogene (4–24 Ma). This tim-
ing is relatively recent, taking into account the old age of the spider divergence
(297.6 Ma) between the Mesothelae and the Opisthothelae (Mygalomorphae and
Araneomorphae).
5.6 The Northern Limits of Arachnida 97
Acari Oribatids are a very ancient group of arachnids and genera present in the
world fauna today have existed unchanged for long periods of geological time. It is
postulated that many of these genera predate the breakup of the supercontinent
Pangaea and the drifting apart of continental landmasses (Wallwork 1979). The dis-
tribution of some genera on the southern landmasses reflects a former continuous
pattern which extended across Gondwanaland. Among the cosmopolitan genera
prevail the “primitive” ones (Inferiores), compared with the derived ones
(Superiores). Wallwork (1979) emphasizes on the zoogeographic principle that ran-
dom dispersal by wind, water, birds, or Man results in dysharmonic faunas. In con-
trast, harmonic faunas are those which present the same proportion of specific or
generic elements in two or more geographically separated areas. Such harmony
implies a previous generic continuity between the faunas in question and, by
inference, previous land connections between the now-separated areas in which
they live. Different groups of Oribatida show harmonic patterns of distribution
either in the Southern Hemisphere (Crotonia or Podacaridae) or in the Northern
Hemisphere (Ameronothridae and others).
5.6 The Northern Limits of Arachnida
If we consider the distributions beyond the northern polar circle, we find their rep-
resentatives of only four arachnid units: Acariformes (superorder) (Acaridida,
Prostigmata, Oribatida), Parasitiformes (superorder) (Mesostigmata, Ixodida),
Opiliones, and Araneae (Cokendolpher 1981).
Opiliones From the catalogue of Cokendolpher and Lee (1993) of the Opiliones of
North America and Greenland, we find the following limited number of species
(parts of these territories are situated south of the NPC):
Alaska. 13 spp. of the genera Dendrolasma (Nemastomatidae), Leiobunum,
Leptobunus, Leuronychus, Liopilio, Mitopus, Nelima, Parodiellus, Phalangium
(Phalangiidae), Sabacon (Sabaconidae)
Greenland. Mitopus morio (Phalangiidae)
Northwest territories (Canada). Mitopus morio (Phalangiidae)
Yukon territory (Canada). Liopilio (Phalangiidae)
Opiliones from Eurasia north of the NPC:
According to Kauri (1980) and Stol (1993, 2003, 2007), a total of five Opilionids
inhabit the Faroe Islands and four inhabit Iceland. North of the NPC are known:
Finland. According to Lehtinen (1964), in Finnish Lappland (mostly Nord of
66°N) are known six species of Opilions of the genera Nemastoma (Nemastomatidae)
and Mitopus, Oligolophus, Phalangium, Euphalangium Platybunus (Phalangiidae).
Similar Opiliones are recorded from Sweden, Norway, and Russia.
In conclusion, north of the NPC are known Opiliones from three families
(Phalangiidae, Sabaconidae, Nemastomatidae). The widest distributed species is
Mitopus morio (Phalangiidae).
What concerns the Pseudoscorpions, Beier (1963) says: “None, except a few
introduced synanthropic species, are found at higher latitudes than ca. 65°N.”
Scorpiones According to Fet (2010), “The northern limit of natural (non-
anthropogenic) scorpion distribution in Europe is in Saratov Province, Russia, at
50°40′54″ N, for Mesobuthus eupeus (Buthidae).” In Western Europe, the northern-
most record of scorpions seems to be in Austria (Euscorpius germanus, North Tyrol,
Walchsee, 47°39′N; Komposch et al. 2001). Davygora and Rusakov (2001) recorded
Mesobuthus eupeus from Orenburg Province, Russia. The finding in Aituar Steppe
(Orenburg Natural Reserve) at 51°13′N, 57°75′E is “the northernmost limit of natu-
ral scorpion distribution in Asia and the Palearctic” (Fet 2010). According to this
author, “only some records of Paruroctonus boreus (Girard 1854) in southern
Alberta, Canada, lie slightly further north.”
Solifugae In Europe – Spain (Gluvia dorsalis), Sicily (Biton ehrenbergi, B. velox),

Macedonia (Galeodes graecus, G. elegans), Greece, Bulgaria (G. graecus), Ukraine
(Galeodes araneoides). In Asia – Mongolia (Karschia, Galeodes, Eusimonia – five
spp.). In North America – two species reach the southern provinces of Canada:
Eremobates docolora Brookhart et Muma and Hemerotrecha denticulata Muma.
Palpigradi In Europe the limit is crossing Slovakia, Hungary, Austria, and France.
Amblypygi In Asia – on the Greek islands of Rhodes and Kos and in southern
Turkey, as well as two spp. of Charinus Simon in Israel and one sp. in Pakistan. In
North America – the genera Phrynus, Paraphrynus, and Acanthophrynus
(Phrynidae) are known from the states of Arizona, California, Florida, and Texas.
Thelyphonida (Uropygi) In Asia – Typopeltis Pocock (? the Far East of Russia).

In North America – Mastigoproctus Pocock (USA – Arizona, Florida, New Mexico,
Oklahoma, Texas)
Schizomida In Asia – India, Taiwan, and Ryukyus. In North America –

Hubbardiidae (Hubbardia Cook - Arizona, California, Stenochrus Chamberlin,
Agasthoschizomus – USA: Texas). After Reddell and Cokendolpher (1995), two
species (Hubbardia idria and H. briggsi) reach in California 36°30′N.
Araneae The huge order of Araneae contains many cold-resisting species (up to
6700 m in Himalaya, 203 spp. above 3500 m. and many inhabitants of Arctic and
Subarctic areas (Beron 2008b, 2016a, b, c; Larsen and Scharff 2003 (Greenland),
Marusik et al. 2005 (Yakutia), Marusik and Koponen 2002; Koponen 1993, 1995,
1996). Even so, the northern end of the world is much less populated by spiders than
the more southern areas and especially in the tropics.
The next table is modified from Koponen (1996).
5.6 The Northern Limits of Arachnida 99
Area Species Families Area (km2)

Mediterranean 4000 50 –
Switzerland 900 39 41,200
Russian plain 940 36 2,000,000
Belgium 650 35 30,500
Great Britain 620 35 230,700
Norway 535 29 323,900
Finland 620 27 338,100
Alaska 339 1,723,300
Iceland 90 10 89,700a
Greenland 70 10 341,700a
a
Ice – free area
The analysis of Koponen (1996) shows that the family Linyphiidae prevails with
45% of all spiders in Finland, 43% in the Sakhalin-Kuril Islands, 48% in Yakutia,
60% in northeastern Siberia, and 62% in Central Siberia (on close latitude). In the
most northern areas, Linyphiidae is characteristic (Koponen 1995) – from 63% of
all spiders in Greenland and 75% among the nine families in Faroe, remains the
only family with five spp. on Yan Mayen. Koponen (1984) announce that 216 spe-
cies of spiders are known from Inari Lapland (68°20′–70°05′N in Finland).
The harsh climate and lack of forests on the Arctic and Subarctic islands are the
main reason for the low number of spider species on them (5 on Jan Mayen, 14 on
Svalbard, 67 on the Faroe, 70 from Greenland, 90 from Iceland). The estimate of
Koponen (1995) is that on the ice-free area of these islands is 450,000 km2, and they
are populated by ca.150 spp. of spiders. The similarity between the spiders of
Iceland and Scandinavia is 93%, but the fauna of Greenland is clearly Nearctic.
Braendegård (1960) identified a collection of 103 spiders (nine species, eight
native), brought from Peary Land (ca. 82°N). These species were Dictyna borealis
Cambridge.
Acari The general paper on Arctic mites (Trägårdh 1904) appeared very early,
since this time some more publications contributed to the better understanding of
the peculiar fauna surviving north of the polar circle. According to Makarova
(2015), “The number of mite species known in Greenland increased from 158 to
290 (59 species were identified only to the generic level). They belong to the orders
Parasitiformes (Mesostigmata 63 species, Ixodida 1) and Acariformes (Prostigmata
94, Endeostigmata 5, Oribatida 110, Astigmata 17). The free-living mite fauna in
general is characterized by a complete taxonomic structure and a full range of eco-
logical diversity. The Greenland fauna of oribatid mites mainly consists of wide-
spread polyzonal species (70%), while specialized cryobiont (arctic and
arctomontane) species are few (4%); the Nearctic species prevail over the European
invaders with a ratio of 1.5:1. On the contrary, the gamasid mite fauna includes only
25% of widespread species; the cryobiont elements comprise up to 45%; the ratio of
the species of Nearctic and Palearctic origin is 1:4.”
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Chapter 6
Review of the Classification of the Orders
of Arachnida
Abstract Review of all orders with maps of their geographical distribution.
6.1 Palpigradi
Order Palpigradi – 102 spp.

Fam. Eukoeneniidae – Europe; North, South, and tropical Africa; Madagascar;
Indonesia; Philippines; Mauritius; Reunion; Sri Lanka; India; Seychelles;
South America; Mexico; Lebanon; New Guinea; Saudi Arabia; Thailand
(ca. 95 spp.)
Eukoenenia Börner, 1901 – Europe, Morocco, Algeria, Indonesia, Singapore,
Madagascar, Angola, Kenya, Gabon, India, Sri Lanka, Chile, Paraguay,
Brazil, Lebanon, Malta, New Guinea, Mexico, etc. (cosmopolitan, ca. 82
spp.)
Allokoenenia Silvestri, 1913 – Guinea (one sp.)
Koeneniodes Silvestri, 1913 – Madagascar, Guinea, Thailand, Philippines,
Indonesia, New Guinea, Mauritius, Reunion, Seychelles, Sri Lanka (eight
spp.)
Leptokoenenia Condé, 1965 – Congo, Brazil, Saudi Arabia (four spp.)
Fam. Prokoeneniidae – Chile, USA, Southeast Asia, Madagascar (seven spp.)
Prokoenenia Börner, 1901 – Thailand, Indonesia (Java, Sulawesi), Madagascar,
Chile, USA (six spp.)
Triadokoenenia Condé, 1991 – Madagascar (one sp.)

https://doi.org/10.1007/978-3-319-74418-6_6
136 6 Review of the Classification of the Orders of Arachnida
6.2 Schizomida
(According to Manzanilla et al. 2008 – ca. 260 spp.)

Fam. Protoschizomidae – Mexico, USA (16 spp.)
Agastoschizomus Rowland, 1971 – Mexico (eight spp.)
Protoschizomus Rowland, 1975 – Mexico (eight spp.), ? Texas
Fam. Hubbardiidae – USA, Mexico, South and Central America, Caribbean,
tropical Africa, South Africa, Australia, Philippines, Burma, Malaysia,
Java, Sumatra, Thailand, Sri Lanka, Ryukyu Island, Taiwan, New Guinea,
Seychelles, Madagascar, Hawaii, greenhouses in Europe
Subfam. Megaschizominae – Mozambique, Republic of South Africa (two
spp.)
Megaschizomus Lawrence, 1969 – Mozambique, Republic of South Africa
(two spp.)
Subfam. Hubbardiinae – South and Central America, Caribbean, tropical
Africa, Australia, Burma, Malaysia, Java, Sumatra, Philippines, Seychelles,
Madagascar (121 sp.)
Adisomus Cokendolpher et Reddell, 2000 – Brazil (one sp.)
Afrozomus Reddell et Cokendolpher, 1995 – Angola, DR Congo (one sp.)
Anepsiozomus Harvey, 2001 – Seychelles, Socotra (two spp.)
Antillostenochrus Armas et Teruel, 2002 – Cuba, Dominican Republic, Puerto
Rico (10 spp.)
Apozomus Harvey, 1992 – Australia, Borneo, New Guinea, Ryukyu Island,
Taiwan (17 spp.)
Artacarus Cook, 1898 – Liberia, Ivory Coast (one sp.)
Attenuizomus Harvey, 2000 – Australia (NT, four spp.)
Bamazomus Harvey, 1992 – Seychelles, Madagascar, Thailand, Hong Kong,
Japan (Ryukyu Island), West Malaysia, Papua New Guinea, Australia,
Hawaii (11 spp.)
Belicenozomus Armas et Viques, 2010 – Belize and Honduras (two spp.)
Brignolizomus Harvey, 2000 – Australia (three spp.)
Bucinozomus Armas et Rehfeldt, 2015 – greenhouse in Germany (one sp.)
Burmezomus Bastawade, 2004 – Burma (one sp.)
Calima Moreno-González et Villarreal, 2012 – Colombia (two spp.)
6.2 Schizomida 137
Caribezomus Armas, 2011 – Jamaica (one sp.)

Clavizomus Reddell et Cokendolpher, 1995 – Java, West Malaysia, Singapore
(one sp.)
Cokendolpherius Armas, 2011 – Cuba (one sp.)
Colombiazomus Armas et Delgado-Santa, 2012 – Colombia (two spp.)
Cubazomus Reddell et Cokendolpher, 1995 – Cuba (three spp.)
Cubacanthozomus Teruel, 2007 – Cuba (one sp.)
Draculoides Harvey, 1992 – W. Australia (six spp.)
Enigmazomus Harvey, 2006 –Somalia, Arabian Peninsula (two spp.)
Gravelyzomus Kulkarni, 2012 – India (one sp.)
Guanazomus Teruel et Armas, 2002 – Cuba (one sp.)
Hansenochrus Reddell et Cokendolpher, 1995 – Venezuela, Central America,
Lesser Antilles (14 spp.)
Heterocubazomus Teruel, 2007 – Cuba (one sp.)
Heteronochrus Armas et Viquez, 2010 – Guatemala (one sp.)
Hubbardia Cook, 1899 – USA (Arizona, California) (nine spp.)
Javazomus Reddell et Cokendolpher, 1995 – Java (one sp.)
Julattenius Harvey, 1992 – Australia (Queensland)(two sp.)
Kenyazomus Armas, 2014 – Kenya (one sp.)
Lawrencezomus Armas, 2014 – Cameroon, Liberia (two spp.)
Luisarmasius Reddell et Cokendolpher, 1995 – Puerto Rico, Cuba (Pinos)
(two spp.)
Mahezomus Harvey, 2001 – Seychelles (one sp.)
Mayazomus Reddell et Cokendolpher, 1995 – Mexico (seven sp.)
Notozomus Harvey, 2000 – Australia (Queensland, 17 spp.)
Orientzomus Cokendolpher et Tsurusaki, 1994 – Philippines (Luzon), Japan,
Bonin Island (two spp.)
Ovozomus Harvey, 2001 – Seychelles, Ceylon, India, Mayotte, Cook Island,
Reunion (two spp.)
Pacal Reddell et Cokendolpher, 1995 – Mexico (three spp.)
Paradraculoides Harvey, Berry, Edward et Humphreys, 2008 – W. Australia
(four spp.)
Piaroa Manzanilla, Giupponi et Tourinho, 2008 – Venezuela, Colombia,
Costa Rica (three spp.)
Reddellzomus Armas, 2002 – Cuba (one sp.)
Rowlandius Reddell et Cokendolpher, 1995 – Venezuela, Central America,
Caribbean, Brazil (55 spp.)
Secozomus Harvey, 2001 – Seychelles (one sp.)
Fig. 6.1 Hubbardia pentapeltis ♀ (Photo and courtesy Marshal Hedin)
Sotanostenochrus Reddell et Cokendolpher, 1991 – Mexico (San Luis Potosí,

Tamaulipas)(two spp.)
Stenochrus Chamberlin, 1922 – USA (Texas), Guatemala, Mexico, Cuba,
Puerto Rico, Dominican Republic, Jamaica, South America (21 spp.) (S.
portoricensis Chamberlin, 1922 is more widely distributed, imported in
Europe)
Stenoschizomus González-Sponga, 1997 – Venezuela (one sp.)
Stewartpeckius Reddell et Cokendolpher, 1995 – Jamaica (one sp.)
Surazomus Reddell et Cokendolpher, 1995 – Costa Rica, South America (six
spp.)
Tayos Reddell et Cokendolpher, 1995 – Ecuador (one sp.)
Trithyreus Kraepelin, 1899 – Burma (Myanmar) (one sp.)
Troglocubazomus Teruel, 2003 – Cuba (one sp.)
Wuayuuzomus Armas et Colmenares, 2006 – Venezuela (one sp.)
Zomus Reddell et Cokendolpher, 1995 – Malaysia (incl. Sarawak), Singapore,
Seychelles, Fiji, Samoa, Rodrigues, England (Kew Garden) (one sp.)
(Fig. 6.1)
6.3 Thelyphonida (Uropygi)
Cooke and Shadab (1973), Gravely (1916), Harvey (2003, 2013d), Haupt (2009b),
Huff and Prendini (2009), Mello-Leitão (1931), Rowland (1973a, b, 2002), Speijer
(1933, 1936), Viquez and Armas (2005), Werner (1935)
6.3 Thelyphonida (Uropygi) 139
The first comprehensive papers on the group were the revisions of Pocock (1894)
and Kraepelin (1897). Important contributions were made also by Thorell, Gravely,
Mello-Leitão, Werner, Speijer, Rowland, Haupt, and others.
Rowland and Cooke (1973) list 85 species in the order. Haupt (2009a) synony-
mized Abaliella Strand, Minbosius Speijer, and Tetrabalius Thorell with Thelyphonus
Latreille. Krehenwinkel et al. (2009) described the new genus Thelyphonoides from
Panay (Philippines). According to the calculations of Harvey (2002b, 2003, 2013d)
and the additions since this time, there are ca. 110 species in the order Thelyphonida
(Uropygi), belonging to 15 genera and one recent family (Blick and Harvey 2011).
They are distributed as follows:
Fam. Thelyphonidae – Southeast Asia, Indonesia, Philippines, Samoa, New
Guinea, Fiji, Solomon Island, Vanuatu, India, Sri Lanka, China, North,
South, and Central America, Caribbean Islands, West Africa (ca.110 spp.)
Subfam. Thelyphoninae – Southeast Asia, Philippines, Samoa, New Guinea,
Fiji, Solomon Island, India (49 spp.)
Ginosigma Speijer, 1936 – Sunda Islands, Thailand, Laos, Cambodia, Vietnam
(two spp.)
Glyptogluteus Rowland, 1973 – Philippines (Panay)(one sp.)
Thelyphonoides Krehenwinkel et al., 2009 – Philippines (Panay)(one sp.)
Thelyphonus Latreille, 1802 (Chajnus Speijer, 1936; Abaliella Strand, 1928;
Minbosius Speijer, 1936; Tetrabalius Thorell, 1889; fide Haupt 2009a) –
Indonesia, Singapore, Philippines, Vanuatu (New Hebrides), Fiji, Rennell,
Solomon Islands, New Guinea, New Britain, Burma, Sri Lanka, India,
Thailand, W. Samoa, Borneo, Moluccas, Cambodia, Vietnam (45 spp.)
Subfam. Hypoctoninae – Burma (Myanmar), South China, Malaysia, West
Africa, India, Sri Lanka, Brazil, Guyana, Venezuela, Dominican Republic (27
spp.)
Etienneus Heurtault, 1984 – Western Africa (one sp.)
Hypoctonus Thorell, 1889 – Burma (Myanmar), South China, Malaysia,
Thailand, Bangladesh, Java, India (19 spp.)
Labochirus Pocock, 1894 – India, Sri Lanka (four spp.)
Thelyphonellus Pocock, 1894 – Brazil, Guyana, Venezuela (two spp.)
Ravilops Viquez et Armas, 2005 – Dominican Republic (one sp.)
Subfam. Mastigoproctinae (Uroproctinae) – Brazil, Cuba, Colombia, El
Salvador, Costa Rica, Honduras, Belize, Venezuela, Mexico, Guatemala,
Haiti, Martinique, USA, Hainan, Philippines (23 spp.)
Uroproctus Pocock, 1894 – India (Assam), Cambodia (one sp.)

Mimoscorpius Pocock, 1894 – Guatemala (one sp.)
Mayacentrum Viquez et Armas, 2006 – Guatemala, El Salvador, Honduras,
Belize (three spp.)
Mastigoproctus Pocock, 1894 (Amauromastigon Mello-Leitão, 1931 –
Brazil) – Cuba, Brazil, Colombia, Venezuela, USA, Mexico, Haiti,
Martinique (17 spp.)
Valeriophonus Viquez et Armas, 2005 – Costa Rica (one sp.)
Subfam. Typopeltinae – China, Taiwan, Hainan, Japan, Thailand, Vietnam,
Laos (11 spp.)
Typopeltis Pocock, 1894 (Teltus Speijer, 1936) – China, Taiwan, Hainan,
Japan, Thailand, Laos, Vietnam; doubtful finding in the Far East of Russia
(11 confirmed spp.)
6.4 Amblypygi
These big and conspicuous, largely cavernicolous, dwellers of the warmer places
have been subject to many articles (and many synonyms) by earlier researchers,
starting with Linnaeus, Lamarck, Herbst, and Gerstaecker, and also the researchers
of nineteenth and early twentieth centuries (Gervais, C.L. Koch, L. Koch, Bilimek,
Karsch, Pocock, Butler, Kraepelin, Simon, Gravely, Thorell, Hansen, and others).
Many contributions have been made in the last 90 years by Franganillo, Mello-
Leitão, Fage, Lawrence, Dunn, Whittick, Caporiacco, Moreno, Ravelo, Kritscher,
Rowland, Armas, González-Sponga, Delle Cave, Harvey, Rahmadi, and other
authors.
The papers of Mullinex and Quintero and especially the revisions of Weygoldt
are the basis of the modern understanding of the order and counting (Harvey 2003,
136 spp.; Blick and Harvey 2011; Miranda and Giupponi 2011; Harvey 2013c, actu-
alized 2017, 184 spp.; Miranda et al. 2018, 220 sp.) of 18 genera and five families.
Suborder Euamblypygi
Fam. Charinidae – Greece (Rhodes, Kos), Turkey, Israel, Egypt, Cuba, Jamaica,
Galapagos Islands, US Virgin Islands, tropical South America (Guiana, Brazil,
Suriname, Venezuela, Peru), Australia, Andaman Islands, India, W. Samoa,
Vanuatu, Oman, Indonesia (Java, Borneo), Singapore, Malaysia, Burma,
Vietnam, Laos, Cambodia, Papua New Guinea (incl. New Britain), Seychelles,
Madagascar, New Caledonia, Solomon Islands, tropical Africa (Somalia,
Tanzania, Guinea, Equatorial Guinea, Sao Tome, Principe, Somalia, Socotra,
Pakistan) (90 spp.)
6.4 Amblypygi 141
Catageus Thorell, 1889 (= Stygophrynus Kraepelin, 1896)– Burma, Solomon

Islands, Thailand, Laos, Cambodia, Vietnam, Indonesia, Malaysia (nine sp.)
(nine sp.)
Weygoldtia Miranda, Giupponi, Prendini et Scharff, 2018 - one sp. W. davidovi
(Fage, 1946) - Laos, Cambodia, Vietnam
Charinus Simon, 1892 (Charinides Gravely, 1911; Enantiosarax Mello-Leitão,
1931; Oligacanthophrynus Caporiacco, 1947; Lindosiella Kritscher, 1959;
Speleophrynus Ravelo, 1975; Tricharinus Quintero, 1986) – Greece (Rhodes,
Kos), Turkey, Israel, Cuba, Jamaica, Guiana, Brazil, Suriname, Venezuela,
Peru, Galapagos Islands, US Virgin Islands, Australia, Oman, Andaman
Islands, India, W. Samoa, Vanuatu, Indonesia (Java, Borneo), Singapore,
Malaysia, Papua New Guinea, Seychelles, Madagascar, New Caledonia,
Solomon Islands, Tanzania, Guinea, Equatorial Guinea, Sao Tome, Principe,
Somalia, Egypt, Socotra, Pakistan (74 spp.)
Sarax Simon, 1892 (Phrynichosarax Gravely, 1915) – Solomon Islands,
Malaysia, Singapore, Philippines (Luzon), Indonesia (Java, Kalimantan),
India, Andaman Islands, Papua New Guinea (incl. New Britain and New
Ireland), Vietnam, Laos, Cambodia, Borneo (17 spp.)
Fam. Charontidae – Australia (incl. Christmas Island), Indonesia (Java, Maluku,
Sumbawa), Malaysia, Burma, Thailand, Vietnam, Palau, Papua New Guinea,
Philippines, Singapore, Solomon Islands (nine spp.)
Charon Karsch, 1879 – Australia (Northern Territory, Queensland, Christmas
Island), Indonesia (Java, Maluku, Sumbawa), Malaysia (incl. Borneo), Palau,
Papua New Guinea, Philippines, Singapore, Solomon Islands (nine spp.)
Fam. Phrynichidae – tropical and Southern Africa, Algeria, Morocco, Yemen,
Saudi Arabia, Oman, Mauritius, Seychelles, Zanzibar, Sao Tome, Madagascar,
India, Sri Lanka, Brazil (33 spp.)
Subfam. Damoninae – tropical and Southern Africa, Algeria, Morocco, Yemen
(12 spp.)
Damon C.L. Koch, 1850 (Titanodamon Pocock, 1894; Nanodamon Pocock,
1894) – South Africa, Malawi, Mozambique, Zambia, Zimbabwe, Ethiopia,
Kenya, Somalia, Tanzania, Yemen, Angola, Namibia, Cameroon, Congo,
DR Congo, Equatorial Guinea, Gabon, Nigeria, Liberia, Ivory Coast, Sao
Tome, Mali, Ghana, Benin, Senegal, Sierra Leone, Togo, Botswana, Sudan
(10 spp.)
Musicodamon Fage, 1939 – Algeria, Morocco (one sp.)
Phrynichodamon Weygoldt, 1996 – Namibia, South Africa (one sp.)

Subfam. Phrynichinae – Brazil, Kenya, Tanzania, Zanzibar, DR Congo,
Madagascar, Malawi, Zimbabwe, Angola, Namibia, Eritrea, Somalia,
Djibouti, Kenya, Sri Lanka, Yemen, Saudi Arabia, Oman, Socotra, Mauritius,
Seychelles (21 spp.)
Euphrynichus Weygoldt, 1995 – Kenya, Tanzania (incl. Zanzibar), DR Congo,
Madagascar, Malawi, Zimbabwe (two spp.)
Phrynichus Karsch, 1879 (Myodalis Simon, 1936) – Eritrea, Somalia,
Djibouti, Kenya, India, Sri Lanka, Yemen, Saudi Arabia, Oman,
Madagascar, Mauritius, Socotra, Seychelles (16 spp.)
Trichodamon Mello-Leitão, 1935 – Brazil (two spp.)
Xerophrynus Weygoldt, 1996 – Angola, Namibia (one sp.)
Fam. Phrynidae – tropical South America, USA (Arizona, California, Florida),
Central America, Bahamas, Cuba, Barbados, St. Vincent, Netherlands Antilles,
Grenades, Haiti, Saint Lucia, Mexico, Indonesia (Flores) (62 spp.)
Subfam. Heterophryninae – tropical South America (14 spp.)
Heterophrynus Pocock, 1894 – Brazil, Peru, Colombia, French Guiana,
Guyana, Venezuela, Ecuador, Suriname (14 spp.)
Subfam. Phryninae – tropical South America, USA (Arizona, California,
Florida, Texas), Mexico, Central America, Bahamas, Cuba, Barbados, St.
Vincent, Netherlands Antilles, Grenades, Haiti, Saint Lucia, Indonesia
(Flores) (48 spp.)
Acanthophrynus Kraepelin, 1899 – Mexico, ? USA (Arizona, California)(one
sp.)
Paraphrynus Moreno, 1940 – USA (Arizona, Florida), Mexico, Cuba,
Guatemala, Colombia, Costa Rica, El Salvador, Nicaragua, Panama,
Honduras, Bahamas (16 spp.)
Phrynus Lamarck, 1801 (Admetus C.L. Koch; Neophrynus Kraepelin, 1895) –
Mexico, Barbados, St. Vincent, Nicaragua, Colombia, Ecuador, Guyana,
Cuba, Dominican Republic, Virgin Islands, Jamaica, Martinique, Cayman
Islands, Antigua, Barbuda, Bermuda, Bahamas, Guatemala, Honduras,
Belize, Costa Rica, USA (Texas, Florida), Trinidad, Panama, Venezuela,
Netherlands Antilles, Grenadines, Puerto Rico, Haiti, Saint Lucia, Brazil,
Indonesia (Flores) (31 spp.)
6.5 Ricinulei 143
Suborder Palaeoamblypygi
Fam. Paracharontidae – Guinea-Bissau (one sp.)
Paracharon Hansen, 1921 – Guinea-Bissau (one sp.)
6.5 Ricinulei
This small (3 genera, 55 spp., according to Harvey 2003; 64 spp. after Cook 2008;
67 after Tourinho et al. 2010; 58 after Prendini 2011) group of arachnids is some-
times considered representing “living fossils” (Selden 1986).
The studies on recent Ricinulei start in 1838 (Guérin-Méneville). Longtime they
are considered very rare, thanks to the technique of collecting. From the 66 living
valid species, recorded up to 2011, 49 have been described after 1950 by Roewer,
Platnick, Shadab, Cooke, Cooreman, González-Sponga, Gertsch, Brignoli, Cor.
Gutierrez, Dumitresco and Juvara-Balş, Merrett, Marquez and Conconi, Goodnight
and Goodnight, Armas, Legg, Naskrecki Cokendolpher, and other researchers. The
increase is mostly to the expense of the American genera Cryptocellus and the
newly described Pseudocellus (44 new species). In 60 years only five new species
have been added to the six African Ricinoides known before 1950 (Legg, 1976,
1977 from Sierra Leone, 1982 from Ivory Coast, Naskrecki, 2008 from Ghana,
Penney et al. from Gambia). Wunderlich (2012) downgraded the suborders
Neoricinulei Selden, 1992, and Palaeoricinulei Selden, 1992, to superfamilial rank,
the Poliocheroidea Scudder, 1884, resp. Ricinoidoidea Ewing, 1929.
Suborder Neoricinulei Selden, 1992 Fam. Ricinoididae (? 66 spp.)
Cryptocellus Westwood, 1874 (Heteroricinoides Dumitresco et Juvara-Balş,
1977) – Colombia, Brazil, Guyana, Panama, Venezuela, Suriname, Costa Rica,
Honduras, Nicaragua, Ecuador, Peru, Tobago (34 spp.)
Pseudocellus Platnick, 1980 – USA (Texas), Mexico, Guatemala, Honduras, El
Salvador, Panama, Cuba (21 spp.)
Ricinoides Ewing, 1929 – Gambia, Guinea, Guinea-Bissau, Nigeria, Ghana,
Cameroon, Sierra Leone, Equatorial Guinea, Benin, Congo, Togo, Ivory Coast
(11 spp.)
6.6 Scorpiones
According to Dupré (2010), there are about 1950 valid species of scorpions, belong-
ing to 152 genera and 7–20 families (18 fam. + Akravidae, after Prendini 2011), but
the latest number of species (after Jan Ove Rein 2017, The Scorpion files) is 2326.
The present arrangement follows generally the systems of Soleglad and Fet (2003)
and Lourenço (2000b), but also takes into account the objections of Prendini and
Wheeler (2005) and the development of the taxonomy after 2005. The number and
contents of higher taxa in the paper of Soleglad and Fet were strongly disputed by
Prendini and Wheeler (2005). Still, as these authors point “The suprageneric clas-
sification of recent (including extant) scorpions is in a state of flux.” This flux is
more obvious than in any other of the arachnid orders. The present author finds this
situation quite unfortunate for the zoogeography, which should be based on good
supraspecific taxonomy for explaining endemism, distribution, and the whole his-
tory of the order. And this is particularly true for scorpions, considered very impor-
tant biogeographically. Vachon (1968) emphasizes that “Scorpions can play an
important part in biogeography. These animals travel slowly…Scorpions thus
evolve in situ and are practically immune to variations in micro-climate. Thus they
constitute excellent biogeographic test and their presence (or absence) provides a
better understanding or interpretation of the population background of a given
region.”
Fet and Soleglad (2014) reject the criticism of Prendini and Wheeler (2005). As
it is not to us to solve here the problem, we shall use the list of families in Prendini
(2011), considering Akravidae recent family, without discussing the higher taxa and
the subfamilies.
Nenilin and Fet (1992) analyzed the zoogeography of the world scorpion fauna
on the level of 1985–1986, but since that time many changes occurred in the nomen-
clature of scorpions and new data were obtained.
Numbers of Families in Classifications of Recent Scorpions Proposed During
the Past 30 Years (Partly from Prendini and Wheeler 2005)
Lamoral (1980) – 7
Stockwell (1989) – 13
Sissom (1990) – 9
Kovařik (1998) – 13
Fet et al. (2000) – 16
Lourenço (2000b) – 20
Soleglad and Fet (2003) – 14
Soleglad et al. (2005) – 13
Prendini and Wheeler (2005) – 18
Prendini (2011) – 18 (+ Akravidae ? extinct)
Present paper – 20
6.6 Scorpiones 145
Superfam. Bothriuroidea
Fam. Bothriuridae – South America, Namibia, Australia, Indian Himalaya (12
gen., 138 spp.)
Superfam. Buthoidea
Fam. Buthidae – all continents, tropical and subtropical countries, and in some
countries with moderate climate (Spain, France, Cyprus)(70 genera, 939 spp.)
Fam. Microcharmidae – Madagascar (one gen., six spp.) (synonymized with
Buthidae by Volschenk et al. 2008 ?)
Superfam. Chaeriloidea
Fam. Chaerilidae – Bangladesh, India, Indonesia, Malaysia, Nepal, Singapore, Sri
Lanka, Vietnam (1 gen., 43 spp.)
Fam. Pseudochactidae – Uzbekistan, Tajikistan, Afghanistan, caves of Laos, and
Vietnam (three gen., five spp.)
Superfam. Chactoidea
Fam. Chactidae – Central and South America, Mexico (Baja California) (9 genera,
178 spp.)
Fam. Euscorpiidae – South Europe, North Africa, Middle East, Central Asia
(south), Mexico, Guatemala (4 gen., 25 spp.)
Fam. Scorpiopidae – Afghanistan, Southeast Asia, India, Pakistan, Indonesia,
Malaysia (5 gen., 48 spp.)
Fam. Troglotayosicidae – Pyrenees (France/Spain)(?), Equatorial Amazonas (?
two genera, three spp.)
Fam. Superstitioniidae – Mexico, Southeast of the USA (four genera)
Superfam. Vaejovoidea
Fam. Iuridae – North and South America, Turkey, Greece (6 gen., 36 spp.)
Fam. Vaejovidae – Mexico, South of the USA (17 gen., 181 spp.)
Superfam. Scorpionoidea
Fam. Hemiscorpiidae – Middle East (2 gen, 13 spp.)
Fam. Heteroscorpionidae – Madagascar (one gen., six spp.)
Fam. Hormuridae (Liochelidae)(incl. Hadogenidae) – Australia, Africa, Central
and South America, Antilles, Southeast Asia, Tibet, Pacific islands, Madagascar,
India, Indian Ocean Islands, Seychelles, Mauritius, Hispaniola, Pacific Cocos
Island (9 gen., 69 spp.)
Fam. Scorpionidae (incl. Diplocentridae) – North America, Central America, the
northern part of South America, Antilles, Middle East, Africa, East, South and
Southeast Asia, Indonesia (12 gen., 248 spp.)
Fam. Akravidae – Israel (one sp.)
Fam. Lisposomidae – South Africa (one genus)
Fam. Urodacidae – Australia (2 gen., 21 spp.)
6.7 Pseudoscorpiones
More than 3538 described species worldwide (Harvey 2002, 2013f, actualized).
After the catalogue of Harvey (1990) and of his revision of 1992 and reviews of
2002, 2011, and 2013f and the additions of Mahnert (several papers) and others, it
is generally accepted that the order includes 2 suborders and 26 recent families with
more than 454 genera.
Suborder Epiocheirata
Fam. Chthoniidae – Cosmopolitic, ca. 650 spp., 28 gen.
Fam. Tridenchthoniidae (Dithidae) – South Africa, Australia, New Caledonia,
Lord Howe Island, Ecuador, Cuba, Jamaica, Mexico, Costa Rica, Trinidad, Peru,
Suriname, Paraguay, Argentina, Brazil, Venezuela, Equatorial Africa, Ethiopia,
Caroline Islands, India, Vietnam, Philippines, Seychelles, Indonesia (Moluccas,
Sumatra, Java), New Guinea, Japan, Palau Island, Bhutan, Nepal, Laos, Marcus
Island, Malaysia, South Africa, Sao Tome, USA (15 gen., 71 spp.)
Fam. Pseudotyrannochthoniidae – Australia, Tasmania, Korea, Eastern USA,
Japan, Chile, China (5 gen., 49 spp.)
Fam. Lechytiidae – Turkey, USA, Mexico, Chile, Peru, Argentina, Paraguay,
Galapagos Islands, Jamaica, Dominican Republic, Trinidad, Guadeloupe,
Martinique, Congo, D.R. Congo, Kenya, Tanzania, Nepal, Vietnam, India, Juan
Fernandez Islands, South Africa, Caroline Islands, Marshall Islands, Marcus
Island (1 gen., 23 spp.)
Fam. Feaellidae – tropical and Southern Africa, Seychelles, Madagascar, India, Sri
Lanka, Australia (1 gen., 12 spp.)
Fam. Pseudogarypidae – North America, Tasmania, Baltic Amber from the
Oligocene (two genera, seven recent species)
Suborder Iocheirata
Fam. Ideoroncidae – Argentina, Brazil, Paraguay, Chile, US Virgin Islands,
Mexico, USA, Afghanistan, Iran, Malaysia, India, Cambodia, Thailand, Vietnam,
Indonesia (Sumatra), Sri Lanka, Kenya, Tanzania, Uganda, Congo, Zimbabwe,
Botswana (13 gen., 70 spp.)
Fam. Bochicidae (Vachoniidae) – Central America as far north as Texas, Cuba,
Grenada, Trinidad, Jamaica, Spain, Portugal (14 gen., 44 spp.)
6.7 Pseudoscorpiones 147
Fam. Hyidae – Philippine Islands, Indonesia (Java, Sumatra), Sri Lanka, India,
Australia (2 gen., 14 spp.)
Fam. Gymnobisiidae – Southern South America, South Africa (4 gen., 11 spp.)
Fam. Neobisiidae – Europe, Corsica, Sicily, Sardinia, Balearic Islands, Canary
Islands, Malta, Georgia, Armenia, Azerbaijan, Azores, Iran, Turkey, Turkmenistan,
Kyrgyzstan, Far East of Russia, Tibet, Nepal, India, Philippines, Mongolia,
Afghanistan, Vietnam, Thailand, Japan, Korea, Burma, China, Taiwan, Canada,
USA, DR Congo, Kenya, Rwanda, Tanzania, Zimbabwe, Algeria, Morocco,
Cyprus (33 genera, 595 spp.)
Fam. Syarinidae – South America, Antillean Islands, Mexico, USA, Canada, Italy,
Greece, Norway, Finland, Austria, Germany, France, Switzerland, Crimea,
Algeria, Balearic Islands, Canary Islands, Lebanon, Malta, Morocco, Portugal,
Sardinia, Sicily, Spain, Bhutan, India, Sri Lanka, Nepal, New Caledonia, Tuvalu,
Philippines, Solomon Island, New Zealand, New Guinea, tropical Africa, South
Africa, Seychelles, Galapagos Island (18 genera, 111 spp.)
Fam. Parahyidae – Caroline Islands, Singapore (one genus, one sp.)
Fam. Garypidae (Synsphyronidae) – Armenia, Algeria, Balearic Island, Crete,
Egypt, Canary Island, Corsica, France, Greece, Ilhas Selvagens, Israel, Italy,
Malta, Libya, Sardinia, Spain, Portugal, Tunisia, ex-Yugoslavia, Bonaire, Aruba,
Mustique Island, Mexico, USA (California, Florida), India, Seychelles, South
Africa, Japan, Indonesia (Krakatau, Nicobar Island), Maldives Island, Sri Lanka,
Kenya, Somalia, Yemen, Namibia, South Africa, Zimbabwe, Madagascar,
Marshall Islands, Ascension Island, Australia (incl. Tasmania), New Zealand (10
genera, 80 spp.)
Fam. Geogarypidae – tropical Africa, South Africa, Aldabra, Seychelles, Indonesia,
Malaysia, India, Sri Lanka, Bhutan, China, Taiwan, Argentine, Chile, Brazil,
Paraguay, Guyana, Grenada, Juan Fernandez Island, Canary Island, Algeria,
Morocco, Australia, Russia, Kazakhstan, Kyrgyzstan, Turkmenistan, Thailand,
Caroline Island, Mariana Island, Marshall Island, Solomon Island, Tuvalu,
Marquesas Island, Marcus Island, Europe, Turkey, Malta, Sardinia, Sicily,
Madeira, Hawaii, Israel, Iran (3 gen., 60 spp.)
Fam. Larcidae – USA, Spain, Italy, Austria, Denmark, Poland, Romania, Sweden
(2 genera, 15 spp.)
Fam. Cheiridiidae – Chile, Guadeloupe, Cuba, Aruba, Curacao, Namibia, tropical
Africa, South Africa, Lesotho, Russia, Estonia, Finland, Azerbaijan, China,
Taiwan, Korea, Nepal, Philippines, Japan, Hawaii, Papua New Guinea, Salomon
Islands, Mariana Islands, Australia, New Zealand; for Ch. museorum Leach,
Europe, India, Mozambique, DR Congo, Russia, Latvia, Kyrgyzstan, Turkey,
South Africa, USA (7 gen., 73 spp.)
Fam. Pseudochiridiidae – Madagascar, Aldabra, tropical Africa, India, Andaman

Island, Burma, Indonesia, Malaysia, Nepal, Philippines, New Guinea, Brazil (2
gen., 12 spp.).
Fam. Olpiidae – Australia, New Zealand, Norfolk Island, Aldabra Islands,
Madagascar, Algeria, Seychelles, USA (Florida), Lesser Antilles, Jamaica, South
America, Turkey, Indonesia, India, Burma, Turkmenistan, Tajikistan, Uzbekistan,
Japan, Cambodia, Vietnam, Cyprus, Yemen, Iran, Iraq, Jordan, Israel, Lebanon,
Corsica, France, Italy, Sardinia, Madeira Islands, Greece, Croatia, Papua New
Guinea, New Caledonia, Salomon Islands, Tuvalu, Samoa, Caroline Islands,
Mariana Islands, Marshall Islands, tropical Africa, Namibia, South Africa, Cabo
Verde, Egypt, Djibouti (36 gen., 268 spp.)
Fam. Garypinidae – Peru, Chile, USA, Mexico, South America, Jamaica,
Galapagos Islands, Solomon Islands, Hawaii, Australia, New Guinea, Israel,
Syria, Turkey, Yemen, Morocco, Cyprus, Crete, Greece, Corfu, Italy, Sicily,
Croatia, Hawaii, South Africa, Yemen, Iran, Indonesia, Thailand, Cambodia,
Vietnam, and others (21 gen., 76 spp.)
Fam. Menthidae – Mexico, USA (California), Argentina, Chile, Israel, Socotra,
Australia (5 gen., 12 spp.)
Fam. Sternophoridae – Ethiopia, Australia, Papua New Guinea, India, Sri Lanka,
Laos, Cambodia, Vietnam, El Salvador, Dominican Republic, USA (Florida),
Mexico, Cayman Islands (3 gen., 20 spp.)
Fam. Withiidae – South Africa, tropical Africa, Seychelles, Cabo Verde, Sao Tome,
St. Helena, New Zealand, Lord Howe Island, Juan Fernandez Islands, USA
(Texas), South America, Mexico, Cuba, Virgin Islands, Vietnam, Indonesia,
Christmas Island, India, Cambodia, Malaysia, Thailand, Philippines, Solomon
Islands, Saudi Arabia, Pakistan, Iran, Nepal, China, Taiwan, Mongolia, Japan,
Yemen, Israel, Morocco, Tunisia, Libya, USA, Mariana Islands, Marshall
Islands, Papua New Guinea, Greece, Bulgaria, Russia, France, Corsica, Italy,
Sardinia, Portugal, Spain, Switzerland, Turkey, Georgia, ex-Yugoslavia; only for
Withius piger, Algeria, Australia, Austria, Balearic Island, Canary Islands, Chad,
Chile, China, Cuba, Denmark, Egypt, France, Ghana, Great Britain, Hungary,
India, Italy, Libya, Malta, Mexico, Morocco, Netherlands (38 gen., 158 spp.)
Fam. Cheliferidae – South Africa, tropical Africa, North Africa, Socotra, Japan,
Europe, Malta, Madeira, Canary Islands, Australia, New Zealand, Papua New
Guinea, USA, Mexico, Guatemala, Virgin Islands, Dominican Republic, Cuba,
Brazil, Ecuador, Mauritius, Reunion, Indochina, India, China, Mongolia,
Indonesia, Afghanistan, Pakistan, Kyrgyzstan, Kazakhstan, Tajikistan,
Uzbekistan, Iran, Iraq, Saudi Arabia, Israel, Jordan, Yemen (58 gen., 273 spp.)
6.8 Opiliones 149
Fam. Atemnidae (Miratemnidae) – Southeast Asia, Central Asia, Mongolia, Japan,

Philippines, Bhutan, Nepal, India, Nicobar Islands, Solomon Islands, Sri Lanka,
Indonesia, China, North Africa, tropical Africa, Yemen, Sao Tome, Comoro
Islands, Seychelles, Madagascar, Europe, Cyprus, Canary Islands, Balearic
Islands, Sardinia, Sicily, Puerto Rico, Brazil, Ecuador, Guyana, Cuba, Dominican
Republic, St. Vincent, Costa Rica, US Virgin Islands, Brazil, Argentina, Peru,
Colombia, Panama, Venezuela, Haiti, Paraguay, St. Vincent, St. Paul Island, Ilhas
Selvagens, USA (Arizona), New Caledonia, Lord Howe Islands, Caroline
Islands, Mariana Islands, Christmas Island, Marquesas Islands, Marshall Islands,
Papua New Guinea (21 gen., 178 spp.)
Fam. Chernetidae (Myrmochernetidae) – Europe; South America; Mexico; USA;
Central America, Antilles; San Ambrosio Island; New Guinea; Solomon Islands;
Mongolia; China, Japan; North, Central, and South Africa; New Zealand;
Australia; etc. (117 gen., 663 spp.)
6.8 Opiliones
The large order Opiliones (46 families, 1649 extant genera, and more than 6000
species, according to Harvey 2002 and Pinto-da-Rocha and Giribet 2007; 6534, acc.
to Kury 2013, 6653 acc. to Kury, April 2017) was traditionally subdivided into three
suborders: Cyphophthalmi, Laniatores (mostly tropical), and Palpatores (mostly
Holarctic). Recently, the “Palpatores” are firmly considered as two suborders –
Eupnoi and Dyspnoi (Giribet and Kury, Pinto-da-Rocha and Giribet 2007; Kury
2011, 2013). The knowledge on the Opiliones was first summarized by Roewer
(1923), who described 2260 valid species and greatly enlarged the knowledge on
Opiliones by many publications since that time. Unfortunately, there is a strong
suspicion that some localities of Roewer’s material have been confused (von
Helversen and Martens (1972), von Helversen (1968)) and should be considered
very carefully when zoogeographical conclusions are made. It is always wiser to use
the data of the modern authors with reliable localities.
Classification of Opiliones After Kury (2013, April, 2017). The Number of
Species in the Families Is Not Updated
Cyphophthalmi (6 fam., ca. 36 genera, 208 spp.). The immense progress in the
study of Cyphophthalmi can be assessed, comparing the present day picture and the
biogeographical and taxonomical outline of the suborder of Juberthie and Massoud
(1979). Their paper starts with the passage: “Les Opilions du sous-ordre des
Cyphophthalmes représentent une petite famille de 15 genres et 56 espèces….répar-

ties sur l’ensemble du globe.”
Fam. Stylocellidae – Southeast Asia (from India to New Guinea) (6 gen., 36 spp.)
Fam. Ogoveidae – Western Equatorial Africa (one gen., three spp.)
Fam. Neogoveidae – South America (Colombia, Brazil, Ecuador, Guyana, French
Guiana, Peru, Suriname, Trinidad and Tobago, Venezuela), USA (Florida,
Georgia, South Carolina), West Africa (Bioko, Gabon, Sierra Leone) (9 gen., 22
spp.) (+many undescr.)
Map 6.1 Distribution of Pettalidae (▼) and Neogoveidae (■) in South America and the southern
part of North America
Superfamily Sironoidea
Fam. Pettalidae – New Zealand, Australia, Chile, South Africa, Sri Lanka (9 gen.,
61 spp.)
Fam. Sironidae – South Europe, Corsica, Carpathians, Asia Minor, USA (Western
and Eastern shore), Japan, ? Kenya (7 gen., 52 spp.) (Fig. 6.2)
6.8 Opiliones 151
Fig. 6.2 Fam. Troglosironidae – New Caledonia (1 gen., 13 spp.)

Troglosiro longifossa Sharma et Giribet (courtesy of G. Giribet)
Map 6.2 Distribution of Sironidae (♦) in Europe and Western Asia

Map 6.3 Distribution of Neogoveidae (●) and Sironidae (♦) in North America
Map 6.4 Distribution of Stylocellidae (►), Pettalidae (▼), and Sironidae (■) in Southeast Asia
6.8 Opiliones 153
Map 6.5 Distribution of Pettalidae (▼– Australia, New Zealand) and Troglosironidae (■ – New
Caledonia)
Suborder Eupnoi (4 fam., ca. 1810 spp.)

Fam. Phalangiidae – Europe; North and South America; Asia Minor; China;
Central Asia; North, Central, and South Africa; Canary Islands; Madeira; Borneo
(56 gen., 486 spp.). Hesperopilio Shear (Australia, Chile) is nested in
Phalangioidea without familial attachment
Fam. Sclerosomatidae – Europe; Asia; Africa; South, Central, and North America;
Antilles; Australia; New Guinea; Solomon Island (153 gen., ca. 1300 spp.)
Fam. Neopilionidae (incl. Monoscutidae) – South Africa, Chile, Argentina, Brazil,
Australia, Tasmania, New Zealand, Auckland, Snares, Campbell Islands (20 gen,
64 spp.)
Fam. Protolophidae – the Western United States (eight spp.)
Suborder Dyspnoi (9 fam., 387 spp.)
Fam. Caddidae – South Africa, Eastern North America, Japan (two gen., nine rec.
spp.)
Fam. Acropsopilionidae – Australia, North America, South America, Japan, New
Zealand (3–4 gen., 14 spp., incl. Hesperopilio Shear)
Fam. Ischyropsalididae (incl. Ceratolasmatidae) – Europe (incl. Abkhazia,
Crimea), Algeria, Tajikistan, USA (Oregon, Washington, California, Idaho),
Mexico, Honduras (3 gen., 45 spp.) (Fig. 6.3)
Fig. 6.3 Ischyropsalis hellwigii (Panzer) (courtesy of Ula N., from A. Kury)
Fam. Sabaconidae – Japan, France, Spain, Alaska, Nepal, China (Sichuan), Siberia,
USA (Alabama, Kentucky, Oregon, California) (1 gen., 35 spp.)
Fam. Taracidae – USA, Canada, Far East of Russia (3 gen., 15 spp.)
Fam. Dicranolasmatidae – Mediterranean, Asia Minor, Hungary, Russia
(Krasnodar), Iraq, Western North Africa (1 gen., 16 spp.)
Fam. Trogulidae – Western and Southern Europe, Western North Africa, the Levant,
Caucasus, Northern Iran (4 gen., 56 spp.), introduced in the USA (Fig. 6.4)
Fam. Nemastomatidae – USA (Washington, California), Mexico, Coronado Island,
Thailand, Japan, Bulgaria, Abkhazia, Crimea, Europe, Algeria, Tajikistan, Asia
Minor, Crete (21 gen., 173 spp.) (Fig. 6.5)
Fam. Nipponopsalididae – Japan, Korea (one gen., three spp.)
Suborder Laniatores
According to the Catalogue of Kury (2003), the suborder Laniatores counts 1317
genera and 3901 spp., including 746 genera and 2372 spp. in the New World. Kury
(2013) defines the number of Laniatores at 4183 spp., Kury (2016) at 4212 spp., and
Kury (status April 2017) at 4248 spp. The Laniatores in Europe are very few – the
6.8 Opiliones 155
Fig. 6.4 Trogulus nepaeformis (Scopoli) (courtesy Ivo Antušek, from A. Kury)
Fig. 6.5 Nemastoma bimaculatum (Fabricius) (courtesy of Jonathan Michaelson, from A. Kury)
families Travuniidae, Cladonychiidae, and Phalangodidae (Roewer 1935; Rambla

1978; Kury and Mendes 2007, and others).
Fam. Nippononychidae – Japan, Korea (four genera, 10 spp.)
Fam. Paranonychidae – USA, Canada, Japan, Korea (7 genera, 26 spp.)
Fam. Travuniidae – Serbia, Croatia, S. France, Sardinia, USA (Washington), Japan
(13 gen., 41 spp.)
Fam. Synthetonychiidae – New Zealand (1 genus, 14 spp.)
Fam. Triaenonychidae – North and South America, Australia, Tasmania, New
Zealand, New Caledonia, Japan, Korea, Madagascar, South Africa (107 gen.,
477 spp.)
Fam. Assamiidae – Africa, Southern Asia; subfam. Dampetrinae to Indonesia, New
Guinea, and Australia (267 gen., 483 spp.)
Fam. Pyramidopidae – West Africa, Tanzania, Canary Island, Belize (15 gen., 47
spp.)
Fam. Epedanidae (incl. Dibuninae, Acrobuninae, Sarasiniciinae, Sarasinellinae) –
Philippines, New Guinea, Thailand, Sulawesi, Borneo, Sumatra, Japan, Malaysia,
India, Taiwan, China, Vietnam (70 gen., 174 spp.)
Fam. Petrobunidae – Southeast Asia (two gen., five spp.)
Fam. Podoctidae (incl. Erecananinae and Ibaloniinae) – Africa, Madagascar,
Reunion, Japan, Palau, Taiwan, Mauritius, Seychelles, Indonesia, Malaysia, New
Guinea, Cuba, India, Philippines, Vanuatu (54 gen., 132 spp.)
Fam. Sandokanidae [Oncopodidae, praeoccup.] – Philippines, Sumatra, Borneo,
Burma, Nepal, India, Malaysia, Singapore, Thailand (6 gen., 72 spp.)
Fam. Tithaeidae – Southeast Asia (6 gen., 39 spp.)
Fam. Agoristenidae – Antilles, South America (27 gen., 76 spp.)
Fam. Cosmetidae – From Argentina to Southern USA (125 gen., 719 spp.) (Fig. 6.6)
Fam. Cranaidae – South America, Trinidad, Panama, Costa Rica (75 gen., 165
spp.)
Fam. Gonyleptidae – From Patagonia to Costa Rica, Guatemala, and Falklands
(272 gen., 829 spp.) (Fig. 6.7)
Fam. Manaosbiidae – South America, Panama, Costa Rica, Trinidad, Windward
Islands (27 gen., 47 spp.)
Fam. Stygnidae – Northern South America, Lesser Antilles (30 gen., 104 spp.)
Fam. Stygnopsidae (Isaeinae, Troglostygnopsinae) – Mexico, Guatemala, Belize,
El Salvador, USA (Texas) (9 gen., 40 spp.)
Fam. Phalangodidae – USA, Japan, Mediterranean (21 gen., 116 spp.)
6.8 Opiliones 157
Fig. 6.6 Sibambea sp. (Cosmetidae, Ecuador) (courtesy of A. Anker, from A. Kury)
Fig. 6.7 Bourguyia bocaina Yamaguti et Pinto-da-Rocha (courtesy J. Wiley & Sons, from
A. Kury)
Fig. 6.8 Biantes sp. (Biantidae, Singapore) (flickr, from A. Kury)
Fam. Biantidae – Madagascar, Comoros, tropical Africa, Seychelles, Nepal, India,

Burma, Bioko, South Africa, Sumatra, Malaya, Socotra, West Indies, South
America (33 gen., 134 spp.) (Fig. 6.8)
Fam. Escadabiidae – Brazil (six gen., eight spp.)
Fam. Kimulidae (Minuidae) – South America (Brazil, Venezuela, Peru), Antilles
(Cuba, Puerto Rico, Dominican Republic)(10 gen., 36 spp.)
Fam. Samoidae – Cuba, Venezuela, Trinidad, Jamaica, Mexico, Haiti, Guatemala,
Fiji, Seychelles, Australia (New South Wales), Samoa, Leeward Islands, Costa
Rica, Cameroon, Panama, Mozambique, Indonesia (24 gen., 47 spp.)
Fam. Stygnommatidae – Mexico, Belize, Ecuador, Jamaica, Puerto Rico,
Colombia, Costa Rica, Panama, Venezuela, ? Malaya (1 gen., 33 spp.) (Fig. 6.9)
Fam. Fissiphalliidae – Colombia (one gen., seven spp.)
Fam. Guasiniidae – Venezuela, Brazil (two gen., three spp.)
Fam. Icaleptidae – Colombia, Ecuador (two gen., two spp.)
Fam. Zalmoxidae (Stygnoleptinae) – Mexico, Costa Rica, El Salvador, Brazil,
Guyana, French Guiana, Bolivia, Peru, Venezuela, Argentina, Cuba, Panama,
Jamaica, Belize, Trinidad, Ecuador, New Guinea, Solomon Islands, Fiji,
Mauritius, Madagascar, Seychelles, Bismarck Archipelago, Philippines, Java,
6.9 Solifugae (Solpugida, Solifugi) 159
Fig. 6.9 Zygobunus sp. (Stygnommatidae, Costa Rica) (courtesy A.P. González, from A. Kury)
Sulawesi, Moluccas, Australia, Marianas, New Caledonia, Marshall Islands,

Caroline Islands, Polynesia (66 gen., 209 spp.)
Fam. Cladonychiidae – Europe (Holoscotolemon Roewer), USA (17 spp.)
Fam. Briggsidae (Pentanychidae, praeoccup.) – USA (Oregon, Washington)(six
spp.)
6.9 Solifugae (Solpugida, Solifugi)
Map 6.6 Distribution. of Solifugae (courtesy of W. Savary)

Fig. 6.10 The order

counts 1113 spp. from 142
genera and 12 families
(Prendini 2011, Blick and
Kury 2011)
Fam. Galeodidae – Africa (North Africa, Somalia), Asia (Turkey, Iran, Israel,
Afghanistan, Central Asia, India, Yemen, Pakistan, Cyprus), Europe (Bulgaria,
Greece, Macedonia, Ukraine, Russia) (9 genera, 201 species) (Fig. 6.10)
Map 6.7 Distribution of fam. Galeodidae

Fam. Karschiidae – Southeast Europe (Greece), Asia (Central Asia, Arabian

Peninsula, Mongolia, Turkey, Iran), North Africa (4 gen., 40 spp.)
Fam. Daesiidae – Europe (Spain, Greece), Central Asia, the Middle East, Africa,
and South America (29 gen., 191 spp.)
Map 6.8 Distribution of fam. Daesiidae

Fam. Solpugidae – Africa, Asia (Iraq, Turkey, Saudi Arabia, etc.) (17 gen, 196 spp.)
Map 6.9 Distribution of fam. Solpugidae

Fam. Ceromidae – Southern Africa (3 gen., 20 spp.)
Map 6.10 Distribution of fam. Ceromidae

Fam. Melanoblossiidae – Southeast Asia, Southern Africa (6 gen., 16 [31] spp.)

Fam. Mummuciidae – South America (10 gen., 20 spp.)
Map 6.11 Distribution of fam. Mummuciidae

Fam. Gylippidae – Central Asia, Turkey, the Near East, and Southern Africa (five
genera, 27 species)
Fam. Eremobatidae – USA, Mexico (8 genera, 192 species)
Map 6.12 Distribution of fam. Gylippidae

Map 6.13 Distribution of fam. Eremobatidae

Fam. Ammotrechidae – USA, Mexico, Central and South America, West Indies
(21 genera, 88 species)
Map 6.14 Distribution of fam. Ammotrechidae

Fam. Hexisopodidae – Southern Africa (2 genera, 23 species)
Map 6.15 Distribution of fam. Hexisopodidae

Fam. Rhagodidae – Northeastern Africa, Southwestern Asia, the Near East (27
genera, 99 species)
Map 6.16 Distribution of fam. Rhagodidae
6.10 Araneae
The order counts 42055 spp. (Platnick 2011) or 46464 spp. (World Spider Catalog,
February 2017) or 47052 (end November 2017) or 47553 (WSC June 2018). The
families of spiders used to be 114, but the efforts of some molecular specialists they
became 117 (WCS, June 2018). The figures for the number of spider species change
very rapidly for the different families.
Maps 6.17 – Distributions of spiders
Suborder Mesothelae – Southeast Asia, China, Japan (97 species)
Fam. Liphistiidae – Southeast Asia, China, Japan (97 species)
6.10 Araneae 171
Map from Xu et al. (2015)
Map 6.17a Distribution of fam. Liphistiidae

Suborder Orthothelae
Infraorder Mygalomorphae
Fam. Actinopodidae – South America, Panama, Australia (48 spp.)
Map 6.17b Distribution of Actinopodidae
Fam. Antrodiaetidae – USA (33 spp.), Japan (two spp.) (35 spp.)
Fam. Atypidae – Europe, Asia, Africa, USA, Mexico (52 spp.)
Fam. Barychelidae – Australia, New Caledonia, Madagascar, Seychelles,
Mauritius, tropical Africa, Southeast Asia, New Guinea, Sri Lanka, Socotra,
Oceania, Fiji, South America, India, Mexico, Belize, Caribbean (296 spp.)
Subfam. Barychelinae – Socotra, Australia, New Caledonia, Africa, Madagascar,
Seychelles, Mauritius, South America
Subfam. Sasoninae – Brazil, Venezuela, Australasia
Subfam. Trichopelmatinae – Caribbean, Central to South America
Fam. Ctenizidae – Mediterranean (Greece, Turkey,) Japan, Taiwan, China,
Thailand, Central Asia, Southern Africa, USA, Mexico, Guatemala, Costa Rica,
Australian region, New Guinea (135 spp.)
Subfam. Ctenizinae – USA to Guatemala and Costa Rica, Thailand, China,
Taiwan, Europe, Central Asia, Australian region, South Africa
Subfam. Pachylomerinae – Australian region, America, Mediterranean, Japan,
Taiwan, Costa Rica, USA
Fam. Cyrtaucheniidae – Africa, Mediterranean, Australia, Central and South
America, USA, Mexico, Turkmenistan, Tajikistan, Afghanistan, Thailand (107
spp.)
6.10 Araneae 173
Subfam. Aporoptychinae – Africa, Australia, Central and South America

Subfam. Cyrtaucheniinae – Mediterranean
Fam. Dipluridae – South America, Cuba, Africa, South Africa, Australia, Oceania,
Southern USA to Costa Rica, Taiwan, Thailand, Central Asia, New Caledonia,
Caribbean, India, Madagascar (188 spp.)
Subfam. Diplurinae – South America, Cuba
Subfam. Euagrinae – Africa, South Africa, Australia, Southern USA to Costa
Rica, Taiwan, Central Asia, New Caledonia
Subfam. Ischnothelinae – Mexico to Argentina, Peru, Caribbean, India, Africa,
Madagascar
Subfam. Masteriinae – Caribbean, Central and South America, Oceania, Australia
Fam. Euctenizidae – USA, Mexico, Guadeloupe (76 spp., up to 2012 subfam. of
Cyrtaucheniidae)
Fam. Hexathelidae – New Zealand, Australia, Tasmania, Chile (Mediothele), North
and Central Africa, Europe, Asia (Macrothele) (113 spp.)
Subfam. Hexathelinae – Australia, New Zealand, Chile, Argentina
Subfam. Macrothelinae – Africa, Europe, Asia, New Zealand
Map 6.17c Distribution of fam. Hexathelidae
Fam. Idiopidae – Australia, New Zealand, Central and South America, Madagascar,
India, Sri Lanka, Seychelles, Southern Africa, Africa, South Asia, Middle East,
Thailand, Canary Islands (325 spp.)
Subfam. Arbanitinae – Australia, New Zealand
Subfam. Genysinae – Central and South America, Madagascar, India, Sri Lanka,
Seychelles
Subfam. Idiopinae – Southern Africa, India, Sri Lanka, South America, Africa,
South Asia, Middle East, Canary Islands
Fam. Mecicobothriidae – USA, Mexico, Brazil, Argentina, Uruguay (nine spp.)
Map 6.17d Distribution of fam. Mecicobothriidae

6.10 Araneae 175
Fam. Microstigmatidae – Panama, Brazil, Argentina, Uruguay, Colombia,

Venezuela, Ecuador, South Africa (17 spp.)
Subfam. Micromygalinae – Panama (one spp.)
Subfam. Microstigmatinae – Brazil, Argentina, Uruguay, Colombia, Venezuela,
Ecuador, South Africa (15 spp.)
Map 6.17e Distribution of fam. Microstigmatidae
Fam. Migidae – Madagascar, tropical Africa, Chile, Argentina, Australia, New

Zealand (97 spp.)
Subfam. Calathotarsinae – Chile, Argentina, Australia
Subfam. Miginae – Chile, New Zealand, Australia, Africa
Subfam. Paramiginae – Madagascar, Africa
Fam. Nemesiidae – South America, USA, Mexico, Burma, India, China, Europe
(Majorca, Spain, Portugal), Russia, Australia, New Zealand, Madagascar, South
Africa, Ryukyu Islands, Southeast Asia (404 spp.)
Fam. Paratropididae – St. Vincent, Panama, Peru, Brazil, Venezuela (11 spp.)
Subfam. Glabropelmatinae – Panama, Venezuela
Subfam. Paratropidinae – St. Vincent, Peru, Brazil, Venezuela
Fam. Theraphosidae (incl. Metinae) – South and Central America, Caribbean,
Australia, South and Southeast Asia, New Guinea, Africa, USA, Mexico, Asia
Minor, Europe (962 spp.)
Suborder Araneomorphae
Fam. Agelenidae – Mediterranean, Europe to Central Asia, USA to Chile, New
Zealand, Socotra, Himalaya, Cuba, Bahamas, Cyprus, China; Tegenaria, world-
wide (1277 spp.)
Fam. Amaurobiidae – Cosmopolitic (286 spp.)
Fam. Ammoxenidae – Australia, Tasmania, Botswana, South Africa, Namibia,
Zimbabwe (18 spp.)
Fam. Amphinectidae – Western South America, Australia, Tasmania, New Zealand,
introduced in the USA (159 spp.)
Map 6.17f Distribution of fam. Amphinectidae

6.10 Araneae 177
Fam. Anapidae (incl. Micropholcommatinae and Holarchaeidae) – Central to

South America, USA, Mexico, New Caledonia, Australia, Tasmania, New
Zealand, New Guinea, Europe, China, Taiwan, Nepal, Korea, Japan, South Africa
(223 spp.)
Map 6.17g Distribution of subfam. Micropholcommatinae
Fam. Anyphaenidae – Australia, New Zealand, India, Polynesia, Europe, Turkey,

Kurile Island, China, USA, Canada, Mexico, Algeria, S. Africa, Central and South
America, Caribbean, Galapagos (558 spp.)
Fam. Araneidae (incl. Nephilinae – pantropical, 75 spp.) – worldwide (3122 spp.)
Fam. Archaeidae – Australia, South Africa, Madagascar (71 spp.)
Fam. Arkyidae – Australia (37 spp.)
Fam. Austrochilidae – Chile, Argentina, Tasmania (10 spp.)
Subfamily Austrochilinae Zapfe, 1955 – Chile, Argentina (nine spp.)
Subfamily Hickmaniinae 1967 (as family Hickmaniidae) (Tasmania, cave)
Fam. Caponiidae – USA, Mexico, Caribbean, Central and South America, South
Africa (110 spp.)
Fam. Chummidae – South Africa (two spp.)
Fam. Cithaeronidae – East Africa, India, Ivory Coast, Ethiopia, Greece, Libya to
Malaysia, Australia, and Brazil (eight spp.)
Fam. Clubionidae – Cosmopolitic (615 spp.)
Fam. Corinnidae – Worldwide (778 spp.)
Fam. Ctenidae – Cosmopolitic (514 spp.)
Fam. Cyatholipidae – Africa, Madagascar, New Zealand, Australia, and one spe-
cies (Pokennips dentipes) in Jamaica (58 spp.)
Fam. Cybaeidae (incl. Argyronetidae) – America, Europe, Japan, Korea, China,
Turkmenistan, Tajikistan, Uzbekistan, Sumatra, Venezuela, Colombia;
Argyroneta, Palearctic (188 spp.)
Fam. Cycloctenidae – New Zealand, Australia, Indonesia (Java) (36 spp.)
Fam. Deinopidae – All continents, except Europe and Antarctica (61 spp.)
Fam. Desidae – Australia, Tasmania, New Zealand, New Caledonia, Chile, USA,
Oceania, Korea, Japan, India, Africa, Galapagos, Paraguay (175 spp.)
Fam. Dictynidae – Europe, China, Mongolia, India, Algeria, Kazakhstan, Canary
Islands, New Zealand, Australia, Sri Lanka, South Africa, USA, Canada, Mexico,
Kyrgyzstan, West Indies, Hawaii, South America, St. Helena, Andaman Islands,
Galapagos Island, New Caledonia, etc. (616 spp.)
Fam. Diguetidae – USA, Mexico, Argentina, Brazil, Peru, Chile (15 spp.)
Fam. Drymusidae – South Africa, Cuba, Brazil, Argentina, Costa Rica, St. Vincent,
Hispaniola (16 spp.)
Fam. Dysderidae – Southern Europe, Sardinia, Corsica, Crete, Syria, Azerbaijan,
Turkmenistan, Israel, Iran, Algeria, Turkey; Dysdera, worldwide (542 spp.)
Subfam. Dysderinae – Europe, Crete, Syria, Azerbaijan, Turkmenistan, Israel,
Algeria, Turkey; Dysdera, worldwide
Subfam. Rhodinae – Mediterranean
Fam. Eresidae – Africa, Madagascar, Brazil, Eurasia (98 spp.)
Subfam. Eresinae – Africa, Madagascar, Brazil, Eurasia
Subfam. Penestominae – South Africa
Fam. Eutichuridae – Holarctic, Africa, South America to Costa Rica, USA,
Mexico, India, Lord Howe Island, Madagascar, Reunion, Samoa, Cape Verde,
Thailand, Indonesia, St. Helena (344 spp.). Formerly (up to 2014) subfamily of
Miturgidae, fam. status by Ramirez (2014)
Fam. Filistatidae –Africa, Madagascar, USA, Mexico, Guatemala, Cuba, Chile,
Brazil, Argentina, Colombia, Galapagos, Australia, New Guinea, Mediterranean
to India, Iran, Israel, Tajikistan, Uzbekistan, etc. (152 spp.)
Subfam. Filistatinae – Mediterranean to India, Iran, Israel, Tajikistan, Uzbekistan,
Mexico
Subfam. Prithinae – Africa, Madagascar, USA, Guatemala, Cuba, Chile, Brazil,
Argentina, Colombia, Galapagos, New Guinea, etc.
Fam. Gallieniellidae – Madagascar, Comoro Island, Kenya, South Africa, Australia,
Argentina (55 spp.)
6.10 Araneae 179
Subfam. Gallienellinae – Madagascar, Comoro Island, Kenya, South Africa

Subfam. Meedoinae – Australia (five gen.), Argentina (one gen.)
Fam. Gnaphosidae – Worldwide (2200 spp.)
Fam. Gradungulidae – New Zealand, Australia (A.C.T., Queensland, New South
Wales, Victoria) (16 spp.)
Fam. Hahniidae – Oceania, Australia, Tasmania, New Zealand, India, Philippines,
Sumatra, Sri Lanka, Europe, Kazakhstan, Kyrgyzstan, Canada to Venezuela,
Brazil, Mexico, USA, Alaska, etc. (252 spp.)
Fam. Hersiliidae – Africa, Australasia, USA, Caribbean, South America, India, Sri
Lanka, Spain, Portugal, Algeria, Central Asia, Australia, Borneo (181 spp.)
Fam. Homalonychidae – USA, Mexico, ?India (? three spp.)
Fam. Huttoniidae –New Zealand (one sp.)
Fam. Hypochilidae – China, USA (12 spp.)
Fam. Lamponidae – Australia, New Zealand, New Guinea, New Caledonia (genus
Centrocalia) (192 spp.)
Subfam. Centrothelinae – Australia, New Caledonia
Subfam. Lamponinae – Australia, New Zealand, New Guinea
Subfam. Pseudolamponinae – Australia
Fam. Leptonetidae – USA, Mexico, Panama, Mediterranean, Portugal, Algeria,
Japan, China (300 spp.)
Fam. Linyphiidae – Worldwide (4547 spp.)
Fam. Liocranidae – Cosmopolitic (272 spp.)
Fam. Lycosidae – Cosmopolitic (2415 spp.)
Fam. Malkaridae – Australia, Chile, Argentina; incl. Pararchaeidae from Australia,
Tasmania, and New Zealand, 34 spp. (46 spp.)
Map 6.17h Distribution of fam. Malkaridae

Fam. Mecysmaucheniidae – Chile, Argentina, Falkland Island, Juan Fernandez

Islands, New Zealand (25 spp.)
Subfam. Mecysmaucheniinae Simon, 1895 – Chile, Argentina, Falkland Island,
Juan Fernandez Islands, New Zealand (22 spp.)
Subfam. Zearchaeinae Forster et Platnick, 1984 – Chile, Argentina, New Zealand
(three spp.)
Fam. Megadictynidae – New Zealand (two spp., formerly in Nicodamidae, family
status reestablished by Dimitrov et al. 2017)
Map 6.17i Distribution of fam. Mecysmaucheniidae
Fam. Mimetidae – Palearctic, Africa, Principe, Central and South America, USA,
Asia, Australia; Ero and Mimetus, worldwide (152 spp.)
Subfam. Gelaninae – Central and South America (18 spp.)
Subfam. Melaenosiinae – Principe, India (two spp.)
Subfam. Mimetinae – Worldwide
Fam. Miturgidae – Thailand, Indonesia, South America to Costa Rica, Holarctic,
Africa, Australia, St. Helena, New Zealand, New Guinea, Sicily, Israel, Pakistan
(157 spp.)
Subfam. Miturginae – Australia, New Guinea, Sicily, Israel, Africa, South
America, USA to Argentina
6.10 Araneae 181
Subfam. Systariinae – USA to Mexico, Costa Rica, Brazil, Pakistan, Southeast

Asia
Subfam. Diaprograptinae – Timor, Australia, New Zealand
Subfam. Zorinae – Central and South America, Australia, Tasmania, New
Zealand, Israel, Guatemala; Zora, Palearctic (73 spp.)
Fam. Mysmenidae – Europe, USA, Mexico, Cuba, Jamaica, Central and South
America, tropical Africa, Algeria, South Africa, Galapagos, China, Nepal, India,
Sri Lanka, New Guinea, Samoa, Hawaii, Philippines, Seychelles, Fiji, New
Caledonia (137 spp.)
Fam. Nesticidae – Europe, North America, Mexico, Japan, Jamaica, Cuba, Tibet,
China, Laos, Philippines, Sri Lanka, Indonesia, Madagascar, Russia, Hawaii,
Fiji, Nepal, New Guinea, Seychelles, South America, Turkey (278 spp.)
Fam. Nicodamidae – Australia, New Guinea (27 spp.)
Fam. Ochyroceratidae – South Asia, China, Seychelles, Angola, Cuba, Mexico to
Peru, Pacific Islands, New Guinea (191 spp.)
Fam. Oecobiidae – Mediterranean, Africa, South Africa, Asia, USA; Oecobius is
cosmopolitan (111 spp.)
Fam. Oonopidae – Central to South America, Mexico, USA to Argentina, Cuba,
Antilles, Yemen, Seychelles, Bhutan, Nepal, China, Africa, Philippines,
Southeast Asia, Australia, Tasmania, New Zealand, New Caledonia, Hawaii,
Socotra, South Africa, Angola, East Africa, Europe, Sierra Leone, Congo,
Algeria, Egypt, St. Helena, Sri Lanka (1777 spp.)
Subfam. Gamasomorphinae – Cuba, Yemen, Seychelles, Bhutan, Nepal, China,
Central to South America, Africa, Philippines, Southeast Asia, Australia, New
Zealand, Mexico, USA to Argentina, Hawaii, Socotra, Seychelles, Antilles
Subfam. Oonopinae – South Africa, Angola, East Africa, Europe, Panama to
Argentina, Seychelles, Hawaii, Sierra Leone, Congo, New Caledonia, Mexico,
Cuba, Tasmania, Algeria, Egypt, St. Helena, Sri Lanka
Fam. Orsolobidae – South Africa, Australia, New Zealand, South Africa, Chile,
Australia, Tasmania, Brazil, Chile, Argentina, Falkland Islands (188 spp.)
Fam. Oxyopidae – Cosmopolitic (457 spp.)
Fam. Palpimanidae – Africa, Socotra, South America, Cuba, South Asia,
Uzbekistan, Mediterranean, Seychelles, Sri Lanka (144 spp.)
Subfam. Chediminae – Africa, Socotra, South Asia, Seychelles, Sri Lanka
Subfam. Otiothopinae – South America, Cuba, Africa
Subfam. Palpimaninae – Mediterranean, Africa, South America, India,
Uzbekistan
Fam. Penestomidae – South Africa, Lesotho (nine spp.)

Fam. Periegopidae – Australia (Queensland), New Zealand (three spp.)
Fam. Philodromidae – Cosmopolitic (539 spp.)
Subfam. Pedinopisthinae – Hawaii
Subfam. Philodrominae – Cosmopolitic
Subfam. Pselloninae – India, New Guinea
Fam. Pholcidae – Worldwide (c. 1611 spp.)
Fam. Phrurolithidae – China, India, Indochina, Lord Howe Island, USA, Mexico,
Canada, Europe, Madeira, Japan, North Africa, Madagascar, Seychelles, Central
Asia, Russia, Taiwan, Philippines, tropical Africa (218 spp.)
Fam. Physoglenidae – (72 spp.)
Fam. Phyxelididae –Madagascar, Kenya, South Africa, Cyprus, Turkey, Sumatra,
Borneo (64 spp.)
Fam. Pimoidae – Europe, North America, Russia, China, Japan, Himalaya (40
spp.)
Fam. Pisauridae – Cosmopolitic (335 spp.)
Fam. Plectreuridae – USA, Mexico, Cuba (31 species)
Fam. Prodidomidae – Africa, South America, Cuba, Australia, USA, Mexico,
Panama, Galapagos, Bahamas, Japan, Spain, Algeria, Canary Islands (309 spp.)
Fam. Psechridae – China, India, Nepal, Sri Lanka, South East Asia, Nicobar
Islands, Philippines, Australia (61 spp.)
Fam. Salticidae – Worldwide (5989 spp.)
Fam. Scytodidae – Worldwide, except of the Far North (239 spp.)
Fam. Segestriidae – America, Africa, Asia, Australia, New Zealand (124 spp.)
Fam. Selenopidae – America, Asia, Africa, Mediterranean, Madagascar, Reunion,
Comoro Islands (257 spp.)
Fam. Senoculidae – South and Central America, Mexico (31 spp.)
Fam. Sicariidae – Worldwide, except of the Far North (149 spp.)
Fam. Sparassidae – Worldwide, except of the Far North (1215 spp.)
Fam. Stenochilidae – Thailand, Malaysia, Singapore, Burma, Philippines, Bali,
New Guinea, Fiji, Borneo, Vietnam, India (13 spp.)
Fam. Stiphidiidae – Australia, Tasmania, New Zealand, New Guinea, Madagascar,
Mauritius, New Caledonia (135 spp.)
Fam. Symphytognathidae – in the tropics of Central and South America and the
Australian region (with Oceania), with three species (Anapistula benoiti, A.
caecula, Symphytognatha imbulunga) found in Africa and one (Anapistula
ishikawai) in Japan. Anapistula jerai occurs in Southeast Asia (71 spp.)
Fam. Synaphridae – Canary Island, Southern Europe, Egypt, Turkmenistan, Spain,
three spp. on Madagascar (13 spp.)
Fam. Synotaxidae – Australia, Tasmania, New Zealand, South America (11 spp.)
Subfam. Pahorinae – endemic to New Zealand
Subfam. Synotaxinae – Australia, South America (Chile, Brazil, Colombia, etc.)
6.10 Araneae 183
Map 6.17j Distribution of fam. Synotaxidae
Fam. Telemidae – Sumatra, Malaysia, China, Vietnam, Japan, France, Spain,

Tanzania, Cameroon, South Africa, Seychelles, New Caledonia, New Guinea,
Guatemala, USA (Oregon, Alaska,) (69 spp.)
Fam. Tetrablemmidae – Southeast Asia, Africa, Borneo, Central America, Brazil,
Colombia, Panama, India, Indonesia (Sulawesi), Samoa, Madagascar, Sri Lanka,
Caroline Islands, Seychelles (166 spp.)
Fam. Tetragnathidae – Cosmopolitic (994 spp.)
Fam. Theridiidae – Cosmopolitic (2400 spp.)
Fam. Theridiosomatidae – Cosmopolitic (111 spp.)
Fam. Thomisidae – Worldwide (2154 spp.)
Fam. Titanoecidae – India, Sri Lanka to China, New Guinea, Marquesas Islands,
Europe, Mexico, South America; Titanoeca, Holarctic (54 spp.)
Fam. Trachelidae – Mexico, etc. (208 spp.)
Fam. Trechaleidae – South and Central America, Mexico, USA, Japan (Shinobius
Yaginuma) (120 spp.)
Fam. Trochanteriidae – Australia, Chile, Sulawesi, Christmas Island, China, India,
Korea, Japan, Africa, Madagascar, South America, New Caledonia (153 spp.)
Fam. Trogloraptoridae – Caves in Oregon and California (one sp.)
Fam. Udubidae – tropical Africa, Madagascar (2 end. genera) (12 spp.)
Fam. Uloboridae – USA, Central and South America, Philippines, Samoa, Fiji,
Vanuatu, New Guinea, Russia, Central Asia to Japan, Mediterranean to
Azerbaijan, Oceania, Australia, New Zealand; Uloborus is spread worldwide,
and Zosis is pantropical (283 spp.)
Fam. Viridasiidae – Madagascar, Brazil (nine spp., until 2015 part of Ctenidae)
Fam. Zodariidae – South America, Mexico, Central America, Australia, New

Zealand, New Guinea, Southeast Asia, Europe (1126 spp.)
Fam. Zoropsidae (incl. Tengellidae and part of Zorocratidae) – Australia, New
Zealand, Sri Lanka, China, Korea, Japan, Cyprus, Mediterranean, North and
South Africa, Madagascar (178 spp.)
Map 6.17k Distribution of subfam. Tengellidae
Acari Traditionally, this numerous group has been considered as containing three
orders: Opilioacarida (41 rec. spp., Beron 2014 and suppl.), Acariformes, and
Parasitiformes (both containing more than 48 200 described species, Harvey 2002,
actually more than 52300 spp. by 2017). The newest general paper on Acari (2009.
Manual of Acarology, Third Edition) provides another subdivision. Following the
recent subdivision of Parasitiformes in four orders (Lindquist et al. 2009),
Holothyrida, Ixodida, and Mesostigmata are treated as separate orders. We prefer to
treat here Opilioacarida as an order.
How Many Are the Acari? For most groups of Acari, our knowledge is far from
the existing taxa in the nature. A very instructive table has been drown by Walter and
Proctor (1998). According to it (we are listing only the species, with minor correc-
tions), less than one tenth of the minimal number of living Acari have been recorded
and described so far.
6.11 Opilioacarida 185
Group Species Minimum (guess) Maximum (guess)

Opilioacarida 41 85 170
Holothyrida 29 160 320
Ixodida 880 1000 1200
Mesostigmata 12,512 97,520 200,500
Endeostigmata 120 1200 2400
Oribatida 11,000 23,000 110,000
Astigmata 4500 90,000 180,000
Trombidiformes 17,050 327,250 637,500
Of course, the calculations of our colleagues are approximate, the real figures for
the extant Acari species will be different, but even this approximation shows how far
we are from the real picture of the number of species, let alone of the distribution of
mites. For most countries there are no checklists and very few taxonomists describe
Acari. We can see clearly that when in one country appear acarologists, the list of
mites becomes much longer. Good example for this is the flood of new information
from countries and regions like Turkey, Iran, Brazil, New Zealand, South Africa,
Crimea, and others. If someday mites will be described using DNA, these figures
will be different, if you judge from the recent multiplication of number of larger
animals like Euscorpius (Scorpiones).
6.11 Opilioacarida
The first to draw attention on the existence of such peculiar mites was C. With,
describing the order, the only family, and the species Opilioacarus segmentatus
(With 1903) and O. italicus (With 1904) – still the only two species known in
Europe (southern parts of Italy and Greece). Shortly after him F. Silvestri (1905)
discovered opilioacarids also in South America. Redikorzev (1937) found the first
and only representative in Central Asia. Chamberlin and Mulaik (1942) not only
described the northernmost American species – Neocarus texanus – but also sepa-
rated it in a new genus, the second in the order. They also separated Opilioacarus
hexophthalmus Redikorzev (1937) in a new genus Paracarus. Followed André
(1947) and Naudo (1963) with descriptions of new species from tropical Africa and
of the fourth genus in the order – Panchaetes.
From 1966 to 1979, L. van der Hammen (1966, 1968, 1969, 1971, 1977) wrote
important papers (one of them together with I. Coineau), adding to the seven species
known prior to them another seven and the new genera Adenacarus, Phalangiacarus,
and Salfacarus.
Another three papers added new species to the system of Opilioacarida: Juvara-
Balş and Baltac (1977) described two new species (the first from caves) from Cuba.
More cave species have been described by Leclerc (1989) from Thailand, including
the new genera Siamacarus and Vanderhammenacarus. Lehtinen (1980) added one
more species from Venezuela. Harvey (1996) made a cladogram of opilioacarid
genera, suggesting that Paracarus and Siamacarus were the sister group to the other
genera.
Modern treatment of the order was enterprised by M. Vásquez and H. Klompen
(2002, 2004, 2009, 2010). They described 13 more species from the Americas and
Madagascar and the new genus Caribeacarus from Central and South America and
left the initial genus Opilioacarus only for the two species O. segmentatus and O.
italicus from Greece, Italy, and Algeria (more details about the distribution in
Brignoli 1967, Beron 1990, and Thaler and Knoflach 2002). Neocarus Chamberlin
et Mulaik became the most numerous genus in the order with 15 species. Harvey
(2002) listed 9 genera and 20 species in Opilioacariformes, since 4 more genera and
17 new species have been described. Amazonacarus setosus and A. paraensis were
described by Vásquez, Araújo, and Feres (2014) from Brazil; also from Brazil
Bernardi et al. (2011, 2012, 2013) described more Opilioacarida. Das and Bastawade
(2007) described Indiacarus pratyushi, new genus and species from India. Finally,
two fossil species (Paracarus pristinus Dunlop, Wunderlich et Poinar, Jr. 2004 and
?Opilioacarus) have been described from Northern Europe (Baltic amber) and the
order was mentioned (without more details) from Australia (Walter and Proctor
1998).
There is still uncertainty concerning the rank and the place in the system of these
strange Opiliones-like mites, inhabiting warmer countries. In the most recent treat-
ment of Kranz et al. (2008), Walter and Harvey use the name Opilioacarida as an
order. We are following this practice (Beron 2014).
Ordo Opilioacarida (Opilioacariformes) – 13 gen., 41 rec. spp.
Fam. Opilioacaridae
Adenacarus Van der Hammen, 1966 – Yemen (one sp.)
Amazonacarus Vásquez, Araújo et Feres, 2014 – Brazil (two spp.)
Brasilacarus Vásquez, Araújo et Feres, 2015 – Brazil (one sp.)
Caribeacarus Vásquez et Klompen, 2009 – Cuba, Dominican Republic, Panama
(four spp.)
Indiacarus Das et Bastawade, 2007 – India (one sp.)
Neocarus Chamberlin et Mulaik, 1942 – USA, Mexico, Nicaragua, Venezuela,
Cuba (14 spp.)
Opilioacarus With, 1902 – Circum-Mediterranean (two spp. + one ? fossil,
Baltic amber)
Panchaetes Naudo, 1963 – Angola, Ivory Coast (two spp.)
Paracarus Chamberlin et Mulaik, 1942 – One sp. (Kazakhstan) and one from
Kaliningrad (fossil!)
Phalangiacarus Coineau et Van der Hammen, 1979 – Gabon (one sp.)
6.11 Opilioacarida 187
Salfacarus Van der Hammen, 1977 – South Africa, Madagascar, Tanzania (nine
spp.)
Siamacarus Leclerc, 1989 – Thailand (two spp.)
Vanderhammenacarus Leclerc, 1989 – Thailand (one sp.)
Map 6.18 Opilioacarida in the Old World

Adenacarus – Δ
Indiacarus – ◄
Opilioacarus – ▲
Panchaetes –
Paracarus – ♦
Phalangiacarus – ▼
Salfacarus – ■
Siamacarus – ►
Vanderhammenacarus –
Map 6.19 Opilioacarida in the Americas

Amazonacarus – ■
Brasilacarus – ▲
Caribeacarus – ▼
Neocarus – ◄
6.12 Trombidiformes 189
Superorder Acariformes The mites of superorder Acariformes are not the favorite
subject of zoogeographic analysis for various reasons. Part of the families includes
only permanent parasites on other animals, which depend on the distribution of
hosts and have no special zoogeography (mange mites, Myobiidae, Listrophoroidea,
endoparasitic acari). Other families consist of adult and nymphal stages which are
soil inhabiting predators and parasitic larvae (Erythraeidae, Trombiculidae,
Trombidiidae s.l.). Many families consist entirely of free living forms (Anystidae,
Caeculidae, Teneriffiidae, Bdellidae, numerous Oribatida). They could provide
some information of zoogeographic interest (Atyeo 1960; Beron 2016; Colloff and
Halliday 1998; Kepka 1968; Robaux 1969; Strandtmann 1974; Travé 1976;
Wallwork 1969, 1973, 1979; Hammer and Wallwork 1979; Vu Kuang Manh 2015).
As we published a world catalogue of Erythraeoidea and Calyptostomatoidea
(Beron 2008a), we shall try to analyze some peculiarities of the distribution of these
mites. We shall analyze also the distribution of some free living mites.
6.12 Trombidiformes
Prostigmata Several families are analyzed as examples.
Erythraeoidea The world catalogue of Erythraeoidea (Beron 2008a) and the sup-
plements (Beron 2017) show that, by October 2017, there are 918 spp. of the fami-
lies Smarididae (10 genera) and Erythraeidae (57 genera). Some particularities in
the distribution of the genera could be the knowledge of these (and many other)
mites is very incomplete and depends largely on the geography of specialists
(Southcott and others). Also, with Smarididae and Erythraeidae, there are two clas-
sifications (of adults and larvae), and they could change with the rearings of larvae.
For the time being, some genera (Smaris, Hirstiosoma) seem distributed worldwide
(more or less cosmopolitic). Other genera seem more confined to one area
(Sphaerotarsus, Surasmaris). Interesting was to find the American genus
Trichosmaris in Papua New Guinea, on top of Mt. Fugilil, 3100 m. No mistake
could be supposed as the mite was found and described by the present author and it
is a native species found in the center of the island, far from the sea or human
settlements.
Fam. Smarididae
Calorema Southcott, 1962 – Panama, Mexico
Fessonia von Heyden, 1826 – India, Australia, Burma, Mexico, USA, Nigeria,
Europe, Guatemala
Kraussiana Southcott, 1961 – South Africa, Malaysia, Caroline Islands
Neophanolophus China, 1976 – Malaysia, India
Pilosoma Southcott, 1961 – France (= Fessonia)
Smaris Latreille, 1796 – Australia, Mexico, Panama, Chile, Europe, Cyprus, ?

South Africa
Surasmaris Southcott, 1995 – Costa Rica
Clavismaris Southcott, 1963 – Costa Rica, Texas
Clipeosoma Southcott, 1948 – Malaysia, Europe, New Guinea
Hirstiosoma Womersley, 1934 – Europe, Mexico, South America, Australia,
Tasmania, New Zealand
Sphaerotarsus Womersley, 1936 – Australia
Trichosmaris Southcott, 1963 – New Guinea, Guatemala, Mexico, USA
Fam. Erythraeidae
Augustsonella Southcott, 1961 – Mexico, USA
Claverythraeus Trägårdh, 1937 – China
Collemboerythraeus Noei, Saboori et Hakimitabar, 2017 – Iran
Curteria Southcott, 1961 – Austria, Switzerland, Greece, Germany, Holland,
Poland, Canary Island, Algeria,
Eatoniana Cambridge, 1898 (= Abalakeus Southcott (1994), China, Niger, Iran,
Spain) – Algeria, Egypt, Greece, Jersey, Spain, Switzerland, Armenia,
Transcaspia, Tunisia, South Africa, ?Argentina
Erythraeus Latreille, 1806 – Europe, Iceland, Israel, Iran, Turkey, Japan, China,
Taiwan, India, Cambodia, Galapagos, Bolivia, Peru, Egypt, Israel, South
Africa, Tanzania, Sudan, Ethiopia, USA, Australia
Erythraxus Southcott, 1961 – Mexico
Erythrellus Southcott, 1946 – Australia
Erythrites Southcott, 1961 – Australia, New Zealand
Erythroides Southcott, 1946 – Australia
Forania Southcott, 1961 – France,
Helladerythraeus Beron, 1988 – Greece
Kakamasia Lawrence, 1944 – South Africa
Lasioerythraeus Welbourn et Young, 1987 – Dominican Republic, USA, Bolivia
Makolia Saboori, Khaustov et Hakimitabar, 2009 – Crimea
Kamertonia Gabrys, 1999 – Poland
Neosmaris Hirst, 1926 – New Zealand
Opserythraeus Fain, 1996 – Rwanda
Paraphanolophus Smiley, 1968 – Mexico, Belize
Podosmaridia Trägårdh, 1937 – China
Rainbowia Southcott, 1961 – Australia
Ramsayella Zhang, 2000 – New Zealand
Taranakia Southcott, 1988 – New Zealand
Tepotztlana Hoffmann et Mendes, 1973 – Mexico
Leptus Latreille, 1796 – Cosmopolitic
Caeculisoma Berlese, 1888 – Congo, Nigeria, China, Bhutan, Sumatra, Australia,
South Africa, New Guinea, New Ireland, Marquesas Islands, New Zealand,
Brazil, Argentina, Paraguay
Callidosoma Womersley, 1936 – USA, Panama, Guatemala, Brazil, Argentina,
Australia, Taiwan, New Guinea, New Zealand, Dominican Republic
6.12 Trombidiformes 191
Carastrum Southcott, 1988 – Australia, Indonesia (Bali)

Dambullaeus Haitlinger, 2001 – Sri Lanka
Harpagella Southcott, 1996 – Australia
Momorangia Southcott, 1972 – New Zealand, Kenya, Brazil
Abrolophus Berlese, 1891 – Europe, Madagascar, China, Vietnam, Iran,
Indonesia, East Africa, USA, Ghana, Ethiopia, South Africa, India, Canary
Island, New Guinea, Australia, New Zealand, Japan, ?Argentina =
Hauptmannia Oudemans, 1910
Andrevella Southcott, 1961 – France, Iran
Cecidopus Karsch, 1879 – South Africa, Nigeria, Sri Lanka, Pakistan, India,
Congo
Charletonia Oudemans, 1910 – Madagascar, tropical Africa, South Africa,
Egypt, Europe, Iran, India, Sri Lanka, Indonesia, Malaya, Vietnam, Cambodia,
China, Japan, Australia, Tasmania, Canada, USA, Brazil, Canary Island
Grandjeanella Southcott, 1961 – Balearic Island, South Europe, Iran, Australia,
Iguatonia Haitlinger, 2004 – Brazil
Neoabrolophus Khot, 1965 – Tanzania, India, Malaysia
Pussardia Southcott, 1961 – Australia
Rudaemannia Haitlinger, 2000 – Europe, Turkey (= Balaustium)
Phanolophus Andre, 1927 – Algeria, Europe, Cyprus, East Africa
Balaustium von Heyden, 1826 – Cosmopolitic
Bursaustium Haitlinger, 2000 – Turkey, Iran
Fozustium Haitlinger, 2005 – Brazil
Italustium Haitlinger, 2000 – Italy
Lomeustium Haitlinger, 2006 – West Africa
Madinahustium Kamran et Alatawi, 2016 – Saudi Arabia
Moldoustium Haitlinger, 2007 – Moldova, Ukraine
Microsmarialla Khot, 1963 – India
Microsmaris Hirst, 1926 – Australia, New Zealand, India
Mypongia Southcott, 1961 – Australia
Neobalaustium Willmann, 1951 – Romania, Bulgaria
Pollux Southcott, 1961 – Australia, India
Wartookia Southcott, 1961 – Australia
Myrmicotrombium Womersley, 1934 – Australia, Greece (subg. Graecotrombium)
The known distribution is rather incomplete. When in one country there are spe-
cialists (Australia, Poland, Iran), the knowledge on the fauna is much closer to the
actual situation.
Conclusion: In the Erythraeoidea, there are no families or subfamilies endemic
to a particular continent or zoogeographical region. Endemism is developed only on
genus and species level.
Calyptostomatoidea
Fam. Calyptostomatidae
Calyptostoma Cambridge, 1875 – Europe, Asia, Africa, New Guinea, Australia
These unmistakable mites are obviously missing in the Americas.
Superfam. Anystoidea
Family Teneriffiidae
Austroteneriffia Womersley, 1935 – Australia, Tajikistan, Iran, Japan, Yemen
(incl. Socotra)
Himalteneriffia Schmölzer, 2002 – India (Ladakh)
Mesoteneriffia Irk, 1939 – Alps (Austria, Switzerland)
Mesoteneriffiola Schmölzer, 1956 – France
Neoteneriffiola Hirst, 1924 – Egypt, Namibia, China, USA, Mexico
Parateneriffia Thor, 1911 – Paraguay
Sinoteneriffia Yin, Bei et Li, 1995 – China
Teneriffia Thor, 1911 – Canary Island, Japan, Malaysia, Mexico
The family is spread on all continents, endemism only on genus level.
Fam. Anystidae
Anystis von Heyden, 1826
Autenriethia Oudemans, 1936
Barellea Oudemans, 1936
Mesoanystis Zacharda, 1985 (fossil)
Scharfenbergia Oudemans, 1936
Snartia Oudemans, 1936
Tencateia Oudemans, 1936
Walzia Oudemans, 1936
Chaussieria Oudemans, 1937
Erythracarus Berlese, 1903
Erythrocheylus Berlese, 1903
Namadia Meyer et Ueckermann, 1987
Paratarsotomus Kuznetsov, 1983
Siblyia Oudemans, 1936
Tarsolarcus Thor, 1912
Tarsotomus Berlese, 1882
Family Pseudocheylidae
Anoplocheylus Berlese, 1910 – Egypt, Iran, Italy, South Africa, Australia, Brazil,
Turkey, Crimea
Neocheylus Trägårdh, 1906 – Australia, South Africa, Uganda
Pseudocheylus Berlese, 1888 – USA, Brazil, Paraguay
Family Stigmocheylidae
Stigmocheylus Berlese, 1910 – USA, Italy, Egypt
Superfamily Caeculoidea
Family Caeculidae
Allocaeculus Franz, 1952 – France, Kazakhstan, Russia, Spain, Austria, Croatia,
Czech Republic, Macedonia, Namibia, South Africa, Morocco, Algeria,
6.13 Sarcoptiformes 193
Tunisia, Pakistan, Kenya, Nigeria, Chad, Mozambique, Sudan, Senegal,

Canary Island
Andocaeculus Coineau, 1974 – Argentina
Caeculus Dufour, 1832 – USA, Mexico, Puerto Rico, Senegal, Europe
Calocaeculus Coineau, 1974 – South Africa
Microcaeculus Franz, 1952 – Austria, Germany, Czech Republic, Italy, France,
Corsica, Greece, Portugal, Spain, Canary Island, Salvages Island, South
Africa, Argentina, Chile, Morocco, Algeria, Namibia, China, Korea, Australia
Neocaeculus Coineau, 1967 – Australia, Tasmania, Lord Howe Island, New
Zealand, Namibia
Procaeculus Jacot, 1936 – Trinidad, Dominican Republic, Hawaiian Island,
Poland (fossil), Mexico, USA, Guatemala
Adamystoidea
Family Adamystidae
Adamystis Cunliffe, 1957 – USA, Iran, Pakistan, Tajikistan, Uzbekistan,
Thailand, South Africa, France, Ukraine
Bovidromus Coineau, Theron et Fernandez, 2006 – South Africa
Rhinodromus Coineau, Theron et Fernandez, 2006 – South Africa
Saxidromus Coineau, 1974 – Mexico, Cuba, South Africa, France
So far the family is known from Europe, Asia, Mexico, Cuba, and South Africa,
but these mites are rarely studied and could be found in other places as well. The
same conclusion is valid also for Pomerantziidae and other small families.
Pomerantzioidea
Family Pomerantziidae
Apomerantzia Quing-Hai Fan et Yan Chen, 2005 – USA
Pomerantzia Baker, 1949 – USA, China, Philippines
6.13 Sarcoptiformes
Acaridida Although the parasitic mites of this suborder follow the distribution of
their hosts, it may be useful to mention a few details of the distribution even of per-
manent parasites as Listrophoroidea or Psoroptoidea. However, it may be noticed
that the large family Atopomelidae is almost absent in Europe and very numerous in
the tropical countries (Beron 2011).
Oribatida The huge group (suborder or part of suborder) of Oribatida is composed

by ca. 6600 spp. of ca.1200 genera and more than 200 families. Several papers out-
line some of the zoogeographical particularities of Oribatida: Wallwork (1969,
1973, 1979), Hammer and Wallwork (1979), Travé (1976a, b), Schatz (1998),
Niedbala (2002, 2004), and others’ interest.
6.14 Holothyrida
These big, conspicuous, and poisonous mites were first noticed by Gervais (1842)
on Mauritius Island. He described Holothyrus coccinella, raised as family by
Thorell (1882), who started the long series of descriptions of New Guinean
Holothyrida (followed by Canestrini 1897). Another early Arachnologist, Butler
(1876), recorded these strange mites from Rodrigues. Thon (1905, 1906) and
Warburton (1913) described three more species from the Seychelles. The series of
early descriptions includes also holothyrids from New Caledonia (Berlese 1923)
and New Zealand (Womersley 1935). So far, all findings came from islands. Finally,
Thon (1909) raised Holothyrida in a suborder.
Van der Hammen (1961, 1965, 1968, 1972, 1983) started the modern stage of the
study of Holothyrida. He described (1972) the second family in the suborder –
Allothyridae – for the two species from Australia and New Zealand, belonging to the
new genera Allothyrus van der Hammen, 1961 and Australothyrus der Hammen, 1983.
From 1981 to 1999 P. Lehtinen contributed greatly to the study of Holothyrida by
describing a new family Neothyridae (1981) from South America with the two new
genera Neothyrus and Diplothyrus (Kontschán and Mahunka 2004 added another
new genus Carabothyrus from the Dominican Republic). Lehtinen (1981, 1995,
1999) described also the new genera Hammenius, Haplothyrus, Indothyrus,
Lindothyrus, and Sternothyrus (from New Caledonia, Lord Howe Island, the
Seychelles, New Guinea, and Sri Lanka) and raised Leiothyrus and Thonius to gen-
era, thus making his own zoogeographical analysis of 1991 out of date. He became
the author of 9 out of the 15 genera in the family. Gerlach, Lehtinen, and Madl
(2010) added three endemic genera from the Seychelles.
Holothyrida (Holothyrina, Holothyroidea, Tetrastigmata) have been considered
by the modern (after 1949) authors either as suborder of Parasitiformes or as order.
It is considered as order by Walter in Kranz and Walter (2009), and we follow this
concept here and in our Catalogue (Beron 2014).
Order Holothyrida – Three fam., 13 gen., 29 spp.
Fam. Allothyridae – Two gen., three spp.
Allothyrus van der Hammen, 1961 – Australia, New Zealand (two spp.)
Australothyrus van der Hammen, 1983 – Australia (one sp.)
Fam. Holothyridae – 8 gen., 20 spp.
Dicrognatus Gerlach, Lehtinen et Madl, 2010 – Seychelles (two spp.)
Hammenius Lehtinen, 1981 – New Guinea (four spp.)
Haplothyrus Lehtinen, 1995 – New Caledonia (two spp.)
Holothyrus Gervais, 1842 – Mauritius (two spp.)
Indothyrus Lehtinen, 1995 – Sri Lanka (one sp.)
Leiothyrus van der Hammen, 1983 – New Guinea (three spp.)
Lindothyrus Lehtinen, 1995 – New Caledonia, Lord Howe Island (two sp.)
6.14 Holothyrida 195
Michaelothyrus Gerlach, Lehtinen et Madl, 2010 – Seychelles (one sp.)

Sternothyrus Lehtinen, 1995 – Seychelles (one sp.)
Thonius Lehtinen, 1981 – New Guinea (four spp.)
Fam. Neothyridae – Three gen., three spp.
Caribothyrus Kontschán et Mahunka, 2004 – Dominican Republic (one sp.)
Diplothyrus Lehtinen, 1999 – Brazil, French Guyana (two spp.)
Neothyrus Lehtinen, 1981 – Peru, Venezuela (one sp.)
Incertae sedes – Rodrigues (one sp.)
Map 6.20 Holothyrida in the Old World (Indian Ocean)

Dicrognatus – ■
Holothyrus – ◘
Indothyrus – ♦
Michaelothyrus – ◄
Sternothyrus – ♣
Map 6.21 Holothyrida in Australasia

Allothyrus – X
Australothyrus – Ω
Hammenius – ●
Haplothyrus –
Leiothyrus –
Lindothyrus – ◇
Thonius –
6.14 Holothyrida 197
Map 6.22 Holothyrida in Americas

Caribothyrus – ▲
Diplothyrus – ▼
Neothyrus – ►
The strange poisonous Holothyrida are 29 valid species from 15 genera and 3
families (Beron 2014), and we think that the guesstimate of 320 existing species and
even the minimum of 160 (Walter and Proctor 2009) are exaggerations. Holothyrids
are large and conspicuous creatures, and it is difficult to miss them where they exist.
It is true that 17 of these 29 species have been described after 1980 (by P. Lehtinen,
L. van der Hammen, J. Kontschán and S. Mahunka, and H. Klompen). A total
restructuring has been done by these authors and by J. Gerlach also in the generic
classification of the order (from one genus Holothyrus to 15 genera). Now is more
credible to analyze the most intriguing distribution of this order, more credible even
in comparison with the relatively recent zoogeographical analysis of Lehtinen
(1991).
Many of the holothyrids are endemic genera, and not because they have not been
looked for elsewhere, but it seems that on their islands they might be really endemic.
The family Neothyridae is entirely American – four species from Dominican
Republic (Caribothyrus Kontschán et Mahunka), French Guyana, Brazil (Diplothyrus
Lehtinen), and Peru and Venezuela (Neothyrus Lehtinen). From Australia have been
described by Van der Hammen a special family (Allothyridae) and two new genera
Australothyrus and Allothyrus. The last one has been recorded also from New
Zealand (Allothyrus australasiae Womersley, 1935), the same species described
from Australia, but such important finding has not been confirmed since.
What concerns the Old World is that there live the remaining 22–23 species of 10
genera. The heavy weight of the whole order seems to be in New Guinea with 11
species of the genera Hammenius Lehtinen, Leiothyrus Van der Hammen, and
Thonius Lehtinen. They all belong to the family Holothyridae, spread from Mauritius
to New Caledonia, and not to the “Australian” Allothyridae. Endemic genus and
species for Sri Lanka is Indothyrus greeni Lehtinen, 1995. New Caledonia and Lord
Howe Island, being between Australia and New Zealand, share another genus of
Holothyridae – Lindothyrus Lehtinen, 1995. Another (monotypic) genus
(Haplothyrus Lehtinen, 1995) is endemic to New Caledonia.
Particularly interesting is the presence of Holothyridae on the islands close to
Africa, but not in Africa itself or Madagascar. On the Seychelles live at least four
species of three endemic genera: Dicrognathus Gerlach, Lehtinen et Mandl, 2010;
Michaelothyrus Gerlach, Lehtinen et Mandl, 2010; and Sternothyrus Lehtinen,
1995. They are part of the many endemisms of the granitic islands.
Finally, Mauritius, the place from where has been described the first Holothyrid
as early as in 1842, remains with two species of the first described genus Holothyrus
Gervais (plus one uncertain species from Rodrigues). The distribution of the holo-
thyrids over so many islands far from each other and their (apparent) absence from
places like Africa, Madagascar, or India remain among the riddles of
zoogeography.
6.15 Ixodida
Fam. Nuttaliellidae
Nuttalliella Bedford, 1931 (one sp.)
This family is (the only described species Nuttalliella namaqua Bedford)
endemic for a narrow area in Southern Africa, from Tanzania to Namibia.
Fam. Ixodidae
The family Ixodidae contains ca. 700 spp. in 14 genera (Filippova 1994, Horak
et al. 2002, Kolonin 1978 1983 1984, 2009, Guglielmone et al. 2010), parasites on
Bibliography 199
vertebrates all over the world. Kolonin (1981, 1983) subdivided the world into
superregions and regions after the tick fauna in them.
Fam. Argasidae
Also parasites of vertebrates (193 spp. in 5 genera).
6.16 Mesostigmata
Many groups are mono- or oligoxenous parasites and their distribution follows the
distribution of their hosts (Beron 2014). Some examples are as follows:
Fam. Spelaeorhynchidae – the only genus Speleorhynchus and seven species are
found in South and Central America and the Antilleans. They follow the distribu-
tion of the bats of families Phylostomatidae and Mormoopidae.
Fam. Hystrichonyssidae – only one species, living on the Malayan porcupine
(Hystrix brachyura)
Fam. Manitherionyssidae – only one species, living on the pangolins (Manis tem-
mincki) in Southern Africa
Fam. Dasyponyssidae – two species, parasites of Cingulata (Dasypodidae) in South
America
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Chapter 7
Regional Arachnogeography
Abstract The arachnofauna of various parts of the Earth is analyzed and the par-
ticularities, endemics, relicts, and the presumed ways of formation of the fauna are
outlined. Also the northern limits of the groups in the Holarctic are indicated, and
the connections in the geological time are analyzed.
Bibliographies
Smaller orders – Harvey (2003)
Scorpiones – Dupré (1998, 2013a, 1913b), Fet et al. (2000)
Pseudoscorpiones – Schawaller (1980), Harvey (1990), Zaragoza (2000)
Opiliones – Giribet (2000), Cokendolpher and Lee (1993), Kury (2011), Schönhofer
(2013)
Palpigradi – Condé (1996), Harvey (2003)
Amblypygi – Harvey (2003)
Uropygi (Thelyphonida) – Harvey (2003), Rowland and Cooke (1973)
Schizomida – Harvey (2003), Reddell and Cokendolpher (1995)
Ricinulei – Harvey (2003)
Solifugae – Harvey (2003), Gromov and Savary (on line)
Opilioacarida – Beron (2014)
Araneae – Roewer (1940), Brignoli (1983), Platnick (1989, 1993, 2014)
Acariformes – Beron (2008, 2011, 2015a, 2015b, 2016b)
Parasitiformes – Beron (2014, 2016a)
7.1 Holarctic
7.1.1 Zoogeography and Arachnozoogeography
From 16 recent orders and 10 suborders of Arachnida, 15 orders and almost all sub-
orders are recorded in the huge Holarctic Kingdom. Lacking from the arachnofauna
of the Holarctic are only Holothyrida. Some others are recorded only on the edge of

https://doi.org/10.1007/978-3-319-74418-6_7
320 7 Regional Arachnogeography
the Kingdom – Schizomida and Ricinulei in the Nearctic part, Thelyphonida

(Uropygi) in the Far East of Russia (?), and Texas in the USA.
Holarctic is usually regarded as Kingdom composed of two regions (Palearctic
and Nearctic). In the scheme of Krzhanovskiy (2002), four regions are included in
the Holarctic Kingdom (Boreal Region, Region of the Ancient Mediterranean, East
Asian Region, Sonoran Region). He does not use the widely accepted terms
Palearctic and Nearctic.
7.2 Europe (General)
7.2.1 Geography, General Zoogeography, and Paleogeography
The European Environment Agency (2002) (EEA) subdivides Europe into six geo-
graphical regions: North Europe, West Europe, Middle Europe, South Europe,
Southeast Europe, and East Europe. Caucasus is a border area between East Europe
and Asia Minor.
The Agency subdivides Europe also into nine biogeographical regions: Arctic,
Boreal, Continental, Mediterranean, Alpine, Atlantic, Pannonic, Steppic, and Black
Sea regions. To them are attached two more regions: Macaronesian (Azores,
Madeira, Ilhas Selvagens, Canary Islands) and Anatolian (Asia Minor).
Arctic Region Island, Svalbard (Spitzbergen), Jan Mayen, the Far North of
Norway and Kola Peninsula, Arkhangelskaya Oblast, Novaya Zemlya, Franz Josef
Land in Russia. Tundra, Boreal forest, cold desert. Several endemic spiders and
mites, no endemic genera.
Boreal Region Most of the Scandinavian Peninsula, the Baltic states, North of
Russia (2,900,000 km2, of which 1,900,000 in Russia). Most of the area is forest
(58%) and cultivated lands.
Continental Region From France to Central Russia. Most is cultivated land (52%),
forest (27%), and meadows (14%).
Mediterranean Region South of France; the Iberian, Apennine, and Balkan

Peninsula; and the Mediterranean Islands. Mountainous area with many caves, rich
in endemic Arachnida and also endemic members of orders not known in the north-
ern regions (Solifugae, Opilioacarida, Scorpiones). The area has 36% of cultivated
land, 25% of forests, 29% of meadows, etc.
7.2 Europe (General) 321
Alpine Region According to this subdivision, it includes not only the Alps but also
the Scandinavian Mountains, the Pyrenees, the Carpathians, the Dinaric Mountains,
the “Rhodopes”(certainly including Rila and Pirin), the Ural Mountains, and the
Caucasus.
From this area 41% are forests, 26% meadows, 8% cultivated land, and 13% with
few or no vegetation.
In many endemic arachnids, particularly remarkable is the scorpion Belisarius
xambeui.
Atlantic Region British Islands, Hebrides, Orkney and Shetland Islands, Faroe
Islands, Southeast of Norway, northern part of Iberian Peninsula, and the area from
the North of France to Denmark. The area consists of 38% cultivated lands, 30%
meadows, 13% forests, etc.
Pannonic Region Mostly the Hungarian and the other plains of Central Europe (in
Serbia, Romania, Slovakia, etc.). Some typical steppe Arachnida.
Steppic Region Dobrogea, south Moldova, Ukraine, European Russia, European

part of Kazakhstan. From this area 75% is cultivated land, 17% meadows, 2% for-
est, etc.
The Black Sea Region The western, southern, and eastern coast areas of Black
Sea and Central Georgia. The meadows are 34%, forest 29%, cultivated land 14%,
etc.
These areas are defined by the European Commission and the Council of Europe
for evaluation and reporting of nature conservation. The real zoogeographical sub-
division of Europe is different. Usually it is divided into Eurosiberian and
Mediterranean provinces. From the map of Udvardy (1975), we can see that the
biogeographical subdivision is much more detailed.
Map 7.1 West Palearctic Biogeographical Realm (Udvardy 1975)
Provinces (on this map):
3 – West Eurasian Taiga
5 – Icelandian
6 – Subarctic Birchwoods
8 – British Islands
9 – Atlantic
10 – Boronemoral
11 – Middle European Forest
12 – Pannonian
13 – West Anatolian
16 – Iberian Highlands
17 – Mediterranean Sclerophyll
18 – Sahara
19 – Arabian Desert
20 – Anatolian-Iranian Desert
21 – Turanian
24 – Iranian Desert
26 – High Arctic Tundra
27 – Low Arctic Tundra
28 – Atlas Steppe
29 – Pontian Steppe
31 – Scottish Highlands
32 – Central European Highlands
33 – Balkan Highlands
34 – Caucasus-Iranian Highlands
7.2.2 Arachnogeography
Ref.: Aakra and Houge (2000), Agnarsson (1996), Armas and Rehfeldt (2015),
Beier (1963), Beron (2011), Blagoev (2002), Blick et al. (2004), Blick and Christian
(2004), Blick et al. (2004), Blick and Komposch (2004), Blick et al. (2004),
Bosmans (2009), Bosmans and Chatzaki (2005), Bosmans and De Keer (1985),
Bosmans and Vanuytven (2001), Bosmans et al. (2013), Bosmans et al. (2013),
Buchar et al. (1995), Buchar and Ružička (1995), Cardoso (2000, 2010), Cardoso
and Morano (2010), Cawley (2002), Chemini (1995), Christophoryová et al. (2012),
Condé (1996), Ćurčić (1974), Ćurčić et al. (2004), Defosse (2004), Deltshev (2005),
Deltshev and Blagoev (2001), Deltshev et al. (2003), Drensky (1936), Farzalieva
and Esyunin (2000), Feider (1965), Fet (2010), Gaidus et al. (1999), Gencheva and
Georgieva (2013), Gardini (1994), Hadži (1973), Klimeš (2000), Komposch (2004),
Komposch and Gruber (2005), Koponen (2005), Kronestedt (2001), Kury and
Mendes (2007), Le Peru (2007 2011), Loksa (1969), Mahnert (2004), Marinu and
Verneau (2002), Martens (1978), Meinertz (1962), Merrett et al. (1985), Mheidze
(1964), Milošević (2002), Morano and Cardoso (2009), Nentwig et al. (2013),
Nikolić and Polenec (1981), Novak (2004, 2005), Novak et al. (2006), Otto (2014),
Ozimec (2000), Pack-Beresford (1926), Pantini et al. (2013), Pantini and Isaia
(2015), Pesarini (1994), Petrov (1997), Prieto (2003, 2008), Proszynski and Starega
(1971), Rafalski (1960, 1961, 1967), Rafalski and Starega (1997), Rambla (1967),
Roberts (1985), Sanu and Szinetar (1994), Sankey and Savory (1974), Scharff and
Gudik – Sørensen (2011), Schmölzer (1995), Simon (1875), Staręga (1976a, 1976b,
1978, 2000), Starkoff (1958), Stoch (2003), Stol (1993 2007), Szalay (1968),
Šilhavý (1956), Telnovs (2000a, 2000b), Topcu et al. (2005), Tumšs (1963),
Vanhercke (1999), Varol (2003), Vilkas (1992), Weiss and Urak (2000), Wiehle
(1953), Wijnhoven (2005), Wintarska (2008), Zaragoza (2007).
In present-day Europe are lacking some “southern” groups – Ricinulei,
Holothyrida, Thelyphonida (Uropygi), Amblypygi, and Schizomida. Some other
warm loving groups are represented only on the southernmost Mediterranean part
of the continent and with small number of taxa – Scorpiones, Solifugae,
Opilioacarida, Opiliones, Cyphophthalmi, and Laniatores.
Well represented are the spiders, pseudoscorpions, opilions (Eupnoi and
Dyspnoi), and most groups of Acari, Palpigradi. Across Europe pass the lines mark-
ing the northern limit of many groups.
Palpigradi In Europe (including Madeira) are registered 28 species of Palpigradi,
all belonging to one genus – Eukoenenia Börner (fam. Eukoeneniidae). They are
recorded (without counting the subspecies) from France (9), Italy (12), Greece (3),
Bulgaria (1), Hungary (1), Austria (2), Bosnia and Herzegovina (1), Croatia (2),
Malta (2), Portugal (2), Romania (4), Slovakia (1), Slovenia (1), and Spain (4) – 14
countries (Blick and Christian 2004, Condé 1996).
Schizomida Not living naturally in Europe, but four species have been found,
imported from other continents in hothouses and other localities in Spain, France,
Great Britain, Germany, Czech Republic, Slovakia, and Poland (Barranco et al.
2014; Armas and Rehfeldt 2015). These are Schizomus crassicaudatus (O.P.-
Cambridge), Bucinozomus hortuspalmarum Armas et Rehfeldt, Zomus bagnalii
(Jackson), and Stenochrus portoricensis Chamberlin.
Solifugae In Europe Solifugae are found only in Spain, Portugal, Sicily, Greece,
Bulgaria, Macedonia, and Ukraine – altogether eight species of the genera Galeodes
Olivier (Galeodidae), Biton Karsch, Gluvia C.L. Koch (Daesiidae), Barrussus
Roewer, and Eusimonia Kraepelin (Karschiidae) (Birula 1912, Blick 2004, Roewer
1934).
Map 7.2 Distribution of Solifugae in Europe

Galeodes – X
Gluvia – ●
Barrussus – ○
Biton – ■
Scorpiones According to Fet (2010 and supplements), in Europe are registered

23 spp. of scorpions, but since several new species have been described and with
more detailed checking of several populations and “subspecies,” this number could
increase to 35. The European scorpions belong to the families Buthidae (Buthus
Leach, 3 spp.; Mesobuthus Vachon, 3 spp.), Euscorpiidae (Euscorpius Thorell, 15
spp.), Iuridae (Iurus Thorell, 1 sp.), and (?)Troglotayosicidae [or Chactidae]
(Belisarius Simon, 1 sp.) (Gantenbein et al. 1999, Kovařik 1999).
Map 7.3 Scorpiones in Europe

■ – Euscorpius
O – Iurus
X – Mesobuthus
♦ – Buthus
! – Belisarius
Pseudoscorpiones The book of Beier (1963) listed 9 families of pseudoscorpions

in Europe (including Canary Islands, Madeira, and Azores), but now (Lissner 2014)
there are 73 genera, belonging to 14 families (Chthoniidae, Neobisiidae, Bochicidae,
Larcidae, Syarinidae, Olpiidae, Garypidae, Geogarypidae, Garypinidae, Atemnidae,
Cheiridiidae, Cheliferidae, Chernetidae, Withiidae) – nearly half of the
Pseudoscorpiones families in the world.
According to Lissner (2014), “The European pseudoscorpion fauna counts to 861
species in 73 genera and 16 families.” These numbers include species and genera
from Asian parts of Russia and Turkey (perhaps 20 species?). The fauna is much
more diverse in the southern parts of Europe. The highest number of species is found
in Italy (226) followed by Spain (213, including the Canary Islands). Blick et al.
(2004) list 102 pseudoscorpion species for Central Europe. Species number decline
to about 20 in southern Scandinavia (Beier 1963). “None, except a few introduced
synanthropic species, are found at higher latitudes than ca. 65°N” (Beier 1963).
Endemic genera of Pseudoscorpiones in Europe:
Fam. Chthoniidae
Neobalkanella Ćurčić, 2013 – Serbia (one sp.)
Troglochthonius Beier, 1939 – Bosnia and Herzegovina, cave (one sp.); ? Sardinia
Fam. Neobisiidae
Insulocreagris Ćurčić, 1987 – Croatia (Vis), Herzegovina (two spp.)
Protoneobisium Ćurčić, 1988 – Croatia (one sp.)
Archaeoroncus Ćurčić et Rada, 2012 – Croatia (one sp.)
Ernstmayria Ćurčić et Dimitrijević, 2006 – Crete (one sp.)
Fam. Syarinidae
Arcanobisium Zaragoza, 2010 – Spain (A. comasi Zaragoza, 2010)
Hadoblothrus Beier, 1952 – Italy, Greece (two spp.)
Pseudoblothrus Beier, 1931 – Italy, Crimea, France, Switzerland (seven spp.)
Microcreagrella Beier, 1961 – Azores, Madeira, Spain, Portugal (one sp.)
Fam. Bochicidae
Titanobochica Zaragoza et Reboleira, 2010 – Portugal (T. magna Zaragoza et
Reboleira, 2010)
Troglobisium Beier, 1939 – Spain [T. racovitzai (Ellingsen, 1912)]
Cyphophthalmi In Europe have been registered 5 genera and 25 species of 1 fam-
ily (Sironidae), mainly in Southern Europe, with 1 species Siro carpathicus Rafalski
going North to Poland and Slovakia. There are two main centers of speciation:
Iberian Peninsula with 5 genera and 5 species and Balkan Peninsula with ?17 spe-
cies of 1 genus (Cyphophthalmus, incl. Tranteeva) (Roewer 1923, Martens 1978,

Murienne and Giribet 2009).
Eupnoi Two families contain the bulk of European Opilions:
Fam. Phalangiidae – Phalangium L., Dicranopalpus Doleschall, Lacinius Thorell,

Mitopus Thorell, Egaenus C.L. Koch, Opilio Herbst, Oligolophus C.L. Koch,
Paroligolophus Lohmander, Dasylobus Simon, Graecophalangium Roewer,
Metaphalangium Roewer, Ramblinus Starega, Rilaena Šilhavý, Paraplatybunus
Dumitrescu, Lophopilio Hadži, Megabunus Meade, Metaplatybunus Roewer,
Platybunus C.L. Koch, Zachaeus C.L. Koch, Stankiella Hadži, Rafalskia Starega,
and others
Fam. Sclerosomatidae – Gyas Simon, Cosmobunus Simon, Micronelima Schenkel,
Leiobunum C.L. Koch, Nelima Roewer, Astrobunus Thorell, Homalenotus
C.L. Koch, Mastobunus Simon, Metasclerosoma Roewer (nine genera)
Dyspnoi The European (Caucasus included) fauna of Dyspnoi consists of five
families:
Nemastomatidae (Nemastoma C.L. Koch, Paranemastoma Redikorzev,
Mitostoma Roewer, Giljarovia Kratochvil et Miller, Carinostoma Kratochvil,
Nemaspela Šilhavý, Acromitostoma Roewer, Saccarella Schönhofer, Hadzinia
Šilhavý, Pyza Staręga, Vestiferum Martens, Histricostoma Kratochvil, Mediostoma
Kratochvil, Centetostoma Kratochvil, Caucnemastoma Martens, Nemastomella
Mello-Leitão), Sabaconidae (Sabacon Simon), Ischyropsalididae (Ischyropsalis
C.L. Koch), Dicranolasmatidae (Dicranolasma Sørensen), and Trogulidae (Trogulus
Latreille, Anarthrotarsus Šilhavý, Anelamocephalus Simon, Calathocratus Simon,
Kofiniotis Roewer (24 genera and 185 accepted spp.) (Schönhofer 2009).
Laniatores In Europe Laniatores are much fewer than the “Palpatores” – only
40 spp. of 12 genera and the families Cladonychiidae (Holoscotolemon Roewer),
Phalangodidae, and Travuniidae. Many genera (Lola Kratochvil, Paralola
Kratochvil, Travunia Absolon et Kratochvil, Trojanella Karaman, Dinaria Roewer)
and species are known only from caves and are considered generally as relict (Kury
and Mendes 2007). However, Martens (1972) denies their relict character and thinks
that this is a normal process of speciation.
Araneae In Europe (including Madeira and Azorean Islands) until the end of
2011 have been registered 4892 species and subspecies of spiders (van Helsdingen
2012). They include 4491 “true” indigenous spiders, 99 introduced species, and
302 nomina dubia. The category of unique records/endemics contains 2041 spe-
cies (45.4%). These spiders belong to the families Atypidae, Ctenizidae,
Nemesiidae, Filistatidae, Sicariidae, Scytodidae, Leptonetidae, Pholcidae,
Dysderidae, Segestriidae, Oonopidae, Palpimanidae, Mimetidae, Eresidae,

Oecobiidae, Uloboridae, Nesticidae, Theridiosomatidae, Anapidae, Mysmenidae,
Araneidae, Linyphiidae, Tetragnathidae, Theridiidae, Lycosidae, Agelenidae,
Pisauridae, Oxyopidae, Zoropsidae, Cybaeidae, Argyronetidae, Desidae,
Hahniidae, Amaurobiidae, Dictynidae, Titanoecidae, Anyphaenidae, Clubionidae,
Liocranidae, Gnaphosidae, Philodromidae, Thomisidae, Salticidae, Corinnidae,
Zodariidae, Prodidomidae, Miturgidae (Zorinae), and Sparassidae. (oshte!). The
most numerous families (figures of 2011) are Linyphiidae (1366), Gnaphosidae
(480), Salticidae (400), Dysderidae (331), Lycosidae (303), Theridiidae (258),
Agelenidae (200), Thomisidae (193), Araneidae (150), Zodariidae (111), and
Philodromidae (106). The best represented what concerns the endemics are
Linyphiidae (534, 42.4%), Agelenidae (93, 50%), and most of all Dysderidae
(227, 70.5%). There are many regional lists of the spiders in European countries
(Decae 2010, Deltshev 2000, Deltshev et al. 2003, Koponen 1991, Le Peru 2011,
Nentwig et al. 2013, Song et al. 2002, Thorell 1869, and others).
Opilioacarida In Europe (the southern parts of Italy and Greece) are known only
two spp. of Opilioacarida: Opilioacarus segmentatus With, 1903, and O. italicus
With, 1904 (Beron 1990, 2014, Brignoli 1967, With 1903, 1904)
7.3 Balkan Peninsula and Aegaeis
7.3.1 G
eography, General Zoogeography,
and Paleogeography
The eastmost large peninsula of Southern Europe meets the Adriatic Sea on the
Northwest, Ionian Sea on the southwest, the Mediterranean and Aegean Sea on the
South and Southeast, and the Black Sea on the East and Northeast. The highest
point of the Balkans is Mount Musalla (2925 m) in Rila mountain in Bulgaria. The
total area of the Balkans is 666,700 square. The Peninsula got its name from the
Balkan Mountains (Stara planina) that stretch from the east of Bulgaria to the very
east of Serbia.
Several attempts have been made to subdivide zoogeographically the most com-
plex area in Europe – on the contact between the Eurosiberian and the Mediterranean
units, with montanmediterranean centers of speciation, with arctic-alpine elements,
with many endemic cave animals, and with influence from the Steppic area and Asia
7.3 Balkan Peninsula and Aegaeis 329
Map 7.4 Balkan Peninsula
Minor. The approach of various authors is different, but usually following the phy-
togeographical subdivision (Asenov 2006, Brunn and Mercier 1971, Dermitzakis
and Papanikolaou 1981, Buresch and Popov, 1973, Gruev 2000, Guéorguiev 1979,
1982, Matvejev 1961, 1969).
The first attempts to analyze the zoogeography of Bulgaria were due to Drenski
(= Drensky) (1936, 1946, 1966). This founder of the zoogeography in Bulgaria was
an arachnologist himself, and some of his analyses were based largely on the distri-
bution of spiders, the sun spider, and some other arachnids.
In several papers, Entomologist V.B. Guéorguiev analyzed the contemporary
data of the distribution of animals in Bulgaria and provided a new subdivision of
Bulgarian territory. The same author analyzed the cave fauna of Balkan Peninsula
(Guéorguiev 1977), taking into account the evolution of the territory in the geologic
past and the influence of elements from Gondwana, the concept of Egeida, etc. His
speleozoogeographic map of the peninsula is unique even to our time.
Gradually, as elsewhere, the biogeographical approach became predominant. In
the second edition of the manual of Gruev and Kuzmanov (1994), we find six “faunis-
tic complexes” in Bulgaria: Northern Holarctic, Siberian, European, Euroasiatic step-
pic, Southwestasian, and Mediterranean. Outlined are five “biogeographic regions”:
North Bulgarian, Middle Bulgarian, South Bulgarian, Pontic, and Mountain regions.
Such units should not bear names of only one state, as the “South Bulgarian
region” stretches also in Turkey, Greece, and the Republic of Macedonia, and in
these countries it would be called differently. When subdividing the peninsula and
other territories, we should disregard the political boundaries.
According to Gruev (2000), the submediterranean elements in the eastern parts of
the submediterranean province “belong to the thermos-mesophilic and the thermos-
xerophilic ecological types in conditions of temperate-continental, transitional-
continental and transitional-mediterranean climate.”
Here are the zoogeographic maps of Bulgaria of Drensky (1946) and Gueorguiev
(1980) and the biogeogeographic map of Gruev (1988):
Map 7.5 Zoogeographical map of Bulgaria after Drensky (1946)

I – Areas with Eurosiberian elements: 1, Danubian Bulgaria; 2, Western Bulgaria; 3, Stara planina;
4, Central High Mountain part. II – Areas with Mediterranean elements: 5, Aegean area; 6, Upper
Maritza plain; 7, Pontian area. III – Areas with Irano-Turanian elements: 8, Dobrudja; 9, Lower
Maritza Thrace
Map 7.6 Zoogeographical map of Bulgaria after Gueorguiev (1980, 1992)

___ Border between the Eurosiberian and Mediterranean territory
----- Borders between the zoogeographical areas
1. Danubian area
2. Area of Stara planina
3. Rilo-Rhodopean area
4. Thracian area
5. Strandja area
6. Black Sea area
7. Struma-Mesta area
Map 7.7 Biogeographical map of Bulgaria after Nikolov (1977)

1. North Bulgarian Steppe and Forest – Steppe Province
1.1 – Danubian Forest-Steppe area
1.2 – Dobrudja Forest-Steppe area
2. Balkan Middle-European forest province
2.1 – North Bulgarian forest area
2.2 – Western-Middle Bulgarian area
2.3 – Osogovo-Maleshevo area
2.4 – South Bulgarian Mountain area
3. Balkan Submediterranean province
3.1 – Struma-Mesta area
3.2 – Upper Thracian area
3.3 – East Rhodopean-Sakar area
3.4 – Strandja area
3.5 – Black Sea area
Drenski (1936) discussed the distribution of spiders in Bulgaria. Guéorguiev (1992)

wrote a special publication on the zoogeography of Opiliones in Bulgaria. Many
papers of Ćurčić and his coauthors analyze the distribution of pseudoscorpions, and
in the book of Deltshev et al. (2003) have been published maps of all species of
Araneae in Serbia. In another monograph (Ćurčić et al. 2004) were outlined all spe-
cies of Pseudoscorpiones in Serbia, Montenegro, and the Republic of Macedonia.
Petrov (1997) wrote a checklist of pseudoscorpions in Bulgaria, completed by
Petrov and Štáhlavský (2007).
According to the analysis of Deltshev et al. (2005), in Bulgaria were recorded by
2005 1091 spp. of Arachnida (outside the Acari): 925 spiders, 61 harvestmen, 51
false scorpions, 2 scorpions, 1 Solifugae, and 1 palpigrade. These authors think that
there are 50 Bulgarian endemics and 68 Balkan endemics, and 240 species are con-
sidered rare. Since 2005 some more species have been published (the scorpions are
now at least four).
The northern limits of some groups (Solifugae, Opilioacarida) cross the penin-
sula. Some others (Amblypygi, Thelyphonida, Schizomida, Ricinulei, Holothyrida)
are absent from it.
Guéorguiev (1992) analyzed zoogeographically the fauna of Opiliones in
Bulgaria (the 42 genera and subgenera known by this time) and concluded that the
“boreal” species are 20 (47,6%), the “meridionals” are 7 (16,7%), and the endemics
are 15 (35,7%). Presently (2017), thanks to the efforts of P. Mitov, in Bulgaria are
known already 63 spp., so another analysis would be interesting.
Gruev (1995) analyzed the “Mediterranean faunistic complex” in Bulgaria,
based only on invertebrates and mostly on Coleoptera, Chrysomelidae. He con-
cluded that Bulgaria belongs biogeographically entirely to the European-West
Siberian area of the Palearctic Region and that none of the territories of the country
belongs to the Mediterranean. It is due mostly to the southern borders of Bulgaria,
running on high mountains and not allowing to the typical biotic societies to be part
of the present-day Bulgarian territory. Only some elements penetrate the areas with
submediterranean climate along the river valley of Struma and to the less extent of
Mesta and Maritza. Speaking of Arachnida, among these elements are Galeodes
graecus (Solifugae), Euscorpius solegladi, E. drenskii, E. popovi, Mesobuthus gib-
bosus (Scorpiones), and some Araneae, Opiliones, and Pseudoscorpiones. They are
part of the submediterranean fauna in Bulgaria, outlined (in Bulgarian) by Gruev
(2000). According to him, the submediterranean biogeographic province includes
(intrazonally) territories from Iberian Peninsula to Caucasus (the middle of the
Iberian Peninsula and the Apennines, South of France, Corsica, parts of the former
Yugoslavia, Hungary, Austria, Switzerland, Northern and Middle Greece, northern
part of Turkey, the south of Crimea, and Northern Caucasus up to the Caspian Sea).
They are characterized (Matvejev 1961) by the following parameters: average
annual temperature of 9,3–11,7 °C, average monthly temperature of the warmest
month 20–23,2 °C, negative winter temperature (average monthly) from 1 to 3
months, and rainfall of 450–650 mm with dry summer. Such climate is found up to
800–1000 m, in places even up to 1500 m and higher. Among the Arachnida in the
paper of Gruev (2000) are listed only four species of spiders (Dysdera bulgarica,
Pardosa pseudostrigilata, P. strigilata, Trachelas maculatus), after Deltshev (1976).
Actually, they are more.
The Republic of Macedonia is also with high mountains on all its borders and has
mostly submediterranean fauna. Albania, Montenegro, Dalmatia, and Slovenia are
exposed to Adriatic (Mediterranean) climatic influence, and are very karstic, the
limestone creating usually comfort to warm-preferring arachnids.
According to Deltshev (2004), the spiders of the “Mediterranean complex” of the
Balkan Peninsula are 13.8% of the total spider fauna of the peninsula. Many Balkan
endemics are also of Mediterranean origin. “The mountain-Mediterranean species
(e.g., Aculepeira talishia, Pardosa incerta) are of particular interest, since they may
be regarded as ancient element.”
Most of the territory of Greece belongs to the typical Mediterranean subregion,
characterized best by the areal of the olive trees (Olea europaea). The fauna of the
Balkan part of Greece contains many solpugids, scorpions, spiders, and other arach-
nids, unknown in the areas north of the country.
The peninsula has many high mountains (up to 2925 m), harboring glacial relicts
and montanmediterranean arachnids.
Palpigradi From the countries of the peninsula (south of Danube) are known seven
Palpigradi species and three subspecies (all from genus Eukoenenia Thorell,
Eukoeneniidae) (Condé 1974, 1976, 1977, 1979, 1990, 1996, Georgescu and Decu
1994, Hansen 1926, Beron 1978, Orghidan et al. 1982).
Bulgaria – Beron (1978) reports three specimens of ?E. austriaca from caves in
NW Bulgaria.
Greece – have been reported E. juberthiei cytheriaca Condé, 1979 (Kythira
Island); E. j. hellenica Condé, 1979 (Attika); E. naxos Condé, 1990 (Iraklia Island)
Bosnia and Herzegovina – E. remyi Condé, 1974 (Herzegovina)
Slovenia – E. austriaca austriaca (Hansen, 1901)
Croatia – E. pretneri Condé, 1977; E. spelaea hauseri Condé, 1974
Northern Dobrudja (Romania) – E. mirabilis (Grassi et Calandruccio, 1885); E.
condei Orghidan, Georgescu, et Sârbu, 1982; E. margaretae Orghidan, Georgescu,
et Sârbu, 1982 (Georgescu & Decu, 1994)
Solifugae The list of European Solifugae of Blick (2004) contains 18 spp., but
geographically half of them live on territories outside Europe (Rhodes, Asiatic
Turkey, Cyprus, Canaries).
On the Balkan Peninsula (continental part, mostly in Greece) are recorded mem-
bers of three families of Solifugae (Drensky 1931, Kraepelin 1899, Roewer 1934):
Fam. Galeodidae (Galeodes Olivier): G. graecus C.L. Koch (Greece, Bulgaria),
G. elegans Roewer (Republic of Macedonia), G. hellenicus Roewer (Greece)
Fam. Karschiidae – Barrussus furcichelis Roewer, Eusimonia nigrescens
Kraepelin (Greece)
Fam. Daesiidae – Biton ehrenbergi Karsch (Greece)

The northern boundary of Solifugae in eastern Europe runs through SW Bulgaria
(Galeodes graecus C.L. Koch, 1842), the Republic of Macedonia (G. elegans
Roewer, 1934), and Russia (G. araneoides Pallas, 1772) (Map 35).
Scorpiones On the Balkan Peninsula (including Slovenia and the islands of the
Aegean, except for the Dodecanese), there is a complicated picture of the number of
taxa of the scorpions, changing very often. We can agree on the current list following
Fet (2010) and many supplements, Crucitti (1995, 1999b), Calinescu and Calinescu
(1930), Æurèiæ (1971, 1972), Daniel (1962), Fet (1985, 2000), Fet and Braunwalder
(2000), Fet et al. (2001), Fet and Soleglad (2007), Fet et al. (2014), Francke (1981),
Kaltsas et al. (2008), Kinzelbach (1975), Kovařik et al. (2010), Michalis and Dolkeras
(1989), Michalis and Kattoulas (1981), Parmakelis et al. (2006), Tropea et al. (2015),
Vachon (1948, 1953), Voulalas and Michalis (1977), and Werner (1937).
Fam. Euscorpiidae
Euscorpius (E.) hadzii Caporiacco, 1950 – SW Bulgaria, Macedonia, Albania,
Bosnia and Herzegovina, Croatia, Greece, Serbia, Montenegro
E. (E.) koschewnikowi Birula, 1900 – NE Greece
E. (E.) “carpathicus” complex – Bulgaria, Albania
E. (E.) sicanus (C.L. Koch, 1837) – Greece
E. (E.) tergestinus (C.L. Koch, 1837) – Albania, Croatia, Slovenia, Montenegro
E. (E.) deltshevi Fet, Graham, Webber, et Blagoev, 2014 – West Bulgaria, East
Serbia
E. (E.) solegladi Fet, Graham, Webber, et Blagoev, 2014 – Southwest Bulgaria,
Northern Greece
E. (E.) drenskii Tropea, Fet, Parmakelis, Kotsakiozi, et Stathi, 2015 – Southeast
Bulgaria
E. (Alpiscorpius) mingrelicus dinaricus Caporiacco, 1950 – Bosnia
E. (A.) mingrelicus caporiaccoi Bonacina, 1980 – Bosnia
E. (A.) beroni Fet, 2000 – Albania
E. (Polytrichobothrius) naupliensis (C.L. Koch, 1837) – Greece
E. (P.) italicus (Herbst, 1800) – Greece
E. birulai Fet, Soleglad, Parmakelis, Kotsakiozi, et Stathi, 2014 – Greece
(Euboea)
E. mylonasi Fet, Soleglad, Parmakelis, Kotsakiozi, et Stathi, 2014 – Greece
(Euboea)
E. popovi Tropea, Fet, Parmakelis, Kotsakiozi, et Stathi, 2015 – Pirin, Slavyanka
(SW Bulgaria, N. Greece)
Fam. Buthidae
Mesobuthus gibbosus (Brullé, 1832) – Albania, Macedonia, Greece, Montenegro,
and European Turkey. The findings in European Turkey, the locality in SW
Bulgaria (Pirin, 1200–1300 m, Teruel et al., 2004), seems doubtful, although
the species may be expected near the border with the Republic of Macedonia.
Fam. Iuridae
Iurus dufoureius (Brullé, 1832) – was restricted by Kovarik, Soleglad, Fet, and
Yağmur (2010) only to Greece (Peloponnese, Kythira, and Crete).
Pseudoscorpiones On the Balkan Peninsula (within the limits outlined above)
have been recorded pseudoscorpions from 44 genera and 13 families: Chthoniidae,
Geogarypidae, Garypidae, Garypinidae, Olpiidae, Larcidae, Neobisiidae,
Syarinidae, Atemnidae, Cheiridiidae, Chernetidae, Cheliferidae, and Withiidae.
Fam. Chthoniidae – Chthonius C.L. Koch, Neobalkanella Ćurčić, Microchthonius

Hadzi,
Mundochthonius Chamberlin, Paraliochthonius Beier, Troglochthonius Beier
Fam. Geogarypidae – Geogarypus Chamberlin
Fam. Garypinidae – Amblyolpium Simon, Garypinus Daday
Fam. Garypidae – Garypus L. Koch
Fam. Olpiidae – Olpium L. Koch, Calocheiridius Beier et Turk, Cardiolpium
Mahnert, Minniza Simon
Fam. Larcidae – Larca Chamberlin
Fam. Neobisiidae – Acanthocreagris Mahnert, Balkanoroncus Ćurčić, Ernstmayria
Ćurčić et Dmitrievic, Insulocreagris Ćurčić, Microbisium Chamberlin,
Neobisium Chamberlin, Protoneobisium Ćurčić, Roncus L. Koch, Archaeoroncus
Ćurčić et Rada
Fam. Syarinidae – Hadoblothrus Beier
Fam. Cheiridiidae – Apocheiridium Chamberlin, Cheiridium Menge
Fam. Atemnidae – Atemnus Canestrini, Diplotemnus Chamberlin
Fam. Chernetidae – Allochernes Beier, Chernes Menge, Dendrochernes Beier,
Dinocheirus Chamberlin, Lamprochernes Töm., Lasiochernes Beier,
Pselaphochernes Beier
Fam. Cheliferidae – Beierochelifer Mahnert, Chelifer Geoffroy, Cheirochelifer
Beier, Dactylochelifer Beier, Hysterochelifer Chamberlin, Rhacochelifer Beier
Fam. Withiidae – Withius Kew
In the countries of Balkan Peninsula have been recorded pseudoscorpions as
follows:
Albania, 18 spp.; Bulgaria, 59 spp.; Serbia (whole, incl. Kosovo), 70 spp.;
Montenegro, 43 spp.(50 spp., according to N. Ćurčić and Dimitriević, 2016);
Republic of Macedonia, 45 spp.; Greece, 121 spp.; European Turkey, 5 spp.;
Croatia (whole), 112 spp.; Slovenia (whole), 31 spp.; and Bosnia and
Herzegovina, 55 spp. And these pseudoscorpions were recorded by several
authors: Beier (1929, 1931, 1939, 1963), Beron (1968, 1985), Ćurčić (many
papers since 1972), Ćurčić and Dimitriević (1984, 2006), Ćurčić et al. (2004),
N. Ćurčić and Dimitriević (2016), Ćurčić et al. (1993), Dumitresco and
Orghidan (1964), Georgescu and Capuşe (1994, 1996), Hadži (1940), Harvey
(2011), Mahnert (1973–1982), Ozimec (2000), Petrov (1997, 2000), Petrov
and Štáhlavský (2007), Redikorzev (1928), and Schmalfuss and Schawaller
(1984). For many species has been used the database of Harvey (2013f) but
have been extracted the subspecies and the repeated names (e.g., Chelifer
cancroides, Ch. cancroides cancroides).
Endemic genera for the peninsula are:
Fam. Chthoniidae
Neobalkanella Ćurčić, 2013 – Serbia (two spp.)
Troglochthonius Beier, 1939 – Bosnia and Herzegovina, cave (two spp.)
Fam. Neobisiidae
Insulocreagris Ćurčić, 1987 – Croatia (Vis Island), Herzegovina (two spp.)
Protoneobisium Ćurčić, 1988 – Croatia (one sp.)
Archaeoroncus Ćurčić, 2012 – Croatia (one sp.)
Ernstmayria Ćurčić et Dimitriević, 2006 – Crete (one sp.)
Besides the endemic genera (all considered relics), particularly interesting is the
finding of Hadoblothrus aegeus Beron in caves on Santorini Island – clearly relict
and the only member of Syarinidae found so far from Balkan Peninsula. The second
species of this genus is known from a cave in Southern Italy.
Opiliones On the Balkan Peninsula (including the entire territories of Slovenia and
Croatia, Northern (Romanian) Dobrudja, Eastern (Turkish) Thrace and the adjacent
islands in Aegean and Ionian seas) have been recorded 60–62 genera of Opiliones
from 9 families (Sironidae, Travuniidae, Phalangodidae, Phalangiidae,
Ischyropsalididae, Nemastomatidae, Trogulidae, Dicranolasmatidae,
Sclerosomatidae) (Boyer et al. 2005, Guéorguiev 1992, Hadži 1928, 1973a, 1973b,
Kratochvil 1937, 1946, Novak 2005b, Novak et al. 1995, 1996), Rambla 1968,
Roewer 1926, 1927b, 1940, 1959, Schönhofer 2009, Schönhofer and Martens 2009,
Schönhofer et al. 2009, Ubick and Ozimec 2005).
The following are a number of registered taxa of Opiliones in the Balkan
countries:
Albania (Mitov 2000, with suppl.; Murányi 2013, 2015) – 46 spp.
Bosnia and Herzegovina (Novak 2005a) – 54 valid species and 2 subspecies
Bulgaria (Beron and Mitov 1996, Juberthie 1991, Kratochvil 1958a, 1958b, Mitov
1994–2011, Roewer 1926, Staręga 1976, Šilhavý 1965) – 63 spp.
European Turkey (Gruber, 1969; Bayram et al., 2010) – 14 spp.
Continental Greece (Juberthie 1968, Martens 1972, Rambla 1968, Thaler 1996) –
the families Sironidae, Trogulidae, Dicranolasmatidae, Nemastomatidae,
Phalangiidae, Sclerosomatidae, Phalangodidae, Phalangiidae, Sclerosomatidae
(Siro, Phalangium L., Dicranopalpus Doleschall, Lacinius Thorell, Mitopus
Thorell, Egaenus C.L. Koch, Opilio Herbst, Oligolophus C.L. Koch,
Paroligolophus Lohmander, Dasylobus Simon, Graecophalangium Roewer,
Metaphalangium Roewer, Ramblinus Staręga, Rilaena Šilhavý, Lophopilio
Hadži, Megabunus Meade, Metaplatybunus Roewer, Platybunus C.L. Koch,
Zachaeus C.L. Koch, Stankiella Hadži, Rafalskia Staręga)
Crete (Martens 1965, 1966, Gruber 1998, Roewer 1927b, 1940; Šilhavý 1967) –
genera Trogulocratus Roewer, Konfiniotis Roewer, Anarthrotarsus Šilhavý,
Platybessobius Roewer, Dicranolasma Sørensen, etc., and the families
Trogulidae, Dicranolasmatidae, Phalangiidae, Sclerosomatidae
Aegean Islands (di Caporiacco 1928, 1948; Ghigi 1929; Gruber 1963, 1978;
Martens 1965; Roewer 1924) – 13 spp., mostly from Rhodes
Croatia (Novak 2004) – 64 valid species and 1 subspecies
Slovenia (Novak, Delakorda, and Novak, 2006) – 64 spp.
Republic Macedonia (Mitov 1995) – Graecophalangium drenskii Mitov – endemic
Northern Dobrudja (Dumitrescu 1970, 1972; Dumitrescu and Orghidan, 1964;
Babalean 1999, 2005) – 15 species; for Southern Dobrudja (in Bulgaria), Mitov
(2008) lists 17 species.
Cyphophthalmi As Karaman (2008) points out, the Balkan region “with New
Zealand and tropical South-East Asia, is one of the richest in cyphophthalmid on the
Earth.” From ca.140 spp. in suborder Cyphophthalmi, ca. 20 have been recorded
from Balkan Peninsula. Most species belong to Cyphophthalmus Joseph (on the
Balkans live 17 species of this genus).
Endemic genus and species Tranteeva paradoxa Kratochvil, 1958, is living in
the caves of Central Stara planina (Bulgaria). Karaman (2009) thinks that genus
Tranteeva should be synonymized with Cyphophthalmus.
The situation for the Balkan Peninsula, with an explosive evolution of only one genus of
Cyphophthalmi, is very different from the one for the Iberian Peninsula. This territory con-
tains four of the eight genera currently recognized in the family Sironidae, a generic diver-
sity and morphological disparity of Cyphophthalmi not found in any other region of the
world so far (Murienne and Giribet, 2009). Although the two European peninsulas have
usually been depicted as glacial refugia, we provide evidence that old endemic lineages in
these two territories have undergone very different diversifications: one – the Balkan
Peninsula – by hosting an old genus with subsequent explosive evolution; and the other –
the Iberian Peninsula – by hosting many ancient genera, each with few species. These dif-
ferences could be related to the very different palaeogeographic histories of the two
peninsulas. (Murienne et al. 2009)
As elsewhere in Europe, the bulk of Balkan Opiliones are the representatives of

“Palpatores.”
Eupnoi
Fam. Phalangiidae – Amilenus Martens, Dasylobus Simon, Dicranopalpus
Doleschall, Egaenus C.L. Koch, Lacinius Thorell, Lophopilio Hadži, Megabunus
Meade, Metaphalangium Roewer, Metaplatybunus Roewer, Mitopus Thorell,
Odiellus Roewer, Oligolophus C.L. Koch, Opilio Herbst, Phalangium L., Platybunus
C.L. Koch, Rilaena Šilhavý, Rafalskia Staręga, Zachaeus C.L. Koch,
Graecophalangium Roewer, Metadasylobus Roewer
Fam. Sclerosomatidae – Astrobunus Thorell, Gyas Simon, Leiobunum C.L. Koch,
Nelima Roewer, Homalenotus C.L. Koch, Mastobunus Simon
Dyspnoi
Fam. Ischyropsalididae – Ischyropsalis C.L. Koch (two spp., Slovenia)
Fam. Dicranolasmatidae – Dicranolasma Sørensen (five spp. from Corfu, Crete,
Bulgaria, Montenegro, and Herzegovina)
Fam. Nemastomatidae – Paranemastoma Redikorzev (11 spp., Bulgaria, Albania,
Herzegovina, Greece)
Fam. Trogulidae – Trogulus Latreille (13 spp., Bulgaria, Greece, Corfu, Bosnia-
Herzegovina, Macedonia, Croatia (Mljet, Hvar, Dubrovnik), Montenegro,
Serbia, Albania), Anarthrotarsus Šilhavý (one sp., Kerkira, Crete), Calathocratus
Simon (six spp., Russia, Crete, Rhodes), Konfiniotis Roewer (one sp., Crete)
Laniatores
Described from Balkan Peninsula is the family Travuniidae, known from Croatia,
Montenegro, Bosnia and Herzegovina, Serbia, Slovenia, Italy, France, Spain,
Switzerland, and Sardinia. From the 15 species in this family, 7 live on the penin-
sula, mostly in caves (Kury and Mendes 2007). From a cave in Serbia (near
Bulgarian border) was described the most interesting new genus and species
Trojanella serbica Karaman, 2005.
The family Phalangodidae is known from Bulgaria with the endemic genus and
species Paralola buresi Kratochvil, 1958 (four caves in western Stara Planina). In
Greece have been described three spp. of Ausobskya (Martens 1972, Thaler 1996).
One new species of Ausobskya was found by us in Belasitsa Mountain (Bulgaria, on
the border with Greece), in leaf litter.
Endemic genera of Opiliones on the Balkan Peninsula are:
[Tranteeva Kratochvil, 1958] – Bulgaria (one sp.) (= Cyphophthalmus?)
Paralola Kratochvil, 1958 – Bulgaria (one sp.)
Lola Kratochvil, 1937 – Croatia (Hvar) (one sp.)
Ausobskya Martens, 1972 – Greece, Bulgaria (four spp.)
Trojanella I. Karaman, 2005 – Serbia (one sp.)
Travunia Absolon et Kratochvil, 1932 – Bosnia and Herzegovina, Croatia,
Montenegro (four spp.)
Dinaria Hadži, 1932 – Bosnia and Herzegovina (one sp.)
Anarthrotarsus Šilhavý, 1967 – Greece (one sp., Kerkira, Crete)
Kofiniotis Roewer, 1940 – Greece (one sp., Crete)
Hadzinia Šilhavý, 1966 – SW Bosnia (one sp.)
Araneae On the Balkan Peninsula (in the dimensions described above) have been
registered over 1409 species of spiders of 337 genera and 48 families: Atypidae,
Ctenizidae, Nemesiidae, Filistatidae, Sicariidae, Scytodidae, Leptonetidae,
Pholcidae, Dysderidae, Segestriidae, Oonopidae, Palpimanidae, Mimetidae,
Eresidae, Oecobiidae, Uloboridae, Nesticidae, Theridiosomatidae, Anapidae,
Mysmenidae, Araneidae, Linyphiidae, Tetragnathidae, Theridiidae, Lycosidae,
Agelenidae, Pisauridae, Oxyopidae, Zoropsidae, Cybaeidae, Argyronetidae,
Desidae, Hahniidae, Amaurobiidae, Dictynidae, Titanoecidae, Anyphaenidae,
Clubionidae, Liocranidae, Gnaphosidae, Philodromidae, Thomisidae, Salticidae,
Corinnidae, Zodariidae, Prodidomidae, Miturgidae (Zorinae), and Sparassidae
(Bayram 2002, Bayram et al. 2014, Delchev 2000, 2004, Deltshev and Blagoev
2001, Deltshev et al. 2005, Schröder et al. 2011, Tatole 2006).
Albania (Deltshev et al. 2011) – 335 spp. of 36 families
Bulgaria (Deltshev and Blagoev 2001; Deltshev 1996, 1999, 2005, 2011) – >
1000 spp.
Bosna and Herzegovina (Komnenov, 2009)
European Turkey (Topçu, Demir, and Seyyar 2005; Demircan and Topçu
2016) – 197 spp.
Continental Greece (Bristowe 1934, Brignoli, many papers, Metzner 1999,
Bosmans and Chatzaki 2005; Chadzaki et al. 2015) – in mid-2015 from the entire
territory of Greece are known 1100 spp. of 50 families.
Crete (Bosmans et al. 2013) – 430 spp. (incl. 57 endemic), 44 families
Croatia – 614 species
Republic of Macedonia (Blagoev 2002; Komnenov 2006) – 558 spp. of 36
families
Serbia (incl. Kosovo) (Deltshev, Ćurčić and Blagoev 2003; Ćurčić et al. 2007) –
638 spp. of 224 genera and 36 families
The first catalogue of spiders of the Balkan Peninsula was the one of Drensky
(1936) with 1066 spp. of 35 families. Modern zoogeographical analyses were made
by Deltshev (1999, 2000, 2004, 2008) and others.
According to Deltshev (1999), the spider fauna of the Balkan Peninsula is repre-
sented by 1409 species of 337 genera and 48 families (now they are more). This
number is relatively high, compared with the 1001 spp. of the huge Russian Plain
and the 925 spp. of Germany. Deltshev is classifying these spiders in 24 zoogeo-
graphical categories in 4 main groups:
1 . Widely distributed species (533, or 38,1% of all), mainly Palearctic
2. Balkan endemics (379 spp., 26,9%), largely due to the many thousands of caves
and the complex orography of the peninsula, with high mountains and many
islands
3. The European complex includes 300 species (21,3%)
4. The Mediterranean complex counts 195 spp. (13,8%), but part of the endemics
are also of Mediterranean origin.
The conclusion of Deltshev is that “…the Balkan Peninsula represents one of the
main centres of speciation in Europe.”
Again after Deltshev (2004), on the Balkan Peninsula, there are 379 endemic spe-
cies of spiders, but now, largely thanks to researchers like Deltshev, Lazarov,
Komnenov, Chadzaki, Bosmans, and others, this figure is increased (Deltshev, in. lit.).
Some endemic genera of spiders on the Balkan Peninsula are:
Fam. Linyphiidae
Antrohyphantes Dumitrescu, 1971
Fageiella Kratochvil, 1934
Fam. Dysderidae
Dysderocrates Deeleman-Reinhold et Deeleman, 1988 (Balkans)
Rhodera Deeleman-Reinhold, 1989 (Crete) (one sp.)

Folkia Kratochvíl, 1970 (Balkans) (seven spp.)
Minotauria Kulczyn’ski, 1903 (Crete) (two spp.)
Stalagtia Kratochvíl, 1970 (Balkans, Greece)
Parastalita Absolon et Kratochvíl, 1932 (Bosnia-Herzegovina) (one sp.)
Stalita Schiödte, 1847 (Balkans)
Stalitella Absolon et Kratochvíl, 1932 (Balkans) (one sp.)
Fam. Liocranidae
Arabelia Bosselaers, 2009 (one sp.), Vankeeria Bosselaers, 2012 (one sp.) – Greece
Fam. Agelenidae
Hadites Keyserling, 1862 – Croatia (one sp.)
Fam. Leptonetidae
Barusia Kratochvil, 1978 (Croatia, Greece, Montenegro)
Sulcia Kratochvil, 1938 (Balkans, Greece)
Fam. Pholcidae
Stygopholcus Absolon et Kratochvil, 1932 – Croatia, Bosnia-Herzegovina, Greece,
Montenegro
Fam. Hahniidae
Cryphoecina Deltshev, 1997 – Montenegro (one sp.)
According to Deltshev (2000), “The endemic taxa of spiders (Araneae) in the
Balkan peninsula are represented by 348 species included in 30 families. Countries
with the highest number of recorded endemic species are Greece (115), Croatia
(68), Bulgaria (55), Bosnia (41), Crete (46).” More than 160 of these species are
from caves. Since 2000 some more taxa have been described. Again from Deltshev
(2000),
This contribution is the result of a critical data available for the endemic spiders of the
Balkan peninsula territory and comprises 348 species from 30 families: Ctenizidae 5,
Nemesiidae 4, Pholcidae 9, Leptonetidae 21, Segestriidae 2, Dysderidae 7, Oonopidae 1,
Palpimanidae 1, Uloboridae 1, Nesticidae 6, Theridiidae 5, Anapidae 1, Mysmenidae 1,
Linyphiidae 109, Tetragnathidae 2, Araneidae 1, Lycosidae 1, Agelenidae 29, Cybaeidae 1,
Hahniidae 5, Dictynidae 1, Amaurobiidae 17, Liocranidae 4, Clubionidae 3, Zodariidae 8,
Gnaphosidae 18, Miturgidae (Zorinae) 1, Philodromidae 2, Thomisidae 4, Salticidae 12.
The established number is high and represents 25% of all spiders of the Balkan peninsula.
The most characteristic families are: Linyphiidae s. l. (31.3%), Dysderidae (21%) and
Agelenidae (8.3%). The genus Troglohyphantes is the most numerous and can be regarded
as a faunistic phenomenon since from all 53 species established in the territory of the
Balkans, 52 are endemics, distributed mainly in caves…The high percentage of endemic
spiders (25%) suggests an important process of autochthonous speciation. So the Balkan
Peninsula can be considered as a main center of speciation in Europe.
The bigger number (32) and percentage (74.41%) of Bulgarian endemics and
certainly the presence of paleoendemic genus Antrohyphantes show the importance
of eastern part of Balkan Peninsula for the speciation in the region. The local ele-
ments comprise paleoendemics (mainly in caves) and neoendemics (mainly in high-
altitude zones). The endemics belong to two principal faunistic complexes:
Mediterranean and European (Deltshev 1996).
In his analysis of the biogeographical connections between the spiders of the

Balkans and the Middle East, Brignoli (1986) concludes that these territories “…
forment une unité biogéographique dont la faune aranéologique est significative-
ment différente de celle de la “Mitteleuropa” lato sensu et, au même temps, est
considerablement semblable à celle de la Méditerranée occidentale.”
Opilioacarida The order is known only from the southernmost part of the penin-
sula (Peloponnese and the islands Karpathos and Kassos), with one species –
Opilioacarus segmentatus With, 1903. The species is known also from Algeria,
Sicily, and Rhodes (Beron 1990 2014, Thaler and Knoflach 2002). These localities
mark the northern limit of the modern Opilioacarida in Europe, but as fossils two
species have been described from Baltic amber.
Acari From Bulgaria are recorded 1673 species of Acari (Acariformes and
Parasitiformes) (Beron 2011, actually about 1700 spp.).
Acariformes
Trombidiformes
Prostigmata
Fam. Erythraeidae
Endemic genera and subgenera:
Helladerythraeus (Beron, 1988) – Rhodes and Kythnos Islands
Myrmicotrombium (Graecotrombium) mirum Beron, 1990 – Greece
Parasitiformes Ticks are of some zoogeographical interest. On Balkan Peninsula
have been recorded five spp. of Argasidae. Ixodidae are recorded in various Balkan
countries like Bulgaria (39 spp.), Turkey (38), Romania (25), Serbia (25), Bosnia
and Herzegovina (20), Montenegro (18), and Croatia (13) (Drenski 1955, Beron
2011, 2014, Feider 1965, Georgieva and Gecheva 2013)
7.4 Apennine Peninsula
The central and the smallest of the three large peninsulas of Southern Europe spans
1000 km from the Po Valley in the north to the central Mediterranean Sea in the
south. The backbone of the peninsula consists of the Apennine Mountains, high up
to 2912 m (Corno Grande del Gran Sasso). The surface of the peninsula is
131,337 km2, and the climate is mediterranean. The mountains are mostly verdant,
although one side of the highest peak, Corno Grande, is partially covered by
Calderone glacier, the only glacier in the Apennines. The Apennines were created in
the Apennine orogeny beginning in the Early Neogene (about 20 mya, the Middle
Miocene) and continuing today (Gueguen et al. 1998, Rook et al. 2006).
7.4 Apennine Peninsula 343
La Greca (1962) outlined several “types of distribution” of elements in the Italian

fauna (enumerated further). This author analyzed many other zoogeographical
aspects of the insects in Italy (La Greca 1955, 1958, 1966).
In the peninsular part of Italy are recorded the orders Palpigradi, Scorpiones,
Pseudoscorpiones, Opiliones, Araneae, Opilioacarida, and different other Acari.
Lacking are the “southern” orders – Ricinulei, Amblypygi, Thelyphonida (Uropygi),
Schizomida, Solifugae, Holothyrida – and the suborders Mesothelae and
Palaeoamblypygi. Some orders are well represented – pseudoscorpions (226 spp.)
and spiders (1620 spp.) – for the entire territory of Italy.
Palpigradi So far from continental Italy have been recorded five spp. of Palpigradi,
including the discovery of the order by Grassi and Calandruccio (1885). They all
belong to Eukoenenia Börner (Condé 1976, Silvestri 1905).
Scorpiones On the continental Apennine Peninsula are recorded, after many

changes, four species of Scorpiones (Crucitti 1993, Fet 2010).
Fam. Euscorpiidae
Euscorpius (Polytrichobothrius) italicus (Herbst, 1800)
E. (Alpiscorpius) germanus (C.L. Koch, 1837) – populations in the Apennines
are relict or introduced (Fet et al., 2004)
E. (Euscorpius) sicanus (C.L. Koch, 1837)
E. (Tetratrichobothrius) flavicaudis (DeGeer, 1778)
Pseudoscorpiones According to the catalogue of Gardini (2000), in peninsular
Italy (without the Alpine part and the islands) were recorded until the end of 1998
pseudoscorpions of the families Chthoniidae, Neobisiidae, Syarinidae, Garypidae,
Geogarypidae, Cheiridiidae, Olpiidae, Atemnidae, Chernetidae, Cheliferidae, and
Withiidae (Beier, many papers, Gardini 2015, Lazzeroni 1970, and other papers).
As a whole, from Italy are known 226 spp. (Lissner 2014) (first place in Europe).
Opiliones According to Stoch (2003), in the total of Italian territory, there are 11
families of Opiliones with 43 genera and 120 species (Brignoli 1968, Canestrini
1872, Chemini 1990, 1995, 1996, Gruber 1965, 1985, Marcellino 1968, 1971, 1982,
1983, 1984, 1986, Staręga 1976, Šilhavý 1969).
Cyphophthalmi
On the Peninsula have been registered Siro valleorum Chemini, 1990 (Lombardia).
Eupnoi
From continental Italy:
Fam. Phalangiidae – Phalangium L., Dicranopalpus Doleschall, Lacinius Thorell,
Mitopus Thorell, Egaenus C.L. Koch, Opilio Herbst, and others
Fam. Sclerosomatidae – Metasclerosoma Roewer, Astrobunus Thorell, Homalenotus
C.L. Koch
Dyspnoi
Fam. Ischyropsalididae – Ischyropsalis C.L. Koch (five spp.)
Fam. Trogulidae – Trogulus Latreille (two spp.)
Fam. Dicranolasmatidae – Dicranolasma C.L. Koch (one sp.)
Fam. Nemastomatidae – Nemastoma C.L. Koch, Saccarella Schönhofer et Martens
(two spp.)
Laniatores
Ptychosoma vitellinum Soerensen, 1873 (Phalangodidae), was reported by Brignoli
(1968) from Southern Italy.
Araneae The list of Stoch (2003) indicates that in the total of Italian territory, there
are 45 families of spiders of 323 genera and 1411 species. According to the cata-
logue of Pantini and Isaia (2015), there are 54 families with 426 genera, 1620 spe-
cies, and 26 subspecies. So, there is a considerable difference. Brignoli (1981)
analyzed the entire Italian fauna of Araneae, known up to this time.
Opilioacarida The two European species of the order Opilioacarida Opilioacarus

italicus With, 1904, and O. segmentatus With, 1903, are represented in Italy
(Brignoli 1967, With 1904).
7.5 Iberian Peninsula and the Pyrenees
On the westernmost of the three big peninsulas in South Europe are situated two
states – Spain and Portugal. Iberian Peninsula has an area of approximately
580,000 km2. On the southern end, near the Moroccan coast of Africa, is situated the
British overseas territory of Gibraltar. To the north the Pyrenees (highest point Pico
de Aneto, 3404 m) forms the border with France (incl. the tiny state of Andorra) and
stretches from the Atlantic Ocean to the Mediterranean Sea over 491 km. In the
southern part (Andalusia) raises the highest mountain of the Peninsula Sierra
Nevada (highest point Mulhasén, 3478 m). About ¾ of the peninsula is taken by a
plateau called Meseta Central.
I had the chance to study the highest parts of Sierra Nevada, “...das südlichste
Hochgebirge Europas und zugleich das höhste Gebirge der Iberischen Halbinsel”
7.5 Iberian Peninsula and the Pyrenees 345
(Franz 1979). It is a very dry mountain, with precipitations at 1800 m of only

6735 mm, compared to the 1130 mm at the same altitude on Vitosha (Bulgaria).
The difference between the treeless Sierra Nevada and the humid, green Pyrenees
is striking. The autoroute ends below the very top of Veleta (3428 m), and one is
among the more humid upper parts of the mountain, among the southernmost snow
fields, glaciers, and mountain lakes in continental Europe (ca. 37° N) (Höllermann
1972). The botanists have discovered here a mixed community of glacial relicts and
betic variants of Alpine plants.
The interesting studies of Janetschek (1957), Franz (1979), as well as my own
observations show that some typically alticolous species of spiders, opilions, pseu-
doscorpions, and mites reach the top of the mountain (Mulhacen, 3482 m).
Situated in the westernmost part of Europe and near Africa, rich in caves and with
some of the highest mountains of the continent, the Iberian Peninsula and the
Pyrenees have varied arachnofauna with many relicts and strange taxa. Such is the
enigmatic scorpion Belisarius Simon, a relative of South American scorpions.
Other such relicts are the pseudoscorpions of the family Bochicidae – two cave-
inhabiting genera (Titanobochica Zaragoza et Reboleira and Troglobisium Beier)
belong to one clearly American family. Remarkable is the presence of three endemic
genera of suborder Cyphophthalmi, forming one of the most important centers of
speciation of this suborder of Opilions. In Spain are registered scorpions of genus
Buthus Leach, unknown in other parts of Europe. Similar is also the situation with
the Solifugae (Daesiidae) – though the peninsula is passing the northern limit of
distribution of this order. Iberian endemics are 236 species of spiders. The paper of
Bacelar (1928) “Aracnidios Portuguèses” contains lists of 330 spp. of spiders, 13
pseudoscorpions, 1 scorpion, 28 opilions, and only 45 spp. of acari of three families.
Now they are many more.
Barranco et al. (2014) recorded from Spain the imported schizomid Stenochrus
portoricensis Chamberlin.
Palpigradi In the peninsula (all from Spain and from the two slopes of the
Pyrenees) have been recorded Eukoenenia bouilloni Condé, 1980; E. brolemanni
Hansen, 1926; E. draco zariquieyi Condé, 1951; E. hispanica Peyerimhoff, 1908; E.
pyrenaella Condé, 1990; E. pyrenaica Hansen, 1926; E. gadorensis Mayoral et
Barranco, 2002; and E. mirabilis Grassi et Calandruccio, 1885. From continental
Portugal is known E. mirabilis (Condé 1951, 1980, 1990, Hansen 1926, Mayoral
and Barranco 2002b, Peyerimhoff 1908).
Solifugae In Iberian Peninsula is known Gluvia dorsalis (Latreille, 1817)

(Daesiidae) (Spain, Portugal) (Pocock 1903, Werner 1925).
Scorpiones In the continental part of Iberian Peninsula (incl. Pyrenees) are

recorded the following scorpions (Simon 1879, Werner 1925, Lourenço and Vachon
2004, Fet 2010):
Fam. Buthidae
Buthus occitanus (Amoreux, 1789) – Spain, Portugal (recorded erroneously also
from Greece and other areas)
B. ibericus Lourenço et Vachon, 2004 – Spain
B. montanus Lourenço et Vachon, 2004 – Spain
Fam. Euscorpiidae
Euscorpius (Tetratrichobothrius) flavicaudis (DeGeer, 1778) – Western
Mediterranean
Fam. ? Troglotayosicidae
Belisarius xambeui Simon, 1879 – Pyrenees in France and Spain (Catalonia)
Pseudoscorpiones In continental Spain, Portugal, and the whole of the Pyrenees
are known pseudoscorpions of 45 genera and the families Chthoniidae, Neobisiidae,
Syarinidae, Bochicidae, Larcidae, Olpiidae, Geogarypidae, Garypinidae, Garypidae,
Cheiridiidae, Atemnidae, Chernetidae, Cheliferidae, and Withiidae (Beier 1939,
1959, 1961, Mahnert 1977, Navás 1925, Nonidez 1917, Vachon 1940, Zaragoza
1986, 2000, full bibliography; 2007, 2010).
With 196 spp. (Lissner 2014, actualized 2017 – 213 spp.), Spain (incl. Canary
Islands) is the second richest country in Europe. From Portugal have been recorded
62 spp. of 13 families.
The new pseudoscorpion genus Lusoblothrus of the family Syarinidae is described from a
cave in the Algarve region, southern Portugal, to accommodate L. aenigmaticus sp. nov.,
whose morphological affinities within the Holarctic syarinid fauna are not clear and resem-
bles the Gondwanan genera. This discovery emphasizes the relevance of the Algarve region
as a hotspot for relictual hypogean fauna within the Iberian Peninsula. (Reboleira et al. 2012)
Endemic genera for the peninsula are:

Titanobochica Zaragoza et Reboleira, 2010 – Portugal (T. magna Zaragoza et
Reboleira, 2010)
Arcanobisium Zaragoza, 2010 – Spain (A. comasi Zaragoza, 2010)
Troglobisium Beier, 1939 – Spain [T. racovitzai (Ellingsen, 1912)]
Lusoblothrus Reboleira et al., 2012 – Portugal [L. aenigmaticus Reboleira et al.,
2012]
Opiliones According to Rambla (1973), by this time the opilionid fauna of the
Iberian Peninsula consisted of 138 spp., including 4 Cyphophthalmi, 8 Laniatores,
and 125 “Palpatores” (Eupnoi and Dyspnoi). Since this list has been completed by
de Bivort and Giribet (2004), now the figures are 5 Cyphophthalmi, 8 Laniatores,
7.5 Iberian Peninsula and the Pyrenees 347
and 125 “Palpatores.” Endemic to the peninsula are three genera. For Cyphophthalmi
the Iberian Peninsula is a hot spot (Murienne and Giribet 2009).
The recent treatments of the Opiliones of the peninsula and the Baleares are
113 spp. and 79 endemic spp. (Mello-Leitão 1936, Prieto 2003, 2008 and suppl.,
based on Juberthie 1956, 1961, 1962, Kraus 1961, Prieto 1990a, 1990b, 2004,
Rambla 1967, 1977b, Rambla and Fontarnau 1984, 1986, etc.).
Cyphophthalmi
Fam. Sironidae
Odontosiro Juberthie, 1961 – Portugal (one sp.) (endemic genus)
Iberosiro de Bivort et Giribet, 2004 – Portugal (one sp.) (endemic genus)
Paramiopsalis Juberthie, 1962 – Portugal, Spain (two spp.) (endemic genus)
Cyphophthalmus [Siro] Joseph, 1868 – Portugal (two spp.)
Parasiro Hansen et Sorensen, 1904 – Spain (two spp.)
Eupnoi
Fam. Phalangiidae
Metaphalangium Roewer, 1911; Odiellus Roewer, 1923; Phalangium L., 1758;
Eudasylobus Roewer, 1911; Mitopus Thorell, 1876; Dicranopalpus
Doleschall, 1852
Fam. Sclerosomatidae
Homalenotus C.L. Koch, 1839; Cosmobunus Simon, 1879; Leiobunum
C.L. Koch, 1839; Mastobunus Simon, 1879; Micronelima Schenkel, 1938
Dyspnoi
On the peninsula are known 25 spp. (Schönhofer, 2009, 2013).
Fam. Ischyropsalididae – Ischyropsalis C.L. Koch, 1839 (eight spp. from Spain,
three from Portugal)
Fam. Sabaconidae – Sabacon Simon, 1879 (six spp.)
Fam. Nemastomatidae – Acromitostoma Roewer, 1951 (two spp., endemic genus)
Centetostoma Kratochvil, 1958 (three spp.)
Fam. Trogulidae – Trogulus Latreille, 1802 (three spp.)
Laniatores
On the peninsula: fam. Phalangodidae with eight spp.:
Ptychosoma Sørensen, 1873 – P. espanoli (Rambla, 1975), Spain
Scotolemon Lucas, 1860 – seven spp. from Spain and Portugal
According to Rambla (1974), on the peninsula have been recorded 138 spp. of
Opiliones, including 56 endemics. From the remaining 82 spp., 24% are of
Mediterranean type and 8% have North African affiliations. Since 1974 several new
taxa have been added to this fauna (Iberosiro de Bivort et Giribet, 2004 and others),
and many taxonomic changes took place (Prieto 2008) but the general picture did
not change too much.
Araneae According to Cardoso and Morano (2010), “At present [end 2009], 1335
species are known from the region, of which 236 are Iberian endemics, in 373
genera and 55 families. Portugal presents 768 species and Spain (including Andorra
and Gibraltar), 1213 species. Although linyphiids present the highest number of
known species (267), dysderids present the highest endemic richness (46 species).”
Speleoharpactea Ribero is an endemic genus in Spain.
Bosmans and de Keer (1985, 1987) prepared a catalogue of the spiders of the
Pyrenees and zoogeographical analysis. Their list contained 860 spp. of 43 families
(238 Linyphiidae, 92 Gnaphosidae, 73 Salticidae, 60 Theridiidae, 53 Lycosidae, 44
Thomisidae), or these 6 families include 560 spp. and the remaining 37 families
include 300 spp. At or above 2200 m have been found at least 170 spp. and above
3000 m at least 22 spp. of 6 families, including 10 Linyphiidae. The highest alti-
tudes are reached (after these authors) by Savignia superstes (3290 m), Gnaphosa
atramentaria (3190 m), and Talavera petrensis (3180 m). However, I have found
spiders on top of the highest summit (Aneto, 3704 m). The highest spiders of Sierra
Nevada have been studied by Denis (1957).
Alderweireldt and Bosmans (2001) completed the araneofauna of Portugal to
649 spp. of 43 families. The linyphiid fauna of Portugal was reviewed by Bosmans
et al. (2010).
7.6 Mediterranean Sea and Its Islands
Map 7.8 Mediterranean Sea and its islands
The sea between Africa, Europe, and Western Asia inherited what was once the
western arm of the Tethys Sea. Its surface is ca. 2.5 million km2. The sea is con-
nected with the Atlantic Ocean by Gibraltar Strait (14 km wide) and is sometimes
7.6 Mediterranean Sea and Its Islands 349
considered part of the Ocean (and Marmara and Black seas are considered parts of
the Mediterranean).
In the Middle Miocene times, the collision between the Arabian microplate and
Eurasia led to the separation between the Tethys and the Indian oceans. This process
resulted in profound changes in the oceanic circulation patterns, which shifted
global climates toward colder conditions. The Hellenic arc, which has a land-locked
configuration, underwent a widespread extension for the last 20 Ma.
The opening of small oceanic basins of the central Mediterranean follows a
trench migration and back-arc opening process that occurred during the last 30 Myr.
This phase was characterized by the anticlockwise rotation of the Corsica-Sardinia
block, which lasted until the Langhian (ca.16 Ma). Subsequently, a shift of this
active extensional deformation led to the opening of the Tyrrhenian basin.
The Betic-Rif mountain belts developed during the Mesozoic and Cenozoic
times, as Africa and Iberia converged. One of the most important events in geologi-
cal history of Mediterranean Sea is the so-called Messinian Salinity Crisis (Hsü
et al. 1973). There are several reasons why the sea, closed in the Miocene, is drying
out during the end of the Messinian phase 5.96 to 5.33 Ma. The sea level drops by
several kilometers below the level of the World Ocean. On the sea bottom remain
only several hypersaline “pools.” After 5.5 Ma the rivers import more water, and the
hypersaline “dead seas” are diluted, until by 5.33 Ma opens the Gibraltar Strait. The
huge waterfall bringing Atlantic water into the Mediterranean bed is called since
1972 the Zanclean Deluge. After some researchers, the waterfall, although in
gradins, has been higher from the highest present-day waterfall in the world (Angel
Falls – 979 m) and more mighty than Iguacu or Niagara. It has been calculated the
filling of the sea during 2 years (may be less) with a water volume 1000 times bigger
than Amazonia, and the level raises by 10 m a day.
The dry bed of the Mediterranean is of crucial importance for the zoogeography.
Many different animals living on the former sea bottom found asylum on the
present-day islands (Malta, Sicily, and Cyprus). Some researchers believe that all
animals have crossed to the islands by swimming or rafting (at least for the ele-
phants, it seems rather unbelievable).
Mediterranean Sea has been since millennia a crossroad of different activities
and intense navigation. Even now approximately 220,000 merchant vessels of more
than 100 tonnes cross the Mediterranean Sea each year – about one third of the
world’s total merchant shipping. This is connected also with dispersion of animal
species between Mediterranean countries and with pollution of the sea and its
shores. Every year between 100,000 and 150,000 t of crude oil are deliberately
released into the sea from shipping activities.
The climate changes reflect on the Mediterranean climate and on the sea level.
Sea level rise for the next century (2100) is expected to be between 30 and 100 cm.
A rise of 30 cm would flood 200 km2 of the Nile Delta and many other low
territories.
The sea is entirely within the Mediterranean Subregion of Palearctic and has many
bigger islands (Sicily, Sardinia, Corsica, Balearic Islands, Crete, Cyprus, many Greek
and Dalmatian islands, and many smaller islands around Italy). The arachnofauna of
these islands is relatively well studied and contains many interesting endemics.
Map 7.9 Secondary subdivision of the Primary Mediterranean Center (De Lattin 1967)
1. Atlantomediterranean Secondary Center; 2. Adriatomediterranean SC; 3. Pontomediterranean
SC; 4. Tyrrhenian SC; 5. Canarian SC; 6. Mauretanian SC; 7. Cyrenaican SC; 8. Cretan SC;
9. Cyprian CS
The climate changes reflect on the Mediterranean climate and on the sea level.
Sea level rise for the next century (2100) is expected to be between 30 and 100 cm.
A rise of 30 cm would flood 200 square kilometers of the Nile Delta and many other
low territories.
In the world there are several “Mediterranean regions” (Eurafrican Mediterranean,
California, Chile, New South Wales in Australia, the Cape area in South Africa).
They are similar in their climate and other patterns. Their biogeographic character-
istics have been outlined by Vitali-di-Castri (1973).
7.7 Balearic Islands
An archipelago in the Western Mediterranean was formed by four largest islands

(Mallorca, 3640 km2; Menorca, 696 km2, highest point 358 m; Ibiza, 572 km2, high-
est point 475 m; and Formentera, 83 km2) and several smaller islands (Cabrera, etc.)
The last two larger islands form a group, called Pityusas. The highest point of the
archipelago is on Mallorca (1445 m in the Serra de Tramuntana) (Colom 1957,
Gautier et al. 1994, Pons and Palmer, 1996).
7.7 Balearic Islands 351
During the Last Glacial Maximum, the Emile Baudot Escarpment and the two
submarine mounts, dels Oliva and Ausiàs Marc, formed a microarchipelago between
the “Pitiusa” and the “Gran Balear” (Mateu et al. 2004).
In their review of the endemic animals of Balearic Islands, Pons and Palmer (1996)
enumerate 1 species of scorpions (a valid species Euscorpius balearicus Caporiacco,
1950, not subspecies of “carpathicus”), 1 sp. of Palpigradi, 13 spp. or ssp. of
Pseudoscorpiones, 4 of Opiliones, 32 spp. of spiders, and 5 spp. of Acari. To them
must be added Anelasmocephalus balearicus Martens et Chemini, 1988 (missed),
and Trogulus balearicus Schönhofer et Martens, 2008 (described later).
Palpigradi Peyerimhoff (1906) described from Mallorca one of the first known
Palpigradi – Eukoenenia draco draco. Another subspecies (E. draco zariquieyi
Condé, 1951) has been described from Catalonia.
Scorpiones The Balearic scorpion was defined as valid autochthonous and

endemic species Euscorpius balearicus Caporiacco, 1950 (Mallorca, Menorca,
Cabrera, Dragonera) (Caporiacco 1950, Gantenbeim et al. 2001).
Pseudoscorpiones On the islands are registered pseudoscorpions of 18 genera and

the families Chthoniidae, Neobisiidae, Syarinidae, Garypidae, Geogarypidae,
Atemnidae, Olpiidae, Cheliferidae, Chernetidae, and Withiidae (Beier 1959, 1961,
Estany 1977, Lagar 1972, Mahnert 1977, 1978b, 1989, 1993, Tullgren 1900,
Zaragoza 2000, Zaragoza and Vadell 2013, Pons and Palmer 1996).
Endemics are:
Chthonius campaneti Zaragoza et Vadell, 2013 (Mallorca, cave)
Chthonius (Ephippiochthonius) cabreriensis Mahnert, 1993 (Cabrera, Ila del
Connils)
Ch. (E.) ponsi Mahnert, 1993 (Cabrera)
Ch. (E.) balearicus Mahnert, 1977 (Mallorca, troglophile)
Ch. (E.) bellesi Mahnert, 1989 (Menorca, troglobite)
Acanthocreagris balearica Beier, 1961 (Menorca)
Neobisium monasterii Mahnert, 1977 (Mallorca)
N. ischyrum balearicum Beier, 1939 (Menorca, Eivissa, Cabrera)
Roncus neotropicus Redikorzev, 1937 (?= R. balearicus Beier, 1961) (Mallorca,
Eivissa)
R. vidali Lagar, 1972 (Mallorca, troglobite)
Allochernes pityusensis Beier, 1961 (Eivissa)
Pselaphochernes balearicus Beier, 1961 (Mallorca)
Dactylochelifer balearicus Beier, 1961 (Menorca, Eivissa)
D. besucheti Mahnert, 1976 (Mallorca)

Opiliones On the Balearic Islands are known nine genera of Opiliones (two
Laniatores and seven “Palpatores”) of eight genera (Ptychosoma, Scotolemon,
Trogulus, Anelasmocephalus, Homalenotus, Odiellus, Phalangium, Paropilio,
Leiobunum) and the families Phalangodidae, Trogulidae, Nemastomatidae,
Phalangiidae, and Sclerosomatidae (Martens and Chemini 1988; Prieto 2003;
Rambla 1972, 1977a, 1977b, 1979; Schönhofer and Martens 2008).
Eupnoi (endemics)
Fam. Phalangiidae
Phalangium clavipus Roewer, 1911 – Mallorca
Eudasylobus ferrugineus (Thorell, 1876) – Mallorca and Ibiza
Metaphalangium abstrusum (L. Koch, 1882) – Baleares
Dyspnoi (endemics)
Fam. Trogulidae
Anelasmocephalus balearicus Martens et Chemini, 1988 – Mallorca
Trogulus balearicus Schönhofer et Martens, 2008 – Ibiza
Laniatores
Endemic (and only) Laniatores (Phalangodidae) are Scotolemon krausi Rambla,
1972, on Ibiza and Ptychosoma balearicum Rambla, 1977, on Mallorca (cave).
Araneae
According to the Iberian spider checklist of Cardoso and Morano (2010), there are
185 species of 122 genera and 37 families of spiders known from Balearic Islands
(Cardoso and Morano 2010, Melic 2001, Morano 2004). The most numerous fami-
lies are Linyphiidae (18 gen., 21 spp.), Salticidae (14 gen., 19 spp.), Araneidae (13
gen., 20 spp.), and Theridiidae (13 gen., 26 spp.). The endemic genus is Chatzakia
Lissner et Bosmans, 2016 (Gnaphosidae)
7.8 Sicily, Sardinia, Corsica, and Elba
The three big islands Sicily, Sardinia, and Corsica and the smaller nearby islands of
the Tuscan Archipelago (Elba, 223.5 km2, Montecristo, Carpalia, Pianos, etc.) as
well as the other small archipelagoes (Aeolian and Pontine islands) are important as
parts of the former microplate, bridge with North Africa (of Sicily) and large enough
area of speciation (Shu 1974, Shu et al. 1973, Shu et al. 1977, La Greco 1957,
1961). Their mountains are high (Volcano Etna in Sicily, 3329 m; Monte Cinto in
Corsica, 2706 m; and Punta La Marmora, 1834 m in Sardinia).
7.8 Sicily, Sardinia, Corsica, and Elba 353
Corsica was formed approximately 250 million years ago with the uplift of a
granite backbone on the western side. It is the most mountainous island in the
Mediterranean, a “mountain in the sea” (with 20 summits of more than 2000 m).
Approximately 3500 km2 of the total surface area of 8680 km2 is dedicated to nature
reserves (Parc Naturel Régional de Corse). The island is 90 km from Italy and
170 km from the Côte d’Azur in France. It is separated from Sardinia to the south
by the Strait of Bonifacio, a minimum of 11 km wide.
Sardinia is the second largest island in the Mediterranean Sea, with an area of
23,821 km2, with Gennargentu Ranges at the center of the island.
Sicily is separated from the Italian region of Calabria by the Strait of Messina,
about 3 km wide in the north and about 250 m deep. The total area of the island is
25,711 km2, while the autonomous region of Sicily (which includes smaller sur-
rounding islands) has an area of 27,708 km2.
Along the northern coast, mountain ranges of Madonie (2000 m), Nebrodi
(1800 m), and Peloritani (1300 m) represent an extension of Apennines (Fig. 7.1).
Sicily and its small surrounding islands have some highly active volcanoes. Etna
covers an area of 1190 km2. The Aeolian Islands in the Tyrrhenian Sea, to the north-
east of mainland Sicily, exhibit a volcanic complex including Stromboli. Currently
active also are the three volcanoes of Vulcano, Vulcanello, and Lipari, usually dor-
mant. The small islands in the Strait of Sicily (Lampedusa, Linosa, and Pantelleria)
have been described in the paper of Massa (Ed.). The only island connected with
Africa back to 18,000 years ago is Lampedusa (20.2 km2).
Fig. 7.1 Mount Etna rising over suburbs of Catania (Wikipedia)

On these islands are recorded 11 orders and 8 suborders of Arachnida. Lacking are the
orders Ricinulei, Amblypygi, Schizomida, Thelyphonida, and Holothyrida and the
suborders Palaeoamblypygi and Mesothelae. Only in Sicily are represented Solifugae
(Biton Karsch, Daesiidae), and there are Opilioacarida and some scorpions.
Palpigradi Rémy (1949) reported from Corsica Eukoenenia mirabilis and E. ber-
lesei. Eukoenenia patrizii Condé, 1956 (endemic), was described from Sardinia.
The first known species of the order E. mirabilis was described from Sicily by
Grassi and Calandruccio (1885) and was recorded from Sardinia by Roewer (1953).
From Sardinia Condé ьха Heurtault (1993) described a second troglobitic
(endemic?) species – E. grafittii.
As a whole, six species of Palpigradi have been recorded from seven islands of
the Mediterranean: Sicily, Sardinia, Mallorca, Iraklia nr. Naxos, Kythira, Corfu, and
Malta (Bertrand 1980, Condé 1956, 1987).
Solifugae Only on Sicily are known two species of Daesiidae (Biton ehrenbergi
Karsch and B. velox Simon) (Chemini 1995).
Scorpiones From Sicily, Sardinia, Corsica, and the smaller archipelagoes around
them are recorded (Caporiacco 1950, Fet 2010, Roewer 1953) the following
scorpions:
Fam. Euscorpiidae
Euscorpius (Tetratrichobothrius) flavicaudis (DeGeer, 1778) – Corsica
E. (E.) oglasae Caporiacco, 1950 – Montecristo Island (Tuscan Archipelago)
(endemic)
E. (E.) sicanus (C.L. Koch, 1837) – Sicily (t.t.), Sardinia
E. (E.) “carpathicus” corsicanus Caporiacco, 1950 – Corsica (probably separate
species)
Pseudoscorpiones From Sardinia are known 72 sure spp. and ssp. of pseudoscor-
pions of 25 genera and 11 families (Beier 1955, 1956, 1956, 1959, 1973, Lazzeroni
1969, Callaini 1983a, 1983b, 1989, Gardini 1981, 1994, 2000, von Helversen 1968).
From Sicily are known 45 sure spp. and ssp. of pseudoscorpions of 18 genera and
8 families (Beier 1961b, 1975; Callaini 1981, 1991; Gardini and Rizzerio 1987).
From Corsica are known 37 sure spp. and ssp. of pseudoscorpions of 18 genera
and 8 families (Beier 1948; Callaini 1981; Gardini 2000; Heurtault 1975; Mahnert
1978c; Schawaller 1981).
Here is the list of the 15 endemic pseudoscorpions known from the three islands:
Fam. Chthoniidae
Chthonius (Ephippiochthonius) corsicus Callaini, 1981 – Corsica
7.8 Sicily, Sardinia, Corsica, and Elba 355
Ch. (E.) aegatensis Calaini, 1981 – Sicily

Ch. (E.) giustii Callaini, 1981 – Corsica
Ch. (E.) grafittii Gardini, 1981 – Sardinia
Ch. (E.) poseidonis Gardini, 1981 – Sardinia
Ch. (E.) remyi Heurtault, 1975 – Corsica
Ch. (E.) siscoensis Heurtault, 1975 – Corsica
Ch. (Ch.) multidentatus Beier, 1963 – Sicily
Spelyngochthonius sardous Beier, 1955 – Sardinia
S. grafittii Gardini, 1994 – Sardinia
S. beieri Gardini, 1994 – Sardinia
? Troglochthonius doratodactylus Helversen, 1968 – published from Sardinia but,
according to Martens and Helversen (1972) and Harvey (1990), mislabeled
Fam. Neobisiidae
Acanthocreagris aelleni Mahnert, 1978 – Corsica
A. corsa Mahnert, 1978 – Corsica
Neobisium mahnerti major Callaini, 1981 – Corsica
N. (N.) pacei Callaini, 1991 – Sicily
Roncus aetnensis Gardini et Rizzerio, 1987 – Sicily
R. carusoi Gardini et Rizzerio, 1987 – Sicily
R. siculus Beier, 1975 – Sicily
R. zoiai Gardini et Rizzerio, 1987 – Sardinia
Fam. Chernetidae
Pselaphochernes scorpioides (Herman, 1804) – Sicily
Lasiochernes siculus Beier, 1961 – Sicily
Opiliones (Juberthie 1958, Marcellino 1970, 1974, 1975a, 1980, 1983, Roewer
1956, Simon 1872)
Cyphophthalmi
Two of the three spp. of the genus Parasiro Hansen et Sørensen, 1904 (Sironidae),
live in Corsica (the third in Spain and continental France).
In the other islands, Parasiro minor was recorded by Brignoli (1968) in Sardinia.
Eupnoi
Mastobunus Simon, Metasclerosoma Roewer (Sardinia, Sicily), Nelima Roewer
(Sicily), etc.
Fam. Phalangiidae
Dicranopalpus brevipes Marcellino – Sicily (endemic)
D. wiehlei Kraus – Sardinia, Sicily
Phalangium L., Metaphalangium Roewer, Opilio Herbst,
Dasylobus argentatus (Canestrini) – Corsica, Sardinia

D. gestroi (Thorell) – Sardinia
Dyspnoi
Fam. Trogulidae
Anelasmocephalus Simon – Corsica, Sardinia, Sicily (A. pusillus Simon)
Calathocratus Simon – Sicily (C. africanus Lucas)
Trogulus Latreille – Corsica (T. acuaticus Simon)
Fam. Dicranolasmatidae
Dicranolasma Sørensen – Sicily (D. wiehlei Kraus, D. soerensenii Thorell)
Fam. Nemastomatidae
Histricostoma Kratochvíl et Miller – Sicily, Corsica, Sardinia (H. argenteolu-
nulatum Canestrini)
Laniatores
Fam. Travuniidae
Buemarinoa Roewer, 1956 – endemic genus, Sardinia (cave)
Fam. Phalangodidae
The genus Scotolemon is represented in Corsica (Scotolemon terricola Simon –
found also in Algeria and Sardinia) and Sicily and Sardinia (S. doriai Pavesi).
Ptychosoma vitellinum Soerensen (Phalangodidae) – Sicily, Sardinia (Brignoli
1968, Marcellino 1970, 1975)
Araneae According to the list of Marinu and Verneau (2002, online), in Corsica
have been recorded 515 spp. of spiders, belonging to 198 genera and 37 families.
The catalogue of spiders in Sardinia (Pantini et al. 2013) includes 43 families, 229
genera, and 495 species (Alicata and Cantarella 2000, Brignoli 1969, 1974, 1978,
Gasparo 1999, Kraus 1955, Thermes 1972, Bosmans and Colombo 2015). In their
catalogue of Sardinian spiders, Pantini et al. (2013) affirm that “…it is interesting to
make a comparison with the fauna of nearby Corsica. In fact, the number of species
in the Sardinian-Corsican area totals 700 species and the number present on each
island is nearly the same, but actually the similarity between the two faunas is not
very high (Sørensen’s index of similarity = 0.63).” Some 319 spp. are shared
between the two islands. Among the Sardinian spiders, 10.9% are endemic.
Endemic genera:
Fam. Dysderidae
Holissus Simon, 1882 – Corsica
Sardostalita Gasparo, 1999 – Sardinia
7.9 Malta 357
Chorological spectrum of Sardinian araneofauna, after Pantini, Sassu, and Serra (2013)
According to Brignoli (1981), “…la région sardo-corse a eu une importance extrême

pour l’histoire du peuplement de toute l’Italie, non seulement parce qu’elle a joué le rôle
de réfuge pour des espèces de climat tempéré (et peut-être aussi pour les thermophiles),
mais aussi parce que, à travers elle (ainsi que à travers la région ligurienne-provençale)
la faune ancienne de la Méditérranée occidentale a peuplé la peninsule.”
Opilioacarida Opilioacarus italicus (With, 1904) was described from Sicily

(Palermo) and is known also from Sardinia (Brignoli 1967).
7.9 Malta
The Maltese archipelago is situated in the center of the Mediterranean, 93 km south
of Sicily and 288 km east of Tunisia. It consists of five islands: Malta (highest point
Ta’ Dmejrek, 253 m), Gozo, and the small Comino, Cominoto, and Filfla. The
whole area of the archipelago is 316 km2. The islands were formed from the high
points of a land bridge between Sicily and North Africa that became isolated. The
archipelago lies on the edge of the African tectonic plate where it meets the Eurasian
plate (Schembri 1993, 2003).
According to the WWF, the territory of Malta belongs to the ecoregion of
“Mediterranean forests, woodlands, and scrub.” The climate is subtropical
Mediterranean.
The small islands with limited number of habitats shelter 1 endemic sp. of Palpigradi,
1 endemic scorpion, 22 pseudoscorpions, 3 suborders of opilions, 74 spp. of spiders,
and some mites. Nothing is very sensational in the zoogeography of Arachnida
(Schembri 2003).
Palpigradi The only species known from Malta is Eukoenenia christiani Condé,
1988, from a cave.
Scorpiones According to the list of Kovaøik (1999), in Malta live four species of
scorpions, all living also on the European continent: Buthus occitanus, Euscorpius
carpathicus, E. italicus, and E. flavicaudis. However, according to Fet (2010), E.
carpathicus is confined to Romania, and the Maltese former subspecies E. carpathi-
cus sicanus (C.L. Koch, 1837) is now a full species (Fet et al. 2003), known from
Malta, Central and Southern Italy, Sardinia, Sicily, Greece, North Africa, and
Madeira. Other sources indicate that in Malta live three species: Buthus occitanus,
B. tunetanus (?), and Euscorpius sicanus.
After Kritscher (1992) and Schembri (2003), on Maltese islands lives only one
scorpion: Euscorpius sicanus.
Pseudoscorpiones In Malta have been recorded 22 species of pseudoscorpions,
belonging to 17 genera and 9 families (Beier 1973, Gardini and Rizzerio 1987,
Mahnert 1975, 1982, Schembri 2003). Endemic species are Chthonius maltensis
Mahnert, 1975; C. girgentiensis Mahnert, 1982; and Roncus melitensis Gardini et
Rizzerio, 1987. Minniza algerica Beier (known also from Lampedusa) is the con-
nection with North Africa and Chernes siciliensis Beier with Sicily.
Opiliones On Maltese islands have been recorded five spp. of Opiliones, but there
are known several others which are unpublished (Schembri 2003). Most species are
known also from Sicily (Marcellino 1974, Thaler 1996).
Eupnoi
Fam. Phalangiidae
Metaphalangium cirtanum (C.L. Koch)
Opilio canestrinii (Thorell)
Phalangium targionii (Canestrini)
Dyspnoi
Fam. Dicranolasmatidae
Dicranolasma soerensenii Thorell
Laniatores
Fam. Phalangodidae
Ptychosoma vitellinum Soerensen
7.10 Central and Northern Europe, Great Britain, Ireland, Island, and Faroe Islands 359
Araneae The literature on Maltese spiders was reviewed first by Baldacchino et al.
(1993) (74 spp. of 21 families) and then by Bosmans and Dandria (1995) (83 spp.
of 22 families). These authors describe several new species, seemingly endemic to
Maltese islands. Most species are known also from Italy, and some are Siculo-
Maltese endemics (Brignoli 1969, Cantarella 1982, Kritscher 1996).
Acari None is zoogeographically interesting.
7.10 C
entral and Northern Europe, Great Britain, Ireland,
Island, and Faroe Islands
7.10.1 G
and Paleogeography
In this area are included the countries north of Spain, peninsular Italy and the Balkan
Peninsula (north of Danube), and east to the borders with Russia, Belarus, and
Ukraine. Included are also the British Isles, Island, Faroes, and other northern
islands. This is one of the best studied parts of the Earth. Dominated by the Alps and
Carpathians, Central Europe has been glaciated, with consequences on its arachno-
fauna (Hulten 1937). However, it is worth noticing the article of Malicky et al.
(1983), advocating the existence of Central European faunal elements (distribution
excluding the Mediterranean region and Asia largely or totally). Some of the exam-
ples are among the spiders and the opilions (by K. Thaler). The idea of the authors
is that part of the fauna has “overwintered” during the Pleistocene glaciations.
Besides the tropical orders, in Central and Northern Europe lack Solifugae and
Opilioacarida, and there are very few scorpions of only one genus (Euscorpius
Thorell). In this area the arachnids of the Alps are especially well studied, with
many papers on high-altitude spiders, mites, opilions, and pseudoscorpions (Beron
2008b, with extensive bibliography on the high mountain Arachnida; 2016, Brinck
1966, Franz 1954, Freytag 1962, Meyer and Thaler 1995, Komposch 2011, Muster
2000, 2001, Schmölzer 1999, 2001, Strand 1906, Thaler, many papers).
Palpigradi So far from mainland Europe (except for the three southern peninsulas)
has been recorded eight species of Palpigradi, all belonging to Eukoenenia Börner.
They are known from France, Austria, Hungary, and Romania (Christian 1998,
Hansen 1926, Peyerimhoff 1902).
Scorpiones The natural distribution of scorpions in Europe, north of the Danube,

is limited to a line across Romania, Austria, and Czech Republic (?) (Braunwalder
2001, Caporiacco 1950, etc.). According to the concept of Fet and Soleglad (2002),
the “widely distributed” before Euscorpius carpathicus (Linnaeus, 1767) is
confined to a small area in Southwest Romania. The small population in Slapy

(Central Bohemia) is considered sometimes introduced or autochthonous (Kovarik
1999). The distribution of this species, still endemic for Romania, is distributed in
two clusters in the Carpathians (Gherghel et al., 2016).
According to Fet (2010), in Europe north of the three southern peninsulas are
recorded four species of scorpions:
Fam. Euscorpiidae
Euscorpius (E.) carpathicus (Linnaeus, 1767) – Romania
E. (Alpiscorpius) alpha Caporiacco, 1950 – Alps (Italy, Switzerland)
E. (A.) germanus (C.L. Koch, 1837) – Austria, Slovenia, Switzerland, NE Italy
E. (A.) gamma Caporiacco, 1950 – Austria, Slovenia, NE Italy
Pseudoscorpiones In the Central and Northern Europe (the area described above)
have been recorded pseudoscorpions of ten families: Chthoniidae, Neobisiidae,
Syarinidae, Geogarypidae, Larcidae, Cheiridiidae, Cheliferidae, Atemnidae,
Chernetidae, and Withiidae (Beier 1952, 1963, Beier and Franz 1954, Gulièka 1977,
Kaisila 1949, Lehtinen 1964, Lohmander 1939, Meinertz 1964, 1962, Rafalski
1967, Uddström and Rinne 2014).
From Czech Republic and Slovakia have been recorded 58 spp. of 22 genera and
8 families (Christophoryová et al. 2012). In Austria live 71 spp. of 24 genera and 10
families (Mahnert 2004, actualized), and from Finland are recorded (Uddström and
Rinne 2014) 17 spp. of 5 families.
Endemic genera of pseudoscorpions in Central and Northern Europe:
Fam. Neobisiidae
Occitanobisium Heurtault, 1977 – France (one sp.)
Roncobisium Vachon, 1967 – France (two spp.)
Simonobisium Heurtault, 1974 – France, Italy (one sp.)
Anthrenochernes Lohmander, 1939 – Sweden (one sp.)
Opiliones In West Europe north of the three southern peninsulas and the Pyrenees
and west of Russia and Belarus are registered Opiliones of ten families: Sironidae,
Cladonychiidae, Phalangodidae, Phalangiidae, Sclerosomatidae, Nemastomatidae,
Trogulidae, Dicranolasmatidae, Ischyropsalididae, and Sabaconidae (Bezdecka
2008, Blick and Komposh, 2004, Cawley 2002, Heinäjoki 1944, Kauri 1980,
Komposch 1999, 2011, Komposch and Gruber 2005, Lehtinen 1964, Pack-Beresford
1926, Rafalski 1956, Roewer 1934, Schuster 1975, Spoek 1975, Spungis 2008,
Staręga 1976a, Stol 1993, Szalay 1968, Tumšs 1963, Vanhercke 1999, Wijnhoven
2005, Winiarska 2008).
According to Stol (2007), a total of 24 Nordic species are currently known, of
which 17 are from Norway, 19 from Denmark, 20 from Sweden, 12 from Finland, 5
from the Faroe Islands, and 4 from Iceland. Komposch (2004) lists for Hungary
33 spp. of Opiliones of 22 genera and 6 families. For Austria Komposch (2005) lists
61 (s) spp. of Opiliones of genera and 8 families. Among them there are two
endemic (sub)species for Austria, Nemastoma bidentatum relictum Gruber et
7.10 Central and Northern Europe, Great Britain, Ireland, Island, and Faroe Islands 361
Martens and N. schuelleri Gruber et Martens, and nine subendemic species. Again
Komposch (2004) lists from Hungary 33 species of 23 genera and 6 families.
From the other countries, the figures are as follows:
Belgium (Vanhercke L. 1999) – 27 (sub)spp. of Opiliones of four families
Netherlands (Spoek 1975; Wijnhoven, 2005) – 27 (sub)spp. of Opiliones of five
families
Switzerland (Martens 1978) – 49 (sub)spp. of Opiliones of six families
Germany (Martens 1978) – 49 (sub)spp. of Opiliones of six families
Poland (Rafalski 1960, 1961; Rafalski and Staręga 1997; Staręga 2000; Winiarska
2008) – 36 (sub)spp. of Opiliones of eight families
Czech Republic (Šilhavý 1956, Klimeš 2000) – 33 spp. of Opiliones of 15 genera
and 5 families
Slovakia (Šilhavý 1956, Klimeš 2000) – 33 spp. of Opiliones of 22 genera and 8
families
Cyphophthalmi
There is only one species (Siro carpaticus Rafalski, 1956), described from Poland,
known also from Slovakia.
The “Palpatores” form the bulk of the European opiliofauna. In Central Europe
are known the families Phalangiidae, Sclerosomatidae, Nemastomatidae, Trogulidae,
Dicranolasmatidae, Ischyropsalididae, and Sabaconidae.
Eupnoi
Fam. Phalangiidae – Phalangium L., Amilenus Martens, Dicranopalpus Doleschal,
Lacinius Thorell, Mitopus Thorell, Odiellus Roewer, Oligolophus C.L. Koch,
Paroligolophus Lohmander, Egaenus C.L. Koch, Opilio Herbst, Dasylobus
Simon, Megabunus Meade, Lophopilio Hadži, Platybunus C.L. Koch
Fam. Sclerosomatidae – Leiobunum C.L. Koch, Gyas Simon, Astrobunus Thorell,
Nelima Roewer
Dyspnoi
Fam. Nemastomatidae – Mitostoma Roewer, Nemastoma C.L. Koch, Paranemastoma
Redikorzev, Carinostoma Kratochvil, Hystricostoma Kratochvil
Fam. Trogulidae – Trogulus Latreille, Anelasmocephalus Simon
Fam. Dicranolasmatidae – Dicranolasma Sørensen
Fam. Ischyropsalididae – Ischyropsalis C.L. Koch
Fam. Sabaconidae – Sabacon Simon
Laniatores Roewer (1935) made a review of the European Laniatores. The species
of the three southern peninsulas and the Mediterranean islands are excluded; in
Central Europe are known to live Laniatores from the genera Arbasus Roewer
(Travuniidae), Scotolemon Lucas (Phalangodidae). From the family Cladonychiidae,
one genus (Holoscotolemon Roewer, 1915) is represented with nine species in
Austria, Romania, Hungary, France, and Italy. The other seven species of four gen-
era live in the USA.
Endemic opilionid genera for Central Europe are:
Fam. Cladonychiidae
Holoscotolemon Roewer, 1915
Araneae The spider fauna of Central and Northern Europe (within the described
limits) is well known and represented by ca. 37 families, after Aakra and Hange
(2000), Agnarsson (1996), Almquist (2005), Blick et al. (2004), Bosmans (2009),
Bosmans and Vanytven (2001, Internet), Buchar (1992), Buchar et al. (1995),
Buchar and Ružicka (2002), Eskov (1994), Gajdos et al. (1999), Hauge (1989),
Komposch (2011), Koponen (1995, 1996, 2005), Kronestedt (Version 2001), Larsen
and Scharff (2003), Marinu and Verneau (2002), Merrett et al. (1985), Merrett and
Millidge (1992), Merrett and Murphy (2000), Mikhailov (1997, 1998, 1999, 2000),
Milošević (2002), Roberts (19895), Thaler (1976, 1980, 1988), Thaler and Buchar
(1994, 1996), Nentwig et al. (2013), Newlands (1978), Platnick (2014), Proszynski
and Staręga (1971), Samu, Szinetar (1999), Vilkas (1992), Weiss and Urak (2000),
and Wiehle (1953): Scytodidae, Sicariidae, Pholcidae, Dysderidae, Oonopidae,
Segestriidae, Eresidae, Oecobiidae, Palpimanidae, Uloboridae, Anapidae, Araneidae,
Linyphiidae, ?Synaphridae, Tetragnathidae, Theridiidae, Theridiosomatidae,
Lycosidae, Oxyopidae, Pisauridae, Miturgidae (Zorinae), Agelenidae, Amaurobiidae,
Anyphaenidae, Cybaeidae, Dictynidae, Hahniidae, Sparassidae, Zodariidae,
Clubionidae, Titanoecidae, Gnaphosidae, Philodromidae, Thomisidae, Salticidae,
Corinnidae, and Liocranidae. This number is not very big in this well-studied terri-
tory (for comparison, only on the Balkan Peninsula are registered 48 families of
spiders). No family is endemic, and many of them are widespread in the world.
In Great Britain are known 658 spp. of spiders (Duffey 2010), in France 1569 spp.
(Le Peru 2007), in Switzerland 875 spp. (Maurer and Hänggi 1990), in Germany
925 spp. (Koponen 1991), in Poland 809 spp. (Rozwalka and Stanska 2008), in
Sweden 704 spp. (Kronestedt 2001), and in Norway 535 spp. (Hauge 1989). Ireland
shares all its spider species (377 names) with Great Britain but possesses only
59.6% of that fauna (van Helsdingen 1995).
What concerns the Czech Republic, according to Buchar (1995), is that it “may be
divided in two zoogeographically and geomorphologically distinct territories –
Moravia and Silesia form division between the Bohemian Highlands and Carpathian
depressions. The Basin of Vienna extends into Moravian territory from south and
builds a way for the immigration of Mediterranean and sub Mediterranean termitophi-
lous species. 761 species of spiders have been collected – 61 species of them are found
in Moravia only 70% of those Moravian spiders occur the very xerotherm habitats.”
A special case is Iceland, situated between Europe and North America. In this
northern country are known the orders Opiliones, Pseudoscorpiones, Araneae,
Ixodida, Mesostigmata, Trombidiformes, and Sarcoptiformes. Nine orders are miss-
ing. What concerns the spiders, Braendegård (1958) recorded 78 species of 9 fami-
lies and confirmed the opinion of Lindroth (1931) that “the Icelandic insect [and
spider] fauna was almost exclusively of palaearctic origine” and that “the boundary
between the nearctic and the palaearctic geographical respect must be the
Danmarkstraede.” From the 78 spp., 54 belong to the family Linyphiidae, contain-
ing more than half of all Arctic spiders, according to the same author.
Acariformes Ref.: Beron (2008b, 2016, Luxton 1995)
Prostigmata Fam. Teneriffiidae – the genus Mesoteneriffia Irk is known from the
Alps.
7.11 Arachnida of the Atlantic Islands (Macaronesia, St. Helena, Ascension) 363
Oribatida Luxton (1995) published checklist of 135 genera and 303 spp. of
Oribatida in the British Isles, with zoogeographical notes on their distribution.
7.11 A
rachnida of the Atlantic Islands (Macaronesia,
St. Helena, Ascension)
7.11.1 Canary Islands, Azores, and Madeira
7.11.1.1 Geography, General Zoogeography, and Paleogeography
Four archipelagoes in the Atlantic Ocean (Canary Islands, Azores, Madeira, Cabo
Verde) are called Macaronesia. The northernmost of them are the Azores. The nine
volcanic islands of the archipelago have a total surface area of 2346 km2, and the
highest peak is Mount Pico (2351 m). During the last several centuries, the forests
(laurisilva) have been destroyed (about 95% of them), and it is believed that more
than half of the insects and other invertebrates having existed originally (until the
discovery of the uninhabited islands in fourteenth century) are gone or heavily
endangered. The Azores are situated 1360 km west of Portugal, about 1510 km
northwest of Morocco, and about 1925 km southeast of Newfoundland.
Another archipelago, a Portugal territory, is Madeira, comprising the islands of
Madeira (741 km2), Porto Santo, Desertas, and Selvagens, 520 km from the
African coast.
The archipelago of the Canary Islands consists of seven main islands, located
between 100 and 500 km from the African coast. They are the Tenerife (2034 km2),
Fuerteventura (1659 km2), Gran Canaria (1560 km2), El Hierro (278 km2), Palma
(706 km2), Lanzarote (846 km2), and Gomera (370 km2).
Pico de Teide on Tenerife is the highest mountain of all Atlantic Ocean islands
(3718 m). All islands emerged in Miocene, Fuerteventura and Lanzarote being the
oldest (Dietz and Sproll 1970b).
For Macaronesia s. str. (Canary Islands, Azores, Madeira, Selvagens), we find on
the EEA (2002) analysis of the geographic regions the following distribution of
land: meadows 25%, forest 15%, cultivated land 14%, and land with few or no veg-
etation 34%.
7.11.1.2 Arachnogeography
In 2001 was published a volume containing database on the terrestrial animals of

the Canary Islands. The Arachnida are distributed as follows (Bacallado 1984,
Schenkel 1938, Kunkel (Ed.) (1976)):
Palpigradi – one sp. (omitted)
Araneae – 453 spp. (292 endemic)
Pseudoscorpiones – 44 (23 endemic) (according to Mahnert (2011), on the islands
are known 56 spp. and subspp. of pseudoscorpions)
Scorpiones – one sp. (introduced)
Opiliones – five spp.

Palpigradi – one sp.
Schizomida – one sp. (introduced)
Solifugae – one sp. (endemic)
Acari – 244 spp.
Total – 799 spp. and 23 ssp., 393 spp. and 11 ssp. endemic (actually, many more
than 800 spp.). Especially the Acari are certainly many more.
Lacking from the islands are the orders Ricinulei, Uropygi, Opilioacarida,
Holothyrida, and non-introduced Scorpiones and Schizomida.
Azorean Islands – Araneae, Opiliones, Pseudoscorpiones, Acari, etc.
Madeira – Palpigradi (one endemic sp.), Araneae, Opiliones, Pseudoscorpiones,
Acari, etc.
Palpigradi Eukoenenia mirabilis (Grassi) was the first Palpigradi, recorded from
the Canary Islands (Tenerife) (Condé 1990). From Madeira is known Eukoenenia
madeirae Strinati et Condé, 1996.
Solifugae The only known species on the Canary Islands is Eusimonia wunderlichi
Pieper, 1977 (Karschiidae). The other 14 spp. of Eusimonia are spread from North
Africa to Central Asia. From Cabo Verde has been described Ammotrechella dias-
pora Roewer, 1934 (Ammotrechidae), a purely American genus and family.
Scorpiones No scorpions on the Azores. On the Canary Islands have been found
several species (incl. Centruroides gracilis), but probably introduced (Fet 2010). Also
according to Fet (2010), “Euscorpius sicanus record from Madeira (Fet et al. 2003)
could be either introduction from the Mediterranean, or a local relict.” Otherwise,
Macaronesia did not seem to harbor native scorpions (Crucitti 2004, Fet et al. 2003,
Fet 2010). Exception is Hottentotta caboverdensis (Buthidae) in Cabo Verde.
Pseudoscorpiones The distribution of Pseudoscorpiones on the Atlantic Ocean

islands is already well known and rich with zoogeographically remarkable taxa
(Beier 1961, 1965, 1970, 1975, 1976, Estany 1979, von Helversen 1965, Mahnert
1980, 1989, 1990, 1993, 1997, 2002, 2011, Pieper 1981, 1991, Schenkel 1938,
Tullgren 1900, Vachon 1961, Zaragoza, 1986, 2000, full bibliography, Zaragoza
et al. 2004).
They are distributed as follows:
Canary Islands: 48 spp. of 13 genera and 8 families (Chthoniidae, Syarinidae,
Geogarypidae, Olpiidae, Cheliferidae, Cheiridiidae, Chernetidae, Withiidae)
Azores: two genera and two families (Neobisiidae, Syarinidae)
Madeira and Selvagens: 23 spp., 5 genera, and 3 families (Chthoniidae, Syarinidae,
Cheliferidae)
7.11 Arachnida of the Atlantic Islands (Macaronesia, St. Helena, Ascension) 365
The distribution of different families and genera is as follows:

Fam. Chthoniidae
Chthonius C.L. Koch – Canary Islands, Madeira
Paraliochthonius Beier – Canary Islands, Madeira
Tyrannochthonius Chamberlin – Canary Islands
Lagynochthonius Beier – Canary Islands (six spp.)
Fam. Geogarypidae
Geogarypus Chamberlin – Canary Islands
Fam. Olpiidae
Calocheirus Chamberlin – Canary Islands (four of the six species in the genus;
the remaining two live in Sudan, Israel, and Saudi Arabia)
Fam. Syarinidae
Microcreagrina Beier – Madeira, Canary Islands (three spp. of Microcreagrina
live on the Canary Islands, one on Madeira, and one is widespread in the
Mediterranean)
Microcreagrella Beier – Madeira, Azores (different subspecies of M. caeca
(Simon))
Pseudoblothrus Beier – Azores (two spp.)
Fam. Cheiridiidae
Cheiridium Menge – Canary Islands
Fam. Chernetidae
Pselaphochernes Beier – Canary Islands
Fam. Cheliferidae
Canarichelifer Beier – Canary Islands (endemic genus)
Pseudorhacochelifer Beier – Madeira, Canary Islands (different spp.)
Rhacochelifer Beier – Canary Islands
Fam. Withiidae
Withius Kew – Canary Islands
Opiliones On the islands are known six genera of Opiliones (Laniatores, Eupnoi)
of the families Pyramidopidae and Phalangiidae (Kauri 1963; Rambla 1956, 1975,
1993; Roewer 1936; Sharma et al. 2011; Spoek 1959; Staręga 1984). Four of them
are endemic.
Endemic on the islands are:
Laniatores
Fam. Pyramidopidae
Maiorerus randoi Rambla, 1993 – Fuerteventura, cave (endemic genus and
species)
Eupnoi
Fam. Phalangiidae (Phalangiinae)
Bunochelis Roewer, 1923 (endemic genus)
Bunochelis canariana (Strand, 1911) (syn. B. altenai Spoek, 1959 – Tenerife) –
Canary Islands
B. spinifera (Simon, 1878) – Canary Islands
Ramblinus Starega, 1984 (endemic genus)
R. spinipalpis (Roewer, 1911) – Madeira
Parascleropilio Rambla, 1975 (endemic genus)
P. fernandezi Rambla, 1975 – Canary Islands
7.11.1.3 Araneae
On the Canaries are known at least 453 spp. (292 endemics) of spiders (Berland
1935a, Denis 1941, Wunderlich 1987, 1991, 1992, 1993, 1995).
On Madeira have been recorded 164 species of spiders (Denis 1962, Wunderlich
1987, Cardoso and Crespo 2008), including 40 single-island endemics.
Analysis of the spider biodiversity patterns on the Azoreans has been done by
Borges and Wunderlich (2008).
Some endemic genera of spiders in Macaronesia:
Fam. Pisauridae
Cladycnis Simon, 1898 – Canaries
Fam. Nesticidae
Canarionesticus Wunderlich, 1992 – Canaries
Fam. Linyphiidae
Afribactrus Wunderlich, 1995 – Azores
Acorigone Wunderlich, 2008 – Azores
Canariellanum Wunderlich, 1987 – Canary Islands (four spp.)
Frontiphantes Wunderlich, 1987 – Madeira (one sp.)
Lomaita Bryant, 1948 – Hispaniola
Fam. Gnaphosidae
Canariognapha Wunderlich, 2011 – Canary Islands
Macarophaeus Wunderlich, 2011 – Madeira, Canary Islands
Fam. Pholcidae
Ossinissa Dimitrov et Ribera, 2005 – Canary Islands (one sp.)
7.12 Cabo Verde 367
Fam. Theridiidae
Eurypoena Wunderlich, 1992 – Canary Islands (one sp.)
Grancanaridion Wunderlich, 2011 – Canary Islands, Gran Canaria (one sp.)
Macaridion Wunderlich, 1992 – Madeira (one sp.)
7.12 Cabo Verde
7.12.1 G
and Paleogeography
Ten bigger and three smaller volcanic islands are located in the central Atlantic Ocean,
570 kilometers off the coast of western Africa (Senegal, Mauretania). The total area
of is 4033 km2 and the highest point is Pico de Fogo (2829 m.), an active volcano.
Islands date from eight million (in the west) to 20 million years (in the east). Climate
is semidesert. The nature is strongly degraded by humane activities after the discov-
ery of the islands in 1456. The archipelago is part of the Macaronesian ecoregion.
On the islands are known members of Solifugae, Scorpiones, Pseudoscorpiones,

and Araneae. The other orders are not recorded. It is to note that the Solifugae are
represented by a member of the American family Ammotrechidae, but the scorpions
and pseudoscorpions are of Afrotropical character.
Solifugae Only one sp. has been recorded: Ammotrechella diaspora Roewer, 1934
(Ammotrechidae). The other members of genus Ammotrechella live in North and
Central America and in West Indies. No Ammotrechidae in the Old World.
Scorpiones Lourenço and Ythier (2006) described a new, and supposedly endemic,
species for Cabo Verde islands (Hottentotta caboverdensis, Buthidae).
Pseudoscorpiones On the archipelago are reported five spp. of pseudoscorpions of

the families Atemnidae, Olpiidae, Garypidae, and Withiidae. The fauna is a mix of
Afrotropical and Mediterranean elements (Ellingsen 1906, Vachon 1956).
Araneae Berland (1936), having studied the spiders collected by A. Chevalier in

1934, brought the number of species of spiders to 48 species of 15 families. Now
from Cabo Verde are reported 58 endemic species of spiders, including Luxuria
lymphatica (jumping spider). Australoechemus Schmidt et Piepho, 1994
(Gnaphosidae); Koinothrix Jocqué, 1981 (Linyphiidae); and Wesolowskana Koçal
et Kemal, 2008 (Salticidae) are endemic genera.
7.13 St. Helena and Ascension
7.13.1 Geography and Paleogeography (General)
St. Helena is an island of volcanic origin in the South Atlantic Ocean. It is part of
the British overseas territory of St. Helena, Ascension, and Tristan da Cunha. St.
Helena measures about 16 by 8 kilometers, with 122 km2 surface area with the high-
est point Diana’s Peak at 818 m. Ascension has a surface area of 91 km2 and Tristan
da Cunha 98 km2.
Important contribution to knowledge on the fauna of St. Helena were the four
volumes, result of the Belgian mission in 1965. The Arachnida have been analyzed
by Benoit (1977 – several papers).
Ashmole P. and M.J. Ashmole (1997, 2000) contributed to the protection of the
seabirds on Ascension and made some important discoveries of blind cave animals
on the island.
The fauna is well studied. It is not very rich as the islands are far from any mainland
and also because of the human impact on the vulnerable environment.
Lacking are Palpigradi, Ricinulei, Holothyrida, Opilioacarida, Amblypygi,
Thelyphonida (Uropygi), and Schizomida. Better known are the pseudoscorpions
and the spiders, with several endemic genera (Ashmole and Ashmole 1997, 2000).
Scorpiones Only one cosmotropical species is known on the islands: Isometrus
maculatus (de Geer) (Benoit 1977a).
Pseudoscorpiones On both islands are represented the families Garypidae,

Withiidae, Olpiidae, and Cheiridiidae (Beier 1951, 1961, 1977a, Mahnert 1993).
Endemic genera for St. Helena are:
Fam. Olpiidae
Hemisolinus Beier, 1977 – one sp.
Fam. Withiidae
Scotowithius Beier, 1977 – one sp.
Sphallowithius Beier, 1951 – one sp.
From Ascension Island have been recorded five (including endemic) species:
Withius ascensionis Beier, 1961; Stenowithius duffeyi Beier, 1961; Garypus titanius
Beier, 1961 (the largest pseudoscorpion); and Apocheiridium cavicola Mahnert, 1993
Araneae
After Benoit (1977a), from St. Helena are recorded 98 spp. of spiders (48 endem-
ics) of 25 families (Merrett and Ashmole 1997, Millidge and Ashmole 1994,
Platnick 1993).
7.13 St. Helena and Ascension 369
Endemic genera for St. Helena:

Fam. Dictynidae
Helenactyna Benoit, 1977 (two spp.)
Fam. Miturgidae
Helebiona Benoit, 1977
Fam. Eutichuridae
Tecution Benoit, 1977 (three spp.)
Fam. Theridiidae
Zercidium Benoit, 1977 (one sp.)
Fam. Linyphiidae
Napometa Benoit, 1977 (two spp.)
Fam. Lycosidae
Dolocosa Roewer, 1960
Fam. Gnaphosidae
Benoitodes Platnick, 1993 (two spp.)
Fam. Thomisidae
Bonapruncinia Benoit, 1977 (one sp.)
Fam. Salticidae
Paraheliophanus Clark et Benoit, 1977 (four spp.)
Endemic genus and species for Ascension is the blind cave spider Catonetria
caeca Millidge et Ashmole, 1994 (Linyphiidae).
Acariformes
Prostigmata – Ref.: Naudo (1977)
Fam. Anystidae – endemic species
Chaussieria benoiti Naudo
Ch. brevis Naudo
Ch. dissimilis Naudo
Ch. sanctaehelenae Naudo
Oribatida
Ref.: Luxton (1995)
Parasitiformes
Benoit (1977) recorded from St. Helena two spp. of Ixodidae, one of Laelapidae and
one of Macronyssidae (all parasitic).
7.14 R
ussia (North of Caucasus), Belarus, Ukraine, Siberia,
Altai, China (North of Yangtze), and Mongolia
7.14.1 G
and Paleogeography
Ukraine, Belarus, and Russia, north of Caucasus, and of Central Asian mountains
are vast plains and taiga forest.
The valley of Yenisei River is considered to be the important frontier between
eastern and western Siberia (Johannsen 1955). The Johannsen’s Line starts from
Severnaya Zemlya Islands and Taymyr Peninsula, follows the east bank of Yenisei
River, and reaches Altai. Another frontier is the Reinig’s Line along the east bank
of Lena River and Aldan River and over the mountains of Stanovoy and Yablonovy
into Tian Shan.
The paleogeography of the glaciation of Northeast Asia was outlined by Glushkova
(1992), the zoogeography of the Far East of Russia by Kolosov (1980), and of North
and Central Asia by Serge (1993).
Map 7.10 The Johannsen’s Line (oblique striated, determined by recent ecological factors) and
Reinig’s Line (vertically striated, determined by historical factors) (from De Latin 1967)
7.14 Russia (North of Caucasus), Belarus, Ukraine, Siberia, Altai, China (North… 371
From this vast area are missing the orders Amblypygi, Schizomida, Ricinulei,
Opilioacarida, and Holothyrida. Thelyphonida (Uropygi) are found only on the very
far east of China (the only record of Uropygi in the Maritime Province of Russia is
considered a chance specimen brought by Man from China – Mikhailov, 2016a).
The northern limits of several groups (Uropygi, Scorpiones, Solifugae) are running
through this area (Ukraine, Siberia, Mongolia, China).
There are several very useful reviews and checklists of the spiders of Russia and
the former Soviet Union (FSU), done by Russian specialists, mainly by
K.G. Mikhailov, who was kind enough to keep me informed about the progress of
the arachnology of Northern Asia.
Solifugae Only one species is known in European Russia and Ukraine: Galeodes
araneoides (Pallas 1772) (Galeodidae) (Birula 1912, 1917, 1922).
In Central Asia, Altai, North China, and Mongolia are known 26 spp. of Karschia
Walter (Karschiidae) (Gromov 2004). They live in Turkmenistan, Uzbekistan,
Tajikistan, Kyrgyzstan, Kazakhstan, China, Iran, Turkey, Armenia, and Azerbaijan.
In Mongolia are recorded five species (Galeodes kozlovi Birula, 1911, and G. mon-
golicus Roewer, 1934 (from family Galeodidae), and Karschia gobiensis Gromov,
2004; K. mongolica Roewer, 1933; and Eusimonia turkestana Kraepelin, 1899
(from family Karschiidae)), marking part of the northern limit of the order in Asia.
As a whole, from the countries of the former USSR are known 18 spp. of Solifugae
(Mikhailov 2016), mostly from Transcaucasia and Central Asia.
Thelyphonida (Uropygi) One species (Typopeltis amurensis Tarnani, 1889) was
described from the Far East of Russia (Primorsky Krai, Olga Bay, acc. to Harvey,
online, nomen dubium, acc. to Mikhailov 2016a, chance specimen brought with
human transport from China).
Scorpiones According to the catalogue of Fet (1988, with suppl.) and Mikhailov
(2016a), in the present territory of Russia (including Crimea) are known five spp. of
scorpions: three from fam. Euscorpiidae – Euscorpius tauricus (C.L. Koch, 1837),
endemic for Crimea (see Fet, 2003); E. italicus (Herbst), east shore of Black Sea,
north of Georgia; and E. mingrelicus (Kessler, 1874) – and two from fam. Buthidae,
Mesobuthus caucasicus (Nordmann, 1840), west shore of Caspian Sea (Birula
1917, Nenilin and Fet 1992), and M. eupeus (C.L. Koch, 1839).
From Mongolia are known three species: Mesobuthus martensii (Karsch, 1879),
M. eupeus (C.L. Koch, 1839), and M. caucasicus (Nordmann, 1840)
Pseudoscorpiones In the area are recorded pseudoscorpions of ca. 20 genera and
the families Chthoniidae (Chthonius, Mundochthonius), Neobisiidae (Neobisium,
Roncus, Bisetocreagris, Microcreagris, Stenohya), Syarinidae, Geogarypidae,
Atemnidae (Atemnus, Diplotemnus), Olpiidae, Cheiridiidae (Cheiridium),
Chernetidae (Megachernes, Wyochernes, Chernes, Allochernes, Pselaphochernes,
Dendrochernes), Cheliferidae (Centrochelifer, Chelifer, Dactylochelifer), and

Withiidae (Withius) (Beier 1966, 1969, 1973, 1979, Ćurčić 1983, 1985, Krumpál
and Kiefer 1982, Redikorzev 1922, 1934, 1949, Schawaller 1985, 1986, 1989).
From Mongolia have been recorded 28 spp. of 14 genera and the families
Neobisiidae, Atemnidae, Cheliferidae, Chernetidae, and Withiidae. The most wide-
spread is Dactylochelifer Beier (seven spp.).
From the huge territory of Russia (17045 million km2) are recorded only 48 spp.
(Mikhailov 2016a).
From South Korea are known 21 spp. of 11 genera and the families Chthoniidae,
Pseudotyrannochthoniidae, Neobisiidae, Syarinidae, Garypidae, Cheiridiidae, and
Chernetidae. There are no information about North Korea (material has been col-
lected, but not yet identified).
Some endemic genera:
Gobichernes Krumpál et Kiefer – Mongolia
Opiliones In this large territory are known harvestmen of more than 30 genera
(Chevrisov 1979, Chemeris 2000, Chemeris and Logunov 2001, Chemeris et al.
1979, 1980, Farzalieva and Esyunin 2000, Gritsenko 1979b, 1980, Kratochvil 1959,
Martens 1989, Šilhavý 1966, Staręga 1964, 1965, 1978, Suzuki 1941c, Tchemeris
et al. 1998, Tsurusaki 1998, Tsurusaki and Crawford 2001, Tsurusaki et al. 2000)
and the families Phalangiidae, Sclerosomatidae, Sabaconidae, Taracidae, Trogulidae,
Nemastomatidae, and Ischyropsalididae. One sp. of Holoscotolemon (Cladonychiidae,
Laniatores) is known from the Ukrainian Carpathians (Chevrisov 1979). Most others
are genera of the former “Palpatores.”
Cyphophthalmi are not known from the area.
Eupnoi
Fam. Phalangiidae – Oligolophus C.L. Koch, Mitopus Thorell, Acanthomegabunus
Tsurusaki et al., Rilaena Šilhavy, Phalangium L., Zachaeus C.L. Koch, Homolophus
Banks, Opilio Herbst, Odiellus Roewer, Metaplatybunus Roewer, Lophopilio Hadzi,
Platybunus C.L. Koch, Egaenus C.L. Koch, Tchapinius Roewer (Kamchatka)
Fam. Sclerosomatidae – Metagagrella Roewer, Psathyropus L. Koch (Far East),
Leiobunum C.L. Koch, Gyas Simon, Nelima Roewer
Dyspnoi
Fam. Sabaconidae – Sabacon Simon (two spp. – S. crassipalpis (L. Koch), Siberia;
S. sergeidedicatus Martens, Altai; S. okadai Suzuki, Mandjuria
Fam. Taracidae – Taracus Simon (one sp. T. birsteini Ljovuschkin, 1971 from the
Far East, cave; the other seven spp. live in the USA)
Fam. Trogulidae – Trogulus Latreille (one sp.), Dicranolasma Sørensen,
Platybessobius Roewer
Fam. Nemastomatidae – Paranemastoma Redikorzev, Nemastoma C.L. Koch
Fam. Ischyropsalididae – Ischyropsalis C.L. Koch
Laniatores
Fam. Cladonychiidae
7.14 Russia (North of Caucasus), Belarus, Ukraine, Siberia, Altai, China (North… 373
Holoscotolemon Roewer – Ukrainian Carpathians, H. jaqueti (Corti, 1905)

Araneae After studying of 505 spp. of spiders in Northern Siberia, Eskov (1986a,
thesis) concludes that the Arctic arachnofauna contains 98 spp. (more than 30% are
limited to the northern parts of the tundra zone). The Yenisei Line, running on the end
of Middle Siberian plateau, is a very clear biogeographical frontier within Holarctic
(a superposition of eastern limits of areas of the European-Western Siberian species
and the western limits of areals of the East Siberian-American species). After the
distribution of spiders in Northern Holarctic are delimitated three base regions,
European, Angaran, and Canadian, and two transitional regions, West Siberian-
Laplandian (from Enisey to Stegman’s Line, connecting Altai with Fennoscandia)
and Beringian (from Kolima to Mackenzie). ) (Eskov 1986b, 1994, Izmailova 1989,
Marusik et al. 1995, Marusik et al. 1992, Marusik et al. 1993, Marusik et al. 1993,
Marusik and Kovblyuk 2011, Marusik et al. 2000). The araneid fauna of the former
Soviet Union has been summarized in the catalogues of Mikhailov (1996, 1997,
1998, 1999, 2000, 2002, 2013). By December 31, 2015, 3374 and 2397 spider species
have been reported from the FSU territories and Russia, respectively (Mikhailov
2016b). From Ukraine are known 1016 spp. (end 2015).
From the relatively small territory of Tuva (170,000 km2), Marusik et al. (2000)
recorded 614 spp. (36% of them in Linyphiidae). From Ural are known ca. 800 spp.
The monograph of Song et al. (1999) enumerates for the entire vast territory of
China (including Taiwan) 2361 spp. of spiders of 450 genera and 56 families (almost
half of the total number of 114 spider families). Two of the families (Dipluridae and
Dysderidae) have not been recorded for continental China (only from Taiwan).
From the families of Chinese spiders, most genera (over 20) have Linyphiidae
(100), Salticidae (71), Araneidae (37), Thomisidae (29), Theridiidae (27), and
Gnaphosidae (26). Since 1999 have been published many new papers (mostly by
Chinese specialists) and added many new genera and species to the list. Meng et al.
(2008) outlined the biogeographical patterns of Chinese spiders, using the parsi-
mony analysis of endemicity. They provided slightly different figures for the fami-
lies (58, Zoridae included) and species (2858) of the Chinese spiders. Most genera
have Linyphiidae (112), Salticidae (88), Araneidae (41), Thomisidae (30),
Theridiidae (36), Thomisidae (30), and Gnaphosidae (30). These figures certainly
have been changed since 2008. Meng et al. (2008) indicate that “Traditionally
[based on vertebrates], China has been divided into two realms and seven biological
provinces. The Palearctic realm includes the northeastern, northern, Inner-Mongolia-
Xinjiang, and Qinghai-Tibetan biological provinces. The Oriental realm includes
the Southwestern, Central, and Southern biological provinces. This division, origi-
nally established by Cheng & Zhang (1959), was later modified by Zhang & Zhao
(1978) and Zhang (1998, 2004).”
No endemic families.
Some endemic genera of spiders in the described area:
Fam. Oonopidae
Spinestis Saaristo et Marusik, 2009 – Ukraine (one sp.)
Fam. Linyphiidae
Typhochrestoides Eskov, 1990 – Northern Russia
Opilioacarida – missing
Acariformes
Oribatida The checklist of Chen, Liu, and Wang (2014) indicates that in China
(incl. Hong Kong and Taiwan) are known 599 species and subspecies of Oribatida
of 275 genera and 97 families (Wang, Wen, and Chen 2002, 2003).
Holothyrida – missing
7.15 Sakhalin and Kuril Islands
7.15.1 G
and Paleogeography
The island of Sakhalin (72,492 km2) is detached from the Asian continent by the
Strait of Tartary (Tartar Strait, 4–20 m deep and 7.5 km wide at the narrowest part,
called Nevelskoy Strait) and from Hokkaido by Soya (La Pérouse) Strait. The island
is forested, with the highest point Mount Lopatin (1609 m). The Strait of Tartary is
freezing often and gives way to migrations by different animals.
The Kuril Islands are a chain of 56 islands stretched on 1300 km between
Kamchatka and Hokkaido. The highest point is Alaid volcano, 2339 m on Atlasov
Island, the archipelago has a surface area of 10,503 km2. Only the southern islands
are forested. On the islands there are over 100 volcanoes (40 active), summits of
stratovolcanoes. The climate is harsh, with long winters.
Some researchers (Sergeev 1993) define Sakhalin (together with Hokkaido) as a
province within the Manchurian Subregion of the Palearctic.
The “southern” orders are not represented in the described chain of islands. The
orders reported from this area are Araneae, Opiliones, Pseudoscorpiones,
Mesostigmata, Ixodida, Trombidiformes, and Sarcoptiformes.
Opiliones Suzuki (1941b, 1956) published papers on the opilions of southern
Sakhalin, Kunashir, and Iturup. In another paper, he (Suzuki 1967b) concluded that
all five species of opilions on Sakhalin belong to the Holarctic fauna. Between
Sakhalin and Hokkaido is situated the Hatta Line. Tsurusaki and Crawford (2001)
analyzed the opilions of Kuril Islands and concluded that on the islands are distrib-
uted 12 species and that Miyabe Line (= Iturup Strait lying between Iturup and
7.16 Caucasus and Transcaucasia 375
Urup) is of certain significance both for the distribution of the flora and the fauna of
the islands. “Along the chain of islands, the number of species per island gradually
decreased from Hokkaido toward the northeast: Hokkaido (20 species), Kunashir
(10), Iturup (nine), Urup to Ketoi (four, except for Simushir where there are five),
and Ushishir to Kharimkotan (two).” According to them, Nipponopsalis yezoensis
and Mizozatus flavidus are the most widespread in the islands. The much larger
Sakhalin has richer, but underexplored, fauna of Opiliones (Suzuki 1967b, Gricenko
1979), Phalangiidae (Oligolophus thienmuschanensis Wang, Phalangium opilio
L.), and others.
Araneae In May 2001 in Sapporo there was a Symposium on Kuril Island

Biodiversity. Some lectures and posters on spiders by Tanaka, Kamura, Logunov,
and Marusik and especially by Marusik and Crawford (2001) outlined the biogeog-
raphy of this group on the islands. Marusik et al. (1993) published a checklist of
spiders from Sakhalin and Kuril Islands.
Endemic genera of spiders on the Kuril Islands:
Fam. Hahniidae: Pacifantistea Marusik, 2011
7.16 Caucasus and Transcaucasia
7.16.1 G
and Paleogeography
The Caucasus Mountains is a mountain system between the Black Sea and the
Caspian Sea. It includes the Greater Caucasus Mountain Range and the Lesser
Caucasus Mountains.
The Greater Caucasus Range extends from the Black Sea nearly to Baku on the
Caspian Sea, while the Lesser Caucasus runs parallel to the greater range, at a dis-
tance averaging about 100 km south. The Meskheti Range is a part of the Lesser
Caucasus system. The Greater and Lesser Caucasus ranges are connected by the
Likhi Range, which separates the Kolkhida Lowland from the Kura-Aras Lowland.
The Lesser Caucasus and the Armenian Highland constitute the Transcaucasian
Highland. The highest peak in the Caucasus range is Mount Elbrus in the Greater
Caucasus, which rises to a height of 5642 meters above sea level. Annual precipita-
tion in the Western Caucasus ranges from 1000–4000 mm. The absolute maximum
annual precipitation is 4100 mm on the Meskheti Range in Ajaria. Snow cover in
several regions (Svaneti and Northern Abkhazia) may reach 5 meters. The Mt.
Achishkho region, which is the snowiest place in the Caucasus, often records snow
depths of 7 meters. The formation of the present structure of the Caucasus and adja-
cent areas is connected with the unidirectional compression of this area and the
rotation of the compressional axis from a northeastern to a sub-meridional direction
in the course of the Cimmerian-Alpine cycle (Gamkrelidze 1986). “The mountain-

ous Crimea and Caucasus Major appeared as islands in the Sarmat Sea 10-5.4 mil-
lion years ago…; Caucasus Minor, Asia Minor and Balkan compounded the single
massif” (Kovblyuk 2004).
The environment north and south of the Caucasus is quite different, and some group
disappear or are strongly reduced north of the mountain (Solifugae, Scorpiones).
Some orders and suborders are missing in the area: Amblypygi, Uropygi,
Schizomida, Opiliones, Cyphophthalmi, Laniatores, Opilioacarida, and Holothyrida.
Palpigradi Christian (2014) described the species Eukoenenia vargovitshi from
Abkhazia as an element bridging the apparent gap between Romania and India.
Solifugae
From Caucasus and Transcaucasia have been recorded five families of Solifugae
(Aliev and Gadzhiev 1983, Birula 1936a, 1936b, Morin 1928).
Fam. Daesiidae – Bitonissus schelkovnikovi (Birula, 1917)
Fam. Karschiidae – Karschia (K.) caucasia (L. Koch, 1878)
Fam. Galeodidae – Galeodes araneoides (Pallas, 1772)
Fam. Gylippidae – Gylippus (Paragylippus) caucasicus caucasicus Birula, 1907
Fam. Rhagodidae – Rhagodes caucasicus Birula, 1905
Scorpiones
In Caucasus and Transcaucasia are known six spp. of scorpions (Birula 1899, 1917a,
Fet 1988, Kinzelbach 1980, etc.) as follows:
Fam. Buthidae
Androctonus crassicauda (Olivier, 1807)
Mesobuthus caucasicus (Nordmann, 1840)
M. eupeus (C.L. Koch, 1839)
Fam. Euscorpiidae
E. (Alpiscorpius) mingrelicus (Kessler, 1874)
Fam. Iuridae
Calchas nordmanni Birula, 1899
Pseudoscorpiones According to Dashdamirov and Schawaller (1992), in Caucasus
have been recorded 66 spp. of pseudoscorpions, belonging to 29 genera and 10
families: Chthoniidae, Neobisiidae, Atemnidae, Cheliferidae, Chernetidae,
7.16 Caucasus and Transcaucasia 377
Geogarypidae, Garypinidae, Olpiidae, Cheiridiidae, and Withiidae (Daday 1889,

Redikorzev 1926, 1930, Schawaller 1983, 1994b, Schawaller and Dashdamirov
1988).
In Armenia are known 18 spp. of 12 genera and the families Chthoniidae,
Neobisiidae, Garypidae, Cheiridiidae, Olpiidae, Cheliferidae, and Chernetidae.
In Georgia are known 48 spp. of 16 genera, and 6 families. Remarkable are the
two troglobitic Neobisium (Blothrus), described by Lapschoff (1940).
In Azerbaijan are known 50 spp. of 26 genera and 10 families (Dashdamirov
1990, actualized). In this paper is done a zoogeographical analysis with 11 zoogeo-
graphical complexes. Asia Minor – Caucasian elements are 23.0% of species;
endemic Caucasian complex includes six species (17.9%); Palearctic type of areal
comprises 15.4%; Mediterranean elements are 10.3%; European complex is 10.3%.
Two species are cosmopolitic.
Opiliones The opilionids of Caucasus were (and still are) explored by several
specialists:
Mcheidze (1964), Martens (2006), Snegovaya (1999, 2011), Snegovaya and
Chemeris (2005, 2016), Staręga (1966), and others.
Cyphophthalmi and Laniatores – not recorded
Eupnoi
Fam. Phalangiidae – Odiellus Roewer, Lacinius Thorell, Phalangium L.,
Metaplatybunus Roewer, Opilio Herbst, Rilaena Šilhavý, etc.
Fam. Sclerosomatidae – Nelima Roewer
Dyspnoi
Fam. Trogulidae – Trogulus Latreille, Calathocratus Simon (= Platybessobius
Roewer)
Fam. Nemastomatidae – Nemastoma C.L. Koch, Paranemastoma Redikorzev,
Histricostoma Kratochvil, Mitostoma Roewer, Mediostoma Kratochvil, Nemaspela
Šilhavý, Giljarovia Kratochvil, Caucnemastoma Martens, Vestiferum Martens
Endemic genera in Caucasus area are Caucnemastoma Martens and Vestiferum
Martens.
Fam. Gylippidae – Gylippus (Paragylippus) caucasicus caucasicus Birula, 1907
Scorpiones
In Caucasus and Transcaucasia are known six spp. of scorpions (Birula 1899, 1917a,
Fet 1988, Kinzelbach 1980, etc.) as follows:
Fam. Buthidae
Androctonus crassicauda (Olivier, 1807)
Mesobuthus caucasicus (Nordmann, 1840)
M. eupeus (C.L. Koch, 1839)
Fam. Euscorpiidae
E. (Alpiscorpius) mingrelicus (Kessler, 1874)
Fam. Iuridae
Calchas nordmanni Birula, 1899
Pseudoscorpiones According to Dashdamirov and Schawaller (1992), in Caucasus
have been recorded 66 sp. of Peudoscorpions, belonging to 29 genera and 10 fami-
lies: Chthoniidae, Neobisiidae, Atemnidae, Cheliferidae, Chernetidae, Withiidae
(Daday 1889, Redikorzev 1926, 1930, Schawaller 1983, 1994b, Schawaller and
Dashdamirov 1988).
In Armenia are known 18 sp. of 12 genera and the families Chthoniidae,
Neobisiidae, Garypidae, Cheiridiidae, Olpiidae, Cheliferidae, Chernetidae.
In Georgia are known 48 sp. of 16 gen, and six families. Remakkable are the two
troglobitic Neobisium (Blothrus), described by Lapschoff (1940).
In Azerbaidjan are known 50 sp. of 26 genera and ten families (Dashdamirov
1990, actualized). In this paper is done a zoogeographical analysis with 11 zoogeo-
graphical complexes. Asia-Minor – Caucasian elements are 23.0% of species;
endemic Caucasian complex includes six species (17.9%); Palearctic type of areal
comprises 15.4%; Mediterranean elements are 10.3%, European complex – 10.3%.
Two species are cosmopolitic.
Opiliones
The opilionids of Caucasus were (and still are) explored by several specialists:
Mcheidze (1964), Martens (2006), Snegovaya (1999, 2011), Snegovaya and
Chemeris (2005, 2016), Staręga (1966), and others.
Cyphophthalmi and Laniatores – not recorded
Eupnoi
Fam. Phalangiidae – Odiellus Roewer, Lacinius Thorell, Phalangium L.,
Metaplatybunus Roewer, Opilio Herbst, Rilaena Šilhavý, etc.
Fam. Sclerosomatidae – Nelima Roewer
Dyspnoi
Fam. Trogulidae – Trogulus Latreille, Calathocratus Simon (= Platybessobius
Roewer)
Fam. Nemastomatidae – Nemastoma C.L. Koch, Paranemastoma Redikorzev,
Histricostoma Kratochvil, Mitostoma Roewer, Mediostoma Kratochvil,
Nemaspela Šilhavý, Giljarovia Kratochvil, Caucnemastoma Martens, Vestiferum
Martens,
Endemic genera in Caucasus area are Caucnemastoma Martens and Vestiferum
Martens.
7.17 Turkey, Cyprus, Sinai, the Dodecanese Islands, Iran, Afghanistan, the Near East,… 379
Araneae Over 1000 species belonging to 46 families are known from the Caucasus
(Buchar and Thaler 1998, Ovcharenko 1978, 1979, Tanasevitch 1987, 1990).
The species richest families are as follows: Linyphiidae (~180), Salticidae (122),
Gnaphosidae (>100), Lycosidae (>100), Theridiidae (80), Dysderidae (70), and
Thomisidae (70). In the different families, endemism values vary from 0 to 100%.
The average level of endemism in the Caucasus is about 22%, and the highest level
of endemism among species-rich families was found in the Dysderidae, being
around 60% (Marusik et al. 2006). According to Mikhailov and Mikhailova (2006),
quantitatively Gnaphosidae is the second most abundant family after the Lycosidae.
According to Marusik and Guseinov (2003), in Azerbaijan have been found over
600 spp. of spiders of 43–44 families – almost the whole number (51) of the spider
families of the former USSR. From Armenia are known 127 spp. and from Georgia
456 spp. (Mikhailov 2002).
7.17 T
urkey, Cyprus, Sinai, the Dodecanese Islands, Iran,
Afghanistan, the Near East, Middle Asia,
and the Arabian Peninsula
7.17.1 Geography, General Zoology, and Paleogeography
The described area includes Sinai, Asiatic Turkey, Cyprus, the Greek islands near
the shore of Asia Minor (Rhodes, Karpathos, Kassos, Kos, Kalymnos, Samos,
Chios, Mitilini, and some smaller islands), Iran, Afghanistan, Syria, Lebanon,
Israel, Palestine, Jordan, Saudi Arabia, Yemen, Oman, Qatar, UAE, Kuwait, Iraq,
Bahrain, Uzbekistan, Tajikistan, Kyrgyzstan, Turkmenistan, and Kazakhstan. The
area is mostly dry, with deserts (Iran) and high mountains like Hindu Kush, Tian
Shan, and Pamir, some volcanoes, etc. (Turkey, Ararat, 5165 m; Iran, Damavand,
5610 m in Alborz Mt.; Afghanistan, Noshaq, 7453 m, second highest summit of
Hindu Kush, after Tirich Mir (7690 m in Pakistan); Kyrgyzstan, Jengish Chokusu
(Pik Pobedy), 7439 m, Tian Shan; Tajikistan, Ismoil Somoni Peak in Pamir, 7495
m.). The highest summit of the Arabian Peninsula is Jabal an Nabi Shu’ayb, at
3666 m (in Yemen), and of Lebanon is Qurnat as Sawda’ (3088 m).
Some sources: Horowitz (1975), Kosswig (1955), Krumsiek (1976), Kuznetsov
(1950, 1957), Kusnezow (1924), Lopatin (1969), Vigna Taglianti et al. (1999)
Anatolia is the region where three of the world’s 35 biodiversity hotspots meet, and interact:
the Caucasus, Irano-Anatolian, and Mediterranean basin hotspots. One of the most distinc-
tive biogeographic features that helps in understanding the biodiversity of Anatolia is the
Anatolian diagonal which has long been recognised as a biogeographic boundary between
the central and eastern Anatolian floras and faunas. (Gür 2016)
Analyzing the zoogeography of the Levant, Por (1975) reached to the conclusion
that the Levant province is a peculiar and complex “subtraction-transition zone”
(following the expression of Darlington, 1957). The Levant is “a stretch of land

about 150 km wide, wedged in between the [Mediterranean] sea and the Syrio-
Arabian desert, stretching from the mouth of the River Orontes and the Amanus and
Taurus mountain ranges in the north, to the Isthmus of Suez in the south” (Por
1975). This “subtraction-transition zone” between the Palearctic and Ethiopian
regions, born by the desertification, is called Paleo-eremic region. Usually the biota
of the Levant is considered to be Palearctic (Mediterranean), but, according to Por
(1975), the inclusion of the Old World deserts in the Holarctic is due to a certain
“Europa-centrism” of the scientists. “According to this scheme, the Ethiopis begins
only south of the Sahara, in the Savannas, where the last of the typical Palaearctic
animals fade out. However, if at all, the connections of the Palaeo-eremic faunal
inventory are much closer to the Ethiopis than to Palaearctis….Our [Israel’s] typical
desert animals, such as scorpions, agamas, gerbils, sand grouse and gazelles, have
Ethiopian rather than Holarctic connections, even if some species are so-called
“Mediterraneans” (Por 1975).
According to the systems of Bodenheimer (1935) and Por (1975), in the Levant
could be distinguished four faunal elements: Palearctic, Paleo-eremic, Ethiopian,
and Oriental, with clear prevalence (especially in the north) of the Palearctics in
most of the animal groups. The line along the foothills of Northern Galilee and the
Golan Heights was called by Por (1975) “Nehring Line.” South of it follows a “tran-
sitional zone,” where Palearctic elements mingle with Paleo-eremic elements.
More to the south, Por (1975) outlines a “Bodenheimer Line,” which is the end
of the transitional zone. Follow Ramon Mountains and the mountains of Sinai. The
“Ethiopian element does not prevail in any area. The Oriental species are even more
scattered, without showing any geographical pattern. The endemic species among
the animals are chiefly of Palaeo-eremic, Ethiopian and also of Oriental origins…
this suggests a higher age of the tropical element, and perhaps a younger and more
expansive character of the Holarctic species” (Por). In conclusion, “The Levant
province is a meeting place and transitional area between the Palaearctic, Oriental
and Ethiopian zoogeographic regions.” The broad “Paleo-eremic desert belt serves
as a filtering barrier between the three faunal regions.”
Turkey is located on cross section of the Boreal, Euro-Siberian, Irano-Turan, and
Mediterranean elements from Central Asia, Mesopotamia, Caucasus, and Balkans
(Akman 1993).
In this large, mostly arid, area are known the orders Palpigradi (only three spp.) and
Solifugae and many genera and species from the families Galeodidae, Daesiidae,
Karschiidae, Gyllipidae, and Rhagodidae, Scorpiones (Pseudochactidae, Akravidae,
Buthidae, Euscorpiidae, Hemiscorpiidae, Scorpionidae, Iuridae), Pseudoscorpiones
(Atemnidae, Cheiridiidae, Cheliferidae, Chernetidae, Chthoniidae, Garypinidae,
Garypidae, Geogarypidae, Menthidae, Lechytiidae, Neobisiidae, Olpiidae,
Withiidae), Opiliones (Sironidae, Phalangiidae, Sclerosomatidae, Trogulidae,

Dicranolasmatidae, Nemastomatidae, Phalangodidae), Amblypygi (six spp. – four
Charinidae and two Phrynichidae), Araneae (50 families), and Opilioacarida (one
sp. of Paracarus from Middle Asia, endemic genus Adenacarus from Yemen). In
the described area, there is only one endemic family (Akravidae).
Six orders of class Arachnida were recorded from Lebanon (El-Hennawy 2013) in
addition to Acari. They are Araneae (38 families, 109 genera, 165 species), Scorpiones
(2 families, 10 genera, 12 species), Pseudoscorpiones (7 families, 8 genera, 10 spe-
cies), Opiliones (3 families, 8 genera, 10 species), Solifugae (1 family, 1 genus, 2
species), and Palpigradi (1 family, 1 genus, 2 species). The total is 52 families, 137
genera, and 201 species. Most of this fauna is clearly Palearctic. The south of the
Arabian Peninsula (Yemen, Oman) is usually considered part of Afrotropical Region
(Oman also with Oriental elements). Missing from it are the orders Ricinulei, Uropygi,
Schizomida, and Holothyrida and the suborders Palaeoamblypygi and Mesothelae
(Gromov and Kopdykbaev 1994, Whittick 1941).
Palpigradi Almost nothing is known about these tiny and rarely collected animals
from the vast and mostly dry area from Turkey and Arabia to Afghanistan (Condé
1965, 1974). The only exceptions are Eukoenenia juberthiei, Condé, 1974, from
Lebanon and the strange Leptokoenenia gerlachi, Condé, 1965, from Farasan Island
near Saudi Arabia (type of a new genus, found next year also in DR Congo).
Solifugae In the vast area of Central Asia and the Near East (Israel, Jordan, Lebanon,
Iraq, Kuwait, Syria, Turkey), the Solifugae are widespread. Recorded are the fami-
lies Galeodidae, Daesiidae, Karschiidae, Gyllipidae, and Rhagodidae (Aliev and
Gadzhiev 1983, Birula 1941, Caporiacco 1948, El-Hennawy 1998 2007, Gromov
2004, Lawrence 1953, Prendini et al. 2006, Roewer 1934, Simon 1872, Turk 1948).
In Iran are recorded 67 spp. of Solifugae from five families (Kraus 1959; Roewer
1952 1960, with suppl., Harvey 2013b). In Turkmenistan are found 31 spp. (Gromov
1998 and others). In Afghanistan 41 spp. of Solifugae are known so far (Lawrence
1956 and suppl.). In Asia Minor are found 37 spp. of Solifugae: Daesiidae (Blossia
Simon, Gnosippus Karsch, Gluviopsilla Roewer, Gluviopsis Roewer, Gluviopsida
Roewer, Biton Karsch), Galeodidae (Galeodes Olivier), Gylippidae (Gylippus
Simon), Karschiidae (Karschia Simon, Eusimonia Kraepelin, Barussus Roewer),
Rhagodidae (Rhagodia Roewer), and Solpugidae (Solpugella Roewer).
In some other countries in the area, the number of Solifugae spp. is as follows:
Israel (65 spp.), Jordan (5 spp.), Saudi Arabia (14 spp.), Syria (17 spp.), and Lebanon
(2 spp.) (Harvey, 2013b). It is clear that this fauna is understudied – the huge number
of Solifugae in the tiny Israel is obviously due to the good specialists in this country.
In Rhodes (Greece) live four spp. of Solifugae of two fam.: Gluviopsilla discolor
Kraepelin, Gluviopsis rufescens Pocock, Gluviella rhodiensis Caporiacco [endemic
genus] (Daesiidae), and Galeodes rhodicola Roewer [endemic] (Galeodidae).
In Cyprus live five spp. of Solifugae of four families (Blick 2004). They are
Biton ehrenbergi Karsch (Daesiidae), Galeodes graecus C.L. Koch (Galeodidae),
Gylippus cyprioticus Lawrence, G. syriacus Simon (Gyllipidae), and Eusimonia

furcillata Simon (Karschiidae). Only G. cyprioticus is endemic.
The number of Solifugae from the republics of Middle Asia is 62 (Mikhailov
2016a, inf. from Gromov in litt.).
In total, from the described area, we have the following:
Endemic genera:
Gluviola Roewer – G. armata Birula, 1905 (Iran)
Gluviopsida Roewer – G. taurica Roewer, 1933 (Turkey)
Haarlovina Lawrence – H. nielsi Lawrence, 1956 (Afghanistan)
Mumaella Harvey – M. robusta Lawrence, 1956 (Afghanistan)
Ceratobiton Delle Cave et Simonetta – C. styloceros Kraepelin, 1899 (Israel and
Jordan)
Gluviella Caporiacco – G. rhodiensis Caporiacco, 1948 (Rhodes)
Galeodumus Roewer – G. colognatoi Roewer, 1960 (Afghanistan)
Roeweriscus Birula – R. paradoxus Birula, 1937 (Iran)
Zombis Simon – Z. pusiola Simon, 1882 (Israel)
Acanthogyllipus Birula – A. judaicus Kraepelin, 1899 (Israel)
Rhagodelbus Roewer – Rh. bucharicus Birula, 1935 (Uzbekistan)
Rhagodax Roewer – Rh. wadidaba Roewer, 194 (Jordan)
Rhagodospus Roewer – Rh. babylonicus Birula, 1935 (Iraq)
Rhagoduja Roewer – Rh. finnegani Roewer, 1933 (Iran)
Rhagodula Roewer – Rh. nigra Roewer, 1941 (Israel)
Scorpiones The list of Vachon (1966) covers most of the countries in this sector. In
contains 78 spp. of the families Buthidae, Euscorpiidae (to use the modern
classification), Scorpionidae (incl. Diplocentridae), and Iuridae. Since this time
many amendments and new data occurred, the most striking being the description of
a new relict species, genus, and family Pseudochactidae Gromov, 1998. The genus
remains endemic, but new genera of the family have been described since from the
caves of Laos and Vietnam (Birula 1917, Cruccitti and Cicuzza 2000, Farzanpay
1988, Fet 1985, Fet et al. 2011, Francke 1981, Gantenbein et al. 2000, Gromov
1998, 2001, Habibi 1971, Khazim Al-Asmari et al. 2013, Kaltsas et al. 2008,
Kovařik 1993, Levy and Amitai 1980, Lourenço 1996, Lourenço and Vachon 1996,
Parmakelis et al. 2006, Vachon 1947, 1951c, 1979a, Warburg et al. 1978).
In the review of Vachon and Kinzelbach (1987) to the 78 spp. in 1966 have been
added another 28, or in total up to 1987 from the Near East have been recorded
106 spp. of scorpions. Have been added the many species described by Levy et al.
(1973) and also by Vachon (1974, 1980) and Francke (1980). Have been described
also the new Kraepelinia Vachon, 1974, and Habibiella Vachon, 1974, from Iran.
From Turkmenistan Fet (1994) are published six genera (Mesobuthus, Orthochirus,
Liobuthus, Anomalobuthus, Pectinibuthus, Kraepelinia) and seven spp. of scorpi-
ons, all from fam. Buthidae. According to Fet (1994), all seven spp. belong to the
Saharo-Gobian (desert Palearctic) genera.
As a whole, from the republics of the former USSR are known 18 spp. of
Scorpiones (Mikhailov 2016a, inf. from Fet, in litt.).
Fet et al. (2009) described a new species, Calchas gruberi, from the Greek
islands Megisti and Samos and from Anatolia. Kovařík et al. (2010) revised the
genus Iurus Thorell (Iuridae), described two new species from Turkey, and con-
cluded that in Turkey live four endemic species of this genus (I. kraepelini, I. kinzel-
bachi, I. asiaticus, and I. kadleci). This and several earlier papers on Turkish
scorpions (Tolunay 1958, Vachon 1947, 1966, Fet and Braunwalder 2000) brought
the number of scorpions in this country to 26 spp. of 12 genera from the families
Euscorpiidae, Buthidae, and Iuridae. The biogeographic analysis of Crucitti (1999a)
says that the Anatolian Peninsula shows its nature of crossroads of different faunis-
tic stocks, as highlighted by its scorpion fauna too.
Yağmur et al. (2011) described from Cyprus the new (and most probably
endemic) species Buthus kunti (Buthidae). According to the monograph of Levy
and Amitai (1980, with suppl.), in Palestine live scorpions of three (modern) fami-
lies: Buthidae, Scorpionidae, and the recently described from dead specimens fam-
ily Acravidae. The present-day fauna of Israel contains 19 recorded spp. of 12
genera: Akrav, Androctonus, Birulatus, Buthacus, Buthus, Compsobuthus,
Hottentotta, Leiurus, Mesobuthus, Orthochirus, Nebo, and Scorpio.
In Jordan Amr and El-Oran (1994) have registered 13 spp. of scorpions of 10
genera and 2 families: Buthidae and Scorpionidae (incl. Diplocentridae). Later this
fauna was completed to 20 spp. of the same genera as in Israel, without Akrav. Nebo
Simon is considered now (not by all specialists) to belong to Scorpionidae.
Extremely interesting is the archaic family Pseudochactidae, from which one spe-
cies (Pseudochactas ovchinnikovi Gromov, 1998) has been described from Tajikistan
and Uzbekistan and another one (P. mischi Soleglad, Kovarik, et Fet, 2012) from
Afghanistan. The genus and the two species are endemic from this area.
In Oman (Vachon 1977, 1980) were found members of the genera Apistobuthus
Finnegan, Butheolus Simon, Buthotus Vachon (Buthidae), and Nebo Simon
(Diplocentridae). Now (2017) the fauna of Oman consists of 39 spp., including sev-
eral endemics, of 16 genera and 3 families. On Rhodes Island live Mesobuthus gib-
bosus (Brullé) of Buthidae and Iurus dufoureius (Brullé) of Iuridae (Gruber 1963).
El-Hennawy (2009) updated the list of scorpions recorded from Saudi Arabia to
23 spp. and 3 subspp., belonging to 3 families: Buthidae (18 spp. of 10 genera),
Hemiscorpiidae (1 sp.), and Scorpionidae (4 spp., 3 subspp.). Another survey
(Khazim Al-Asmari et al., 2013) provides the following figures: 22 spp., 3 subspp.,
and 3 families – Buthidae (18 spp. of 10 genera), Hemiscorpiidae (1 or 2 spp.), and
Scorpionidae (1 sp., 2 subspp.) (Table 7.1).
Endemic genera for this area are:
Fam. Pseudochactidae
Pseudochactas Gromov, 1998 (Tajikistan, Uzbekistan, Afghanistan)
Fam. Buthidae
Neohemibuthus Lourenço, 1996 (Iran)
Paraorthochirus Lourenço et Vachon, 1995 (Iran)
Table 7.1 Scorpiones in Asia Minor, the Near East and Central Asia
Asia
Country Minor Syria Lebanon Jordan Israel Iraq Afgan. Uzb. Kaz. Kir. Tadz.
Number of sp. 26 20 10 19 19 20 29 8 5 2 6
Taxa
Fam. Akravidae – – – – + – – – – – –
Akrav Levy – – – – 1 – – – – – –
Fam. Buthidae + + + + + + + + + + +
Afghanobuthus – – – – – – 1 – – – –
Lour.
Androctonus 1 3 3 4 3 1 2 – – – –
Ehr.
Anomalobuthus – – – – – – – 1 1 – –
Kr.
Apistobuthus – – – – – – – – – – –
Finn.
Birulatus – 1 – 1 1 – 1 – – – –
Vachon
Buthacus Birula – 3 1 1 2 2 1 1 – – –
Buthus Leach – – – 2 1 – – – – – –
Compsobuthus 2 5 1 5 4 2 3 – – – –
Vach.
Hottentotta 2 1 1 1 1 3 5 – – – 1
Birula
Iranobuthus – – – – – – – – – – –
Kovarik
Isometrus Ehr. – 1 – – – – – – – – –
Kraepelinia – – – – – – – – – – –
Vachon
Leiurus 1 2 1 2 1 – – – – – –
Ehrenberg
Liobuthus Birula – – – – – – – 1 1 – –
Mesobuthus 4 2 1 – – 3 2 2 2 2 2
Vachon
Odontobuthus – – – – – 2 – – – – –
Vachon
Orthochirus 1 – – 1 1 2 9 2 1 – 1
Karsch
Polisius Fet – – – – – – – – – – –
et al.
Psammobuthus – – – – – – – – – – 1
Birula
Razianus – – – – – – – – – – –
Farzanpay
Sassanidiothus – – – – – – 2 – – – –
Far.
(continued)
Table 7.1 (continued)
Asia
Country Minor Syria Lebanon Jordan Israel Iraq Afgan. Uzb. Kaz. Kir. Tadz.
Vachoniolus – – – – – – – – – – –
Levy et al.
Fam. + – – – – (+) – – – – –
Euscorpiidae
Euscorpius 6 – – – – (+) – – – – –
Thorell
Scorpiops Peters – – – – – – – – – – –
Fam. – – – – – + – – – – –
Hemiscorpiidae
Hemiscorpius – – – – – 1 – – – – –
Peters
Fam. – + + + + + – – – – –
Scorpionidae
Nebo Simon 1 1 1 1 – 1 – – – – –
Scorpio L. – 1 1 1 1 1 – – – – –
Fam. Iuridae + – – – – + – – – – –
Calchas Birula 4 – – – – 2 – – – – –
Iurus Thorell 1 – – – – – – – – – –
Neocalchas 1 – – – – – – – – – –
Yagmur et al.
Protoiurus 3 – – – – – – – – – –
Soled.et al.
Country Turkm. Saudi Arabia Qatar UAE Bahrain Yemen Oman
Number of 7 29 6 14 4 37 39
species
Taxa
Fam. Buthidae + + + + + + +
Androctonus Ehr. – 4 1 1 1 2 2
Anomalobuthus 1 – – – – – –
Kraepelin
Apistobuthus – 1 1 1 – 1 1
Finn.
Babycurus – – – – – 1 1
Karsch
Buthacus Birula – 3 2 3 2 – 1
Butheolus Simon – 4 – 1 – 2 2
Buthus Leach – – – – – 1 –
Compsobuthus – 5 1 4 – 6 4
Vachon
Femtobuthus – – – – – – 1
Lowe
Hottentotta – 2 – 2 – 4 4
Birula
(continued)
Country Turkm. Saudi Arabia Qatar UAE Bahrain Yemen Oman

Kraepelinia 1 – – – – – –
Vachon
Leiurus – 4 – – – 3 2
Ehrenberg
Liobuthus Birula 1 – – – – – –
Mesobuthus 1 – – – – – –
Vachon
Microbuthus – – – – – 1 3
Kraep.
Odontobuthus – – – – – – 2
Vach.
Orthochiroides – – – – – 2 –
Kov.
Orthochirus 2 2 – – – 1 4
Karsch
Parabuthus – 1 – – – 1 –
Pocock
Pectinibuthus Fet 1 – – – – – –
Picobuthus Lowe – – – – – – 2
Vachoniolus – 1 – 1 – – 3
Levy et al.
Fam. – – – – – (+) –
Euscorpiidae
Euscorpius – – – – – (+) –
Thorell
Fam. – + – + – + +
Hemiscorpiidae
Hemiscorpius – 1 – 1 – 3 3
Peters
Fam. – + + – – + +
Scorpionidae
Heteronebo – – – – – 2 –
Pocock
Nebo Simon – – – – – 4 3
Pandinus Thorell – – – – – 1 –
Scorpio Linnaeus – 1 1 – – 1 –
Fam. Akravidae
Akrav Levy, 2007 – Israel
Amblypygi From the described area are known (Delle Cave 1986, 1989, Delle
Cave et al. 2009, Kovařík and Vlasta 1996, Kritscher 1959, Pocock 1895, Weygoldt
et al. 2002, Whittick 1941, Miranda et al. 2016) only six species of Amblypygi:
Fam. Charinidae
Charinus ioanniticus Kritscher, 1959 – Turkey, Israel, the Dodecanese islands
Kos and Rhodes (known also from Egypt)
Charinus israelensis Miranda et al., 2016 – Israel
Charinus omanensis Delle Cave, Gardner, et Weygoldt, 2009 – Oman (cave)
Charinus dhofarensis Weygoldt, Pohl, et Polák, 2002 – Oman
[Sarax mediterraneus Delle Cave, 1986 – Kos and Rhodes] – misidentification
(Seiter et al., 2015)
Fam. Phrynichidae
Phrynichus dhofarensis Weygoldt, Pohl, et Polák, 2002 – Oman
Ph. jayakari Pocock, 1894 – Hadramaut
These are among the species outlining the northern limit of Amblypygi. Actually,
Yemen and Oman and its province Dhofar are part of the south of Arabian Peninsula,
considered usually as part of the Afrotropical Region.
Schizomida From the area described Schizomida occurs only in the Arabian
Peninsula (Enigmazomus eruptoclausus Harvey in Oman) (Harvey 2006b).
Pseudoscorpiones From the whole of Turkey have been published 102 spp. of
pseudoscorpions from 34 genera and the families Chthoniidae, Neobisiidae,
Garypinidae, Lechytiidae, Geogarypidae, Cheiridiidae, Atemnidae, Olpiidae,
Cheliferidae, Chernetidae, and Withiidae (Beier 1949, 1963, 1965a, 1967, 1969c,
1973d, Harvey 2011, Kunt et al. 2008).
From Afghanistan (Beier 1959, 1960, 1961, 1967, 1971, Harvey 2011, 2013,
actualized) have been recorded 32 spp. of 22 genera and the families Neobisiidae,
Olpiidae, Garypinidae, Ideoroncidae, Atemnidae, Cheliferidae, and Chernetidae.
From the Dodecanese and the other islands near Asia Minor have been published
pseudoscorpions by Caporiacco (1948), Mahnert, Beier (1962a), and others.
From Iraq and Kuwait have been published only three spp. of three genera and
the families Cheliferidae and Olpiidae.
Pseudoscorpions from Middle Asia have been published by Dashdamirov and
Schawaller (1992b, 1993a, 1993b, 1995) and others. In Kazakhstan, Uzbekistan,
Turkmenistan, Tajikistan, and Kyrgyzstan are registered scorpions of 30 genera and
10 families: Chthoniidae, Pseudotyrannochthoniidae, Atemnidae, Cheliferidae,
Chernetidae, Garypinidae, Ideoroncidae, Neobisiidae, Cheiridiidae, and Olpiidae.
According to Mikhailov (2016a), from the former USSR are known 144 spp. of
pseudoscorpions.
The pseudoscorpions of Iran have been identified by Beier (1971), Nassirkhani
and Shoushtari (2015), and Nassirkhani (several papers). Now (2017) from this
country are known to live 41 spp. of 23 genera and 10 families: Chthoniidae,
Atemnidae, Cheliferidae, Chernetidae, Garypinidae, Ideoroncidae, Neobisiidae,
Cheiridiidae, Menthidae, and Olpiidae.
From Arabian Peninsula (Saudi Arabia, the Emirates, Bahrain, Qatar, Oman,
Yemen – without Socotra) are known pseudoscorpions of the families Chthoniidae,
Ideoroncidae, Olpiidae, Garypidae, Garypinidae, Menthidae, Atemnidae, Chernetidae,
Cheliferidae, and Withiidae (Mahnert 1980, 1991). According to Mahnert (1991),
“the results obtained in pseudoscorpions [of Arabian Peninsula] seem to be well in
concordance with those of other animal groups: clear affinities to the Afrotropical
region are recognizable in the southwestern part of the peninsula; the species of the
central and northern parts are highly influenced by palearctic elements. The south-
eastern corner yielded oriental elements.”
In Israel, Palestine, Lebanon, and Syria are known 37 spp. of Pseudoscorpiones
from the families Chthoniidae, Neobisiidae, Geogarypidae, Atemnidae, Chernetidae,
and Cheliferidae (Beier 1955, 1963b, Harvey 2011, Mahnert 1974). Endemic gen-
era and species are Paramenthus shulovi Beier, 1963, and Ayalonia dimentmani
Ćurčić, 2008. According to Beier (1963b), Israel’s fauna is predominantly Palearctic,
but with a few species (Lamprochernes savignyi, Nudochernes spalacis,
Myrmecowithius wahrmani) reminding the Afrotropical fauna and with genus
Apolpium reminding the Neotropical fauna.
In Cyprus have been found 12 species of 11 genera and 8 families, and the
endemic genus and species is Mesatemnus cyprianus (Beier et Turk, 1952)
(Atemnidae).
Opiliones From the Dodecanese are registered the families Phalangiidae (Eupnoi),
Trogulidae, Dicranolasmatidae, and Nemastomatidae (Dyspnoi) (Roewer 1924;
Caporiacco 1926, 1929, 1948; Gruber 1963, 1966, 1978; Martens 1965).
From Turkey are known Opiliones of three of the four suborders (Gruber 1969,
1998; Bayram et al. 2010; Mitov 2012, Kurt et al. 2010; Kurt et al. 2013, and
others).
According to Bayram et al. (2010), 50 species plus 3 subspecies of Opiliones
from 25 genera in 6 families inhabit Turkey (Sironidae, Phalangiidae,
Sclerosomatidae, Trogulidae, Dicranolasmatidae, Nemastomatidae). A more recent
checklist (Kurt 2014) enumerates 88 spp. and 7 ssp. of 35 genera and 7 families,
adding the family Ischyropsalididae. From this number 14 spp. are known from
European Turkey.
According to Mikhailov (2016a), from the former USSR are known 178 spp. of
Opiliones.
Opiliones in the described area:
Cyphophthalmi
Fam. Sironidae
Cyphophthalmus Joseph is known from Asia Minor (Gruber 1969).
Eupnoi
Fam. Phalangiidae – Phalangium L., Metaphalangium Roewer, Buresilia Šilhavý,
Dasylobus Simon, Egaenus C.L. Koch, Homolophus Banks, Lacinius Thorell,
Redikorcevia Snegovaya et Staręga, Mitopus Thorell, Oligolophus C.L. Koch,
Opilio Herbst, Platybunoides Šilhavý, Platybunus C.L. Koch, Rafalskia Staręga,

Rilaena Šilhavý, Zachaeus C.L. Koch
Fam. Sclerosomatidae – Leiobunum C.L. Koch
Endemic genera are:
Redikorcevia Snegovaya et Staręga, 2008 (Phalangiidae) – Kazakhstan (one sp.)
Dyspnoi
Fam. Trogulidae – Trogulus Latreille (one sp., Turkey), Trogulocratus Roewer (one
endemic sp., Rhodes – T. rhodiensis Gruber, 1963)
Fam. Dicranolasmatidae – Dicranolasma C.L. Koch (four spp., Turkey, Iraq)
Fam. Nemastomatidae – Giljarovia Kratochvil, Histricosoma Kratochvil,
Mediostoma Kratochvil, Nemastoma C.L. Koch, Paranemastoma Redikorzev,
Vestiferum Martens, Pyza Staręga (two spp., Turkey), Starengovia Snegovaya
(one sp. from Kyrgyzstan)
Endemic genera are:
Fam. Nemastomatidae
Starengovia Snegovaya, 2010 (one sp. from Kyrgyzstan)
Laniatores
From Laniatores is known the family Phalangodidae.
Endemic genera are:
Fam. Phalangodidae
Haasus Roewer, 1949 – Israel (one sp.)
Other sources: Caporiacco (1925), Gruber (1968, 1976, 1979), Logunov (2013),
Mitov (2013), Roewer (1949), Šilhavý (1967), Snegovaya (2004, 2008), Snegovaya
and Staręga (2008)
Araneae In Turkmenistan 335 species of spiders belonging to 162 genera and 38
families are found (Michailov and Fet 1994, Wesolowska 1996).
With mountain uplift and aridization in the Pliocene, Kopet Dagh became a sublatitudinal
dispersal pathway for mesophilic species. About half of araneofauna is represented by
widely distributed species; of the other half, Iranian, Iranian-Turkestanian, European,
European-Caucasian, and European-Mediterranean comprise the majority of the mountain
araneofauna. Turanian desert species are predominant in the lowland deserts (Michailov
and Fet 1994).
The extensive field work of E.M. Andreeva in Tajikistan resulted in a general

survey of the spiders (Andreeva 1975b), preceded by a zoogeographical analysis
(Andreeva 1975a). By this time from Tajikistan have been recorded 260 species and
subspecies of spiders (223 species), from them 50% being Central Asiatic endem-
ics, 13.2% of Mediterranean elements. Others are widely distributed species (8.9%
Palearctic, 7.3% Holarctic, and 2.3% cosmopolitan elements). The figure of
Mediterranean elements (13.2%) curiously coincides with the number of the
Mediterranean elements on Balkan Peninsula (13. 8%, acc. to Deltshev, 1999,

2004). Other supposition of Andreeva (1975a) is that “…it seems highly probable
that the Central Asiatic spider fauna has originated locally from the widespread
genera living there during the Palaeogene period. The high percentage of Central
Asiatic endemics, about half the number of species occurring there, appears to be an
important argument for this assumption.” Spasskiy (1952) presumed that numerous
forms arrived into the Turanian Province from northern areas (connected with
Palaeogene Turgaian mesophilous forests).
The first substantial paper of the spiders of Afghanistan was due to Denis (1958)
on the collection of 74 spp. (30 newly described) of the 3rd Danish Expedition to
Central Asia.
According to Zamani et al. (2017), the spider fauna of Iran is composed of 608
recorded spp./subspp. of 259 genera and 48 families.
The actualized checklist of all Turkish spiders (Bayram et al. 2014) contains
1013 spp. of 330 genera and 53 families. Papers on the Turkish spiders were pub-
lished also by Brignoli (several), Bayram (2002), Seyyar et al. (2008), Topçu et al.
(2005), and others.
Some recent papers on the spiders of some Greek islands near Asia Minor show
how incomplete are the lists available for Turkey and other countries in the area: on
Chios have been recorded 315 spp. (Russel-Smith et al. 2011) and on Lesbos
299 spp. (Bosmans et al. 2009). Even much less is known the spider fauna of Cyprus
(some 70 spp., after Logunov 2013).
From the described area are known spiders of 50 families (no endemic
families).
Some endemic genera in the area are:
Fam. Zoropsidae
Akamasia Bosselaers, 1997 – Cyprus (one sp.)
Fam. Oonopidae
Megabulbus Saaristo, 2007 – Israel
Megaoonops Saaristo, 2007 – Israel
Semibulbus Saaristo, 2007 – Israel (one sp.)
Fam. Tetragnathidae
Zygiometella Wunderlich, 1995 – Israel (one sp.)
Fam. Caponiidae
Iraponia Kranz-Baltensperger et al., 2009 – Iran (one sp.)
Fam. Miturgidae
Israzorides Levy, 2003 – Israel (one sp.)
Fam. Palpimanidae
Levymanus Zonstein et Marusik, 2013 – Israel (one sp.)
7.18 Socotra 391
Fam. Dysderidae
Tedia Simon, 1982 – Israel, Syria (two spp.)
Other sources: Andreeva (1975a, 1975b, 1976), El-Hennawy (2002), Mozaffarian
and Marusik (2001), Mirshami et al. (2015), Simon (1890), Tanasevitch (1989),
Saaristo (2007), and others
Opilioacarida The only species of Opilioacarida in the former Soviet Union
(Kazakhstan) was described by Redikorzev (1937) and assigned by Chamberlin and
Mulaik (1942) to a new genus. Paracarus hexophthalmus (Redikorzev, 1937) is
endemic for Central Asia, but a new (fossil) species (Paracarus pristinus Dunlop,
Wunderlich, et Poinar, 2004) was described from Baltic amber (? Kaliningrad). This
is also the northernmost Opilioacarida in the Eastern Hemisphere.
Acariformes
Oribatida In Iran have been recorded 380 spp. of Oribatida of 191 genera and 86
families (Akrami 2015).
7.18 Socotra
7.18.1 G
and Paleogeography
Socotra (Soqotra) is an archipelago of four islands (Socotra, Abd al Kuri, Samhah,

and Darsah) in the Indian Ocean, at some 240 km from the Horn of Africa and 380 km
from the Arabian Peninsula (politically part of Yemen). The surface area is 3796 km2
and the highest point is 1503 m. It is an isolated part of Gondwana, detached in
Miocene, and has three geographical parts: the narrow coastal plain, a limestone
plateau with karstic caves, and the Haghier Mountains. It has a tropical desert and
semi-desert climate and a mean annual temperature over 18oC (De Geest 2005).
Several endemic birds, no native mammals (except bats). High endemism in phyto-
geography, from 825 plant species 307 are endemic (37%). Several endemic birds, no
native mammals (except bats).
So far absent from the island are the orders Thelyphonida (Uropygi), Schizomida,
Holothyrida, and Opilioacarida and the suborders Palaeoamblypygi, Cyphophthalmi,
Eupnoi, Dyspnoi, and Mesothelae.
Solifugae There is one endemic species Gluviopsis balfouri (Pocock) known from
the archipelago (Socotra, Samha, Abd al Kuri) (Pocock 1889).
Scorpiones At least five species of scorpions are known from the Archipelago.
Two belong to the Family Buthidae, the relatively large and most common yellow-
ish Hottentotta socotrenis (Pocock, 1889) (recorded from Socotra and Samha), and
the dark coloured and apparently more rare Orthochirus bicolor insularis (Pocock,
1889) (Socotra). Widespread on Socotra is the brownish Hemiscorpius socotranus
Pocock, 1889. The specimens recorded on Samha and Darsa are more light in colour
and need further studies to clarify their status. The remaining two forms belong to
the Family Diplocentridae (Heteronebo forbesii Pocock, 1889; H. granti Pocock,
1889), and were supposed to be restricted to Abd al Kuri. However during the last
expeditions we found Heteronebo on Socotra too. It is a small genus, but of great
taxonomic and zoogeographical interest, because it has a peculiar distribution, with
the two (or three ?) forms on the Archipelago, and the next relatives on Caribbean
islands. (Pocock 1898)
Pseudoscorpiones On the island are known 19 spp. of pseudoscorpions of 12 gen-

era and the families Chthoniidae, Ideoroncidae, Garypidae, Geogarypidae, Olpiidae
(7 spp.), Menthidae, Withiidae, and Cheliferidae (Mahnert 2007, he described 17
new species, for the time being considered endemic). According to him, the fauna
has clear affinities with the East African pseudoscorpions and more discrete affini-
ties with the Arabian fauna.
Endemic genus:
Pseudomenthus Mahnert, 2007 (three spp.), fam. Menthidae (Socotra is consid-
ered by Mahnert a hot spot for this rare family)
Opiliones
Laniatores From the island are known Opiliones of the family Biantidae (Hirst
1911, Loman 1902, Pocock 1903). The endemic harvestmen Hinzuanius flaviven-
tris (Pocock, 1903) (Biantidae) is conspicuous in reddish color.
Amblypygi From Giniba Cave in Socotra was described the blind endemic spe-
cies Charinus stygochtobius Weygoldt et Van Damme, 2004. It was added to the two
other Socotran species Charinus soqotranus Weygoldt, Pohl et Polak, 2002, and
Phrynichus heurtaultae Weygoldt, Pohl, et Polák, 2002.
Araneae From Socotra have been recorded 42 species of spiders (32) from 36
genera (4 endemics) and 22 families. Salticidae is the best represented, with 15 spp.
(13 endemics) (Blackwall 1877, Hirst 1911, Pocock 1903, Lehtinen 1967, Saaristo
and Van Harten 2002, Wesolowska and Van Harten 2002, Jocqué and van Harten
2015).
7.19 Karakorum, Hindu Kush, Pamir, Tian Shan, Himalaya, and Tibet 393
Endemic genera are:

Fam. Barychelidae
Atrophothele Pocock, 1903 (one sp.)
Fam. Oonopidae
Socotroonops Saaristo et van den Harten, 2002 (one sp.)
Fam. Palpimanidae
Scelidomachus Pocock, 1899 (one sp.)
Fam. Linyphiidae
Agelenella Lehtinen, 1967 (one sp.)
Fam. Thomisidae
Bassaniodes Pocock, 1903 (one sp.)
Dimizonops Pocock, 1903 (one sp.)
Parasitiformes
Fam. Ixodidae (Hyalomma truncatum Koch, 1844; Boophilus annulatus Say,
1821)
7.19 K
arakorum, Hindu Kush, Pamir, Tian Shan, Himalaya,
and Tibet
7.19.1 G
and Paleogeography
This area includes some of the highest mountains and plateaus on Earth. The culmi-
nant point of Hindu Kush Tirich Mir (in Pakistan) reaches 7690 m. The glaciers in this
mountain cover 6200 km2, and the snow line is at 4650 m on the northern slope and
at 5400 m on the southern slope. On the southeast slopes of Hindu Kush, the upper
forest limit reaches 3300–4000 m; above this altitude spread mountain pastures.
Himalaya sensu stricto is called the system of Tertiary ranges south and west of
Tsangpo (Brahmaputra) and east of the Indus River. North of them lie the ranges of
“Transhimalaya” (Zanskar‚ Ladakh, and Karakorum) and Tibet‚ and in the south are
situated the Siwalik and other lower mountains.
Himalaya is the highest mountain system in the world‚ containing more than 500
summits higher than the Montblanc. This system includes also 10 of the 14 summits
higher than 8000 m. Himalaya is a very young mountain. In the Pleistocene its altitude
should have been only half of the present 8848 m. The collision between the Indian
subcontinent and Eurasian continent started in Paleogene time and continues today.
The Indian plate continues to move northward relative to Asia about 5 cm per year.
With an average elevation exceeding 4500 m, the Tibetan Plateau is the world’s
highest and largest plateau, with an area of 2,500,000 square kilometers. The Tibetan
Plateau is surrounded by massive mountain ranges. The plateau is bordered to the
south by the Himalayan Range, to the north by the Kunlun Range which separates
it from the Tarim Basin, and to the northeast by the Qilian Range which separates
the plateau from the Hexi Corridor and Gobi Desert. To the east and southeast, the
plateau gives way to the forested gorge and ridge geography of the mountainous
headwaters of the Salween, Mekong, and Yangtze rivers in western Sichuan (the
Hengduan Mountains) and southwest Qinghai (Bassoullet et al. 1977, Beron 2008a,
Colchen 1981, Dobremez 1972, 1976, 1978, Le Fort 1996, Mani 1968, Reinig 1930,
Schweinfurth 1957, Troll 1957, Vtorov 1966, Wissmann 1959).
Caporiacco (1934–1935) initiated the studies on the arachnofauna of the highest

parts of the planet, mainly in Karakorum. Beron (2008a, with suppl. Beron, 2016)
summarized the information on the high-altitude Arachnida of these mountain sys-
tems (above 2200 m) (see also Martens 1983, 1984, 1987, 1993, Schmölzer 2001).
The expeditions of J. Martens resulted in hundreds of new taxa and in several analy-
ses (Martens 1981 and others).
Solifugae From the area described have been recorded six families of Solifugae
(Birula 1938, Hirst 1907).
Fam. Rhagodidae – Rhagoderma Roewer, Rhagodia Roewer, Rhagodixa Roewer

Fam. Karschiidae – Karschia tibetana Hirst, 1907 was described from altitude
4570 m in Tibet
Fam. Daesiidae – Daesiola Roewer, Haarlovina Lawrence, Mumaella Harvey
Fam. Galeodidae – Galeodes setulosus Birula, 1938 is known from Tajikistan
(3000 m)
In Pakistan are found 29 spp. of Solifugae of the families Galeodidae (Galeodes
Olivier, 22 spp.) and Rhagodidae (Rhagodes Pocock, 4spp.; Rhagodia Roewer, 1
sp.; Rhagoduna Roewer, 1 sp.; Rhagoderma Roewer, 1 sp.)
Amblypygi
Fam. Charinidae – Charinus pakistanus Weygoldt, 2005 (Pakistan)
Scorpiones In the described area are known scorpions of six families (Fet 1988,
Lourenço 1997, 2003, Lourenço et al. 2005, Lourenço and Qi 2006, Mani 1959, Shi
and Zhang 2005, Tikader and Bastawade 1983, Qi et al. 2005, Zhu et al. 2004, Di
et al. 2013).
Fam. Scorpiopidae – Scorpiops Peters (endemic high-mountain species in Tibet,

Kashmir, and Himalaya), Euscorpiops Vachon (E. montanus Karsch, Himalaya)
7.19 Karakorum, Hindu Kush, Pamir, Tian Shan, Himalaya, and Tibet 395
Fam. Buthidae – Himalayotityobuthus Lourenço (endemic genus, H. martensi

Lourenço, Kashmir), Mesobuthus Vachon
Fam. Scorpionidae – Heterometrus Ehrenberg
Fam. Bothriuridae – Cercophonius Peters (Indian Himalaya)
Fam. Chaerilidae – Chaerilus Pocock (Himalaya)
Fam. Hormuridae – Tibetiomachus Lourenço et Qi (Tibet)
Pseudoscorpiones In Nepal and Bhutan are known 41 spp. of pseudoscorpions of
31 genera and 14 families: Chthoniidae, Lechytiidae, Tridenchthoniidae,
Neobisiidae, Syarinidae, Geogarypidae, Olpiidae, Cheiridiidae, Pseudochiridiidae,
Sternophoridae, Atemnidae, Cheliferidae, Chernetidae, and Withiidae (Beier 1959,
1959, 1974, 1976, Curčić 1980, Mahnert 1977, Mani 1959, Martens 1975,
Schawaller 1983, 1987, 1988, 1991).
Dashdamirov (2004) recorded from northern Pakistan 30 spp. of 20 genera and 9
families: Chthoniidae, Tridenchthoniidae, Geogarypidae, Olpiidae, Neobisiidae,
Cheiridiidae, Atemnidae, Cheliferidae, and Chernetidae. To them is to be added
Withiidae (Nannowithius pakistanicus Beier, 1978, described from Kashmir). The
preliminary conclusion from this study is that “the false-scorpion fauna of the north-
ern mountainous provinces of Pakistan can be concluded to be Palaearctic, with
only few species derived from the Oriental region.”
As a whole, from the 25 species of pseudoscorpions living in the Old World
above 3500 m, 15 are known from Himalaya and Karakorum (Beron 2008a).
Endemic genera from the described region are:
Fam. Neobisiidae
Nepalobisium Beier, 1974 – Nepal (one sp.)
Fam. Tridenchthoniidae
Rheodithella Dashdamirov et Judson, 2004 – N. Pakistan
Fam. Chernetidae
Bipeltochernes Dashdamirov, 2004 – N. Pakistan
Opiliones
As a result of the research mainly of Martens (1972, 1973, 1977, 1978, 1979, 1980,
1984, 1987, 1993), Roewer (1957), and Suzuki (1966a, 1966b, 1970), the extremely
rich fauna of Opiliones of the Himalaya was revealed.
Cyphophthalmi – not recorded
Eupnoi
Fam. Phalangiidae – endemic genera: Himalphalangium Martens (subendemic:
five end. spp. in the described area, one sp. in Japan, one in Korea; other endem-
ics: Homolophus Banks (= Euphalangium Roewer) thienshanense (Šilhavý,

1967) and several other endemic spp., including H. nordenskioeldi (L. Koch), the
highest living Opilion at 5600 m in Karakorum; Egaenus C.L. Koch (several
endemic species)
Fam. Sclerosomatidae – endemic genera: Pseudastrobunus Martens and
Granulosoma Martens (Nepal); Gyoides Martens (6 spp., Nepal), Diangathia
Roewer (1 sp., Nepal), Globulosoma Martens (2 spp., Nepal), Harmanda Roewer
(15 spp. from Nepal and India); Himaldroma Martens (2 end. spp. from Nepal);
Himalzaleptus Martens (1 sp., Nepal), Metaverpulus Martens (7 spp., Nepal,
Bhutan), Nepalkanchia Martens (2 spp., Nepal); Octozaleptus Suzuki (1 sp.,
Nepal); Pokhara Suzuki (7 spp., Nepal); Rongsharia Roewer (3 spp., Nepal);
Zaleptiolus Roewer (4 spp.); other endemics: Gagrella Stolička (several endemic
species), Opilio Herbst (several endemic species in Nepal and Karakorum)
(Gricenko 1975, Janetschek 1990, Staręga and Snegovaya 2008).
Dyspnoi
Fam. Sabaconidae – Sabacon Simon (six endemic species from Nepal, up to above
5000 m)
Laniatores In the described mountain area are represented Laniatores from the
families Sandokanidae (Oncopodidae), Podoctidae, Phalangodidae, Epedanidae,
Assamiidae, and Biantidae.
Fam. Sandokanidae (Oncopodidae) – Gnomulus Thorell

Fam. Podoctidae – in India five species
Fam. Phalangodidae (? Epedanidae) – endemic genus: Dhaulagirius Martens
(Nepal)
Fam. Assamiidae – endemic genera: Assaphala Martens (Nepal), Micrassamula
Martens (Nepal), Nepalsia Martens (Nepal), Nepalsioides Martens (Nepal)
Fam. Biantidae – Biantes Simon (many endemic species from Nepal and India)
Araneae To the important paper of Caporiacco (1934) on the Arachnida of Karakorum
have been added further studies of this mountain system (Beron 2008a, 2016).
Andreeva (1975, 1976) summarized the information on the spiders of Tajikistan.
According to Tikader (1970), the spider fauna of Sikkim consists of 65 species
(45 of them described as new and presumably endemic) of 11 families.
The spiders of the Himalaya above 2200 m belong to the families Anapidae,
Oonopidae, Zodariidae, most of all Linyphiidae, Tetrablemmidae, Amaurobiidae,
Lycosidae, Araneidae, Sparassidae, Clubionidae, Drassodidae, Dictynidae,
Scytodidae, Hahniidae, Gnaphosidae, Sicariidae, Thomisidae, and Salticidae. The
families found above 3500 m are underlined. Spiders (indet.) have been observed up
to 6700 m, Acantholycosa baltoroi (di Caporiacco) – up to 6100 m.
Ixodida Ref.: Clifford et al. (1975), Beron (2008a) Argas himalayensis Hoogstraal
et Kaiser parasitizes Lerwa lerwa up to 4575 m. Ixodes berlesei Birula in Nepal is
found up to 5488 m. Almost all genera of Himalayan Argasidae and Ixodidae are
7.20 Indian Peninsula 397
widespread. Special is Anomalohimalaya Hoogstraal, Kaiser, et Mitchell, high

mountain inhabitant of Pamir, Tajikistan and Himalaya, up to 3800 m.
7.20 Indian Peninsula
7.20.1 G
and Paleogeography
Geographically, the Indian subcontinent is a peninsular region in South-Central

Asia, delineated by the Himalaya in the north, the Hindu Kush in the west, and the
Arakanese in the east and extending southward into the Indian Ocean with the
Arabian Sea to the southwest and the Bay of Bengal to the southeast. This region
rests on the Indian Plate and is isolated from the rest of Asia by mountain barriers
(Alegre et al. 1984, Dobremez 1972, 1976, 1978, Hocutt 1987, Katz and Premoli
1979, Mani (Ed.) 1974, Sahni 1984, Sahni and Kumar 1974). Upland plain (Deccan
Plateau) and plains near Ganges River are typical for the peninsula, and to the north
India shares with Nepal the third highest mountain in the world – Kanchenjunga
(8,586 m). Andaman and Nicobar Islands also belong to India.
The onset of the collision of India with Asia occurred near the Paleocene-Eocene
boundary (Gaetani and Garzanti 1991), some 55 million years ago. To quote Briggs
(1989): “If India broke its contact with the other continents sometime around 148
m.y.a. and, if it existed as an isolated, oceanic continent until the early Miocene, its
fossil terrestrial and shallow marine biotas should demonstrate the evolutionary
effects of more than 100 m.y. of isolation. This means that India should have devel-
oped a peculiar biota with a high percentage of distinct genera and families. But,
with the possible exceptions ...the expected preponderance of peculiar organisms
has simply not been found.”
The northern part of India is the chain of Himalaya where there is a transition
between the Palearctic and the Indomalayan regions and even between two
Kingdoms – the Holarctic and the Paleotropic. It is difficult to differentiate the gen-
era and species of Arachnida in this area (Blasco 1981, Briggs 1989).
In Peninsular India are known the orders Palpigradi, Solifugae (21 spp.),
Scorpiones (117 spp.), Pseudoscorpiones (165 spp.), Opiliones (1 endemic genus
in Cyphophthalmi, Dyspnoi are lacking), Amblypygi (4 endemic species), Uropygi
(endemic genus), Schizomida (several species, 1 endemic genus), Araneae
(1442 spp. of 59 families), and Opilioacarida (1 endemic genus) and many different
mites. Lacking are also the orders Holothyrida and Ricinulei and the suborders
Palaeoamblypygi and Dyspnoi. In India are not recorded endemic families of

Arachnida.
In the western edge of the peninsula (Thar Desert) is passing the band separating
Holarctic and Paleotropic Kingdoms (Map 8.2).
Palpigradi From India have been recorded three taxa: Eukoenenia angusta hindua
Condé, 1989; E. a. tamula Remy, 1960; and E. singhi Condé, 1989.
Solifugae From India are known 21 spp. of Solifugae, belonging to the genera
Galeodes Olivier, 14 spp. (Galeodidae), Gluviopsis Roewer (Daesiidae) and
Rhagodеrma Roewer, Rhagodomma Roewer, Rhagodima Roewer, and Rhagodopa
Roewer (Rhagodidae) (Pocock 1895).
Endemic genera for India are:
Rhagodomma Roewer, 1933 – one sp.
Rhagodima Roewer, 1933 – one sp.
Scorpiones In India are known 117 spp. of Scorpiones of the families Bothriuridae,
Buthidae, Chaerilidae, and Scorpiopidae (see the list) (Ambalaparambil et al. 2010,
Bastawade 2006, Lourenço 1996, 1997, 2003, Tikader and Bastawade 1983,
Vachon 1982).
Pseudoscorpiones From India are known about 160 spp. (and several subspecies)
of pseudoscorpions of 61 genera. Altogether from India, Nepal, and Bangladesh are
known 71 genera and the families Chthoniidae, Hyidae, Geogarypidae, Feaelidae,
Atemnidae, Cheliferidae, Lechytiidae, Chernetidae, Garypidae, Geogarypidae,
Ideoroncidae, Olpiidae, Cheiridiidae, Pseudotyrannochthoniidae, Sternophoridae,
Syarinidae, and Withiidae (Beier 1973, 1974, Murthy and Ananthakrishnan 1977,
Vachon 1982).
Endemic genera for Peninsular India are:
Ectoceras Stecker, 1875 – India (two spp.)
Hygrochelifer Murthy et Ananthakrishnan, 1977 – India (two spp.)
Opiliones In Peninsular India have been registered the families Stylocellidae
(Cyphophthalmi), Phalangiidae, Sclerosomatidae (Eupnoi), and Epedanidae,
Podoctidae, Sandocanidae, and Assamiidae (Laniatores), with several endemic gen-
era (Roewer 1929a, 1929b, Giribet et al. 2007).
Cyphophthalmi
From India (Arunachal Pradesh) has been described an endemic genus and species
Meghalaya annandalei Giribet, Sharma, et Bastawade, 2007 (Stylocellidae)
Eupnoi
Fam. Phalangiidae – Euphalangium Roewer
Fam. Sclerosomatidae – Metagagrella Roewer
7.20 Indian Peninsula 399
Dyspnoi – not represented

Laniatores
Aboriscus Roewer, Buparus Thorell
Fam. Podoctidae (incl. Erecananinae and Ibaloniinae) – Eupodoctis Roewer,
Pumbaraius Roewer, Tandikudius Roewer, Vandaravua Roewer
Fam. Sandokanidae [= Oncopodidae, praeoccup.] – Gnomulus Thorell
Fam. Assamiidae – Assamia Sørensen
Amblypygi On the peninsula are known four species (endemics) (Bastawade et al.
2005, Gravely 1911, 1915, Pocock 1894).
Fam. Charinidae
Charinus Simon – Ch. bengalensis (Gravely, 1911)
Sarax Simon – S. cochinensis (Gravely, 1915)
Fam. Phrynichidae
Phrynichus Karsch – Ph. andhraensis Bastawade, Rao, Javed, et Krishna, 2005;
Ph. phipsoni (Pocock, 1894)
Thelyphonida (Uropygi) From India are known two spp. of Labochirus Pocock,
one of Hypoctonus Thorell, one of Thelyphonus Latreille, and Uroproctus assamen-
sis (Stoliczka). According to Bastawade (2006), the Indian fauna of Uropygi con-
sists of four genera and six species (Butler 1872, Pocock 1900, Stoliczka 1869,
Gravely 1912 Harvey 2013 ).
Schizomida From India (Maharashtra) has been described the endemic (so far)
genus Neozomus with the species N. tikaderi (Cokendolpher, Sissom, et Bastawade,
1988). Also from India are known other species of “Schizomus” kharagpurensis
Gravely, Ovozomus lunatus Gravely, and others (see Reddell and Cokendolpher
1995), requiring further study. Together with the species described by Bastawade
(2002, 2004, 2006), in India are known seven spp. of Schizomida. A new genus
(Gravelyzomus Kulkarni) has been described for “Schizomus” chaibassicus
Bastawade, 2002, and a new sp. Schizomus arunachalicus Bastawade was found in
Arunachal Pradesh (Bastawade 1985, 2002, 2004, 2006 Cokendolpher et al. 1988,
Gravely 1911, 1912, 1915, 1925, Kulkarni 2012, Pocock 1900, Reddell and
Cokendolpher 1995).
Araneae According to the list of Indian spiders (Siliwal, Molur and Biswas 2005)
in the whole of India are recorded 1442 species of 361 genera and 59 families (see
also Brignoli 1972, Pocock 1900, Tikader 1970, 1987). The genera with more than
50 spp. are Araneidae (147), Gnaphosidae (134), Lycosidae (126), Oxyopidae (65),
Salticidae (181), Sparassidae (80), Tetragnathidae (51), Theraphosidae (51),
Theridiidae (52), and Thomisidae (154). Of the 1442 spp., 1002 are endemic to
mainland India. In the updated list of Siliwal and Molur (2007) are 1447 spp., 365
genera, and 60 fam. (20 genera are endemic to India). No endemic families.
Opilioacarida The only representative of Opilioacarida in Indian subcontinent is

Indiacarus pratyushi Das et Bastawade, 2007 – India (endemic genus and species).
Acariformes
Prostigmata
Fam. Erythraeidae
Microsmarialla coniferana Khot, 1963 (endemic genus and species)
7.21 Ceylon (Sri Lanka)
7.21.1 G
and Paleogeography
The Island of Ceylon, on which is situated the state Sri Lanka, has been covered
with luxuriant tropical forests, but most of them have been replaced by tea planta-
tions. The island is close to India and even is almost connected with the continent
by several islands called Adam’s Bridge. The surface of the island is 65,610 km2.
Geologists differ concerning how and when arose the 30 km-long Adam’s Bridge
(Pathirana 1980). For us there is no doubt that Ceylon had a land connection with
the continent.
Despite the quasi-total loss of forests, replaced by tea plantations, this mountainous
island still has quite interesting arachnofauna. Represented are all main orders and
suborders (except Ricinulei, Palaeoamblypygi, Dyspnoi, and Mesothelae).
Especially interesting is the genus Pettalus (from fam. Pettalidae, known from far-
away southern areas like Chile, South Africa, and New Zealand). In India live
Cyphophthalmi of different family (Stylocellidae). Remarkable is the holothyrid
endemic genus Indothyrus Lehtinen (Holothyrida are not known from the nearby
India). The Palpigradi indicate connections with Madagascar and Southeast Asia.
Palpigradi The three species found in Sri Lanka (Remy 1961b) show connections
with South Asia (Eukoenenia angusta in Thailand and India), but also with Madagascar
and the Mascarene (Eukoenenia chartoni, Koeneniodes madecassus). Remy (1961b)
was a supporter of the ideas of Jeannel for the existence of continental interconnec-
tions between Madagascar, the Seychelles, Sri Lanka, Mauritius, and India.
7.21 Ceylon (Sri Lanka) 401
Thelyphonida (Uropygi) Labochirus proboscideus Butler has been described

from Sri Lanka and is an endemic species to this island (Butler 1872, Pocock 1900).
Schizomida From Sri Lanka are known the first schizomid ever described: Schizomus
crassicaudatus Pickard-Cambridge, 1972, and also several other “Schizomus” needing
further study (buxtoni Gravely, formicoides Fernando, greeni Gravely, peradeniyensis
Gravely, perplexus Gravely, suboculatus Pocock, vittatus Gravely) (Cokendolpher
et al. 1988, Fernando 1957, Gravely 1911, 1912, 1915, Pickard-Cambridge 1972,
Pocock 1900, Reddell and Cokendolpher 1995, Harvey 2013g).
Scorpiones On the island are recorded 15 spp. of the genera Buthoscorpio Werner,
Charmus Karsch, Hottentotta Birula, Isometrus Ehr., Lychas C.L. Koch (Buthidae),
and Chaerilus Simon (Chaerilidae) (Lourenço 1997, Tikader and Bastawade 1983,
Vachon 1982). Endemic species of scorpions in Sri Lanka is Lychas srilankensis
Lourenço, 1997 – low level of endemicity.
Pseudoscorpiones On Sri Lanka have been recorded 43 spp. of Pseudoscorpiones,

belonging to 31 genera and 14 families: Chthoniidae (2 spp.), Pseudotyrannochthoniidae
(2), Atemnidae (11), Feaellidae (1), Garypidae (1), Geogarypidae (3), Hyidae (1),
Ideoroncidae (1), Sternophoridae (1), Olpiidae (5), Syarinidae (1), Cheliferidae (5),
Chernetidae (7), and Withiidae (2) (Beier 1973a, Batuwita and Benjamin 2014).
Some endemic species for the island are:
Fam. Chthoniidae – Lagynochthonius brincki Beier, Afrochthonius ceylonicus Beier,
A. reductus Beier
Fam. Syarinidae – Ideobisium ceylonicum Beier
Fam. Olpiidae – Olpium ceylonicum Beier
There are no genera, recorded only from Sri Lanka.
Opiliones On Sri Lanka have been recorded 26 spp. of Opiliones from Pettalidae,
Sclerosomatidae, Podoctidae, and Assamiidae (Boyer and Giribet 2007, Giribet
2008, Karsch 1891, Roewer 1910, 1912, 1929, Šilhavý 1974a, 1974b, Sharma et al.
2009, Sharma and Giribet 2006, Thorell 1876) (after Kury, online).
Cyphophthalmi On Sri Lanka were recorded four species of the remarkable

Gondwanan endemic genus Pettalus Thorell, a base of the family Pettalidae, spread
over Sri Lanka, New Zealand, Australia, South Africa, and Chile.
Eupnoi From Sri Lanka is known the family Sclerosomatidae (Gagrella,

Gagrellula)
Dyspnoi – not represented
Laniatores From Sri Lanka are known Laniatores from the families:
Podoctidae – Eusitalces Roewer, Eurytromma Roewer, Neopodoctis Roewer

Assamiidae – Assamia Soerensen, etc.

Trionyxellinae – India and Sri Lanka
The endemic genera are in bold:
Solifugae From Sri Lanka is known one sp. of Solifugae (Pocock 1895).
Fam. Rhagodidae
Rhagodes phipsoni (Pocock) (endemic sp.)
Araneae In Sri Lanka have been recorded 383 spp. of spiders of 213 genera and 46
families (Benjamin et al. 2012, Brignoli 1972, Huber and Benjamin 2005, Pocock
1900, Siliwal et al. 2005, Tikader 1970, 1987). These figures include 271 spp. and
22 genera endemic for Sri Lanka. Benjamin, Channa and Bambaradeniya (2006)
affirm that the known spiders on the island are 501 spp. of 46 families, including
106 spp. of Salticidae.
Spider families in Sri Lanka: Agelenidae, Araneidae, Barychelidae, Clubionidae,
Corinnidae, Ctenidae, Dictynidae, Dipluridae, Eresidae, Eutichuridae, Hahniidae,
Hersiliidae, Idiopidae, Linyphiidae, Liocranidae, Lycosidae, Mimetidae, Mysmenidae,
Nesticidae, Ochyroceratidae, Oonopidae, Oxyopidae, Palpimanidae, Philodromidae,
Pholcidae, Phrurolithidae, Pisauridae, Psechridae, Salticidae, Scytodidae, Segestriidae,
Sparassidae, Stenochilidae, Tetrablemmidae, Tetragnathidae, Theraphosidae,
Theridiidae, Theridiosomatidae, Thomisidae, Titanoecidae, Trachelidae, Ubudidae,
Uloboridae, Zodariidae, and Zoropsidae
Some endemic genera are:
Fam. Barychelidae
Plagiobothrus Karsch, 1891 (one sp.)
Fam. Ctenidae
Diallomus Simon, 1897 (two spp.)
Fam. Dictynidae
Atelolathys Simon, 1892 (one sp.)
Rhion O. P.-Cambridge, 1870 (one sp.)
Fam. Tetrablemmidae
Gunasekara Lehtinen, 1981 (one sp.)
Fam. Zoropsidae
Devendra Lehtinen, 1967 (three spp.)
Fam. Linyphiidae
Labullinyphia van Helsdingen, 1985
7.22 Andaman and Nicobar Islands 403
Fam. Hersiliidae
Promurricia Baehr et Baehr, 1993 (one sp.)
Fam. Tetragnathidae
Atelidea Simon, 1895 (one sp.)
Schenkeliella Strand, 1934 (one sp.)
Fam. Mysmenidae
Phricotelus Simon, 1895 (one sp.)
Fam. Udubidae
Campostichomma Karsch, 1892 (one sp.)
Fam. Pholcidae
Wanniyala Huber et Benjamin, 2005
Fam. Thomisidae
Peritraeus Simon, 1895 (one sp.)
Holothyrida Only Indothyrus greeni Lehtinen, 1995, is known from Sri Lanka
(endemic genus and species).
Ixodida According to Dilrukshi (2006), in Sri Lanka are registered 27 spp. of
Ixodidae of 9 genera, mostly widespread (Nosomma monstrosum Nuttall et
Warburton, 1908 is endemic to Ceylon).
7.22 Andaman and Nicobar Islands
7.22.1 G
and Paleogeography
The territory comprises two island groups in the Indian Ocean, the Andaman Islands
and the Nicobar Islands, separated by the 10° N parallel, with the Andaman to the
north of this latitude and the Nicobar to the south. The total land area of the Andaman
Islands is approximately 6408 km2 and of Nicobar Islands – 1841 km2. The islands
are situated in the Bay of Bengal, and geographically are part of Southeast Asia,
150 km north of Aceh in Indonesia and separated from Thailand and Burma by the
Andaman Sea. The Andaman Islands were linked to the mainland during the low sea
levels in glaciation time.
So far Pseudoscorpiones, Scorpiones, Amblypygi and Araneae are recorded from

these isolated and rarely visited islands. Some species are known from Madagascar
and Aldabra. Low-level endemism.
Pseudoscorpiones
From the Andaman Islands, Beier (1981) recorded three species: Xenolpium mada-
gascariense Beier, known from Madagascar and Aldabra; Anagarypus oceanusindi-
cus Chamberlin, known from Aldabra and Chagos Archipelago; and Pseudochiridium
clavigerum (Thorell), known from India and Indonesia. From the Nicobar Islands
are known Catatemnus nicobarensis With, 1906, Garypus nicobarensis Beier, 1930.
Scorpiones From the Andaman Islands (Little Andaman) is known the endemic
Chaerilus andamanensis Lourenço, Duhem, et Leguin, 2011. Before that only the
widespread Isometrus maculatus (De Geer) and Liocheles australasiae (Fabricius)
have been recorded (Tikader and Bastawade 1983).
Amblypygi From the Andaman Islands is known the family Charinidae –

Charinus Simon, Sarax Simon.
Araneae In the paper of Tikader (1977) and other papers are listed 20 families of
spiders of 41 genera, found in the Andaman and Nicobar Islands. There are also 26
newly described (? endemic) species, among the 58 in the list. The families are
Scytodidae, Filistatidae, Uloboridae, Amaurobiidae, Dictynidae, Theridiidae,
Pholcidae, Linyphiidae, Tetragnathidae, Araneidae, Hersiliidae, Oxyopidae,
Pisauridae, Lycosidae, Gnaphosidae, Heteropodidae, Thomisidae, Clubionidae,
Salticidae, and Psechridae.
7.23 S
outheast Asia (Burma, Thailand, Indochina,
Peninsular Malaysia, China South of Yangtze,
and Hainan)
7.23.1 G
and Paleogeography
The continental part of Southeast Asia is bordering India to the west and China to
the north. On it are situated the countries Burma (Myanmar), Thailand, Laos,
Cambodia, Vietnam, western Malaysia, and Singapore (connected with bridge to the
7.23 Southeast Asia (Burma, Thailand, Indochina, Peninsular Malaysia, China South… 405
continent). The development of the peninsula has been analyzed many times by Hall
(1997, 1998, 2001, 2002), Hall and Holloway (Eds) (1998), and other authors.
Indochina is (was) covered by lush rain forest and is very fast dwindling. The penin-
sula is crossed by big rivers, coming from the northern mountains. The northern part
of Burma includes some parts of Himalaya (Peak Hkakabo Razi, 5881 m). In the
other parts of the peninsula, the highest summit is Fansipan in Vietnam (3143 m).
The loss of habitats, deforestation, and replacement of the rain forest by oil palm
plantations are very serious treats to the rich and varied wildlife of the peninsula.
The paleogeography of South China and the adjacent territories is presented in
the Atlas of Wang Hongzhen, Ed. (1985).
The Hainan Island had been connected with the mainland until Miocene, when
faulting caused subsidence and brought about the formation of the Qiongzhou Strait
(Wang Hongzhen, Ed., 1985). The biogeographic analysis of Hua Zhu (2016) indi-
cates the low endemism of the flora (7 seven endemic genera out of 1283), and the
conclusion is that “Hainan Island could have been adjacent to northern Vietnam and
the Guangxi at least in Eocene.”
“Today, the waters of SE Asia contain the highest marine faunal diversity in the world, and
the islands of the region contain some of the most diverse collections of plant and animal
species found on Earth. The period 30-0 Ma is of most interest to biogeographers since
before then the separation between Asia and Australia was greater and for almost all land
plants and animals it was probably not possible to cross this barrier… There were never
continuous land links between Sundaland and Australia”. (Hall, 2001)
Other sources: Blair Hedges (2006), Dao Van Tien (1978), Golonka et al. (2006),
Hoffmann (2001), Lydekker (1896), Metcalfe (2002), Turner et al. (2001), Wallace
(1860), Weber (1899)
The Malayan part of the Indomalayan Region has a rich and varied arachnofauna.
They are represented by Palpigradi (both families); Solifugae (one genus and spe-
cies, the easternmost member of the order); scorpions of the families Buthidae,
Scorpionidae, Chaerilidae (the center of speciation), Euscorpiidae, Hormuridae,
and Pseudochactidae (the most remarkable recently described relict genera);
Pseudoscorpiones; Opiliones; Araneae; Opilioacarida (two endemic genera); and
other Acari. It is to mark the importance of the cave habitat for certain groups
(Opilioacarida, Pseudochactidae). The area is also center of such groups as
Stylocellidae (Opiliones) and Mesothelae (Araneae) (Zhang and Zhao 1978).
Palpigradi Eight species have been recorded from Thailand: Prokoenenia asiatica
Condé, 1994, (Prokoeneniidae) and six spp. of Eukoeneniidae – Eukoenenia angusta
Hansen, 1901; E. deleta Condé, 1992; E. lyrifer Condé, 1992; E. siamensis Hansen,
1901; E. thais Condé, 1988; Koeneniodes leclerci Condé, 1992; and K. spiniger
Condé, 1984. All of them are known only from this country (endemics). Condé
(1992) recorded Koeneniodes madecassus from Hong Kong (the first Palpigradi
known from China).
Solifugae The only member of Solifugae in Southeast Asia is Dinorhax rostrump-

sittaci (Simon, 1877) (Vietnam, Melanoblossiidae).
Scorpiones In the Indomalayan Region are recorded scorpions of 34 genera and

six families Pseudochactidae. Lourenço (2007) added to the only known species of
Pseudochactas Gromov from Central Asia another endemic genus and species
Troglokhammouanus steineri Lourenço, 2007 from a cave in Laos. A third endemic
genus and the species Vietbocap canhi Lourenço et Ðình Sắc Pham, 2010, V. thien-
duongensis Lourenço et Ðình Sắc Pham, 2012, and V. lao Lourenço, 2012
(Vietbocapinae, end. subfam.) have been found in caves in Vietnam and Laos. They
are all relicts (Table 7.2).
According to Di et al. (2011), Yunnan Province has the biggest scorpion biodi-
versity in the whole of China (nine spp.). From the Hainan Island have been recorded
three widespread species (Di et al. 2011) and two endemic taxa.
Pseudoscorpiones In Southeast Asia are recorded pseudoscorpions of 18 families:
Chthoniidae, Lechytiidae, Tridenchthoniidae, Sternophoridae, Atemnidae,
Ideoroncidae, Hyidae, Neobisiidae, Syarinidae, Pseudocheiridiidae, Geogarypidae,
Garypinidae, Olpiidae, Parahyidae, Cheiridiidae, Cheliferidae, Chernetidae, and
Withiidae (Beier 1951, 1967, Ćurčić 1983, Dashdamirov 1997, 2007, Ellingsen 1911,
Gao and Zhang 2013a, 2013b, Mahnert 2003, 2009, Redikorzev 1938, Schawaller
1994b, 1995, Thorell 1889, With 1906). Nguyen Thi Dinh (20) published a list of the
pseudoscorpions of Vietnam, containing 62 spp., of 32 genera and 13 families.
Parahyidae is an endemic family (Singapore, Caroline Islands).
Number of pseudoscorpion species in some of the countries in the area: Burma
(Myanmar), 13; Thailand, 43; Vietnam, 62; Laos, 9; Cambodia, 15 (Harvey, 2013f)
Endemic pseudoscorpion genera in Southeast Asia are:
Fam. Garypinidae
Caecogarypinus Dashdamirov, 2007 – Vietnam (one sp.)
Fam. Ideoroncidae
Shravana Chamberlin, 1930 – Thailand (one sp.)
Fam. Cheliferidae
Tetrachelifer Beier, 1967 – Vietnam (two spp.)
Table 7.2 Scorpions in the Indomalayan Region

Country India Nepal Bhutan Bangladesh Burma Sri Lanka Mal. Indon.
Number of species 117 11 4 5 11 15 27 29
Taxa
Fam. Bothriuridae + – – – – – – –
Cercophonius Peters 1 – – – – – – –
Fam. Buthidae + + – + + + + +
Androctonus Ehrenb. 2 – – – – – – –
Buthacus Birula 1 – – – – – – –
Buthoscorpio 4 – – – – 1 – –
Werner
Charmus Karsch 3 – – – – 2 – –
Compsobuthus 3 – – – – – – –
Vachon
Hemibuthus Pocock 1 – – – – – – –
Himalayotityobuthus 2 1 – – – – – –
Lourenço
Hottentotta Birula 6 – – – – 1 – –
Isometrus Ehr. 13 1 – 1 – 6 3 7
Lychas C.L. Koch 15 4 – – 4 1 5 6
Mesobuthus Vachon – – – – – – 1 –
Odontobuthus 1 – – – – – – –
Vachon
Orthochirus Karsch 6 – – – – – – –
Thaicharmus 2 – – – – – – –
Kovařik
Vachonus Tikader et 1 – – – – – – –
Bastawade
Fam. Chaerilidae + + – + – + + +
Chaerilus Simon 6 2 – 1 – 1 11 8
Fam. Euscorpiidae + + + + + – + –
Alloscorpiops – – – – 1 – – –
Vachon
Dasyscorpiops – – – – – – 1 –
Vachon
Euscorpiops Vachon 4 – 2 1 4 – – –
Neoscorpiops 4 – – – – – – –
Vachon
Parascorpiops – – – – – – 1 –
Banks
Scorpiops Peters 10 2 2 1 – – – –
Fam. Hormuridae + – – + + – – –
Chiromachetes 2 – – – – – + +
Pocock
Hormurus Thorell – – – – – – – 2
(continued)
Country India Nepal Bhutan Bangladesh Burma Sri Lanka Mal. Indon.
Hormiops Fage – – – – – – 1 –
Iomachus Pocock 4 – – – – – – –
Liocheles Sundevall 2 – – 1 1 – 1 3
Fam. Scorpionidae + + – – + + + +
Heterometrus Ehr. 23 1 – – 1 3 3 3
Rugodentus 1 – – – – – – –
Bastawade et al.
Country Thailand Laos Vietnam Cambodia Philippines China Hainan
Number of species 20 15 25 5 14 [50] 5
Taxa
Hottentotta Birula – – – – – 2 –
Isometrus Ehrenberg 3 1 3 1 2 3 2
Lychas C.L. Koch 3 4 1 1 3 2 1
Mesobuthus Vachon – – – – – 6 1
Orthochirus Karsch – – – – – 1 –
Thaicharmus 1 – – – – – –
Kovařik
Razianus Farzanpay – – – – – 1 –
Fam. Chaerilidae + + + + + + –
Chaerilus Simon 2 1 6 1 3 8 –
Fam. Euscorpiidae + + + – – + –
Alloscorpiops 2 2 – – – – –
Vachon
Euscorpiops Vachon 3 2 5 – – 11 –
Scorpiops Peters 1 1 2 – – 11 –
Fam. Hormuridae + – + – + + +
Hormiops Fage – – 1 – – – –
Hormurus Thorell – – – – 2 – –
Liocheles Sundevall 1 – 1 – 1 1 1
Tibetiomachus – – – – – 1 –
Lourenço et al.
Fam. Scorpionidae + + + + + + –
Heterometrus Ehr. 4 2 4 2 3 3 –
Fam. – + + – – – –
Pseudochactidae
Troglokhammouanus – 1 – – – – –
Lourenço
Vietbocap Lour. et – 1 2 – – – –
Pham
Opiliones
Cyphophthalmi In Eastern Palearctic the suborder Cyphophthalmi is known

only from Japan (Suzukielus sauteri Roewer) – endemic genus Suzukielus Juberthie.
In the Southeast Asia are represented five of the six genera of fam. Stylocellidae,
two of them endemic (Thorell 1882, 1890, Rambla 1991, 1994, Giribet 2000,
Clouse 2012, Clouse et al. 2011, Schwendinger and Giribet 2005, Sharma and
Giribet 2009, 2011, Clouse and Giribet 2010, Shear 1993, Clouse et al. 2010, Kury
et al. 2009):
Subfam. Fangensinae (endemic)
Fangensis Rambla, 1994 – Thailand (three spp.)
Giribetia Clouse, 2012 – Thailand (one sp.)
Subfam. Stylocellinae
Leptopsalis Thorell, 1882 – Malaya (three spp.)
Stylocellus Westwood, 1874 – Malaya (one sp.)
Miopsalis Thorell, 1890 – Malaya (two spp.)
The species are also endemic, mostly known from the type localities.
Map 7.11 Distribution of Stylocellidae (After Clouse 2012)

Three genera of fam. Stylocellidae are recorded in Malay Archipelago (19 spp.):
Leptopsalis Thorell, 1882 – Sumatra (three spp.), Java (two spp.), Sulawesi (four
spp.)
Stylocellus Westwood, 1874 – Sumatra (one sp.)
Miopsalis Thorell, 1890 – Borneo (nine spp.)
Two species of Stylocellus Westwood have been found in the western part of
New Guinea, crossing, thus, the Lydekker’s Line (Clouse and Giribet 2007).
In Sri Lanka live three species of the endemic genus Pettalus Thorell, 1876 (fam.
Pettalidae, known also from Madagascar, Chile, South Africa, Australia, and New
Zealand) (Sharma and Giribet 2006, Sharma et al. 2009). From India (Arunachal
Pradesh) has been described an endemic genus and species Meghalaya annandalei
Giribet, Sharma, et Bastawade, 2007 (Stylocellidae). The family Pettalidae is not
known so far from the Asiatic continent (Giribet 2000 and suppl.).
Dyspnoi Eastern Palearctic is inhabited by members of the families Sabaconidae,
Nemastomatidae, and Nipponopsalididae; present are also a few Ischyropsalididae
and Trogulidae. Two more families (Dicranolasmatidae and Taracidae) are dwellers
of Western Palearctic and the USA. (Schönhofer 2013).
Fam. Sabaconidae – the only genus Sabacon Simon is widespread (USA, Europe,
Japan, China, Siberia, Altai; several species have been described from Nepal –
the high Himalaya up to above 5000 m, which form part of the boundary between
Palearctic and Indomalayan regions). Tsurusaki and Daxiang Song (1993b) pub-
lished two new species of Sabacon from Sichuan Province (China).
Fam. Ischyropsalididae – found easternmost to Tajikistan.
Fam. Nemastomatidae – in the Eastern Palearctic, the genera Mediostoma Kratochvil
(Iran, Tajikistan) and Starengovia Snegovaya (Kyrgyzstan) reach the mountains
of Central Asia.
Martens (2016) published as new genus and sp. the first nemastomatine from
South China (Yunnan, 3800 m) – Sinostoma yunnanicum. This was quite surprising
finding of a Palearctic family far from the easternmost localities of the family
(Tajikistan, Kyrgyzstan). Also, this is the maximal altitude reached by a member of
Nemastomatinae (relict).
From Southeast Asia are recorded only two other spp. of Dyspnoi from the fam-
ily Nemastomatidae. The finding of Cladolasma [Dendrolasma] angka
(Schwendinger et Gruber, 1992) (Ortholasmatinae) in Thailand is the second spe-
cies of the genus Cladolasma Suzuki; the other species is C. parvulum Suzuki from
Japan. Another Cladolasma – C. damingshan Zhang et Zhang (Zhang and Zhang
2013) – was described from Guangxi, China (the first representative of
Nemastomatidae in China).
Fam. Nipponopsalididae – three species of genus Nipponopsalis Martens et Suzuki
from Korea and Japan (including Ryukyus and the Kuril Islands) (Schönhofer
2013)
Fam. Trogulidae – easternmost to the northern Iran (Schönhofer 2013)

Practically the Dyspnoi are missing in the Indomalayan Region.
Tsurusaki and Daxiang Song (1993a) recorded the occurrence of Crosbycus
dasycnemus in China (already known from Japan and North America). According to
Schönhofer (2013), Crosbycus should belong to the family Taracidae, and its amphi-
pacific populations “require investigation for specific distinction.” Suzuki (1972)
analyzed some other cases of discontinuous distribution of opilions.
Eupnoi In Eastern Palearctic have been found members of the families Caddidae,
Phalangiidae, and Sclerosomatidae (Gricenko 1979a, Gritsenko 1979b, 1980;
Starêga 1978; Tsurusaki et al. 2000).
Fam. Caddidae – only one species of genus Caddo Banks (otherwise North
American) is known from Japan. Even the species Caddo agilis Banks is shared
between Japan and North America (Suzuki and Tsurusaki 1983).
Fam. Phalangiidae – genera, represented in Eastern Palearctic: Lacinius Thorell
(China), Mitopus Thorell (Japan, Mongolia), Oligolophus C.L. Koch (China),
Egaenus C.L. Koch (Karakorum, Iran, Mongolia, Siberia, Tajikistan, Uzbekistan),
Homolophus Banks (Altai, Korea, Siberia, Mongolia, China, Kazakhstan), Opilio
Herbst (China, Mongolia, Iran, Japan, Kuril Islands, Karakorum, Kazakhstan),
Scleropilio Roewer (= Scutopilio Roewer) (Central Asia), Acanthomegabunus
Tsurusaki et al. (Siberia), Liropilio Gritsenko (Kazakhstan, Russia), Phalangium L.
(? China), Rilaena Šilhavý (Iran), Thapinius Roewer (Kamchatka), Pamirphalangium
Starêga (Pamir, nomen nudum). Many of these genera are known also from Europe.
Himalphalangium was described by J. Martens from Nepal Himalaya, where the
Eastern Palearctic meets the Indomalayan Region. Other important papers on the
Opilions of Himalaya are due to Martens (1977 and others).
Fam. Sclerosomatidae – genera, represented in Eastern Palearctic: Gagrellula
Roewer (China, Japan), Harmanda Roewer (Nepal), Harmandina (China),
Psathyropus L. Koch (Japan, Far East of Russia), Pseudogagrella Redikorzev
(China), Systenocentrus Simon (Japan), Leiobunum C.L. Koch (Japan), Nelima
Roewer (Japan), Pseudohomalenotus Caporiacco (Karakorum), Pygobunus Roewer
(Japan). Many of these genera are known also from Europe. Some genera
(Himaldroma, Nepalgrella, Nepalkanchia, Gyoides) were described by J. Martens
from Nepal Himalaya (Martens 1983, 1984 and others). After Roewer and Suzuki,
Tsurusaki (1991) reported opilions from Taiwan.
From Southeast Asia are recorded many species of Eupnoi from the families
Phalangiidae and Sclerosomatidae, particularly Gagrellinae (many papers of
Thorell, Roewer, Suzuki, and other authors).
Some endemic genera are Mitopiella Banks; Adungrella Roewer, Akalpia
Roewer, Altobunus Roewer, Antigrella Roewer; Aurivilliola Roewer; Bakerinulus
Roewer; Bastia Roewer; Baturitia Roewer; Biceropsis Roewer; Bonthainia
Roewer; Bullobunus Roewer; Carinobius Roewer; Carmichaelus Roewer
Ceratobunellus Roewer; Ceratobunoides Roewer; Cervibunus Roewer; Chasenella
Roewer; Chebabius Roewer; Coonoora Roewer; Dentobunus Roewer, Diangathia
Roewer, Echinobunus Roewer; Euceratobunus Roewer, 1923; Eugagrella Roewer;

Eusclera Roewer, Euzaleptus Roewer; Gagrella Stoliczka; Gagrellenna Roewer;
Gagrellina Roewer; Gagrellissa Roewer; Gagrellopsis Sato et Suzuki; Gagrellula
Roewer; Globulosoma Martens; Hamitergum Crawford; Harmanda Roewer;
Harmandina Schenkel; Hehoa Roewer; Heterogagrella Roewer; Hexazaleptus
Suzuki; Himaldroma Martens; Himalzaleptus Martens; Hologagrella Roewer;
Hypogrella Roewer; Koyamaia Suzuki; Marthana Thorell; Melanopa Thorell;
Melanopella Roewer; Melanopula Roewer; Metadentobunus Roewer; Metahehoa
Suzuki; Metasyleus Roewer; Metaverpulus Roewer; Metazaleptus Roewer,
Microzaleptus Roewer, Neogagrella Roewer, Nepalgrella Martens, Nepalkanchia
Martens, Obigrella Roewer, Octozaleptus Suzuki, Oobunus Kishida, Orissula
Roewer, Padangrella Roewer, Palniella Roewer, Paradentobunus Roewer,
Paragagrella Roewer, Paragagrellina Schenkel, Paraumbogrella Suzuki,
Pergagrella Roewer, Pokhara Suzuki, Prodentobunus Roewer, Psathyropus
L. Koch, Pseudarthromerus Karsch, Pseudogagrella Redikorzev, Pseudomelanopa
Suzuki, Pseudosystenocentrus Suzuki, Sarasinia Roewer, Sataria Roewer,
Scotomenia Thorell, Sericicorpus Martens, Sinadroma Roewer, Syleus Thorell,
Syngagrella Roewer, Systenocentrus Simon, Tetraceratobunus Roewer, Toragrella
Roewer, Umbogrella Roewer, Umbopilio Roewer, Verpulus Simon, Verrucobunus
Roewer, Xerogrella Martens, Zaleptiolus Roewer, Zaleptulus Roewer, Zaleptus
Thorell
Laniatores From the Southeast Asia and Malayan Archipelago are recorded
Laniatores from more than 50 genera and the families Assamiidae, Biantidae,
Podoctidae, Epedanidae, Sandokanidae (= Oncopodidae), Phalangodidae,
Zalmoxidae, Tithaeidae, and Petrobunidae. The families in bold are endemic for
the Indomalayan Region. Most of the others are not found in the Palearctic Region,
and other families (Phalangodidae) are considered by some researchers to be relicts
in the Palearctic. Other specialists (Martens 1972) disagree with the relict character
of European Phalangodidae.
Among the authors having worked on the Southeast Asian Laniatores, we should
not miss Tsurusaki (1995), Schwendinger (1992, 2006), Sharma et al. (2012),
Sharma and Giribet (2011), Suzuki (1969, 1977a, 1977b, 1982, 1985), and the
numerous papers by Roewer (1912, 1927, 1931, 1935, 1938, 1940, 1949), Thorell,
and other authors.
Endemic genera are:
Fam. Assamiidae
Assamiinae: Assamiella Roewer, 1912, Burma (one sp.); Neassamia Roewer,
1935, Thailand (one sp.); Pechota Roewer, 1935, Malacca (one sp.);
Popassamia Roewer, 1940, Burma (one sp.); Tavoybia Roewer, 1935, Malacca
(one sp.).
Dampetrinae: Cadomea Roewer, 1940, Malaysia (one sp.); Dongmolla Roewer,
1927, Vietnam (one sp.); Mermerus Thorell, 1876, Java, Borneo (two spp.);
Nothippulus Roewer, 1923, Vietnam (one sp.); Nothippus Thorell, 1890,
Sumatra, Malacca (three spp.); Pahangius Roewer, 1935, Malacca (one sp.);
Paradampetrus Giltay, 1930, Sumatra (one sp.); Sudaria Roewer, 1923,
Sumatra, Simalur, Sulawesi (four spp.)
Fam. Podoctidae
Dongmoa Roewer, 1927, Vietnam; Heteroibalonius Goodnight et Goodnight,
1947 (one sp.); Mesoceratula Roewer, 1949 (one sp.); Podoctellus Roewer,
1949, Malaysia (Johore) (one sp.); Podoctis Thorell, 1890, Pinang (one sp.);
Sibolgia Roewer, 1923, Malaya (one sp.); Stobitus Roewer, 1949, Malaya
(one sp.)
Fam. Petrobunidae
Petrobunus Sharma et Giribet, 2011 – Philippines (three spp.)
Fam. Tithaeidae – 38 spp
Istithaeus Roewer, 1949, Borneo; Kondosus Roewer, 1949, Borneo;
Metatithaeus Suzuki, 1969, Borneo; Sterrhosoma Thorell, 1891, Sumatra;
Tithaeomma Roewer, 1949, Burma; Tithaeus Thorell, 1890, Burma,
Thailand, Malaya, Singapore, Sumatra, Krakatau, Java, Borneo, Sarawak,
Timor
Fam. Epedanidae
Epedaninae: Alloepedanus Suzuki, 1985, Thailand (one sp.); Caletorellus
Roewer, 1938, Thailand (one sp.); Epedanidus Roewer, 1945, Malaysia
(Perak) (one sp.); Euepedanus Roewer, 1915, Thailand, Malacca (seven
spp.); Heteroepedanus Roewer, 1912 (two spp.); Paratakaoia Suzuki, 1985,
Thailand (two spp.); Plistobunus Pocock, 1903, Hong Kong, Hainan Island
(two spp.); Pseudoepedanus Suzuki, 1969 (one sp.); Pseudomarthana P. D.
Hillyard, 1985, Malaysia (one sp.); Thyreotus Thorell, 1889, Burma (two
spp.); Toccolus Roewer, 1927, Vietnam (“Tonking”), etc. (three spp.);
Zepedanulus Roewer, 1927, Malacca, Thailand, etc. (four spp.)
Acrobuninae: Heterobiantes Roewer, 1912, Hong Kong (one sp.); Paracrobunus
Suzuki, 1977 (two spp.)
Sarasinicinae: Gintingius Roewer, 1938, Pahang (Malaya) (one sp.); Panticola
Roewer, 1938 (placement is uncertain), Malacca (one sp.); Pasohnus Suzuki,
1976, was in Phalangodidae (one sp.); Sembilanus Roewer, 1938, Malacca
(one sp.); Siponnus Roewer, 1927, Pulau Pinang (one sp.); Sungsotia
Tsurusaki, 1995, Vietnam (one sp.); Tonkinatus Roewer, 1938, Vietnam
(Tonking) (one sp.)
Incertae sedis Buparellus Roewer, 1949 – Burma, Thailand (four spp.)
Fam. Zalmoxidae
Zalmoxis Soerensen is not endemic, but this is the only genus of Zalmoxidae in
the Old World, with ca. 15 spp. in South Asia, including the Philippines
(Sharma et al. 2012).
Fam. Sandokanidae (Oncopodidae) – 71 spp.

Sandokan Thorell, 1876, Southeast Asia (10 spp.); Gnomulus Thorell, 1890,
Southeast Asia, India, South China (53 spp.); Caenoncopus Martens et
Schwendinger, 1998, Sumatra (three spp.); Palaeoncopus Martens et
Schwendinger, 1998, Sumatra (three spp.); Biantoncopus Martens et
Schwendinger, 1998, Leyte, Philippines (one sp.); Martensiellus
Schwendinger, 2006, Borneo (one sp.)
The distribution of Sandokanidae appears to be governed principally by limitations of this

group’s dispersal ability. Four other laniatorid families —Assamiidae, Epedanidae,
Podoctidae, and Zalmoxidae—are distributed throughout Sundaland, but all of these have
greater range than Sandokanidae, and frequently demonstrate clear dispersal events (Giribet
and Kury 2007). The restriction of Sandokanidae to Sundaland and the Philippines is sug-
gestive of diversification in accordance with breakup of Sundaland’s components.” (Sharma
and Giribet 2009)
The opilionids of Kuril Islands have been analyzed by Tsurusaki and Crawford
(2001), the ones from the Ryukyus – by Suzuki (1971, 1973).
Amblypygi
The Amblypygi are represented in Southeast Asia by at least 22 spp. of 5 genera
(Harvey).
Fam. Charinidae
Catageus Thorell, 1889 (= Stygophrynus Kraepelin, 1895) – Burma, Thailand,
Laos, Cambodia, Vietnam, Indonesia, Malaysia (C. pusillus Thorell, 1889=
Stygophrynus cavernicola (Thorell, 1889), C. berkeleyi (Gravely, 1915),
C. brevispina (Weygoldt, 2002), C. cerberus (Simon, 1901), C. dammermani
(Roewer, 1928), C. longispina (Gravely, 1915), C. moultoni (Gravely, 1915),
C. orientalis (Seiter et Wolf, 2017), C. sunda (Rahmadi et Harvey, 2008)
Charinus Simon, 1892 – Ch. bengalensis (Gravely, 1911); Ch. pakistanus
Weygoldt, 2005
Sarax Simon, 1892 – S. brachydactylus Simon, 1892 (Malaysia, Cambodia); S.
buxtoni (Gravely, 1915) (Malaya, Singapore); S. davidovi Fage, 1946
(Vietnam, Laos, Cambodia); S. rimosus (Simon, 1901) (Malaya); S. singapo-
rae Gravely, 1911 (Singapore); S. cavernicola Rahmadi et al., 2010; S. mar-
dua Rahmadi, 2010; S. sangkulirangensis Rahmadi et al., 2010; S. yayukae
Rahmadi et al., 2010 (the last four from Kalimantan)
Weygoldtia Miranda, Giupponi, Prendini et Scharff, 2018 – one sp. W. davidovi
(Fage, 1946) – Laos, Cambodia, Vietnam
Fam. Charontidae
Charon Karsch, 1879 – Ch. grayi (Gervais, 1842) (s. lato) (Singapore, Malaysia)
Map 7.12 Distribution of Amblypygi in Asia and East Africa

■ – Phrynichus
● – Damon
□ – Sarax
X – Charinus
– Catageus
! – Phrynus
♦ – Stygophrynus
◊ – Charon
Ο – Euphrynichus
Thelyphonida (Uropygi) The early papers of Oates (1889, 1890), Gravely (1912),
and Hirst and Pocock (1902) described from Burma 13 out of the 17 Asiatic species
in Hypoctonus Thorell. Teltus vanoorti Speijer, 1936, was described as endemic
genus and species for Hainan, but Haupt (1996) synonymized the genus Teltus with
Typopeltis Pocock. From South China have been recorded three species, Hypoctonus
carmichaeli Gravely, H. granosus Pocock, and Typopeltis cantonensis Speijer, and
from Taiwan and the Ryukyu T. crucifer Pocock. Another Typopeltis, living on the
Ryukyus, is T. stimpsoni (Wood) (Yoshikura 1973; Haupt and Daxiang Song 1996a).
Other Southeast Asian Uropygi are known from Bangladesh (3 spp.), Burma (12
spp.), Thailand (8 spp.), Laos (2 spp.), the Philippines (8 spp.), Indonesia (25 spp.),
Malaysia (7 spp.), Cambodia (3 spp.), Singapore (3 spp.), and Vietnam (5 spp.)

(Haupt 2004, Kraepelin 1897, Rowland 1973, Tarnani 1894, 1900, Thorell 1888,
Harvey 2013d).
Haupt (2009a) synonymized Abaliella Strand, Minbosius Speijer, and Tetrabalius
Thorell with Thelyphonus Latreille. The newest calculation (Harvey 2013d) lists for
Southeast Asia (east of India and south of Yangtze, but including Bangladesh, the
Philippines, Taiwan, and the Ryukyus) the genera Uroproctus Pocock (1 sp.),
Hypoctonus Thorell (15 spp.), Typopeltis Pocock (12 spp.), Ginosigma Speijer (2
spp.), Glyptogluteus Rowland (1 sp.), Thelyphonoides Krehenwinkel et al. (1 sp.),
and Thelyphonus Latreille (30 spp.) or in total 62 of the 110 spp. of Thelyphonida.
Map 7.13 Distribution of order Thelyphonida (Uropygi) in Asia

● – Glyptogluteus Rowland
■ – Typopeltis Pocock
∆ – Labochirus Pocock
▼– Uroproctus Pocock
▲ – Hypoctonus Thorell
◊ – Thelyphonus Latreille
O – Thelyphonoides Krehenwinkel et al.
– Minbosius Speijer
– Ginosigma Speijer
Schizomida The list of Schizomida of South Asia is relatively long (34 spp. of 11

genera), but several of them require additional study for more accurate generic
placement (Bastawade 2004, Brignoli 1974, Gravely 1911, 1912, 1924, Hansen and
Soerensen 1905, Harvey 2013 , Kraepelin 1912, Reddell and Cokendolpher 1995,
Cokendolpher et al. 2010, Rémy 1946, Shimojana 1981, Thorell 1889). It includes:
Apozomus brignolii Reddell et Cokendolpher, 2000) – Marshall Islands (Ailuk)

A. buxtoni Gravely, 1915 – Sri Lanka
A. daitoensis (Shimojana, 1981) – Ryukyus (Minami-Daitô-jima) (endemic sp.)
A. sauteri (Kraepelin, 1912) – Taiwan, Vietnam, Ryukyu Islands
A. termitarium Cokendolpher, Sissom, et Reddell, 2010 (Malaysia, Taman Negara)
A. yamasakii (Cokendolpher, 1988) – Taiwan
A. zhensis Chen and Song, 1996 – China (Zhejiang)
Bamazomus siamensis (Hansen, 1905) – Thailand, Hong Kong, Ryukyu Islands,
Hawaii (Oahu)
B. pileti (Brignoli, 1974) – W. Malaysia (endemic sp.)
Burmezomus cavernicola (Gravely, 1912) – Burma
Clavizomus claviger (Hansen, 1905) – Singapore, W. Malaysia
Javazomus oculatus Reddell et Cokendolpher, 1995 – Java
Neozomus tikaderi Reddell et Cokendolpher, 1995 – India
Oculozomus biocellatus (Sissom, 1980) – Sumatra
Orientzomus luzonensis (Hansen, 1905) – Philippines (Luzon)
O. ralik Cokendolpher et Reddell, 2000 – Marshall Islands (Jibu)
O. sawadai (Kishida, 1930) – Bonin Islands
Ovozomus lunatus (Gravely, 1911) – India (Kolkata), Réunion, Seychelles
O. peradenyiensis (Gravely, 1911) – Sri Lanka
“Schizomus” modestus (Hansen, 1905) – W. Malaysia [also in Papua New Guinea,
incl. New Britain]
“S.” cambridgei (Thorell, 1889) – Burma
“S.” procerus (Hansen, 1905) – Singapore
“S.” peteloti (Rémy, 1946) – S. Vietnam (endemic sp.)
“S.” vittatus Gravely, 1911 – Sri Lanka
“S.” greeni Gravely, 1912 – Sri Lanka
“S.” chaibassicus Bastawade, 2002 – India
“S.” chalakudicus Bastawade, 2002 – India
“S.” formicoides Fernando, 1957 – Sri Lanka
“S.” crassicaudatus (O.P.-Cambridge, 1872) – Sri Lanka
“S.” kharagpurensis Gravely, 1912 – India (West Bengal)
“S.” perplexus Gravely, 1915 – Sri Lanka
Trithyreus grassii (Thorell, 1889) – Burma (endemic sp.)
T. sijuensis (Gravely, 1924) – India (Meghalaya)
Zomus bagnalii (Jackson, 1908) – Singapore, W. Malaysia, Sarawak [also Kew
Gardens in England]
Map 7.14 Distribution of order Schizomida in Asia

Apozomus – ■
Bamazomus – ▲
Burmezomus – ▼
Clavizomus – ●
Javazomus –
Neozomus – ♣
Oculozomus – ☺
Orientzomus – ♠
Ovozomus – ♦
Schizomus – ◊
Trithyreus – □
Zomus –
Araneae From continental Southeast Asia are known spiders from 58 families
(none of them endemic) (Deeleman-Reinhold 1995, 2000, Jäger 2005, Meng et al.
2008, Tang and Li 2010a, b, Wang et al. 2010).
From the Peninsular Malaysia, Norma-Rashid and Li (2009) list 425 species in
42 families and 238 genera of spiders.
From the tiny Singapore, Song et al. (2002) announced the existence of 310 spe-
cies of spiders of 186 genera and 39 families.
From the list of Chinese spiders of Song et al. (1999) and the papers published
afterward, considerable part consist of South Chinese species.
Fam. Liphistiidae
Liphistius Schiødte, 1849 – Southeast Asia (47 species)
Nanthela Haupt, 2003 – Hong Kong, Vietnam (two spp.)
Fam. Theraphosidae
Ornithoctonus Pocock, 1892 – Myanmar, Thailand (three spp.)
Fam. Dipluridae
Leptothele Raven et Schwendinger, 1995 – Thailand (one sp.)
Fam. Theridiosomatidae
Chthonopes Wunderlich, 2011 – Laos (two spp.)
Luangnam Wunderlich, 2011 – Laos (one sp.)
Fam. Nemesiidae
Atmetochilus Simon, 1887 – Myanmar (two spp.)
Fam. Cyrtaucheniidae
Angka Raven et Schwendinger, 1995 – Thailand
Fam. Pisauridae
Ilipula Simon, 1903 – Vietnam (one sp.)
Fam. Liocranidae
Jacaena Thorell, 1897 – Thailand, Laos, Myanmar, China (11 spp.)
Fam. Tetrablemmidae
Bacillemma Deeleman-Reinhold, 1993 – Thailand (one sp.)
Fam. Dictynidae
Penangolyna Wunderlich, 1995 – Malaysia (one sp.)
Fam. Araneidae
Perilla Thorell, 1895 – Myanmar, Vietnam, Malaysia (one sp.)
Fam. Agelenidae
Acutipetala Dankittipakul et Zhang, 2008 – Thailand (two spp.)
Fam. Linyphiidae
Thainetes Millidge, 1995 – Thailand (one sp.)
Thaiphantes Millidge, 1995 – Thailand (two spp.)
Fam. Oonopidae
Bannana Tong et Li, 2015 – Yunnan (two spp.)
Fam. Selenopidae
Siamsporops Dankittipakul et Corronca, 2009 – Thailand, Malaysia
Fam. Zodariidae
Euryeidon Dank. et Jocqué, 2004 – Thailand (six spp.)
Heliconilla Dank et al., 2012 – Thailand, Vietnam, Malaysia, Singapore,
Myanmar, Laos, China (nine spp.)
Heradion Danki et Jocqué, 2004 – Thailand, Vietnam, Malaysia, Laos, China
(ten spp.)
Malayozodarion Ono et Hashim, 2008 – Malaysia (one sp.)
Fam. Gnaphosidae
Allomicythus Ono, 2009 – Vietnam
Aracus Thorell, 1887 – Myanmar
Fam. Lycosidae
Zantheres Thorell, 1887 – Myanmar
Fam. Thomisidae
Boliscodes Simon, 1909 – Vietnam (one sp.)
Haplotmarus Simon, 1909 – Vietnam (one sp.)
Ocyllus Thorell, 1887 – Myanmar (two spp.)
Pothaeus Thorell, 1895 – Myanmar (one sp.)
Tagulinus Simon, 1903 – Vietnam (one sp.)
Fam. Salticidae
Angustaea Szombathy, 1915 – Singapore
Ceclusa Thorell, 1895 – Myanmar
The subfamily Onomastinae is endemic for the Indomalayan Region.
Acariformes
Prostigmata. Ref.: Mąkol and Gabryś (2005)
Fam. Eutrombidiidae
Caecothrombium deharvengi Mąkol et Gabryś, 2005 – Vietnam
Oribatida According to the monograph of Vu Quang Manh (2015), in Vietnam
have been recorded 320 spp. of Oribatida, zoogeographically zonated as follows:
Oriental (60.3%), Palearctic (12.2%), Cosmopolitan (10.6%), Afrotropical (6.9%),
Australian (5.0%), Neotropical (3.8%), Nearctic (0.9%), and Pacific (0.3%).
Important contributions are made by Aoki (1965, 1967).
7.24 Taiwan 421
Parasitiformes
Ixodida Petney et al. (2007) announced the presence of 104 spp. of ticks (Ixodida)
of 12 genera in Southeast Asia.
7.24 Taiwan
7.24.1 G
and Paleogeography
Separated from the Asian continent by the 160 km-wide Taiwan Strait, the main
island of the group is 394 km long and 144 kilometers wide. The southern end of the
Ryukyu Islands of Japan is directly to the east; the Batanes Islands of the Philippines
lie to its south across the Bashi Channel. The mountainous island spans the Tropic
of Cancer and is covered by tropical and subtropical vegetation. Other minor islands
and islets of the group include the Penghu Islands (Pescadores), Green Island, and
Orchid Island. The area of the main islands is 35,801 km2 (land 32,260 km2, together
with the smaller islands – 35,980 km2). Taiwan’s highest point is Yushan at 3952
meters, and there are five other peaks over 3500 meters.
This island, recently detached from the continent (Wang Hongzhen, Ed., 1985),
counts among its inhabitants many animals like mammals, snakes and others,
clearly inherited from the mainland (Zhang and Zhao 1978). The high mountains
(almost 4000 m) and the preserved vegetation in the national parks harbor many
interesting arachnids – Thelyphonida (Uropygi), Schizomida, Pseudoscorpiones,
Opiliones, and Araneae – and mites. Lacking (or not yet found) are Palpigradi,
Ricinulei, Amblypygi, Holothyrida, and Opilioacarida.
One amblypygid (Charon grayi) is known from a cave on the small Orchid Island
or Botel Tobago (Kano, 1937).
Thelyphonida (Uropygi) Whip scorpions from Taiwan are known as early as from
the paper of Schwangart (1906).
Fam. Thelyphonidae – Typopeltis crucifer Pocock, 1894
Schizomida Only two species (one of them endemic) are known so far from
Taiwan (Cokendolpher 1988, Kraepelin 1911, Yamasaki and Shimojana 1974).
Fam. Hubba Apozomus sauteri rdiidae

Apozomus sauteri (Kraepelin, 1912) – Taiwan, Vietnam, Ryukyu Islands
yamazakii (Cokendolpher, 1988) – Taiwan (endemic sp.)
Pseudoscorpiones In Taiwan are recorded ten spp. of pseudoscorpions of eight
genera and the families Chthoniidae, Neobisiidae, Geogarypidae, Olpiidae,
Cheiridiidae, Cheliferidae, Chernetidae, and Withiidae.
Opiliones The opilionid fauna of Taiwan consists of 13 genera and the families
Sclerosomatidae, Epedanidae, Phalangodidae, and Podoctidae (Roewer 1911, 1915,
1938, Suzuki 1944, 1974, 1977b – list of species, Tsurusaki 1991).
Cyphophthalmi – not found
Eupnoi
Fam. Phalangiidae – Oligolophus C.L. Koch
Fam. Sclerosomatidae – Gagrella Stoliczka, Pseudomelanopa Suzuki, Leiobunum
C.L. Koch, Pseudogagrella Redikorzev, Psathyropus L. Koch (incl.
Metagagrella), Metadentobunus Roewer, Nelima Roewer
Dyspnoi – not known
Laniatores
Fam. Phalangodidae – Tokunosia Suzuki
Fam. Epedanidae – Funkikoa Roewer, Takaoia Roewer, Kilungius Roewer, Mosfora
Roewer (endemic genera in bold)
Fam. Podoctidae – Metapodoctis Roewer, Lomanius Roewer
Araneae From Taiwan are registered (Lee 1964, 1966; Chu and Okuma 1974,
1975, Chen 1996, Song D. X. and M. S. Zhu 1997, Song D. X. et al. 1999, Ono
2007) spiders of 37 families. All of them are known also in continental China,
except for Dipluridae and Dysderidae. From the Taiwan spiders, there are five fami-
lies of Mygalomorphae (Atypidae, Ctenizidae, Hexathelidae, Dipluridae, and
Theraphosidae). The remaining 32 families are members of Araneomorphae.
Endemic genera:
Fam. Agelenidae: Femoracoelotes Wang, 2002 (two spp.)
Fam. Anapidae: Enielkenie Ono, 2007 (one sp.)
Fam. Selenopidae: Pakawops Crews et Harvey, 2011 (one sp.)
Acariformes
Oribatida – Tseng (1982, 1984)
Parasitiformes
Ixodida – The checklist of Robbins (2005) announced the existence in Taiwan of
33 spp. of Ixodida (4 Argasidae and 29 Ixodidae). Ixodes kuntzi Hoogstraal et Kohls,
1965, seems endemic to Taiwan.
7.25 The Malay Archipelago 423
7.25 The Malay Archipelago
7.25.1 G
and Paleogeography
Situated between the Indian and Pacific Oceans, the group of over 25,000 islands is
the largest archipelago by area in the world. It includes Indonesia, the Philippines,
Singapore, Brunei, East Malaysia, and East Timor. The islands of New Guinea are
not included in definitions of the Malay Archipelago.
In the present study, the volume of the notion “Malay Archipelago” includes only
Indonesia (without Papua), East Timor, and northern Borneo (Sarawak, Sabah, and
Brunei). The biggest islands are as follows:
Borneo – area 743,330 km2, highest point Kinabalu (4095 m)
Sumatra – area 473,481 km2, highest point Kerinci (3805 m)
Sulawesi – area 174,600 km2, highest point Rantemario (3478 m)
Java – area 132,187 km2, highest point Semeru (3676 m)
Bali – area 5,633 km2, highest point Agung (3142 m)
Lombok – area 4,725 km2, highest point Rinjani (3726 m)
Flores – area 13,540 km2, highest point Poco Mandasawu (2370 m)
Timor – area 30,777 km2, highest point Tatamailau (2963 m)
We have to consider the analysis of Moss and Wilson (1998) concerning the
biogeographical implications of the events on Sulawesi and Borneo in the Tertiary.
According to them Wallacea is a biogeographical area situated between Asiatic and
Australian flora and fauna with organisms of high level of endemicity. The land con-
nection between Borneo and continental Southeast Asia might have existed during
important section of Tertiary and could have allowed migrations of species. Western
Sulawesi has been connected with East Borneo by Late Cretaceous and Early
Eocene (more than 50 Ma) with option of dispersion of fauna. The ophiolites of east
Sulawesi have been accreted to Sulawesi in the Late Oligocene, which resulted in a
more extensive land of the large (174,600 km2) island.
Microcontinental fragments accreted onto eastern Sulawesi in the Miocene to Pleistocene
may have been emergent as they drifted towards Sulawesi and allowed island hopping or
rafting for biota of Australian affinity. Island hopping routes for the dispersal of organisms
between Borneo-Sulawesi and the Philippines may have existed along volcanic arcs, such
as the long-lived North Sulawesi arc. (Moss and Wilson 1998)
Here is the timing of events in the distribution of the elements of land in the area
(according to Audley-Charles, 1984):
1 . Australia/New Guinea splits from Antarctica (ca.53 Ìa).
2. Postulated formation of the Philippines by collision of an Asian continental frag-
ment with an island arc (Îligocene).
3. Possible land connection(s) across Makassar Strait (mid-Miocene).
4. Collision between New Guinea and a Tertiary island arc (ca. 15 Ìa).
5. Submarine collision between Gondwana (Sula Peninsula) and Laurasia at or

near east Sulawesi (ca. 15 Ìa).
6. Island chain established between east Sulawesi and Australia (Late Miocene to
Late Pliocene).
7. Collision between parts of Gondwanic Outer Banda Arc and Laurasian (volca-
nic) Inner Banda Arc (Late Miocene to Early Pliocene).
8. Gulf of Bone opens (about the same time).
9. Probable land connection(s) across south Makassar Strait (from Late Pliocene).
Some sources: Golonka et al. (2006), Groves (1975), Keast (1983), Katili (1971,
1975, 1978), Mayr (1944), Wallace (1869), Weber (1899), Whitmore (Ed.) (1987),
Wilson and Moss (1999)
Zoogeographically one of the most remarkable areas is the so-called Wallacea, the
transitional islands with elements of the Indomalayan and Papuan-Australian fauna.
Its “mainland” is Sulawesi (Celebes), explored by the Swiss Fritz and his cousin
Paul Sarasin in 1893–1902 (and later by many others). Roewer described the opil-
ions collected by the remarkable naturalists.
The Malay Archipelago is the richest area of fauna in the world. What concerns
the arachnids, in the archipelago are recorded all the orders of Arachnida, except for
Ricinulei, Holothyrida, and Opilioacarida. Remarkable is the absence of Holothyrida,
well represented in the nearby New Guinea.
Palpigradi Koeneniodes berndti Condé, 1988, was described from Malaysia
(Borneo). From Indonesia are known seven species from Java, Sulawesi, and
Sumatra (Condé 1989, 1990, 1992, 1994). Some of them are endemic (Eukoenenia
maros, E. lienhardi (also in Brunei and Singapore), E. paulinae, Prokoenenia cele-
bica, P. javanica), and others are known from Madagascar (Koeneniodes madecas-
sus, K. frondiger). They belong to the families Eukoeneniidae (Eukoenenia Börner)
and Prokoeneniidae (Prokoenenia Börner).
Scorpiones In Indonesia are registered 29 spp. of 6 genera of 4 families of scorpi-

ons (see Table 7.2) – Buthidae, Chaerilidae, Hormuridae, and Scorpionidae.
An eyeless forest litter scorpion was described from Halmahera (Chaerilus tel-
novi Lourenço, 2009), followed by another species of the same genus Chaerilus
spinatus Lourenço et Duhem, 2010, from the cave Batu Lubang on the same island.
Both are endemic to Halmahera.
Solifugae
Fam. Melanoblossiidae, subfam. Dinorhacinae

The genus Dinorhax Simon, 1879, is the only solpugid forming the eastern limit
of the order. The species Dinorhax rostrumpsittaci (Simon, 1877) is known from
Vietnam and Indonesia (Maluku).
Pseudoscorpiones In Indonesia (incl. W. New Guinea) have been registered
88 spp. of pseudoscorpions and in Malaysia (incl. Peninsular), 38 spp. Altogether,
in Malay Archipelago are recorded 45 genera of 14 families: Chthoniidae,
Chernetidae, Cheliferidae, Atemnidae, Garypidae, Garypinidae, Geogarypidae,
Hyidae, Olpiidae, Parahyidae, Pseudochiridiidae, Syarinidae, Tridenchthoniidae,
and Withiidae (Beier 1932, 1935, 1952, 1953, 1954, Muchmore 1972, Simon 1899,
Thorell 1883, Tullgren 1907, With 1906).
Endemic genera for Malay Archipelago are:
Fam. Cheliferidae: Aporochelifer Beier, 1953 – Flores (one sp.)
Fam. Chernetidae: Chiridiochernes Muchmore, 1972 – Sulawesi (one sp.)
Opiliones Clouse (2012), Loman (1892), Martens and Schwendinger (1998),
Roewer (1913, 1914a, 1927, 1931, 1938, 1949), Schwendinger (2006), Shear
(1993a), Thorell (1882, 1890, 1891)
Cyphophthalmi Three genera of fam. Stylocellidae are recorded in Malay

Archipelago (19 spp.)
Leptopsalis Thorell, 1882 – Sumatra (three spp.), Java (two spp.), Sulawesi (four
spp.)
Stylocellus Westwood, 1874 – Sumatra (one sp.)
Miopsalis Thorell, 1890 – Borneo (nine spp.)
Eupnoi (Roewer, 1913, 1931)
Fam. Phalangiidae – Mitopiella Banks, 1930
Fam. Sclerosomatidae – Altobunus Roewer, 1910; Dentobunus Roewer, 1910;
Echinobunus Roewer, 1912; Eugagrella Roewer, 1910; Euzaleptus Roewer, 1911;
Gagrella Stoliczka, 1869; Gagrellina Roewer, 1913; Gagrellissa Roewer, 1931;
Gagrellula Roewer, 1910; Marthana Thorell, 1891; Melanopella Roewer, 1931;
Metazaleptus Roewer, 1912; Neogagrella Roewer, 1913; Paragagrella Roewer,
1912; Zaleptus Thorell, 1876
Endemic genera are:
Mitopiella Banks, 1930, Borneo (one sp.); Gagrellina Roewer, 1913, Sulawesi (one
sp.)
Gagrellissa Roewer, 1931 – Sunda Islands (one sp.)
Dyspnoi – not known
Laniatores They are represented in Malay Archipelago (s.str.) by the families

Biantidae, Podoctidae, Epedanidae, and Sandokanidae (= Oncopodidae preocc.).
Endemic genera are:
Fam. Epedanidae
Epedaninae
Epedanulus Roewer, 1913 – Sulawesi (one sp.)
Epedanus Thorell, 1876 – Borneo, Java, Sarawak, Pinang, Sumbawa (eight
spp.)
Heteroepedanus Roewer, 1912 – (two spp.)
Lobonychium Roewer, 1938 – Borneo (one sp.)
Metepedanulus Roewer, 1913 – Sulawesi, Borneo (two spp.)
Metepedanus Roewer, 1912 – Borneo (two spp.)
Nanepedanus Roewer, 1938 – Borneo (one sp.)
Parepedanulus Roewer, 1913 – Sulawesi (two spp.)
Pseudoepedanus Suzuki, 1969 – (one sp.)
Acrobuninae
Acrobunus Thorell, 1891 – Sumatra, Borneo (three spp.)
Anacrobunus Roewer, 1927 – Riau Islands (one sp.)
Harpagonellus Roewer, 1927 – Sumatra (one sp.)
Paracrobunus Suzuki, 1977 – Palawan, Philippines (two spp.)
Sarasinicinae
Acanthepedanus Roewer, 1912 – Sumatra (one sp.)
Albertops Roewer, 1938 – Borneo (one sp.)
Delicola Roewer, 1938 – Sumatra (one sp.)
Koyanus Roewer, 1938 – Borneo (one sp.)
Kuchingius Roewer, 1927 – Borneo (one sp.)
Padangcola Roewer, 1963 – should possibly be in Epedaninae [Sumatra] (one
sp.)
Punanus Roewer, 1938 – Borneo (one sp.)
Sarasinica Strand, 1914 – Borneo, Sulawesi (four spp.)
Sinistus Roewer, 1938 – Borneo (two spp.)
Siponnus Roewer, 1927 – Pulau Pinang (one sp.)
Incertae sedis
Beloniscellus Roewer, 1912 – Indonesia (six spp.)
Beloniscops Roewer, 1949 – Sumatra (two spp.)
Belonisculus Roewer, 1923 – Simalur Island (Sumatra) (one sp.)
Beloniscus Thorell, 1891 – Singapore, Sumatra, Simalur Island, Java, “Sunda
Islands” (14 spp.)
Parabupares Suzuki, 1982 – Indonesia (one sp.)
Sotekia Suzuki, 1982 – Indonesia (one sp.)
Fam. Sandokanidae [= Oncopodidae, praeoccup.]

Caenoncopus Martens et Schwendinger, 1998 – Sumatra (three spp.)
Martensiellus Schwendinger, 2006 – Borneo (one sp.)
Palaeoncopus Martens et Schwendinger, 1998 – Sumatra (three spp.)
Fam. Podoctidae (incl. Erecananinae and Ibaloniinae)
Baramella Roewer, 1949 – Borneo (one sp.)
Baramia Hirst, 1912 – Sarawak, Indonesia (four spp.)
Baso Roewer, 1923 – Sumatra (one sp.)
Basoides Roewer, 1949 – Sumatra (one sp.)
Dino Loman, in Weber 1892 – Sumatra (one sp.)
Gargenna Roewer, 1949 – Indonesia (one sp.)
Heteroibalonius Goodnight et Goodnight, 1947 – Papua (one sp.)
Mesoceratula Roewer, 1949 – (one sp.)
Orobunus Goodnight et Goodnight, 1947 – Papua (one sp.)
Pentacros Roewer, 1949 – Indonesia (one sp.)
Gaditusa Roewer, 1949 – Borneo (one sp.)
Idjena Roewer, 1927 – Java (one sp.)
Japetus Roewer, 1949 – Borneo (one sp.)
Lejokus Roewer, 1949 – Borneo (one sp.)
Lundulla Roewer, 1927 – Borneo (one sp.)
Podoctis Thorell, 1890 – Pinang (one sp.)
Podoctomma Roewer, 1949 – Java (one sp.)
Podoctops Roewer, 1949 – Sumatra (one sp.)
Trencona Roewer, 1949 – Borneo (one sp.)
Trigonobunus Loman, 1894 – Borneo (one sp.)
Amblypygi From Malay Archipelago have been recorded until 2016 11 (may be
more; see Harvey and West 1998) species of Amblypygi from 4 genera and 3 fami-
lies (Gravely 1915, Harvey 2002, Kraepelin 1895, Rahmadi and Harvey 2008,
Roewer 1928, Thorell 1888). Many of them were found in caves. No endemic gen-
era, some endemic species. Quite puzzling was, however, to find in the cave Batu
Cermin on the Flores Island the new species Phrynus exsul Harvey, 2002 – the first
representative of the American family Phrynidae in the Old World. Rahmadi et al.
(2010) added four new species of Sarax from Borneo.
Fam. Charinidae
Sarax Simon – S. cavernicola Rahmadi, Harvey, et Kojima, 2010; S. sangkuli-
rangensis Rahmadi, Harvey, et Kojima, 2010; S. mardua Rahmadi, Harvey, et
Kojima, 2010; S. yayukae Rahmadi, Harvey, et Kojima, 2010; S. sarawaken-
sis (Thorell, 1888) (all Borneo or Kalimantan); S. javensis (Gravely, 1915)
(Java)
Fam. Charontidae
Charon Karsch – Ch. grayi (Gervais, 1842) (syn.? C. beccarii Thorell, 1888, and
C. subterraneus Thorell, 1888), Indonesia (Ambon, etc.)
Stygophrynus Kraepelin – S. (S.) dammermani Roewer, 1928 (Java); S. (S.)
sunda Rahmadi et Harvey, 2008 (Java); S. (Neocharon) moultoni Gravely,
1915 (Sumatra, Kalimantan)
Fam. Phrynidae: Phrynus Lamarck – Ph. exsul Harvey, 2002 (Flores)
Thelyphonida (Uropygi) From Indonesia and Malaysian Borneo have been
described or recorded 15 spp. of Thelyphonus Latreille, Hypoctonus gastrostictus
Kraepelin, Tetrabalius nasutus Thorell, T. seticauda (Doleschall), Minbosius
manilanus (Koch), Ginosigma lombokensis Speijer, and Chajnus renschi Speijer
(endemic genus for the Sunda Islands). In conclusion, from the species from the
Philippines and the three Labochirus and one Uroproctus from India and Sri Lanka,
ca. 60 spp. of Uropygi are represented in the “Indomalayan Region” (without
Melanesia – New Guinea, the Solomon Islands, and Vanuatu). Haupt (2009a) syn-
onymized the genera Minbosius, Tetrabalius, and Abaliella with Thelyphonus, so
now we know from Malayan Archipelago 21 spp. of Thelyphonida from the genera
Thelyphonus Latreille (18 spp.), Hypoctonus Thorell (1), Ginosigma Speijer (1),
and Chajnus Speijer (1). All of them (except Thelyphonus manilanus) are endemic
for the various islands of the archipelago (Kraepelin 1897, Speijer 1931, 1936,
Tarnani 1901, Thorell 1888).
Schizomida Schizomida from the Malay Archipelago (s.s.):
Clavizomus sp. – Java

Javazomus oculatus (Cokendolpher et Sites, 1988) – Java (end. genus)
Oculozomus biocellatus (Sissom, 1980) – Sumatra (end. genus)
Zomus bagnalii (Jackson, 1908) – Sarawak [also Singapore, W. Malaysia, Kew
Gardens in England]
According to Reddell and Cokendolpher (1995), unidentified Schizomida are
known also from Ambon, Kalimantan, Sulawesi, and Krakatau Islands.
Araneae From Malay Archipelago is registered rich fauna of spiders (Deeleman-
Reinhold 1995, 2001, Prószyn’ski and Deeleman-Reinhold 2010, and others).
According to the checklist of Stenchly (2011), from Indonesia and New Guinea
have been registered 58 families of spiders with 505 genera and 1954 spp., 499 spp.
being found only in New Guinea. Six of the 58 families (Dipluridae, Lamponidae,
Micropholcommatidae, Nicodamidae, Stiphidiidae, Titanoecidae) are found only in
New Guinea.
Some endemic genera of spiders from the Malay Archipelago (s.str.)

Fam. Theraphosidae
Phormingochilus Pocock, 1895 – Borneo, Sumatra (five spp.)
Fam. Liocranidae
Sudharmia Deeleman-Reinhold, 2001 – Sumatra (three spp.)
Fam. Miturgidae
Tamin Deeleman-Reinhold, 2001 – Borneo, Sulawesi (two spp.)
Fam. Oonopidae
Myrmopopaea Reimoser, 1933 – Sumatra (one sp.)
Plectoptilus Simon, 1905 – Java (one sp.)
Xyphinus Simon, 1893 – Borneo, Singapore, Malaysia
Fam. Oxyopidae
Tapponia Simon, 1885 – Malaysia, Sumatra, Borneo (one sp.)
Fam. Phrurolithidae
Plynnon Deeleman-Reinhold, 2001 – Sumatra, Borneo (three spp.)
Fam. Phyxelididae
Vytfutia Deeleman-Reinhold, 1986 – Sumatra, Borneo (two spp.)
Fam. Tetrablemmidae
Borneomma Deeleman-Reinhold, 1980 – Borneo (two spp.)
Lamania Lehtinen, 1981 – Borneo, Bali, Malaysia, Sulawesi (seven spp.)
Maijana Lehtinen, 1981 – Java (one sp.)
Sabahya Deeleman-Reinhold, 1980 – Borneo (two spp.)
Singalangia Lehtinen, 1981 – Sumatra (one)
Fam. Tetragnathidae
Mitoscelis Thorell, 1890 – Java (one sp.)
Neoprolochus Reimoser, 1927 – Sumatra (one sp.)
Fam. Anapidae
Borneanapis Snazell, 2009 – Borneo (one sp.)
Fam. Cycloctenidae
Galliena Simon, 1898 – Java (one sp.)
Fam. Clubionidae
Scopalio Deeleman-Reinhold, 2001 – Borneo (one)
Fam. Eutichuridae
Calamoneta Deeleman-Reinhold, 2001 – Java, Sumatra (two spp.)
Summacanthium Deeleman-Reinhold, 2001 – Sulawesi (two spp.)
Fam. Symphytognathidae
Anapogonia Simon, 1905 – Java (one sp.)
Fam. Linyphiidae
Caenonetria Millidge et Russell-Smith, 1992 – Borneo
Dumoga Millidge et Russell-Smith, 1992 – Sulawesi (two spp.)
Eordea Simon, 1899 – Sumatra (one sp.)
Erigophantes Wunderlich, 1995 – Borneo (one sp.)
Phyllarachne Millidge et Russell-Smith, 1992 – Borneo
Piesocalus Simon, 1894 – Java (one sp.)
Fam. Araneidae
Actinacantha Simon, 1864 – Sumatra, Java
Friula O.P.-Cambridge, 1896 – Borneo (one sp.)
Plicatiductus Millidge et Russell-Smith, 1992 – Sulawesi
Racata Millidge, 1995 – Krakatau (one sp.)
Mitrager van Helsdingen, 1985 – Java
Fam. Theridiidae
Borneoridion Deeleman et Wunderlich, 2011 – Borneo (one sp.)
Deelemanella Yoshida, 2003 – Borneo (one sp.)
Helvidia Thorell, 1890 – Sumatra (one sp.)
Fam. Thomisidae
Musaeus Thorell, 1890 – Sumatra (one sp.)
Narcaeus Thorell, 1890 – Java (one sp.)
Nyctimus Thorell, 1877 – Sumatra, Sulawesi (one sp.)
Pseudamyciaea Simon, 1905 – Java (one sp.)
Reinickella Dahl, 1907 – Java
Scopticus Simon, 1895 – Java (one sp.)
Fam. Salticidae
Stichius Thorell, 1890 – Sumatra (one sp.)
Thianella Strand, 1907 – Java (one sp.)
Tisaniba Zhang et Maddison, 2014 (six spp.)
Taivala Peckham et Peckham, 1907 – Borneo (one sp.)
7.26 The Philippines 431
Map 7.15 Outline of the Malayan Archipelago
7.26 The Philippines
7.26.1 G
and Paleogeography
The Philippines is an archipelago of 7107 islands with a total land area, including
inland bodies of water, of approximately 300,000 km2. It is located between 116°
40′ and 126° 34′ E and 4° 40′ and 21° 10′ N and is bordered by the Philippine Sea
to the east, the South China Sea to the west, and the Celebes Sea to the south. The
island of Borneo is located a few hundred kilometers southwest, and Taiwan is
located directly to the north. Most of the mountainous islands are covered in tropical
rain forest and volcanic in origin. The highest mountain is Mount Apo (in Mindanao,
2954 m). Forest cover declined from 70% of the country’s total land area in 1900 to
about 18.3% in 1999. Important paper on the zoogeography of the Philippines was
written by Dickerson (1928), who introduced the word Wallacea.
The archipelago is rich in Arachnida (the orders Palpigradi, Scorpiones,

Pseudoscorpiones, Opiliones, Amblypygi, Thelyphonida, Schizomida, and Araneae
and many mites). Missing are the orders Ricinulei, Solifugae, Opilioacarida, and
Holothyrida and the suborders Palaeoamblypygi and Dyspnoi. Remarkable is the
rich fauna of Uropygi.
Palpigradi There is one species recorded: Koeneniodes deharvengi Condé, 1981.
Scorpiones The scorpion fauna of the Philippines consists of 14 species of the

genera Isometrus Ehr. (Buthidae, 2 spp.), Lychas C.L. Koch (Buthidae, 3 spp.),
Chaerilus Simon (Chaerilidae, 3 spp.), Liocheles Sundevall (Hormuridae, 1 sp.),
Hormurus Thorell (Hormuridae, 2 spp.), and Heterometrus Ehr. (Scorpionidae, 3
spp.). There are no endemic genera (Kovaøik 2012).
Pseudoscorpiones From the Philippines are recorded 41 spp. of pseudoscorpions

of 29 genera and 13 families (Beier 1931, 1937, 1966, 1967), Chamberlin (1930),
Harvey (2013f).
Fam. Chthoniidae – Lagynochthonius Beier (two spp.), Tyrannochthonius

Chamberlin (one sp.)
Fam. Tridenchthoniidae – Compsaditha Chamberlin (one sp.), Ditha Chamberlin
(one sp.), Dithella J.C. Chamberlin et R.V. Chamberlin (one sp.)
Fam. Neobisiidae – Bisetocreagris Curcic (one sp.), Microcreagris Balzan (two
spp.)
Fam. Syarinidae – Alocobisium Beier (one sp.)
Fam. Geogarypidae – Geogarypus Chamberlin (one sp.)
Fam. Hyidae – Hya Chamberlin (one sp.)
Fam. Pseudochiridiidae – Pseudochiridium With (one sp.)
Fam. Olpiidae – Olpium L. Koch (one sp.)
Fam. Atemnidae – Anatemnus Beier (two spp.), Atemnus Canestrini (one sp.),
Oratemnus Beier (four spp.), Metatemnus Beier (one sp.), Paratemnoides Harvey
(three spp.), Stenatemnus Beier (one sp.)
Fam. Cheiridiidae – Apocheiridium Chamberlin (one sp.), Cryptocheiridium
Chamberlin (one sp.)
Fam. Cheliferidae – Lissochelifer Chamberlin (one sp.), Lophochernes Simon (two
spp.), Metachelifer Redikorzev (one sp.)
Fam. Chernetidae – Adelphochernes Beier (two spp.), Megachernes Beier (three
spp.), Ceriochernes Beier (one sp.)
Fam. Withiidae – Metawithius Chamberlin (one sp.), Withius Kew (one sp.)
Endemic genera:
Fam. Chernetidae: Adelphochernes Beier, 1937 – Mindanao, Mindoro (two spp.)
Opiliones From the Opiliones of the Philippines are recorded 98 spp. of 3 subor-
ders (Martens and Schwendinger 1998, Roewer 1926, 1954, 1955, Shear 1993a,
Suzuki 1977a, 1982).
Cyphophthalmi
Fam. Stylocellidae – Miopsalis Thorell is recorded from Palawan, an island with
special position within the Philippine Archipelago, also from Mindanao.
Eupnoi Following Suzuki (1977a), we can see that on the archipelago is known
only one family of “Palpatores,” with 20 genera and 63 species.
7.26 The Philippines 433
Gagrellinae – Altobunus Roewer (1), Bakerinulus Roewer (1), Bastia Roewer
(1), Bonthainia Roewer (1), Bullobunus Roewer (8), Ceratobunellus Roewer
(1), Dentobunus Roewer (2), Eugagrella Roewer (2), Gagrella Stoliczka (21),
Gagrellula Roewer (3), Harmanda Roewer (1), Hologagrella Roewer (1),
Marthana Thorell (5), Metagagrella Roewer (2), Melanopula Roewer (1),
Paragagrella Roewer (1), Pentazaleptus (2), Prodentobunus Roewer (1),
Zaleptanus Roewer (2), Zaleptus Thorell (6)
Dyspnoi – missing
Laniatores From the (very conservative for the many monotypical genera) esti-
mate of Suzuki (1977a and suppl.), the following 34 spp. of Laniatores have been
recorded on the Philippine archipelago:
Fam. Sandokanidae (Oncopodidae)

Biantoncopus Martens et Schwendinger, 1998 – two spp. (Leyte, endemic
genus)
Gnomulus Thorell, 1876 (Pelitnus Thorell, 1891) – six spp. (Luzon, Mindanao,
Leyte)
Fam. Epedanidae
Paracrobunus Suzuki, 1977 – two spp. (Palawan)
Balabanus Suzuki, 1977 – one sp. (Balabac)
Dumaguetes Roewer, 1927 – one sp.
Fam. Podoctidae
Oppodoctis Roewer, 1927– one sp. (Luzon, endemic genus)
Hoplodino Roewer, 1915– one sp. (Dak)
Ibalonius Karsch, 1880 – five spp. (Luzon, Bohol, Mindanao)
Bonea Roewer, 1914 – two spp. (Luzon, Palawan)
Fam. Biantidae
Dibunus Loman, 1906 – ten spp. (Luzon, Bohol, Negros, Mindanao)
Fam. Assamiidae
Simalurius Roewer, 1923 – one sp. (Palawan)
Amblypygi As a result of his visit to the Philippines in 1890, Simon (1892)
described the first Amblypygi from the islands, including the new genus Sarax
(Charinidae). As a whole, on the archipelago are represented three spp. of two gen-
era and two families (Giupponi and Miranda 2012, Harvey 2003, 2013).
Fam. Charinidae: Sarax Simon, 1892 – S. brachydactylus Simon, 1892; S. curioi

Giupponi et Miranda, 2012 (endemic)
Fam. Charontidae: Charon Karsch, 1879 – Ch. grayi (Gervais, 1842)

Thelyphonida (Uropygi) From the Philippines we used to know five genera of
Uropygi: Glyptogluteus Rowland (Panay, one sp.), Minbosius Speijer (one sp.),
Mimoscorpius Pocock (one sp.), Abaliella Strand (one sp.), and Thelyphonus
Latreille (three spp. on Luzon and Mindanao). As Abaliella and Minbosius were
synonymized under Thelyphonus by Haupt (2009a) and a new genus and species
were described from Panay (Thelyphonoides panayensis Krehenwinkel Curio,
Takud et Haupt, 2009), the situation changed, and now in the Philippines are known
four genera with eight spp., including seven species endemic to the islands.
“Minbosius” manilanus (C.L. Koch, 1843) is found also in New Guinea and the
Moluccas. The fauna is mostly of Southeast Asiatic (Indomalayan) extraction, but
with high endemism. The former genera Abaliella and Minbosius have mostly
Papuan affinities (Kraepelin 1897, Krehenwinkel et al. 2009, Rowland 1973b,
Speijer 1936).
Schizomida The record for the Philippines consists only of Orientzomus luzonicus
(Hansen, 1905), endemic sp. for Luzon, and “Trithyreus sp.” Undetermined material
is known from Mindanao and Negros (Hansen and Sørensen 1905, Reddell and
Cokendolpher 1995). The genus Orientzomus Cokendolpher et Tsurusaki is found
also on the islands Mariana and Bonin.
Araneae In the Philippines are represented ca. 40 families of spiders (Simon 1892,
Huber 2011, Barrion and Litsinger 1995, Elias and Nuneza 2016).
Endemic genera of spiders:
Fam. Linyphiidae
Apobrata Miller, 2004
Fam. Selenopidae
Amamanganops Crews et Harvey, 2011 (one sp.)
Fam. Uloboridae
Astavakra Lehtinen, 1967
Fam. Theridiidae
Achaearyopa Barrion et Litsinger, 1995
Landoppo Barrion et Litsinger, 1995
Fam. Thomisidae
Pycnaxis Simon, 1895 (one sp.), Taypaliito Barrion et Litsinger, 1995 (one sp.)
Acariformes The mite fauna of the Philippines is well known, thanks to
L. Corpuz-Raros (2005 and many other papers). There are many endemic genera
of various families.
7.27 Japan (the Ryukyus Excluded) 435
7.27 Japan (the Ryukyus Excluded)
7.27.1 G
and Paleogeography
Japan is a stratovolcanic archipelago extending along the Pacific coast of Asia.

Measured from the geographic coordinate system, Japan is 36° north and 138° east.
The major islands are (from north to south) Hokkaido, Honshu (the “mainland”),
Shikoku, and Kyushu. There are 2456 islands, including the Bonin Islands, Daitô
Islands, Minami-Tori-shima, Okinotorishima, Ryukyu Islands, and Volcano Islands.
The climate varies from tropical in south to cool temperate in Hokkaido. The Japanese
islands are the summits of mountain ridges uplifted near the outer edge of the conti-
nental shelf. About 73 percent of Japan’s area is mountainous. The highest point is
Mount Fuji, 3776 m. Three mountain chains – the Hida, Kiso, and Akaishi Mountains –
form the Japanese Alps (Nihon Arupusu), several of those peaks are higher than 3000
meters. The highest point in the Japanese Alps is Mount Kita at 3193 meters.
The Bonin Islands, known in Japan as the Ogasawara Guntô, are an archipelago
of over 30 subtropical and tropical islands, some 1,000 kilometers south of Tokyo.
The total area of the islands is 73 km2.
In Japan, as in most other places, the zoogeography was based mostly on mam-
mals. The conclusions of Dobson (1994) are that:
Distribution patterns among the terrestrial mammal species of Sakhalin and the main
islands of Japan are shown to fall into 12 clear groups. The most fundamental distributional
break (Blakiston’s Line) is that separating Hokkaido and Sakhalin to the north, with their
boreal fauna typical of northern Eurasia, from ‘Hondo’ (Honshu, Shikoku, Kyushu) to the
south, which demonstrates a high degree of endemism and supports a small number of
Indo-Malayan elements. Distribution patterns may be explained by considering the
Quaternary geohistory of the area, particularly the formation of land bridges and the
changes in climatic conditions during this period. Hondo underwent two main periods of
land bridge connection to the Asiatic mainland. The first, prior to the Pleistocene, allowed
immigration of forms which have since developed into distinctive elements of the endemic
fauna (‘Old Hondo Endemics’). The second, during the Middle Pleistocene, brought in
widespread Palaearctic species as well as components from South-East Asia (‘Early
Colonists’), some of which have since undergone vicariant speciation (‘New Hondo
Endemics’); it also allowed several of the Old Hondo Endemics to extend their range to the
mainland and Hokkaido (‘Expanding Hondo Endemics’). Sakhalin and Hokkaido have
been more intimately connected to the mainland (most recently until less than 10,000 years
ago), such that endemism is very restricted. Species groups here are the ‘Late Colonists’,
cold-adapted tundra species which expanded with the glacial advances, but which are now
restricted in distribution, and ‘Recent Colonists’, postglacial forest species which recolo-
nized before the severance of land bridges. Moving the other way were ‘Expanding
Northern Endemics’, which arose in Hokkaido or Sakhalin during the last glacial and colo-
nized the adjacent mainland before severance of land links.
The northern Ryukyu Islands are separated from Kyushu by the so-called Myake
Line.
From Japan (Ryukyu excluded) are missing the orders Palpigradi, Solifugae,
Ricinulei, Opilioacarida, and Holothyrida. As the country is a chain of islands from
the Indomalayan Ryukyus to the harsh climate of Hokkaido, the fauna varies from
one area to another (Haupt 2003).
The study of Haupt (1993) was “based on four groups of ground-living arachnids:
whip scorpions and spiders (Hexathelidae, Ctenizidae, Mesothelae). A close zoogeo-
graphical connection was found between the Yaeyama Islands and nearby Taiwan. The
central and northern parts of the Ryukyu Islands are faunistically closest to Kyushu.
No distinct border between an Oriental and a Palearctic fauna can be detected.”
Scorpiones In Japan are recorded only two widespread species of scorpions:
Isometrus maculatus (Buthidae) and Liocheles australasiae (Hormuridae).
Pseudoscorpiones In Japan (including Ryukyus) are recorded 68 pseudoscorpi-

ons, belonging to 13 families (none of them are endemic) (Beier 1952,
J.C. Chamberlin 1929, 1938, Čurčić 1979, Ellingsen 1907, Kishida 1966, Morikawa
1955, 1957, 1960; 17 papers of Japanese pseudoscorpions from 1952 to 1972),
Sakayori, 1999, 2000, 2002, Sato (1978, 1979, 1982, 30 papers of Japanese pseudo-
scorpions from 1976 to 1988).
Endemic genera for Japan:
Fam. Neobisiidae: Pararoncus J.C. Chamberlin, 1938 – seven spp.
Fam. Cheliferidae: Kashimachelifer Morikawa, 1957 – one sp. (Honshu)
Fam. Olpiidae: Nipponogarypus Morikawa, 1955 – one sp. (Enoshima and the
Ryukyus)
Opiliones Several authors (Martens and Suzuki 1966, Miyosi 1957, Sato and
Suzuki 1939, Shear 2010a, Suzuki, papers from 1939 and 1991, Suzuki and
Tsurusaki 1983) have contributed to our understanding that in Japan (Ryukyus and
Bonin excluded) are recorded Opiliones of the families Sironidae, Caddidae,
Phalangiidae, Nemastomatidae, Sabaconidae, Nipponopsalididae, Travuniidae,
Triaenonychidae, Phalangodidae, and Podoctidae.
Cyphophthalmi From Japan is known only one (endemic) genus and species.
Fam. Sironidae: Suzukielus Juberthie, 1970
Eupnoi
Fam. Phalangiidae – Opilio Herbst, 1798
Dyspnoi Fam. Caddidae (Caddo Banks, known also from eastern North America)
7.27 Japan (the Ryukyus Excluded) 437
Fam. Nemastomatidae (Cladolasma Suzuki, known also from Thailand and China)
Fam. Sabaconidae – Sabacon Simon (nine spp.)
Fam. Nipponopsalididae (one genus Nipponopsalis Martens et Suzuki, known from
Japan, the Kuril Islands, and Korea)
Laniatores In mainland Japan Laniatores are represented by eight genera and
three families (Travuniidae, Phalangodidae, Podoctidae).
Endemics for the mainland Japan are the genera:
Fam. Travuniidae
Yuria Suzuki, 1964 – one sp. (inc. sedis)
Nippononychinae (endemic subfamily)
Nippononychus Suzuki, 1975 – one sp.
Metanippononychus Suzuki, 1975 – four spp.
Izunonychus Suzuki, 1975 – one sp.
Paranonychinae (USA, Canada, Japan)
Kainonychus Suzuki, 1975 – one sp.
Fam. Phalangodidae
Proscotolemon Roewer, 1916 – one sp.
Fam. Podoctidae
Iyonus Suzuki, 1964 – one sp.
Idzubius Roewer, 1949 – one sp.
Thelyphonida (Uropygi) Typopeltis stimpsonii (Wood, 1862) from the islands
Tokunoshima to Amakusa and the south of Kyushu, the only Thelyphonida from the
mainland Japan (Yoshikura 1973)
Araneae The araneofauna of Japan (total), according to Ono (2009), counts more
than 1500 spp.
In Japan (Ryukyus excluded) are known at least 50 families of spiders (Yaginuma
1961, Chikuni 1989, Kamura and Hayashi 2009, Kamura and Irie 2009, Komatsu
1961, Nishikawa 2009, Ono (ed.) 2009, Saito and Ono 2001, Tanikawa 2009,
Tanikawa and Ono 2009).
Some endemic genera:
Fam. Leptonetidae: Masirana Kishida, 1942, Falcileptoneta Komatsu, 1970
Acariformes
Oribatida. According to the checklist of Fujikawa et al. (1993, with suppl.), in
Japan (entire) are known oribatid mites of 286 genera and 735 spp.
7.28 Ryukyu Islands
7.28.1 G
and Paleogeography
Ryukyu Islands (Ryûkyû-shotô), known in Japanese as the Nansei-shotô, lit.

“Southwest Islands,” and also known as the Ryukyu Arc are a chain of more than
100 volcanic Japanese islands that stretch 1100 km southwest from Kyushu to
Taiwan: the Ôsumi, Tokara, Amami, Okinawa, and Sakishima Islands (further
divided into the Miyako and Yaeyama Islands), with Yonaguni the southernmost.
The largest of the islands is Okinawa. The surface of the archipelago is 4642 km2,
and the highest point is at 1936 m (Mt. Miyanoura-dake). The two largest islands
are Okinawa (1,204 square km) and Amami Great Island (712 square km).
The islands have a subtropical climate with mild winters and hot summers.
Precipitation is very high and is affected by the rainy season and typhoons. Except
the outlying Daitô Islands, the island chain has two major geologic boundaries, the
Tokara Strait between the Tokara and Amami Islands and the Kerama Gap between
the Okinawa and Miyako Islands.
Between 1.6 and 1.3 Ma, the East China Sea area, including most of the Okinawa
Trough, may have been subaerial. At that time, the Ryukyu Arc region may have
been a part of the Eurasian continent. Extensive subsidence may have occurred at
the second stage, at about 1.3 Ma, in the Early Pleistocene. The present Ryukyu Arc
(Ryukyu Ridge) has been formed since then. The Ryukyu Arc may have been nearly
connected to the Chinese continent, through Taiwan as a land bridge, sometime dur-
ing the two major development periods (such as sometime during 1.6–1.0 Ma and
0.2–0.025 Ma). The Paleo-land may have been submerged step by step since
0.03 Ma by both crustal movement and sea level rising after the last Ice Age.
Submarine stalactite caves at 10–35 m deep off the Ryukyu Islands were discov-
ered. The caves have subsided since the Würm Ice Age. Stone tools were also recov-
ered inside one of them (Kimura 2000).
Watase’s Line, which crosses the Tokara Islands, marks a major biogeographic
boundary. The north of the line belongs to the Palearctic region, while the southern
portion is the northern limit of the Oriental region. Yakushima in Ôsumi is the
southern limit of the Palearctic region. It is featured with millennium-old cedar
trees. The island is part of Kirishima-Yaku National Park and was designated as a
World Heritage Site by UNESCO in 1993.
The south of Watase’s Line is recognized by ecologists as a distinct subtropical
moist broadleaf forest ecoregion. The flora and fauna of the islands have much in
common with Taiwan, the Philippines, and Southeast Asia and are part of the
Indomalayan ecozone.
Mammals endemic to the islands include Iriomote cat, the Ryukyu flying fox, the
Ryukyu long-tailed giant rat, the Ryukyu mouse, and the Ryukyu shrew.
Approximately one half of the amphibian species of the islands are endemic.
7.28 Ryukyu Islands 439
The northern Ryukyu Islands are separated from Kyushu by the so-called Myake
Line.
Other sources: Dobson (1994), Kimura (2002)
Map 7.16 Map of the Ryukyus
On the Ryukyus are absent the orders Ricinulei, Opilioacarida, and Holothyrida.
The study of Haupt (1993) was “based on four groups of ground-living arach-
nids: whip scorpions and spiders (Hexathelidae, Ctenizidae, Mesothelae). A close
zoogeographical connection was found between the Yaeyama Islands and nearby
Taiwan. The central and northern parts of the Ryukyu Islands are faunistically clos-
est to Kyushu. No distinct border between an Oriental and a Palearctic fauna can be
detected. Instead, only climatic and other ecological factors appear to be responsi-
ble for distributional boundaries of various organisms observed in different regions
of southern Japan. Similarities to the fauna of continental China can be explained by
land connections between the Ryukyus and the continent during the Pleistocene.”
Among the Opiliones the “Oriental” elements dominate (Suzuki 1973).
Scorpiones Fam. Hormuridae – Liocheles australasiae (Fabricius) (Takashima

1941)
Pseudoscorpiones
Fam. Olpiidae – Nipponogarypus enoshimaensis okinoerabensis Morikawa, 1960
(endemic for the Ryukyus) (Morikawa 1960)
Opiliones
According to the paper of Suzuki (1971), “The Opiliones fauna of the Ryukyus is
composed of eleven genera and thirteen species (eleven species of Oriental origin
and two of Pale- or Holarctic one).” A later paper (Suzuki 1973) listed already 16
genera and 26 species (Phalangiidae, Podoctidae, Epedanidae, Travuniidae,
Sclerosomatidae (sub-“Leiobunidae”), and Ischyropsalididae (also Suzuki (1964)).
Cyphophthalmi – not recorded
Eupnoi
Fam. Phalangiidae
Fam. Sclerosomatidae – Gagrella Stoliczka, Metagagrella Roewer, Pseudogagrella
Redikorzev, Gagrellula Roewer, Leiobunum C.L. Koch, Nelima Roewer)
Dyspnoi
Fam. Nipponopsalididae (Nipponopsalis Martens et Suzuki)
Fam. Travuniidae (Peltonychia)
Fam Phalangodidae (Proscotolemon Roewer, Parabeloniscus Suzuki, Tokunosia
Suzuki)
Fam. Podoctidae (Dongmoa Roewer)
Fam. Epedanidae (Zepedanulus Roewer, Epedanellus Roewer, Kilungius Roewer,
Pseudobiantes Hirst)
Thelyphonida (Uropygi) In Japan live two species of Thelyphonida: Typopeltis
crucifer Pocock, 1894, in the southernmost Ryukyu Islands (from Iriomote Shima
to Ibeya Shima, also in Taiwan) and T. stimpsonii (Wood, 1862) from the islands
Tokunoshima to Amakusa and the south of Kyushu, the only Uropygi from the
mainland Japan (Yoshikura 1973) (Map 7.15). According to Karasawa et al. (2015),
“…the border of distribution of the two species lies between the Central and
Southern Ryukyus, i.e., the Kerama Gap.”
Schizomida The order is known only on the smaller islands: Orientzomus sawadai
(Kishida, 1930) – endemic sp. on Ogasawara-shotô (= Bonin Islands), Tokyo Pref.,
and three species on the Ryukyu Islands (Okinawa Pref.) (Cokendolpher 1988,
Kishida 1930, Shimojana 1981). There are no endemic genera for Japan.
Araneae The spider fauna of Ryukyus has been studied by several arachnologists
(Yaginuma, Komatsu, Shimojana, Haupt, and others). A preliminary report of cave
spiders (Shimojana 1977) contains 42 species of 36 genera and 23 families. To them
has been added the blind spider Coelotes okinawensis Shimojana et Nishihira, 2000.
7.29 Korea (North and South) 441
The conclusion of Shimojana (1977) is that “The cave spider fauna of the Ryukyu
Archipelago is much different from the Japanese Islands.”
Endemic genera from the Ryukyus:
Fam. Liphistiidae: Ryuthela Haupt, 1983 – Ryukyu Islands, Okinawa (seven spp.)
Fam. Corinnidae: Humua Ono, 1987 – one sp.
7.29 Korea (North and South)
7.29.1 G
and Paleogeography
Korea is a 1100-kilometer-long peninsula located in the easternmost part of the

Asian continent. To the northwest, the Amnok River (Yalu) separates Korea from
China, and to the northeast, the Duman (Tumen) River separates Korea from China
and Russia. Notable island is Jeju. The highest mountain in Korea is Mount Paektu
or Paektusan (2744 m), through which runs the border with China.
Mountains cover 70 percent of Korea. Most of the 3579 islands off the peninsula
are found along the south and the west coasts. Unlike most ancient mountains on the
mainland, many important islands in Korea were formed by volcanic activity in the
Cenozoic orogeny. Jeju Island, situated off the southern coast, is a large volcanic
island whose main mountain Mount Halla or Hallasan (1950 m) is the highest in
South Korea.
The climate of Korea differs dramatically from north to south. The southern
regions experience a relatively warm and wet climate similar to that of Japan,
affected by warm ocean waters including the East Korea Warm Current. The north-
ern regions experience a colder and to some extent more inland climate, in common
with Manchuria.
According to the World Wide Fund for Nature, Korea consists of several ecore-
gions. The South Korea evergreen forests occupy the southernmost portion of the
peninsula, as well as the island of Jeju. The Central Korea deciduous forests occupy
the more temperate central portion of the peninsula. Manchurian mixed forests
occupy the northern lowlands and low hills of the peninsula and extend north into
Manchuria as far as the Amur River on the Russia-China border. The Changbai
Mountains mixed forests include the higher elevation mountain region along the
North Korea-China border, where forests are dominated by conifers, with alpine
meadows and rock slopes on the highest peaks (World Wide Fund for Nature).
The sea level of the South Sea area was about 150–160 m. below present sea level about
15,000 years ago. This drop allowed a land bridge to form between the Korea peninsula and
Japan, which remained above sea level for about 6000 years. The sea level rose to approxi-
mately 60 m. (below present sea level) about 9000 years ago, remained at this depth until
about 4000–5000 years ago, and then rose to 10–20 m. below the present level about 3000–
4000 years ago. (Yeon Gyu Lee et al. 2008)
The arachnofauna fauna of North Korea is not well studied. On the peninsula are
known the orders Scorpiones, Pseudoscorpiones, Opiliones, Araneae, Uropygi,
Ixodida, Mesostigmata, Sarcoptiformes, and Trombidiformes. Lacking are
Palpigradi, Ricinulei, Amblypygi, Opilioacarida, Holothyrida, and Cyphophthalmi.
To be noted is the presence of the Opilion subfamily Kaolinonychinae
(Triaenonychidae, shared with Japan).
Our collection of Arachnida from North Korea (1982, 1987) is still in Sofia,
unidentified.
Scorpiones
Only two scorpion species have been recorded so far (Zhu et al. 2004).
Fam. Buthidae – Mesobuthus martensii
Fam. Hormuridae – Liocheles australasiae
Pseudoscorpiones
From South Korea have been recorded 19 spp. of 9 genera and the families
Chthoniidae, Pseudotyrannochthoniidae, Neobisiidae, Syarinidae, Cheiridiidae,
and Chernetidae (Morikawa 1970; Lee 1981, 1982, Harvey 2014). No information
about North Korea (material has been collected, but not yet identified).
Opiliones The total number of the Opiliones from Korea is 17 spp. of the families
Phalangiidae, Sclerosomatidae, Sabaconidae, Nipponopsalididae, and
Triaenonychidae (Kharitonow 1957, Roewer 1927, Staręga 1964, 1965, Suzuki
1941b, 1966, 1975, Kim et al. 2006, Ban Kwon and Kim 2010). According to Kury
(online), in Korea there are seven endemic species.
Cyphophthalmi are not recorded from Korea.
Eupnoi
Fam. Phalangiidae – Euphalangium Roewer, Mitopus Thorell, Oligolophus
C.L. Koch, Opilio Herbst, Himalphalangium Martens
Fam. Sclerosomatidae – Metagagrella Roewer, Nelima Roewer
Dyspnoi
Fam. Sabaconidae – Sabacon Simon
Fam. Nipponopsalididae – Nipponopsalis Martens et Suzuki
7.29 Korea (North and South) 443
Laniatores From the described area are known Laniatores from the family
Triaenonychidae (according to some authors, the subfamily Kaolinonychinae
should belong to Travuniidae).
Endemic genera are:
Fam. Triaenonychidae
Kaolinonychinae – Korea, Japan (two spp.)
Kaolinonychus Suzuki, 1975 – Korea (one sp.)
Fam. Thelyphonidae – according to Harvey (2013d), whip scorpions are not known
from Korea, but they certainly live there, as in the neighboring China and Japan
(Typopeltis?). Butler (1872) describes from “Korea” the new species Thelyphonus
lucanoides (= now considered a dweller of Sarawak and Indonesia).
Amblypygi – not known
Araneae The list of spider fauna of Korea (Paik 1967) contains 242 spp. of 120
genera and 33 families. Zoogeographically Korean spider fauna “comprises 136
northern species, 61 southern species, 6 cosmopolitan species, and 20 endemic spe-
cies.” Most species (125) are known also from Japan. Paik Kap-Yong (1914–1996)
contributed largely to the study of Korean spiders and also to the publishing of the
first pictorial encyclopedia (1978). Namkung Joon published in 2001 even bigger
pictorial encyclopedia of Korean spiders with taxonomic accounts on 546 spp. of
220 genera and 43 families. Namkung et al. (2009) increased this number to 681 spp.
of 258 genera and 46 families, including 131 endemic species (19.2%). The spider
fauna is close to the Japanese, influenced by northern elements, including 35
Holarctic and 67 Palearctic species. Most species are in Linyphiidae (82), Theridiidae
(77), Araneidae (70), and Salticidae (65).
Endemic genera:
Fam. Liphistiidae: Ryuthela Haupt, 1983
Fam. Leptonetidae: Longileptoneta Seo, 2015 (five spp.)
Fam. Gnaphosidae: Shiragaia Paik, 1992
Map 7.17 Map of Africa (political)
7.30 Africa, North of 20oN
7.30.1 G
and Paleogeography
In the narrow sense, here are included the territories of Mauretania, Western Sahara,
Morocco, Algeria, Tunisia, Libya, and Egypt, roughly north of 20oN. The
Mediterranean coast with its Mediterranean climate, especially in the Maghreb, is
followed south by Atlas Mountains, rising in Morocco up to 4167 m (Djebel
Toubkal). More to the south start the Saharan sands.
The classical schemes in zoogeography include North Africa in the Palearctic
region (under different names). We have to notice that
7.30 Africa, North of 20oN 445
Maghreb, by its origin, is a territory of Tirrenis and relates much more to Europe, than to
the rest of Africa; its autochthonic elements are Tyrrenic and are in common with the other
lands of the Western Mediterranean, and only by the end of the Miocene (in the Pontian)
were received various African elements (firstly from the Saharan-Ethiopian area) and
Asiatic elements, originating from Palestine and Egypt. (La Greca 1961)
Cox (2001) has left the African part of his “Eurasian” (generally called Palearctic)
region only a narrow band in Maghreb North of Atlas Mountains. After him, the
Sahara is an area where the former tropical flora of North Africa has disappeared and
logically is considered a part of the African (usually called “Afrotropical”) region.
One zoogeographical subdivision of North Africa was done by Joleau (1928).
The Atlas Mountains is a mountain range across the northwestern stretch of Africa
extending about 2,500 km through Algeria, Morocco, and Tunisia. The Atlas ranges
separate the Mediterranean and Atlantic coastlines from the Sahara Desert.
Map 7.18 The location of the Atlas Mountains across North Africa
In the Paleogene and Neogene periods (~66 million to ~1.8 million years ago),
the mountain chains that today comprise the Atlas were uplifted as the landmasses
of Europe and Africa collided at the southern end of the Iberian Peninsula. Such
convergent tectonic boundaries occur when two plates slide toward each other form-
ing a subduction zone (if one plate moves underneath the other) and/or a continental
collision (when the two plates contain continental crust). In the case of the Africa-
Europe collision, it is clear that tectonic convergence is partially responsible for the
formation of the High Atlas, as well as for the closure of the Strait of Gibraltar and
the formation of the Alps and the Pyrenees.
The Sahara is the world’s hottest and largest desert (over 9,400,000 km2). The
Sahara stretches from the Red Sea, including parts of the Mediterranean coasts, to
the Atlantic Ocean. To the south, it is delimited by the Sahel, a belt of semiarid
tropical savanna that composes the northern region of central and western sub-
Saharan Africa.
Sahara’s boundaries are the Atlantic Ocean on the west, the Atlas Mountains and
the Mediterranean on the north, the Red Sea on the east, and the Sudan (region) and
the valley of the Niger River on the south. The Sahara is divided into Western
Sahara, the central Ahaggar Mountains, the Tibesti Mountains, the Aïr Mountains (a
region of desert mountains and high plateaus), Ténéré Desert, and the Libyan Desert
(the most arid region). The highest peak in the Sahara is Emi Koussi (3,415 m) in
the Tibesti Mountains in northern Chad.
The southern border of the Sahara is marked by a band of semiarid savanna
called the Sahel; south of the Sahel lies Southern Sudan and the Congo River Basin.
Most of the Sahara consists of rocky hamada; ergs (large areas covered with sand
dunes) form only a minor part.
Sahara used to be a much wetter place than it is today. Over 30,000 petroglyphs of
river animals such as crocodiles survive, with half found in the Tassili n’Ajjer in
southeast Algeria. The modern Sahara, though, is not lush in vegetation, except in the
Nile Valley, at a few oases, and in the northern highlands, where Mediterranean plants
such as the olive tree are found to grow. The region has been this way since about
1600 BCE, after shifts in the Earth’s axis increased temperatures and decreased pre-
cipitation. Then, due to a climate change, the savanna changed into the sandy desert.
The Sahara covers large parts of Algeria, Chad, Egypt, Libya, Mali, Mauritania,
Morocco, Niger, Western Sahara, Sudan, and Tunisia. Several deeply dissected
mountains and mountain ranges and many volcanoes rise from the desert, including
the Aïr Mountains, Ahaggar Mountains, Saharan Atlas, Tibesti Mountains, Adrar
des Iforas, and the Red Sea hills.
Most of the rivers and streams in the Sahara are seasonal or intermittent, the chief
exception being the Nile River. Underground aquifers sometimes reach the surface,
forming oases. The central part of the Sahara is hyperarid, with little vegetation.
To the north, the Sahara reaches to the Mediterranean Sea in Egypt and portions
of Libya, but in Cyrenaica and the Maghreb, the Sahara borders Mediterranean for-
est, woodland, and scrub ecoregions of northern Africa, which have a Mediterranean
climate characterized by a winter rainy season. According to the botanical criteria
of Frank White, the northern limit of the Sahara corresponds to the northern limit of
date palm cultivation. The northern limit also corresponds to the 100 mm isohyet of
annual precipitation.
To the south, the Sahara is bounded by the Sahel, a belt of dry tropical savanna
with a summer rainy season that extends across Africa from east to west. According
to climatic criteria, the southern limit of the Sahara corresponds to the 150 mm
isohyet of annual precipitation (this is a long-term average, since precipitation var-
ies annually).
The Sahara comprises several distinct ecoregions, and with their variations in
temperature, rainfall, elevation, and soil, they harbor distinct communities of plants
and animals.
The Atlantic coastal desert is a narrow strip along the Atlantic coast, where fog
generated offshore by the cool Canary Current provides sufficient moisture to sus-
tain a variety of lichens, succulents, and shrubs. It covers 39,900 km2 in Western

Sahara and Mauritania.
The North Saharan steppe and woodlands is along the northern desert, next to
the Mediterranean forests, woodlands, and scrub ecoregions of the northern
Maghreb and Cyrenaica. It covers 1,675,300 km2 of Algeria, Egypt, Libya,
Mauritania, Morocco, Tunisia, and Western Sahara.
The Sahara Desert ecoregion covers the hyperarid central portion of the Sahara
where rainfall is minimal and sporadic. Vegetation is rare, and this ecoregion con-
sists mostly of sand dunes (erg, chech, raoui), stone plateaus (hamadas), gravel
plains (reg), dry valleys (wadis), and salt flats. It covers 4,639,900 km2 of Algeria,
Chad, Egypt, Libya, Mali, Mauritania, Niger, and Sudan.
The South Saharan steppe and woodlands ecoregion is a narrow band running
east and west between the hyperarid Sahara and the Sahel savannas to the south.
Movements of the equatorial Intertropical Convergence Zone (ITCZ) bring summer
rains during July and August which average 100 to 200 mm but vary greatly from
year to year. This ecoregion covers 1,101,700 km2 of Algeria, Chad, Mali,
Mauritania, and Sudan.
In the West Saharan montane xeric woodlands, several volcanic highlands
provide a cooler, moister environment that supports Sahara-Mediterranean wood-
lands and shrublands. The ecoregion covers 258,100 km2, mostly in the Tassili
n’Ajjer of Algeria, with smaller enclaves in the Aïr of Niger, the Dhar Adrar of
Mauritania, and the Adrar des Iforas of Mali and Algeria.
The Tibesti-Jebel Uweinat montane xeric woodlands ecoregion consists of the
Tibesti and Jebel Uweinat highlands. Higher and more regular rainfall and cooler
temperatures support woodlands and shrublands of palms, acacias, myrtle, olean-
der, tamarix, and several rare and endemic plants. The ecoregion covers 82,200 km2
in the Tibesti of Chad and Libya and Jebel Uweinat on the border of Egypt, Libya,
and Sudan (http://en.wikipedia.org/wiki/Sahara-cite_note-34).
The Saharan halophytics is an area of seasonally flooded saline depressions
which is home to halophytic (salt-adapted) plant communities. The Saharan halo-
phytics cover 54,000 km2, including the Qattara and Siwa depressions in northern
Egypt, the Tunisian salt lakes of central Tunisia, Chott Melghir in Algeria, and
smaller areas of Algeria, Mauritania, and Western Sahara.
The Tanezrouft is one of the harshest regions on Earth and the driest in the
Sahara, with no vegetation and very little life. It is along the borders of Algeria,
Niger, and Mali, west of the Ahaggar Mountains.
The flora of the Sahara is highly diversified based on the biogeographical char-
acteristics of this vast desert. Floristically, the Sahara has three zones based on the
amount of rainfall received – the Northern (Mediterranean), Central, and Southern
Zones. There are two transitional zones – the Mediterranean-Sahara transition and
the Sahel transition zone.
According to Udvardy (1975), North Africa is divided biogeographically into three

parts (from north to south): No 17 (Mediterranean sclerophyll), 28 (Atlas steppe),
and 18 (Sahara). The arachnofauna of the first two “provinces” (Udvardy) is very
close to the South European fauna, even as the same species (Ptychosoma vitellinum
Sør.). Here (especially in the caves) are found such hygrophilous animals as
Palpigradi or some pseudoscorpions of Neobisiidae.
As expected, the animals of the desert and hot climate as Solifugae and Scorpiones
are very numerous and quite different from the European-Mediterranean fauna. The
spider fauna is well studied up to the very summit of Atlas (4167 m) (Deltshev 2015,
collected by P. Beron).
Palpigradi Barranco and Mayoral (2007) described from the cave of Kef Aziza the
new species Eukoenenia maroccana – the third Moroccan Palpigradi, after E. mirabi-
lis (Grassi et Calandruccio, 1885) and E. hanseni (Silvestri, 1913) (Remy 1948, 1952).
Solifugae In North Africa are distributed Solifugae from 19 genera and 5 families:
Galeodidae, Karschiidae, Daesiidae, Solpugidae, and Rhagodidae (Borelli 1924,
Lawrence 1966).
Endemic genus of Solifugae for North Africa is Barrus Simon, 1880 – Egypt
(one sp.).
In the North African countries, the number of Solifugae species is as follows
(Table 7.3):
Scorpiones
According to the lists of Vachon (1952) and Dupré (2013a), updated (Lourenço and
Duhem 2007), from the North African countries are recorded scorpions of 24 genera
and 4 families.
From Tassili n’Ajjer, Vachon (1958) identified ten species of scorpions of seven
genera and the families Buthidae and Scorpionidae. According to this author, in the
“saharo-mountainious” region (Tassili n’Ajjer, Air, Adrar Iforas, and Tibesti) is seen
one altitudinal separation of species of Palearctic origin, having “climbed” above
1000 m with the establishment of drought in North Africa, and species, widespread
in Sahara.
This list has been completed by Lourenço (2009) with the description of
Compsobuthus tassili and by Qi Jian-Xin and Lourenço (2007) who described from
Mauritania the new genus and species Mauritanobuthus geniezi.
From Tunisia Vachon (1951) identified seven spp. of scorpions of the genera
Euscorpius, Scorpio, Buthus, Androctonus, Buthacus, and Buthiscus. Among the
scorpions of Tunisia, this author distinguishes two species of European origin
(“Buthus” occitanus and Euscorpius sicanus) and five North African elements.
The scorpions of Egypt (then 12 spp., now they are 28) have been listed as early
as 1910 by E. Simon. The list of Moroccan scorpions (50 spp.) is more recent – of
Nickel et al. (2009). Teruel (2007) described from Atlas the new genus and species
Cicileiurus monticola (relict at 2000 m) (Table 7.4).
Table 7.3 Distribution of Solifugae in North Africa

Country Mauritania Morocco Algeria Tunisia Libya Egypt
No. of species 4 25 30 23 19 27
Fam. Daesiidae + + + + + +
Biton Karsch – 1 6 5 5 4
Blossia Simon 1 5 1 1 – 2
Gluviopsilla – – 1 – – –
Roewer
Gnosippus – – – – – 1
Karsch
Tarabulida – – – – 1 –
Roewer
Fam. Galeodidae + + + + + +
Galeodes Olivier 1 9 7 9 7 1
Galeodopsis – – – – 1 –
Birula
Othoes Hirst 1 – 1 – – –
Paragaleodes 1 1 2 1 – 2
Kraepelin
Fam. – + + – + +
Karschiidae
Barrus Simon – – – – – 1
Eusimonia – 4 3 – 1 1
Kraepelin
Fam. – + + + + +
Rhagodidae
Rhagodes – – 1 1 1 3
Pocock
Rhagoditta – – 2 1 – 1
Roewer
Rhagodira – – 2 1 – 1
Roewer
Rhagodeya – – – – 1 –
Roewer
Fam. Solpugidae – + + + + +
Oparba Roewer – 1 – – – –
Oparbella – 2 4 4 1 2
Roewer
Solpuga – – 1 – 1 –
Lichtenstein
Zeria Simon – 1 1 – – –
Table 7.4 Scorpions in northern Africa

Country Morocco Algeria Tunisia Libya Egypt Mauritania
Number of Species 50 30 17 16 28 16 28
Ananteroides – – – – – 1 –
Borelli
Androctonus 7 7 4 3 5 5 5
Ehrenberg
Buthacus Birula 4 5 2 3 2 2 2
Butheoloides Hirst 5 1 – – – 1 –
Buthiscus Birula – 1 1 1 – – –
Buthus Leach 15 4 4 3 6 1 6
Cicileiurus Teruel 1 – – – – – –
Cicileus Vachon – 1 – – – – –
Compsobuthus 2 2 – 1 4 1 4
Vachon
Egyptobuthus – – – – 1 – 1
Lourenço
Hottentotta Birula 3 3 – 2 – – –
Isometrus – 1 – – – – –
Ehrenberg
Leiurus Ehrenberg – – – – 1 – 1
Lissothus Vachon – 1 – 1 – 1 –
Mauritanobuthus – – – – – 1 –
Qi et Lourenço
Microbuthus 1 – – – 1 – 1
Kraepelin
Orthochirus 3 2 1 1 2 1 2
Karsch
Parabuthus Pocock – – – – 2 – 2
Pseudolissothus – 1 1 – – – –
Lourenço
Saharobuthus 1 – – – – – –
Lourenço et
Duhem
Fam. + + + – – – +
Euscorpiidae
Euscorpius Thorell 1 2 3 – – – –
Fam. + + + + + + +
Scorpionidae
Nebo Simon – – – – 1 – 1
Scorpio L. 6 2 2 1 1 1 1
Fam. – – – – + – +
Hemiscorpiidae
Hemiscorpius – – – – 1 – 1
Peters
Endemic genera for the area are:

Mauritanobuthus Qi et Lourenço, 2007 – Mauritania (M. geniezi)
Saharobuthus Lourenço et Duhem, 2009 – Western Sahara
Lissothus Vachon, 1948 – Mauritania, Libya, Algeria (three spp.)
Cicileiurus Teruel, 2007 – Morocco
Pseudoscorpiones From North Africa (Egypt, Libya, Tunisia, Algeria, Morocco,
and Mauritania) and Tibesti in northern Chad are known Pseudoscorpiones of nine
families (Chthoniidae, Garypidae, Garypinidae, Geogarypidae, Olpiidae,
Atemnidae, Chernetidae, Cheliferidae, Withiidae) (Beier 1955, 1961, El-Hennawy
1988, Heurtault 1970a, 1970b, 1970c, 1971, 1990, Leclerc 1989, Mahnert 1983,
Vachon 1954). General papers of the pseudoscorpions have been published by
Callaini on Algeria (1983) and Morocco (1988).
According to Harvey (online), the number of species known from these countries
is Egypt, 14; Libya, 15; Tunisia, 26; Algeria, 43; Morocco, 38; Mauritania, 2; and
Tibesti in northern Chad (after Heurtault 1970 a, b, c, 1971), 8.
Endemic species are:
Chthonius kabylicus Callaini, 1983 – Algeria
Neobisium (Blothrus) peyerimhoffi Heurtault, 1990 – Algeria
N. dumitrescoae Heurtault, 1990 – Algeria
Roncus numidicus Callaini, 1983 – Algeria
Roncus (Parablothrus) comasi Mahnert, 1985 – Tunisia
Roncus (Parablothrus) gardinii Heurtault, 1990 – Algeria
Rhacochelifer massylicus Callaini, 1983 – Algeria
Geogarypus mirei Heurtault, 1970 – Chad (Borkou)
Goniochernes vachoni Heurtault, 1970 – Tibesti
Endemic genus for this (mostly desert) area is only Plesiowithius Vachon,
1954 – Mauritania (one sp.) (Withiidae).
Opiliones In North Africa are registered Opiliones of the families Phalangiidae,
Trogulidae, Nemastomatidae, and Phalangodidae. The Cyphophthalmi, so numer-
ous in South Europe, are not known so far from North Africa.
In Egypt (Cokendolpher 1990) are known six spp. of Opiliones: Trogulus gyp-
seus, Metaphalangium cirtanum, M. orientalis, Phalangium aegyptiacum, Ph. savi-
gnyi, and Ph. copticum.
Eupnoi
Fam. Phalangiidae – Metaphalangium Roewer, Phalangium L.
Dyspnoi
Fam. Nemastomatidae – Nemastomella M.L. (N. maarebensis Simon, 1913, ¿
“Nemastoma” lilliputanum Lucas, Algeria)
Fam. Trogulidae – Trogulus Latreille (one sp., Egypt)
Laniatores
Fam. Phalangodidae: Ptychosoma vitellinum Soerensen, 1873 – Algeria
Amblypygi
Musicodamon atlanteus Fage, 1939 (Phrynichidae), is an endemic genus and spe-
cies from Morocco and Algeria.
El-Hennawy and Hisham (2002) recorded from Egypt Charinus ioanniticus
(Kritscher) (Charinidae), known also from Turkey, Israel, and the Greek islands Kos
and Rhodes.
Charinidae and Phrynichidae are the only families of Amblypygi, known from
the Palearctic region.
Araneae Bosmans (many papers on Atlas spiders), Denis (1961, 1967),
El-Hennawy (1990 2006), Deltshev (2015), Caporiacco (1936)
Some endemic genera of spiders in Africa, north of 20oN:
Fam. Linyphiidae
Cherserigone Denis, 1954 – Algeria (one sp.)
Fam. Lycosidae
Phonophilus Ehrenberg, 1831 – Libya
Fam. Dictynidae
Chaerea Simon, 1884 – Algeria (one sp.)
Fam. Agelenidae
Pseudotegenaria Caporiacco, 1934 – Libya (one sp.)
Fam. Araneidae
Poecilarcys Simon, 1895 – Tunisia (one sp.)
Fam. Corinnidae
Scorteccia Caporiacco, 1936 – Libya (one sp.)
Opilioacarida One species (Opilioacarus segmentatus With, 1903) is known from
Algeria (also in South Europe)
Parasitiformes In Egypt have been registered Ixodida by Hoogstraal and Kaiser

(1958).
7.31 Africa Between 20oN and Zambezi-Kunene (Tropical or Intertropical Africa) 453
7.31 A
frica Between 20oN and Zambezi-Kunene (Tropical or
Intertropical Africa)
7.31.1 G
and Paleogeography
Across Sahara around the 20°N, is situated the usually accepted southern border of
the Palearctic (Holarctic), a frontier between one kingdom and another (Paleotropica).
South of the Sahara follow the belts of semidesert (Sahel) and the different types
of savanna. In the described area, there are also high mountains with afroalpine
vegetation and mountain rain forest.
Tropic rain forests are tropical moist forests of semi-deciduous varieties distrib-
uted across nine West African countries. In the first half of the 1980s, an annual
forest loss of 7200 square kilometers was noted down along the Gulf of Guinea, a
figure equivalent to 4–5% of the total remaining rain forest area. By 1985, 72 percent
of West Africa’s rain forests had been transformed into fallow lands, and an addi-
tional 9 percent had been opened up by timber exploitation. It is generally believed
that firewood provides 75 percent of the energy used in sub-Sahara Africa. With the
high demand, the consumption of wood for fuel exceeds the renewal of forest cover.
The rain forests which remain in West Africa now merely are how they were
hardly 30 years ago. In Guinea, Liberia, and the Ivory Coast, there is almost no
primary forest cover left unscathed; in Ghana the situation is much worse, and
nearly all the rain forest are cut down. Guinea-Bissau loses 200 to 350 km2 of forest
yearly, Senegal 500 km2 of wooded savanna, and Nigeria 6,000,050,000 of both.
Liberia exploits 800 km2 of forests each year. Tropical Africa is about 18% of the
world total covering 20 million km2 of land in West and Central Africa. Recent esti-
mates show that the annual pace of deforestation in the region can vary from 150 km2
in Gabon to 2900 km2 in Ivory Coast. The remaining tropical forests still cover
major areas in Central Africa but are abridged by patches in West Africa.
The tropical environment is rich in terms of biodiversity. Tropical African forest
is 18 percent of the world total and covers over 3.6 million square kilometers of land
in West, East, and Central Africa. This total area can be subdivided to 2.69 million
square kilometers (74%) in Central Africa, 680,000 square kilometers (19%) in
West Africa, and 250,000 square kilometers (7%) in East Africa.
In West Africa, a chain of rain forests up to 350 km long extends from the eastern
border of Sierra Leone all the way to Ghana. In Ghana the forest zone gradually
dispels near the Volta River, following a 300 km stretch of Dahomey savanna gap.
The rain forest of West Africa continues from east of Benin through southern Nigeria
and officially ends at the border of Cameroon along the Sanaga River.
The variety of the African rain forest flora is also less than the other rain forests.
This lack of flora has been credited to several reasons such as the gradual infertility
since the Miocene, severe dry periods during Quaternary, or the refuge theory of the
cool and dry climate of tropical Africa during the last severe Ice Age of about 18000
years ago.
These analyses of the prominent French entomologist are based mostly on the
distribution and the affinities of pselaphids and some other groups of Coleoptera.
When studying the distribution of Arachnida, it seems useful to compare the conclu-
sions of Jeannel with the data extracted of the recent profound research on many
groups of arachnids. It is clear, for example, that such group as Ricinulei in Africa
(only in the western part) suggest former very old “Afro-Brazilian” distribution.
There are similar examples also by other groups (the family Neogoveidae –
Opiliones, Cyphophthalmi).
Jeannel (loc. cit.) indicates that the equatorial forest in Central Africa was estab-
lished in the Pliocene and is only a residue of a much more extensive forest, coming
from the Malaysian area and broken by the vast steppic and desert regions of India,
Arabia, and East Africa, deprived of forest by climatic changes and human activi-
ties. So, when defining a group, family, or genus of arachnids (e.g., in Madagascar)
as having an African origin, we should think of a former origin of these (now East
African) elements from Southeast Asia. Some of the elements in the present-day
intertropical fauna of Africa actually originate of the southern “sudamadian” areas.
Usually tropical Africa is divided into two subregions: East Africa and West
Africa. Other sources: Hurni (1989), Scott (1958)
Old and rich fauna, with all orders of Arachnida (Holothyrida only on the islands)
Palpigradi – one endemic genus; Solifugae – six families, some endemic genera,
no endemic families
Ricinulei – one endemic genus in West Africa
Thelyphonida (Uropygi) – one endemic (relict?) genus in West Africa
Amblypygi – three families; one endemic and relict suborder in West Africa;
Schizomida – three endemic genera
Scorpiones – three families
Pseudoscorpiones – 16 families
Opiliones – eight families, many endemic genera
Araneae – many endemic genera
It is to notice the relict elements of orders, represented only in West Africa
(Ricinulei, Thelyphonida, or Uropygi, suborder Palaeoamblypygi, fam.
Neogoveidae).
Missing are the suborders Dyspnoi and Mesothelae.
Palpigradi Very few Palpigradi have been described from tropical Africa:
Eukoenenia pauli Condé, 1979 (Gabon), E. angolensis (Remy, 1956), E. machadoi
(Remy, 1950) (Angola), E. hesperia (Remy, 1953) (Ivory Coast), E. kenyana Condé,
1979 (Kenya), Koeneniodes notabilis Silvestri, 1913, Leptokoenenia scurra
Monniot, 1966 (Congo), and Allokoenenia afra Silvestri, 1913 (Guinea) (Condé
1979a, 1979b, Monniot 1966, Remy 1950, 1953, 1956, Silvestri 1913).
The genus Allokoenenia is endemic for tropical Africa. This number does not
reflect the real picture of the distribution of Palpigradi between 20oN and Kunene-
Zambezi. These tiny and fragile creatures are rarely collected, mostly by the few
specialists on them in person.
Table 7.5 Solifugae in the countries of Afrotropical Region

Countries Ethiopia Eritrea Somalia Djibouti
Number of species 46 5 48 8
Fam. Ceromidae + – – –
Ceroma Karsch 1 – – –
Fam. Daesiidae + + + +
Biton Karsch 7 2 8 2
Bitonota Roewer 1 – – –
Blossia Simon – – 2 –
Blossiana Roewer 1 – – –
Eberlanzia Roewer – – 1 –
Gluviopsilla Roewer – – 1 –
Gluviopsis Roewer 1 – 5 1
Gnosippus Karsch – 1 – –
Hemiblossia Roewer 1 – – –
Triditarsula Roewer 1 – – –
Fam. Galeodidae + – + +
Galeodes Olivier 7 – 5 1
Gluviema Caporiacco – – 1 –
Paragaleodes Kraepelin 2 – – –
Fam. Rhagodidae + + + +
Rhagodes Pocock – – 6 –
Rhagodessa Roewer – 1 – –
Rhagodia Roewer 1 – – –
Rhagodinus Roewer 1 – – 1
Rhagodippa Roewer – – 1
Rhagoditta Roewer 1 1 1 –
Rhagodoca Roewer 5 – 5 1
Rhagoduna Roewer 1 –
(continued)
Countries Ethiopia Eritrea Somalia Djibouti
Fam. Solpugidae + – + +
Ferrandia Roewer – – 2 –
Solpuga Lichtenstein 1 – – –
Solpugassa Roewer 2 – 1 2
Solpugeira Roewer 1 – – –
Solpugella Roewer – – – –
Solpugyla Roewer 1 – 1 –
Zeria Simon 8 – 3 –
Zeriassa Pocock 3 – 5 –
Countries Sudan Senegal Gambia Guinea Guinea-Bissau Chad Mali Niger
Number of sp. 21 1 1 1 5 1 2 6
Fam. + – – – + – – +
Daesiidae
Biton Karsch 4 – – – – – – 1
Blossia Simon 1 – – – – – – –
Gluviopsis – – – – – – – 1
Roewer
Gnosippus – – – – 1 – – –
Karsch
Fam. + – – + + + + +
Galeodidae
Galeodes 5 1 – – 1 1 1 2
Olivier
Othoes Hirst 1 – – – – – – –
Fam. + – + – – – – –
Rhagodidae
Rhagodalma 1 – – – – – – –
Roewer
Rhagodessa 3 – – – – – – –
Roewer
Rhagodeya 1 – – – – – – –
Roewer
Rhagodolus – – 1 – – – – –
Roewer
Rhagoduna 1 – – – – – – –
Roewer
Fam. + – – + + – – +
Solpugidae
Oparbella – – – – – – – 1
Roewer
(continued)
Countries Sudan Senegal Gambia Guinea Guinea-Bissau Chad Mali Niger

Solpugassa 1 – – – – – – –
Roewer
Zeria Simon 2 – – 1 2 – – –
Zeriassa 1 – – – – – – –
Pocock
Countries Nigeria Togo Cameroon Malawi Ruanda Kenya Tanzania
Number of 4 9 2 4 1 37 33
species
Fam. Ceromidae – – – + – + +
Ceroma Karsch – – – 2 – 1 4
Fam. Daesiidae – + – – – + +
Biton Karsch – 2 – – – 3 –
Bitonupa Roewer – – – – – 1 –
Blossia Simon – – – – – 2 3
Hemiblossia – – – – – 1 –
Roewer
Fam. Galeodidae + + + – – + –
Galeodes Olivier 1 2 1 – – 1 –
Fam. + – – – – + –
Rhagodidae
Rhagodes Pocock – – – – – – 2
Rhagodoca – – – – – 7 –
Roewer
Rhagodolus 1 – – – – – –
Roewer
Fam. Solpugidae + + + + + + +
Oparba Roewer 1 3 – – – – –
Oparbella Roewer – 1 – – – – –
Solpuga Licht. – – – – – 1 –
Solpugassa – – – – – – 1
Roewer
Solpugella – – – – 1 – –
Roewer
Solpugiba Roewer – – – – – 1 –
Solpugistella Turk – – – – – 1 –
Solpugylla – – – – – – 4
Roewer
Zeria Simon 1 1 1 2 – 11 15
Zeriassa Pocock – – – – – 7 3
(continued)
Countries Congo DR Congo Angola Zambia Mozambique
Number of sp. 1 30 31 4 13
Taxa
Fam. – + + – –
Ceromidae
Ceroma Karsch – 1 1 – –
Fam. – – + + –
Hexisopodidae
Chelypus – – 1 1 –
Purcell
Hexisopus – – 1 – –
Karsch
Representatives of six families are known from the described area: Ceromidae,
Daesiidae, Galeodidae, Hexisopodidae, Rhagodidae, and Solpugidae (El-Hennawy
2008, Lawrence 1960, Roewer 1934, Simonetta and Delle Cave 1968, Harvey 2013b).
None of the families is endemic or typical for tropical Africa.
Endemic genera for this area:
Blossiana Roewer (Ethiopia)
Bitonota Roewer (Ethiopia)
Bitonupa Roewer (Kenya)
Solpugistella Turk (Kenya)
Rhagodalma Roewer (Sudan)
Rhagodippa Roewer (Djibouti)
Benoit (1960) enumerated 29 species from the former Belgian Congo and
Ruanda-Urundi. They all live far from the dense tropical forest of Congo and are
concentrated mainly in Katanga.
Ricinulei All Ricinulei in Africa (and the Old World) are known from west tropical
Africa (11 spp., all of genus Ricinoides Ewing, 1929) (Guerin-Meneville 1838,
Hansen 1921, Hansen and Soerensen 1904, Judson and Hardy 2001, Legg 1976,
1978, 1982, Naskrecki 2008, Tuxen 1974).
Map 7.19 Distribution of Ricinulei (Genus Ricinoides Ewing) in Africa
Scorpiones
In tropical Africa from 20oN to Zambezi-Kunene are known 21 genera of scorpions
of the families Buthidae (13), Scorpionidae (three), Hemiscorpiidae (one), and
Hormuridae (four) (Dupré 2013, Kovaøik 2001, 2003, Lamoral and Reynders 1975,
Lourenço et al. 2010, Probst 1973, Vachon 1968) (Tables 7.6 and 7.7).
Table 7.6 Scorpions in Western and Central Tropical Africa
460
Countries Senegal Gambia Guinea Guinea-Bissau Liberia Ghana Burkina Faso Mali Congo DR Congo Gabon Centr. AR Angola
Number of species 11 2 15 3 3 5 55 8 18 14 9 4 24
Fam. Buthidae + + + + + + + + + + + +
Ananteroides – – 1 1 – – – – 1 1 – – –
Borelli
Androctonus 1 – – – – – – 2 4 4 4 1 3
Ehrenberg
Babycurus Karsch 2 – 3 – – 3 – – 1 – – – –
Buthacus Birula 2 – – – – – – – – – 1 – –
Butheoloides Hirst 1 – – – – 1 1 1 2 1 1 2 1
Buthus Leach 2 – 1 – – – – – 1 – – – –
Compsobuthus – – – – – 1 1 1 – – – 1 –
Vachon
Hottentotta Birula 1 1 1 1 – – 1 1 2 2 – – 1
Uroplectes Peters – – – – 1 – – – – – – – 5
Fam. + + + + + + ++ + – – 1 1 –
Scorpionidae
Pandinus Thorell 1 1 4 1 1 1 1 2 + + + + +
Scorpio L. 1 – – – – – 1 – – – – – 5
Fam. Hormuridae – – + – + – – – 2 3 1 1 –
Hormurus Thorell – – 2 – – – – – + + + + +
Liocheles – – 1 – – – – – 1 – – – 1
Sundevall
Opisthacanthus – – 2 – – – – – 1 1 2 1 1
Peters
7 Regional Arachnogeography
Countries Togo Benin Nigeria Niger Cameroon Equatorial Guinea
Number of sp. 5 4 7 14 17 5
Taxa
Fam. Buthidae + + + + + +
Akentrobuthus Lam. – 1 – – – –
Androctonus Ehrenberg 1 – – 3 – –
Babycurus Karsch 2 1 2 1 3 2
Buthacus Birula – – 1 2 – –
Butheoloides Hirst – – 1 1 – –
Buthus Leach – – – 1 1 –
Cicileus Vachon – – – 1 – –
Congobuthus Lour. – – – 2 – –
Darchenia Vachon – – – – 1 –
Hottentotta Birula 1 1 1 1 2 –
Leiurus Ehrenberg – – – – 1 –
Lychasioides Vachon – – – – 1 –
Uroplectes Peters – – 1 – 1 –
Fam. Scorpionidae + + + + + +
Pandinus Thorell 1 1 1 1 1 1
Scorpio L. – – – 1 1 –
Fam. Hormuridae – – – – + +
Hormurus Thorell – – – – 1 –
Opisthacanthus Peters – – – – 2 2
7.31 Africa Between 20oN and Zambezi-Kunene (Tropical or Intertropical Africa)
461
Table 7.7 Scorpions in Eastern Tropical Africa

Countries Sudan Ethiopia Erythraea Djibouti
Number of sp. 23 43 17 12
Taxa
Fam. Buthidae + + + +
Androctonus Ehr. 1 1 – –
Babycurus Karsch 4 4 1 –
Buthacus Birula 1 – 2 –
Butheoloides Hirst 2 – – –
Butheolus Simon – – 1 –
Buthus Leach 3 2 1 1
Compsobuthus Vachon 3 4 1 1
Gint Kovarik et al. 1 2 – –
Hottentotta Birula 6 3 2 2
Isometrus Ehrenb. 1 1 – –
Lanzatus Kovařik – 1 – –
Lychas C.L. Koch 1 2 – –
Microbuthus Kraep. 1 2 1 2
Neobuthus Hirst 2 1 1 1
Odonturus Karsch – 1 – –
Orthochiroides Kov. – 1 – –
Orthochirus Karsch – 1 – 1
Parabuthus Pocock 4 6 4 2
Sabinebuthus Lour. – 1 – 1
Somalibuthus Kov. – 1 – –
Somalicharmus Kovařik – 1 – –
Uroplectes Peters 1 3 – –
Uroplectoides Lour.
Fam. Scorpionidae + + + +
Pandinus Thorell 10 14 3 1
Fam. Hemiscorpiidae – + + –
Hemiscorpius Peters – 2 1 –
Fam. Hormuridae + – – –
Iomachus Pocock 1 – – –
Countries Kenya Tanzania Uganda Malawi
Number of sp. 2 29 6 7
Taxa
Fam. Buthidae + + + +
Babycurus Karsch 4 5 1 –
Hottentotta Birula 4 3 4 1
Isometrus Ehrenb. – – 1 –
Lychas C.L. Koch 2 – 3 1
Odonturus Karsch 1 – 1 –
Parabuthus Pocock 2 – 3 –
(continued)
7.31 Africa Between 20 oN and Zambezi-Kunene (Tropical or Intertropical Africa) 463
Countries Kenya Tanzania Uganda Malawi
Riftobuthus Lour. et al. 1 – – –
Uroplectes Peters 1 5 – –
Uroplectoides Lour. 1 – – –
Fam. Scorpionidae + + + +
Opistophthalmus – – 2 1
C.L. Koch
Pandinus Thorell 3 2 1 1
Fam. Hormuridae + + + +
Chiromachus Pocock – 1 – 2
Iomachus Pocock 1 1 1 1
Opisthacanthus Peters – 1 – –
Some endemic genera of Scorpiones in the area:

Fam. Buthidae: Riftobuthus Lourenço, Duhem, et Cloudsley-Thompson (Kenya);
Lanzatus Kovařik (Somalia)
Pseudoscorpiones In tropical Africa (the described area) have been recorded pseudo-
scorpions of 16 families: Chthoniidae, Tridenchthoniidae, Feaellidae, Garypidae,
Geogarypidae, Olpiidae, Ideoroncidae, Neobisiidae, Syarinidae, Cheiridiidae,
Pseudochiridiidae, Sternophoridae, Chernetidae, Cheliferidae, Atemnidae, and Withiidae
(Beier 1944, 1955, 1959, 1967, 1972, 1979, Heurtault 1970a, 1970b, 1970c, 1983,
Mahnert 1981, 1982a, 1982b, 1982c, 1983a, 1983b, 1985, 1988, Redikorzev 1924).
There are no endemic families of pseudoscorpions in tropical Africa.
Some endemic genera of pseudoscorpions are:
Fam. Chthoniidae
Congochthonius Beier, 1959 – DR Congo (one sp.)
Fam. Ideoroncidae
Afroroncus Mahnert, 1981 – Kenya (two spp.)
Nannoroncus Beier, 1955 – Kenya, Uganda (one sp.)
Fam. Withiidae
Aisthetowithius Beier, 1967 – Kenya, Tanzania (one sp.)
Cryptowithius Beier, 1967 – Kenya (one sp.)
Pogonowithius Beier, 1979 – DR Congo (one sp.)
Trichotowithius Beier, 1944 – Ethiopia, Kenya (two spp.)
Fam. Cheliferidae
Chamberlinarius Heurtault, 1983 – Ivory Coast (one sp.)
Fam. Atemnidae
Synatemnus Beier, 1944 – Tanzania (two spp.)
Opiliones Giribet and Prieto (2003), Goodnight and Goodnight (1959), Santos and
Prieto (2010), Sharma et al. (2011)
Map 7.20 Map of Cyphophthalmi of Africa and Madagascar

Ogovea – ◊
Parogovia– ♦
Marwe – ▲
Manangotria – ●
Purcellia – ►
Ankaratra – ◄
Parapurcellia – ■
Cyphophthalmi From tropical Africa are known three genera: Ogovea Hansen et
Soerensen, 1914, from the Gulf of Guinea (Cameroon, Congo, Bioko, three spp.);
Parogovia Hansen, 1921, from Bioko, Gabon, and Sierra Leone (three spp.); and the
aberrant genus Marwe Shear, 1985, from Kenya (one sp.). The family Ogoveidae is
endemic for the Gulf of Guinea. Fam. Neogoveidae has one genus in Africa
(Parogovia Hansen); the other genera live in South America. Marwe belongs (?) to
Sironidae (Hansen 1921, Hansen and Soerensen 1914, Shear 1985).
Eupnoi
According to the catalogues of Staręga (1984, 1992, with suppl.), in the described
area are recorded Eupnoi of the fam. Phalangiidae.
Fam. Phalangiidae (Phalangiinae) – Camerobunus Staręga et Snegovaya (1 sp.),
Cristina Loman (13 spp.), Rhampsinitus Simon (47 spp.), Guruia Loman,
Coptophalangium Staręga (1 sp.), Odontobunus Roewer, Dacnopilio Roewer,
Megistobunus Hansen, ?Hindreus Kauri
Endemic (known only from one country) are the genera:
Fam. Phalangiidae
Camerobunus Staręga et Snegovaya, 2008 – Cameroon
Coptophalangium Staręga, 1984 – Ethiopia
Dyspnoi – not recorded
Laniatores (Kauri 1985, Roewer, 1912, 1927, 1935, 1949a, 1950b, 1951a, 1952,
1954, 1957)
According to the catalogues of Staręga (1984, 1992, with suppl.), in the described
area are recorded Laniatores from 35 genera and 5 families.
Fam. Assamiidae – Aberdereca Goodnight et Goodnight (1 sp.), Bambereca Kauri
(1), Bwitonatus
Roewer (1), Comereca Roewer (1), Ereca Soerensen (23), Erecella Roewer (9),
Erecula Roewer (7),
Eusidama Roewer (1), Hypoxestus Loman (10), Metereca Roewer (= Leleupereca
Roewer),
Lygippulus Roewer (5), Metarhagdopygus Roewer, Randilea Roewer (1),
Sesostris Soerensen (5),
Sesostrellus Roewer (3), Spinixestus Roewer (6)
Fam. Pyramidopidae – Guinea-Bissau, Gambia, Equatorial Guinea, DR Congo,
Tanzania, Ghana,
Togo, Ivory Coast, São Tomé and Príncipe, Fuerteventura (Canary Islands) (13
genera, ca. 40 spp.)
Fam. Podoctidae (incl. Erecananinae and Ibaloniinae) – Erecanana Strand
Fam. Biantidae – Hinzuanius Karsch, Metabiantes Roewer, Monobiantes Lawrence,
Proconomma Roewer
Fam. Samoidae – Microconomma Roewer, Cameroon; Tetebius Roewer,

Mozambique
Some endemics (known only from one country) are the genera (almost all
described by Roewer and monotypic):
Microconomma Roewer – Cameroon
Tetebius Roewer – Mozambique
Zairebiantes Kauri
Prolacurbs Roewer – DR Congo
Monobiantes Lawrence – Kenya
Ivobiantes Lawrence – Ivory Coast
Heterolacurbs Roewer – Togo
Eubiantes Roewer – Tanzania
Clinobiantes Roewer – Cameroon
Biantomma Roewer – Bioco, Equ. Guinea
Biantella Roewer – Cameroon
Vilhena Lawrence – Angola
Uviranus Kauri – DR Congo
Umbonimba Roewer – Ivory Coast
Typhloburista Lawrence – Ivory Coast
Typhlobunus Roewer – Kenya
Tusipulla Roewer – Tanzania
Tubereca Kauri – Rwanda
Thomecola Roewer – São Tomé
Sokodea Roewer – Togo
Simienatus Roewer – Ethiopia
Seuthessus Kauri – DR Congo
Seuthesplus Roewer – Ivory Coast
Selencula Roewer – Bioco (Equatorial Guinea)
Scabrosidama Lawrence – Tanzania
Santhomea Roewer – São Tomé
Roewereca Lawrence – Tanzania
Randilea Roewer – Uganda (Elgon)
Pulchandrus Kauri – DR Congo
Podaucheniellus Roewer – Cameroon
Parasesostris Roewer – Tanzania
Parakodaika Goodnight et Goodnight – Angola
Palmanella Roewer – Sao Tomé
Numipedia Kauri – DR Congo
Nkogoa Roewer – Congo
Orsimonia Roewer – Ethiopia
Neocoryphus Lawrence – Ivory Coast
Neobaeorix Lawrence – Tanzania
Mwenga Kauri – DR Congo
Mutadia Kauri – DR Congo
Musola Roewer – Bioko (Equatorial Guinea)

Montereca Lawrence – Tanzania
Metasidama Roewer – Tanzania
Metasesostris Roewer – Tanzania
Merucola Roewer – Tanzania
Maracandinus Roewer – Ethiopia
Mandaria Roewer – Cameroon
Machadoessa Lawrence – Angola
Lygippus Roewer – Angola
Lucandamila Roewer – Tanzania
Lubudia Roewer – DR Congo
Lossidacola Roewer – Tanzania
Lossida Roewer – Tanzania
Lisposidama Lawrence – Tanzania
Leleupiolus Roewer – DR Congo
Kungwea Roewer – Tanzania
Kituvia Kauri – DR Congo
Karangania Kauri – DR Congo
Kasaina Lawrence – DR Congo
Kakontwea Roewer – DR Congo
Jaundea Roewer – Cameroon
Ivocoryphus Lawrence – Ivory Coast
Irumua Roewer – DR Congo
Irnia Roewer – Ethiopia
Humbea Roewer – Angola
Henriquea Roewer – Island Principé
Gulufia Roewer – Ethiopia
Gomezyta Roewer – Ethiopia
Fizibius Roewer – DR Congo
Fakoa Roewer – Cameroon
Eusidama Roewer – Tanzania
Eregonda Roewer – DR Congo
Erecops Roewer – Tanzania
Erecongoa Roewer – DR Congo
Erecabia Roewer – Tanzania
Erebalda Roewer – Tanzania
Ereala Roewer – DR Congo
Edeala Roewer – Cameroon
Dongila Roewer – Congo
Djemia Roewer – Ethiopia
Cangonia Roewer – DR Congo
Cogonella Roewer – DR Congo
Congolla Roewer – DR Congo
Comereca Roewer – DR Congo
Cereodiscus Roewer – Cameroon
Cereipes Roewer – Cameroon

Celimba Roewer – Tanzania
Cassinia Roewer – Guinea-Bissau
Bwitonatus Roewer – DR Congo
Buniabia Roewer – DR Congo
Bundukia Lawrence – Tanzania
Buemba Roewer – DR Congo
Bueana Roewer – Cameroon
Bolama Roewer – Guinea-Bissau
Blantyrea Roewer – Malawi
Binderia Roewer – Chad
Bibundina Roewer – Cameroon
Banconyx Lawrence – Ivory Coast
Bancoella Lawrence – Ivory Coast
Bambereca Kauri – DR Congo
Angopygoplus Lawrence – Angola
Angolyppa Lawrence – Angola
Amhara Pavesi – Ethiopia
Allereca Roewer – Rwanda
Afroassamia Caporiacco – Ethiopia
Adamauna Roewer – Cameroon
Acanthophrysella Strand – Tanzania
Acaca Roewer – Ethiopia
Aburistella Roewer – Ivory Coast
Aburista Roewer – Ghana
Aberdereca Goodnight et Goodn. – Kenya
Abanatus Roewer – DR Congo
Tonkouinatus Roewer – Ivory Coast
Scufia Roewer – Togo
Proconomma Roewer – DR Congo
Opconommula Roewer – Cameroon
Opconomma Roewer – São Tomé
Nimbadus Roewer – Ivory Coast
Micronimba Roewer – Ivory Coast
Metaconomma Kauri – DR Congo
Kwangonia Kauri – DR Congo
Aburiplus Roewer – Ghana
Amblypygi The first list of African Amblypygi (Lawrence 1969a) contained

16 spp. Without the North African Musicodamon and the island Charinus spp. from
Socotra, the Seychelles, and Madagascar, 13 spp. remain from tropical Africa (Delle
Cave 1986; Fage 1939, 1954; Hansen 1921; Lawrence 1958; Prendini et al. 2005;
Weygoldt 1972, 1995, 1999, 2000).
Meanwhile several nomenclatorial changes took place. The new genus
Euphrynichus Weigoldt, 1995, has been described; Titanodamon Pocock was syn-
onymized with Damon C.L. Koch, Myodalis Simon, and with Phrynichus Karsch.
Paracharon caecus Hansen should be added to the list.
“Hemiphrynus” machadoi Fage is now Xerophrynus Weigoldt, 1996 (endemic).
Now (2017) in tropical Africa are known 26 spp. of Amblypygi of seven genera
and three families (Charinidae, Phrynichidae, Paracharontidae).
Fam. Charinidae
Charinus Simon – widespread in the world, in continental tropical Africa known
from Guinea, Equatorial Guinea, São Tomé, and Somalia (six spp.).
Fam. Phrynichidae
Damon C.L. Koch – widespread and endemic in tropical Africa (11 spp., nine in
tropical Africa, north of Zambezi-Kunene). In his revision of the genus,
Weygoldt (2006) concluded that there are four species confined to West Africa
(which evolved mostly in rain forests) and six species (mostly smaller and
capable to survive in dryer habitats) distributed from East Africa to South
Africa and Namibia.
Euphrynichus Weygoldt – Kenya, Tanzania, Zimbabwe, Angola, Malawi (two
spp. – E. amanica (Werner, 1916), E. bacillifer (Gerstaecker, 1873)
Phrynichus Karsch – Cameroon, Eritrea, Somalia, Djibouti, Rwanda, Kenya,
Tanzania, Congo (seven spp.)
Xerophrynus Weigoldt – Angola (one sp. X. machadoi with unclear position),
Namibia
Fam. Paracharontidae
Paracharon Hansen – P. caecus Hansen, 1921
The most interesting amblypygid in tropical Africa is Paracharon Hansen from
Guinea-Bissau (one sp., P. caecus Hansen, 1921), the only member of the family
Paracharontidae and the suborder Palaeoamblypygi (“living fossil”).
Map 7.21 Amblypygi and Uropygi in Africa

Charinus – x
Paracharon –
Musicodamon – ▲
Euphrynichus – ▼
Etienneus (Uropygi) – ♣
Damon – ●
Phrynichus – ■
Xerophrynus – ◊
Phrynichodamon – □
Thelyphonida (Uropygi) The enigmatic Etienneus africanus (Hentschel, 1899) from

Senegal, Gambia, Guinea, and Guinea-Bissau is now considered autochthonous and
relict (Huff and Prendini 2009). Nowhere else in Africa live Uropygi (Cooke and Shadab
1973, Harvey 2013d, Hentschel 1899, Heurtault 1984, Huff and Prendini 2009).
Schizomida From tropical Africa are known five genera of Schizomida (four
endemic) of seven species (all endemic), all of family Hubbardiidae (Hubbardiinae)
(Lawrence 1969b, Reddell and Cokendolpher 1995, Harvey 2013g, Armas 2014).
Fam. Hubbardiidae
Enigmazomus Harvey, 2006 – Somalia, Arabian Peninsula (two spp.)
Afrozomus Reddell et Cokendolpher, 1995 – Angola, Congo (one sp.)
Artacarus Cook, 1899 – Liberia, Ivory Coast (one sp.)
Kenyazomus Armas, 2014 – Kenya (one sp.)
Lawrencezomus Armas, 2014 – Cameroon, Liberia (two spp.)
Map 7.22 Distribution of Schizomida in Africa, Socotra, Seychelles, and Madagascar

Megaschizomus – X
Enigmazomus – ▼
Kenyazomus – ▲
Lawrencezomus –
Mahezomus –
Afrozomus – ►
Artacarus – □
Bamazomus – ■
Anepsiozomus – ◊
Ovozomus –
Secozomus –
Araneae The spiders of Africa south of Sahara have been (and are) studied by
many arachnologists, and among them are Benoit (many papers), Griswold 1991,
Holm 1962, Scharff 1992, and others.
According to Scharff (1990), in Africa south of Sahara are considered valid 365
species of Linyphiidae of 67 genera.
Some endemic genera of spiders in tropical Africa from 20oN to
Zambezi-Kunene:
Fam. Theridiidae
Pycnoepisinus Wunderlich, 2008 – Kenya (one sp.)
Fam. Theraphosidae
Bacillochilus Gallon, 2010 – Angola (one sp.)
Batesiella Pocock, 1903 – Cameroon (one sp.)
Encyocratella Strand, 1907 – Tanzania (one sp.)
Eucratoscelus Pocock, 1898 – Kenya, Tanzania (two spp.)
Loxomphalia Simon, 1889 – Zanzibar (one)
Loxoptygus Simon, 1903 – Ethiopia (three spp.)
Pelinobius Karsch, 1885 – Kenya, Tanzania (one sp.)
Fam. Liocranidae
Cteniogaster Bosselaers et Jocqué, 2013 – Tanzania, Kenya (seven spp.)
ToxoniellaWarni et Jocqué, 2002 – Kenya (two spp.)
Fam. Mimetidae
Kratochvilia Strand, 1934 – Principe (one sp.)
Fam. Mysmenidae
Kilifina Baert et Murphy, 1992 – Kenya (one sp.)
Leviola Miller, 1970 – Angola (one sp.)
Fam. Oecobiidae
Urocteana Roewer, 1961 – Senegal (one sp.)
Fam. Oonopidae
Anophthalmoonops Benoit, 1976 – Angola
Blanioonops Simon et Fage, 1922 – East Africa (one sp.)
Caecoonops Benoit, 1964 – Congo (two spp.)
Hypnoonops Benoit, 1977 – Congo (one sp.)
Kijabe Berland, 1914 – Kenya, East Africa (two spp.)
Termitoonops Benoit, 1964 – Congo (five spp.)
Zyngoonops Benoit, 1977 – Congo (one sp.)
Fam. Palpimanidae
Badia Roewer, 1961 – Senegal (one sp.)
Fam. Phyxelididae
Kulalania Griswold, 1990 – Kenya (one sp.)
Fam. Pisauridae
Cispinilus Roewer, 1955 – Central Africa (one sp.)
Conakrya Schmidt, 1956 – Guinea (one sp.)
Tapinothele Simon, 1898 – Zanzibar (one sp.)
Vuattouxia Blandin, 1979 – Ivory Coast (one sp.)
Fam. Prodidomidae
Katumbea Cooke, 1964 – Tanzania (one sp.)
Plutonodomus Cooke, 1964 – Tanzania (one sp.)
Theumella Strand, 1906 – Ethiopia (two spp.)
Fam. Sparassidae
Berlandia Lessert, 1921 – East Africa (two spp.)
Sarotesius Pocock, 1898 – East Africa (one sp.)
Stasinoides Berland, 1922 – Ethiopia (one sp.)
Fam. Tetrablemmidae
Anansia Lehtinen, 1981 – Angola (one sp.)
Cuangoblemma Brignoli, 1974 – Angola (one sp.)
Hexablemma Berland, 1920 – Kenya (one sp.)
Fam. Tetragnathidae
Parazilia Lessert, 1938 – Congo
Fam. Anapidae
Forsteriola Brignoli, 1981 – Congo, Burundi, Rwanda (two spp.)
Fam. Ctenidae
Arctenus Polotow et Jocqué, 2014 – Kenya (one sp.)
Fam. Dictynidae
Hoplolathys Caporiacco, 1947 – Ethiopia (one sp.)
Mashimo Lehtinen, 1967 – Zambia (one sp.)
Fam. Zodariidae
Mastidiores Jocqué, 1987 – Kenya (one sp.)
7.32 Differences Between the Arachnofaunas of East and West Africa 475
Fam. Thomisidae
Felsina Simon, 1895 – West Africa (one sp.)
Gnoerichia Dahl, 1907 – Cameroon (one sp.)
Haedanula Caporiacco, 1941 – Ethiopia (one sp.)
Heriaesynaema Caporiacco, 1939 – Ethiopia (one sp.)
Hewittia Lessert, 1928 – Congo (one sp.)
Mystaria Simon, 1895 – West Africa (two spp.)
Ostanes Simon, 1895 – West Africa (one sp.)
Parasmodix Jezequel, 1966 – Ivory Coast (one sp.)
Pasiasula Roewer, 1942 – Bioco (one sp.)
Opilioacarida The known opilioacarids of tropical Africa belong to four species:
Phalangiacarus brosseti Coineau et Van der Hammen, 1979, from Gabon;
Salfacarus tanzaniensis Van der Hammen, 1977, from Tanzania; and two Panchaetes
from Ivory Coast and Angola. The genera Panchaetes Naudo and Phalangiacarus
Coineau et Van der Hammen are known only from tropical Africa (Beron 2014).
Acariformes: Beron (2008a), Niedbala (2002)
Prostigmata
Fam. Erythraeidae
Lomeustium Haitlinger, 2006 – one sp. (Togo, Ghana, Benin) (larval, endemic
genus)
Opserythraeus Fain, 1996 – one sp. (Rwanda) (larval, endemic genus)
7.32 D
ifferences Between the Arachnofaunas of East
and West Africa
In many manuals in zoogeography (Geptner 1936), delimitation of tropical Africa is

made between the eastern part (mostly savanna, including also the highest moun-
tains) and western part (mostly rain forest, or what remains from it, and savanna
forest). Usually they are considered at level of subregions.
Eastern part. Geptner 1936: East Africa plus Sahel and the area between DR
Congo and Southern Africa
Western part. Geptner 1936: From Senegal to (including) DR Congo and
Northern Angola
Palpigradi – endemic genus in West Africa, Allokoenenia (Guinea); in East Africa,
none. The same family.
Ricinulei – in the Old World only in West Africa (end. genus Ricinoides in Gambia,
Guinea, Guinea-Bissau, Nigeria, Ghana, Cameroon, Sierra Leone, Equatorial
Guinea, Benin, Congo, Togo, Ivory Coast
Amblypygi – in West Africa (Guinea-Bissau) endemic suborder

(Palaeoamblypygi) with one family, genus, and species
Thelyphonida (Uropygi) – in Africa only in the western part (end. genus Etienneus
in Senegal, Guinea-Bissau, Gambia, Guinea). Relicts?
Schizomida – endemic genera in West Africa: Afrozomus, Artacarus,
Lawrencezomus; in East Africa, Enigmazomus, Kenyazomus. No endemic
families.
Scorpiones – endemic genera in West Africa
In East Africa: Somalibuthus, Riftobuthus, Somalicharmus, Lanzaia. No endemic
families.
Pseudoscorpiones – endemic genera in West Africa: Congochthonius,
Pogonowithius, Chamberlinarius;
In East Africa: Afroroncus, Nannoroncus, Aisthetowithius, Cryptowithius,
Trichotowithius, Synatemnus. No endemic families.
Opiliones
Cyphophthalmi – endemics in West Africa, Ogoveidae (endemic family), Ogovea,
Paragovia; in East Africa, Marwe (Kenya)
Opilioacarida – in West Africa two endemic genera (Panchaetes, Angola, Ivory
Coast; Phalangiacarus, Gabon); in Tanzania the genus Salfacarus, known also from
Southern Africa and Madagascar
Holothyrida – not living on the African continent, but found in the Seychelles
near East Africa
Conclusion In West Africa there is one endemic suborder (Palaeoamblypygi), one
order living in Africa only in the western part (Ricinulei), and two endemic fami-
lies, living only in the western part of Africa. In East Africa there are no orders
or suborders living only there (Holothyrida on the Seychelles). In both parts
there are many endemic genera of various orders. The differences between both
parts are mainly in the landscape and the flora.
7.33 Tropical South America
7.33.1 G
and Paleogeography
The South American continent between Panama and the 30oC is the undisputed core
of Neotropica in the classical subdivision of the world. Its center is the great
Amazonian selva and its backbone are the Andes. It contains also the cold plains of
Bolivia (the Altiplano) and many other specific features.
The chain of the Andes reaches 6961 m (Aconcagua), and many of the world
records of high-altitude arachnids (Solifugae, etc.) are registered in Peruvian and
7.33 Tropical South America 477
other parts of the Andes. In Peru we visited once the highest high forests in the
world (Polylepis forest, with trees as tall as 15 meters at an altitude of 5500 m).
The tropical zone of South America consists of three fundamentals of vegetal formations:
the wet tropical forest; the Serrados et similar formations of savanna type, corresponding to
open non xerophytic formations; the Caatingas, open xerophytic formations. The wet tropi-
cal forest includes two vast regions: Hileia and Atlantic forest. (Lourenço 1986)
What concerns the climatic regions of South America, after Eidt (1968), is that
we follow the Köppen system – four major types of climate: A, B, C, and E. The A
climates, or those in which average monthly temperatures are all above 18oC, are
divided into two primary types: the ever-humid or tropical rain forest variety (Af)
and the “winter” dry season or savanna type (Aw). The four zones of Af are the
Pacific coast of Colombia and Ecuador, Amazon Basin, Bahia-Victoria coast of
Brazil, and Rio de Janeiro-Santos coast of Brazil. Monthly and annual temperature
ranges are less than 3oC throughout the entire area with the exception of some of the
highest valleys in the southwest. In the Andes this zone coincides with the so-called
Tierra Caliente and Tierra Templada. The second type of A climate, the savanna of
Aw variety, is with a definite dry season with insufficient moisture to support a true
rain forest vegetation.
The Neotropical region has been subdivided many times (Cabrera and Willink
1973, 1980, Cabrera and Yepes 1940, Sclater and Sclater 1999, Eidt 1968, Harrington
1962, Martin 1968, Mello-Leitão 1931, 1939, Morrone 2001, 2006, 2010, 2015,
Müller 1974, Roig-Juñent et al. 2006). Here is one biogeographic subdivision by
Morrone (2006):
The Nearctic region, inhabited by Holarctic insect taxa, comprises five provinces:
California, Baja California, Sonora, Mexican Plateau, and Tamaulipas. The Mexican transi-
tion zone comprises five provinces: Sierra Madre Occidental, Sierra Madre Oriental,
Transmexican Volcanic Belt, Balsas Basin, and Sierra Madre del Sur. The Neotropical
region, which harbors many insect taxa with close relatives in the tropical areas of the Old
World, comprises four subregions: Caribbean, Amazonian, Chacoan, and Parana. The
South American transition zone comprises five provinces: North Andean Paramo, Coastal
Peruvian Desert, Puna, Atacama, Prepuna, and Monte. The Andean region, which harbors
insect taxa with close relatives in the Austral continents, comprises three subregions:
Central Chilean, Subantarctic, and Patagonian.
The most important of the biogeographic subdivisions of Morrone is the dividing

of South America into two regions: Neotropical (mostly lowlands) and Andean
(mostly mountainous, but including the southernmost plains of Patagonia). The sepa-
ration is along the meridian and not across the continent. The “Andean region”
includes very long band, from Cape Horn to Venezuela. The “Caribbean” Subregion
of Morrone almost separates from the main area of the “Amazonian, Chacoan, and
Parana” subregions, which is not very practical. Most classical zoogeographers con-
sider the whole of South America as an entire Neotropical region (some of them sepa-
rating the far south with its relations to Australian or/and Neozealandian faunas).
South America, one of the richest zoogeographical territories in the world, was
longtime considered to represent one (the only) region (Neotropical) of the Neogean
Kingdom. However, the difference – climatic and paleogeographical – of Patagonia
with the tropical part of the continent has been noticed since longtime. The presence
of many elements in the Patagonian fauna close to the fauna of Australia and New
Zealand was incited by some authors (Lopatin 1980) to include the south of Chile
and Argentina in the Notogean Kingdom. We divided conditionally South America
in Patagonian (south of 30oS) and tropical parts, in order to examine their arachno-
fauna separately. The very active work of Brazilian, Argentinian, and Mexican
arachnologists made possible to elucidate many aspects in the arachnofauna of the
continent – see Mello-Leitão (1937) and Ringuelet (1956, 1978).
In tropical South America are recorded so far most of the orders and suborders of
Arachnida: Palpigradi (no endemics above species), Solifugae (2 families, one of
them endemic for tropical South America), Ricinulei (17 spp. of 1 genus, only end.
species), Scorpiones (3 end. genera; ?1 end. family in caves); Brazil is one of the
richest countries in scorpions (165 spp. of 26 genera): Pseudoscorpiones (16 fami-
lies, 18 endemic genera), Opiliones (Cyphophthalmi, 5 endemic genera; Eupnoi, 1
endemic genus; Laniatores, huge variety, with 15 fam., 13 of them endemic),
Amblypygi (many species, 1 endemic genus, the only genus in the family
Phrynichidae living in the Western Hemisphere), Uropygi (2 endemic genera),
Schizomida (8 genera), and Araneae (spiders of 77 families are known south of
Panama (72% of all spider families)). No family is endemic for the Southamerican
continent. There are Opilioacarida (two spp.), Holothyrida (two endemic genera),
many mites. Absent are the suborders Palaeoamblypygi, Dyspnoi, and Mesothelae.
As a whole, the level of endemism is too low to characterize a kingdom.
The regions, subregions, and provinces of Morrone (2001) are characterized by
many taxa, but very few of them (and not the most typical) are among the
arachnids.
In their analysis of the biogeographic history of South American arid lands,
Roig-Juñent et al. (2006) outlined 21 areas of endemism south of 5oS. Among the
taxa used were spiders (Nemesiidae: Acanthogonatus Karsch and Diplothelopsini)
and scorpions. Their main conclusion was that “That area of endemism of Patagonia
form a natural group, showing that this biota evolved as a unit, as well the biota that
occurs in the areas of Central Chile.”
Map 7.23 Zoogeographical subdivision of South America

(I) (after Cabrera and Yepes 1940) – 1. Sabánica, 2. Amazonica, 3. Tropical, 4.

Subtropical, 5. Tupi, 6. Pampasica, 7. Patagónica, 8. Subandino, 9. Chileno,
10. Andino, 11. Incásico
(II) (after Mello-Leitão 1931) – 1. Guayano-Amazonica, 2. Andina, 3. Bororô-
Cariri, 4. Tupi-Guarani, 5. Patagónica
(III) (after Mello-Leitão 1937) – 1. Caribea, 2. Hyléa, 3. Gê, 4. Bororo, 5. Tupi, 6.
Guarani, 7. Andino-Patagonica
(IV) (Sclater and Sclater 1999) – 1. Colombiana, 2. Amazonica, 3. Sud-brasileira,
4. Patagónica
Map 7.24 Regions of the world, with indication of the transition zones in gray (After Morrone
2015b) 1, Mexican; 2, Sahar-Arabian; 3, Chinese; 4, South American; 5, Indomalayan
According to the biogeographical subdivision in Morrone (2001), the sierra in

Ecuador belongs to the Andean region and the other three areas (Costa, Oriente, and
Insular) – to the Neotropical region.
Palpigradi Very few Palpigradi have been recorded from South America – see
Condé (1979, 1986, 1993), Hansen (1901), and Mello-Leitão and Arlé (1935).
Condé (1996) enumerates Eukoenenia improvisa Condé, 1979, and Koeneniodes
notabilis Silvestri, 1913, from French Guiana; E. grassii (Silvestri in litt., Hansen,
1901) from Paraguay; and E. janetscheki Condé, 1993, and E. roquettei (Mello-
Leitão et Arlé, 1935) from Brazil. The first troglobitic Palpigradi from South
America is Eukoenenia spelunca Souza et Ferreira, 2010, again from Brazil.
Followed several new species from the caves of Brazil (Sousa and Ferreira 2011,
2012a, 2012b) and two Leptokoenenia (the first in the New World) have been
described from an iron ore mine in Brazil (Sousa and Ferreira 2013). The wide-
spread E. florenciae is known also from Colombia.
Solifugae In tropical (north of Chile and Argentina and south of Panama) South
America are known representatives of two families of Solifugae: Ammotrechidae
and Mummuciidae (?14) (Kraepelin 1900, Maury 1982, Mello-Leitão 1937, Muma
1976, Roewer 1934, 1957).
Endemic genera are:
Fam. Mummuciidae (and five genera)
Gauchella Mello-Leitão, 1937 – Bolivia (one sp.)
Metacleobis Roewer, 1934 – Brazil (one sp.)
Mummucina Roewer, 1934 – Chile, (five spp.)
Mummuciona Roewer, 1934 – Venezuela (one sp.)
Mummucipes Roewer, 1934 – Paraguay (one sp.)
Fam. Ammotrechidae
Ammotrechinae
Campostrecha Mello-Leitão, 1937 – Ecuador (one sp.)
Saronominae
Saronomus Kraepelin, 1900 – Colombia, Venezuela (one sp.)
Subfamily not assigned
Eutrecha Maury, 1982 – Venezuela (one sp.)
Xenotrecha Maury, 1982 — Venezuela (one sp.)
Ricinulei
So far 17 spp. have been recorded from South America, the southernmost localities
being in Peru and Brazil. They belong to the single genus Cryptocellus Westwood
(Ricinoididae), which is found also in Central America and Cuba (Cokendolpher
2000, Cooke 1967, Cooreman 1977, Dumitresco and Juvara-Balş 1977, Ewing
1929, González-Sponga 1998, Platnick 1977, 1980, 1981, 1988, Platnick and Paz
1979, Roewer 1952, Tourinho and Salette de Azevedo 2007, Tourinho et al. 2010,
2014, Pinto-da-Rocha and Andrade 2012, Pinto-da-Rocha and Bonaldo 2007).
Scorpiones
In South America are represented eight families of scorpions: Buthidae, Chactidae,
Hormuridae, Bothriuridae, Troglotayosicidae, Scorpionidae, Caraboctonidae, and
Euscorpiidae (Mello-Leitão 1942, 1945, 1949, Lourenço 1981, 1986, 1987,
Lourenço et al. 2004, Lourenço and Duhem 2009, Lourenço and Monod 2000,
Ringuelet 1953)
In Paraguay, according to Maury (1984), have been found until then seven spp.
of Buthidae and five spp. of Bothriuridae.
In Bolivia Acosta and Ochoa (2002) have listed 24 spp. of scorpions from the
families Bothriuridae, Buthidae, and Iuridae.
The Scorpiones of Ecuador, according to Lourenço (1995a), are 36 spp. of 8
genera and 4 families (Bothriuridae, Buthidae, Chactidae, and Iuridae). According
to him, “Predominant distributional patterns show an Amazonian origin for some
Buthidae (Ananteris and Tityus) and Chactidae (genus Chactas), whereas several
other Buthidae (genus Tityus), Chactidae (genus Teuthraustes) and Iuridae (genus
Hadruroides) represent a typical andine fauna.” Now the genus Hadruroides is con-
sidered a member of fam. Caraboctonidae (Soleglad and Fet 2003), and the number
of Scorpiones in Ecuador is already 48 (or 47 on the continent and two on Galapagos,
according to Brito and Borges 2015).
Acosta (online) compiled a list of Argentinian scorpions, based on Acosta and
Maury (1998). It contains the families Buthidae and Bothriuridae (Table 7.8).
So far the following genera of scorpions are known only from tropical South
America:
Brazilobothriurus Lourenço et Monod, 2000; Troglorhopalurus Lourenço,
Baptista, et Giupponi, 2004; Troglotayosicus Lourenço, 1981
Fam. Troglotayosicidae – if the European Belisarius Simon does not belong to
this family, the family would be endemic for tropical South America.
Pseudoscorpiones The checklist of Ceballos and Florez (2007) lists for Colombia
only 22 spp. of Pseudoscorpiones of 18 genera and 6 families. Mahnert and Adis
(2002) enumerate from Amazonia 34 genera of 12 families.
According to the catalogue of Harvey (1990) and Harvey (2011, 2013f) and the
supplementary publications from tropical South America (between Panama and
30oS) are recorded pseudoscorpions from 17 families: Chthoniidae, Lechytiidae,
Tridenchthoniidae, Cheiridiidae, Feaellidae, Bochicidae, Syarinidae, Ideoroncidae,
Geogarypidae, Garypidae, Garypinidae, Gymnobisiidae, Olpiidae, Atemnidae,
Cheliferidae, Chernetidae, and Withiidae (Beier 1959a, Heurtault 1986c, Andrade
and Mahnert, 2003, Mahnert 1979, 1984, 1985b, 1985c, 1994, 2001, 2009, Mahnert
and Adis 1985, Mahnert et al. 1986, Harvey et al. 2016).
The following are the number of pseudoscorpion species in different countries:
Brazil, 164; Ecuador, 59; Peru, 42; Colombia, 22; Venezuela, 62; Guyana, 9;
French Guiana, 1; Suriname, 5; and Bolivia, 6. Some of these countries are clearly
understudied.
From this area endemic genera are:
Cryptoditha J.C. Chamberlin et R.V. Chamberlin, 1945 – Brazil (one sp.)
Haploditha Caporiacco, 1951 – Venezuela (one sp.)
Neoditha Feio, 1945 – Brazil (one sp.)
Sororoditha J.C. Chamberlin et R.V. Chamberlin, 1945 – Brazil (one sp.)
Spelaeobochica Mahnert, 2001 – Brazil (one sp.)
Fam. Feaellidae
Iporangella Harvey, Andrade, et Pinto-da-Rocha, 2016 – Brazil (one sp.)
Fam. Ideoroncidae
Ideoroncus Balzan, 1887 – Brazil, Paraguay (eight spp.)
Fam. Cheliferidae
Lophodactylus J.C. Chamberlin, 1932 – Brazil (one sp.)
Table 7.8 Scorpions in South America
484
Country Brazil Colom. Venezuela Guyana Suriname French Guiana Uruguay Paraguay
Number of species 165 80 230 26 14 25 6 14
Таxa
Fam. Buthidae + + + + + + + + +
Ananteris Thorell 23 13 31 8 2 8 1 – 1
Centruroides Marx 1 4 3 – – – 1 – 1
Isometrus Ehrenberg 1 – 1 – 1 – – – –
Mesotityus G.-Sponga – – 1 1 – 1 – – –
Microananteris Lour. – – – – – –
Microtityus K.-Waer. 1 2 – 9 – 1 – – –
Physoctonus M.-Leitão 1 – – – – – – – –
Rhopalurus Thorell 10 2 2 3 – 1 – – –
Tityus C.L. Koch 60 32 81 8 5 4 1 1 5
Troglorhopalurus Lour. et al. 1 – – – – – – – –
Zabius Thorell 1 – – – – – 1 – 1
Fam. Chactidae + + + + + + – + –
Auyantepuia G.-Sponga 3 – 1 – 1 5 – – –
Broteochactas Pocock 4 – 10 1 – 1 – – –
Brotheas C.L. Koch 10 1 15 2 3 1 – – –
Chactas Gervais 2 15 26 – – – – – –
Chactopsis Kraepelin 5 – 3 – – – – – –
Chactopsoides Ochoa et al. 1 – 3 – – – – – –
Guyanochactas Lour. 3 – – – – 1 – – –
Hadrurochactas Pocock 4 – 3 1 1 1 – – –
Megachactops Ochoa et al. – – 2 – – – – – –
Neochactas Sol. et Fet 6 1 27 1 1 6 – – –
Country Brazil Colom. Venezuela Guyana Suriname French Guiana Uruguay Paraguay
Teuthraustes Simon 4 2 5 – – – – – –
Vachoniochactas G.–Sponga 2 1 3 1 – – – – –
Fam. Hormuridae + + + – – – – – –
Opisthacanthus Peters 3 1 2 – – 1 – – –
Fam. Bothriuridae + – – – – – + + +
Bothriurus Peters 13 – – – – – 3 3 3
Brachistosternus Pocock 1 – – – – – 1 – 1
7.33 Tropical South America
Brazilobothrius Lour. et Monod 1 - – – – – – – –

Thestylus Simon 3 – – – – – – – –
Timogenes Simon – – – – – – 1 – 2
Urophonius Pocock 1 – – – – – – 1 –
Fam. Troglotayosicidae – + – – – – – – –
Troglotayosicus Lour. – 2 – – – – – – –
Fam. Scorpionidae – + + – – – – – –
Tarsoporosus Francke – 3 3 – – – – – –
Fam. Euscorpiidae (+) – – – – – – – (+)
Euscorpius Thorell (introduced) 1 - – – – – (+) – 1
Country Аrgentina Bolivia Chile Peru Еcuador (cont.)
Number of species 66 27 53 62 47
Тaxa
Fam. Buthidae + + + + +
Ananteris Thorell 1 3 – 1 3
Centruroides Marx 1 1 1 1 2
(continued)
485
486
Country Аrgentina Bolivia Chile Peru Еcuador (cont.)

Isometrus Ehrenb. 1 – – 1 –
Tityus C.L. Koch 8 9 1 12 16
Zabius Thorell 2 – – – –
Fam. Chactidae – + – + +
Brotheas C.L. Koch – 1 – – –
Chactas Gervais – – – 2 3
Chactopsis Kraepelin – – – 2 –
Teuthraustes Simon – – – – 12
Fam. Caraboctonidae – – + + +
Caraboctonus Pocock – – 1 1 –
Hadruroides Pocock – – – 16 6
Fam. Hormuridae – – – + –
Opisthacanthus Peters – – – 1 –
Fam. Troglotayosicidae – – – – +
Troglotayosicus Lour. – – – – 1
Fam. Atemnidae
Brazilatemnus Muchmore, 1975 – Brazil (one sp.)
Caecatemnus Mahnert, 1985 – Brazil (one sp.)
Fam. Chernetidae
Attaleachernes Mahnert, 2009 – Brazil (one sp.)
Atherochernes Beier, 1954 – Venezuela (one sp.)
Calidiochernes Beier, 1954 – Venezuela (one sp.)
Ceratochernes Mahnert, 1994 – Venezuela (two spp.)
Corosoma Karsch, 1879 – Brazil (one sp.)
Dasychernes J.C. Chamberlin, 1929 – Colombia, Panama (four spp.)
Spelaeochernes Mahnert, 2001 – Brazil (eight spp.)
Attaleachernes Mahnert, 2009 – Brazil (one sp.)
Fam. Withiidae
Tropidowithius Beier, 1955 – Peru (one sp.)
Opiliones South America is extremely rich in Opilions, mostly Laniatores (Acosta
2002, 2006, Benavides and Giribet 2007, 2013, DaSilva et al. 2010, Pinto-da-Rocha
et al. 2012, González-Sponga – 19 papers from 1981 to 2005; 1987, 1992, Goodnight
and Goodnight 1943, 1980, Hinton 1938, Kury 2003, Kury and Maury 1998, Kury
and Pérez 2002, Kury and Pinto-da-Rocha 2002, Martens 1969, 1988, Mello-Leitão
1932, 1935, 1941, 1945, Pinto-da-Rocha 1997, Rambla 1976, Ringuelet 1959,
Roewer 1913, 1932, 1943, 1949a, 1949b, 1949c, 1956, 1957, 1961, 1963, Rosas
Costa 1950, Shear 1979, 1993b, H. E. M. Soares and Avram 1981, H. E. M. Soares
1979, Soares B.A.M. and H.E.M. Soares 1948). According to the statistics of Kury
(2002 and onward), only in Brazil have been recorded 987 spp. of Opiliones (the
richest country in the world). In the Neotropical region are known 20 fam. of
Opilions, including 13 families endemic to the Neotropics (all Laniatores).
Cyphophthalmi Typical for tropical South America is the family Neogoveidae

(Colombia, Brazil, Ecuador, Guyana, French Guiana, Peru, Suriname, Trinidad and
Tobago, Venezuela), which has also one genus in tropical Africa (Parogovia Hansen,
1921) and to the north goes as far as Southern USA (Metasiro). Five genera in South
America:
Brasilogovea Martens, 1969 – Brazil, Colombia (two spp.) (endemic)

Canga DaSilva, Pinto-da-Rochaet Giribet, 2010 – Brazil (one sp., caves) (endemic)
Huitaka Shear, 1979 – Colombia (seven spp.) (+ two undescr.) (endemic to
Colombia)
Metagovea Rosas Costa, 1950 – Colombia, Ecuador, Brazil (Manaus) (four spp.)
(+17 undescr.) (endemic)
Neogovea Hinton, 1938 – Brazil, Guyana (five spp.) (+12 undescr.) (endemic)
One genus of the family Pettalidae (Chileogovea Roewer, 1961) is known from
Chile (two spp.); Ch. oedipus Roewer, 1961, from Concepción to Chiloé; and Ch.
jocasta Shear, 1993, from Malleco and Arauco Provinces.
Eupnoi
Fam. Phalangiidae (Dicranopalpinae – Dicranopalpus Doleschall, 1852) – Venezuela
Fam. Neopilionidae (Thrasychiroides Soares et Soares, 1947 – Brazil, one sp., the
only endemic genus among the Eupnoi of tropical South America)
Dyspnoi Fam. Acropsopilionidae (Acropsopilio Silvestri, 1904 – Venezuela)
Laniatores According to the catalogue of Kury (2003), in tropical South America

(south to 30oS), this suborder contains genera from the families Agoristenidae,
Cosmetidae, Cranaidae, Escadabiidae, Fissiphaliidae, Gonyleptidae, Guasiniidae,
Icaleptidae, Manaosbiidae, Kimulidae (Minuidae), Samoidae, Stygnidae,
Stygnommatidae, Stygnopsidae, and Zalmoxidae. The 13 underlined families are
entirely Neotropical. Many of the genera are monotypic and have been partly
revised and synonymized by Pinto-da-Rocha et al. (2012). According to Kury
(2003), up to 2002 in the New World have been recorded 2372 spp. of 746 genera
and 21 families. Of course, with the activities of the South American specialists in
the last 15 years, many of these figures are altered, the number of species is
increased, but still the most numerous families remain: Gonyleptidae (in 2002 823
spp.) and Cosmetidae (710 spp.). In South America N of 30oN, there are at least 470
endemic genera of Laniatores (including 226 genera of Gonyleptidae).
Endemic genera (from one or few countries, Argentina, Uruguay, and Chile
excluded) in the described area are:
Fam. Agoristenidae
Leiosteninae
Barinas González-Sponga, 1987; Barlovento González-Sponga, 1987; Ocoita
González-Sponga, 1987; Paravima Caporiacco, 1951; Vimina González-Sponga,
1987 – Venezuela
Leptostygnus Mello-Leitão, 1940 – Colombia, Venezuela
Sabanilla Roewer, 1913 – Colombia?, Venezuela
Vima Hirst, 1912 – Guyana
Zamorinae
Palcabius Roewer, 1956, Ramonus Roewer, 1956 – Peru
Zamora Roewer, 1927 – Ecuador
Fam. Cosmetidae
Acantholibitia Mello-Leitão, 1928; Elleriana Kury, 2003; Metavononoides
Roewer, 1927;
Bodunius Mello-Leitão, 1935; Fortalezius Roewer, 1947; Paragryne Roewer,
1912; Roquettea Mello-Leitão, 1931 – Brazil
Cocholla Roewer, 1927 – Brazil, Peru
Protus Simon, 1879 – Brazil, Ecuador

Neocynorta Roewer, 1915 – Brazil, Venezuela
Acrita Sorensen, 1932; Anduzeia González-Sponga, 1992; Baria González-
Sponga, 1993;
Caurimare González-Sponga, 1992; Corosalia González-Sponga, 1998;
Cynortetta Roewer, 1947;
Forfexia González-Sponga, 1992; Gnaricia González-Sponga, 1992; Gnatopia
González-Sponga, 1992; Litoralia González-Sponga, 1992; Oligovonones di
Caporiacco, 1951; Puerilia González-Sponga, 1992; Szczurehia González-
Sponga, 1992 – Venezuela
Ambatoiella Mello-Leitão, 1943; Cynortellina Roewer, 1915; Pygocynorta
Roewer, 1925;
Rhaucoides Roewer, 1912; Vononella Roewer, 1925; Sibambea Roewer,
1917 – Ecuador
Cynortellula Roewer, 1925 – Ecuador, Peru
Brachylibitia Mello-Leitão, 1941; Cumbalia Roewer, 1963; Cynortosoma
Roewer, 1947; Libitiella Roewer, 1947; Megarhaucus Mello-Leitão, 1941;
Messatana Strand, 1942; Neorhaucus Pic.-Cambridge, 1905; Zaraxes
Roewer, 1947 – Colombia
Caracarana Roewer, 1956; Chinchipea Roewer, 1952; Chirinosbius Roewer,
1952; Chusgonobius Roewer, 1952; Frizellia Mello-Leitão, 1941; Gnidiella
Roewer, 1957; Moselabius Roewer, 1956;
Opisthopristis Roewer, 1952; Pebasia Roewer, 1947; Praelibitia Roewer, 1956;
Tobotanus Roewer, 1957; Vononissus Roewer, 1956 – Peru
Eulibitia Roewer, 1912 – Ecuador, Colombia
Pelechucia Roewer, 1947; Syncynorta Roewer, 1947 – Bolivia
Ferkeria Roewer, 1947 – Bolivia, Peru
Maniapure González-Sponga, 1992 – Bolivia, Venezuela
Rhaucus Simon, 1879 – Colombia, Venezuela
Fam. Cranaidae
Cranainae
Acanthocranaus Roewer, 1913; Carsevennia Roewer, 1913 – French Guiana
Aguaytiella Goodnight et Goodnight, 1943; Aucayacuella S. Avram et H. E.
M. Soares, 1983;
Cenipa Goodnight et Goodnight, 1943 – Peru
Alausius Roewer, 1932; Angistrisoma Roewer, 1932; Angistrius Roewer, 1932;
Balzabamba Mello-Leitão, 1945; Baustomus Roewer, 1932; Bucayana
Mello-Leitão, 1942; Bunicranaus Roewer, 1913; Callcosma Roewer, 1932;
Cayabeus Roewer, 1932; Chetronus Roewer, 1932; Clinocippus Roewer,
1932; Diptyonius Roewer, 1932; Guayaquiliana Mello-Leitão, 1935;
Licornus Roewer, 1932; Puna Roewer, 1925; Spinicranaus Roewer, 1913;
Spirunius Roewer, 1932; Thaumatocranaus Roewer, 1932; Ventripila Roewer,
1917 – Ecuador
Allocranaus Roewer, 1915; Homocranaus Roewer, 1915; Megacranaus Roewer;

Metacranaus Roewer, 1913; Panalus Goodnight et Goodnight, 1947;
Paracranaus Roewer, 1913; Quindina Roewer, 1914; Sibundoxia Roewer,
1963; Tetracranaus Roewer, 1963; Ventrifurca Roewer, 1913; Ventrisudis
Roewer, 1963 – Colombia
Cranaostygnus Caporiacco, 1951; Stygnicranella Caporiacco, 1951; Timotesa
Roewer, 1943 – Venezuela
Deriacrus Roewer, 1932; Isocranaus Roewer, 1915; Microcranaus Roewer,
1913; Peripa Roewer, 1925 – Ecuador, Colombia
Digalistes Roewer, 1932; Idomenta Roewer, 1932; Ladantola Roewer, 1932;
Mecritta Roewer, 1932; Parkocranaus Mello-Leitão, 1949 – Brazil
Neocranaus Roewer, 1913 – Colombia, Venezuela
Tripilatus Roewer, 1932 – Bolivia
Heterocranainae – Ecuador, Colombia
Heterocranaus Roewer, 1913 – Ecuador, Colombia
Prostygninae
Binamballeus Roewer, 1952; Cutervolus Roewer, 1957 – Peru
Chiriboga Roewer, 1959; Globibunus Roewer, 1912; Lisarea Roewer, 1943;
Peladoius Roewer, 1919; Prostygnellus Roewer, 1919; Troya Roewer, 1919;
Yania Roewer, 1919 – Ecuador
Globitarsus Roewer, 1913; Micropachylus Roewer, 1913; Prostignidius Roewer,
1915; Prostygnus Roewer, 1913; Sclerostygnellus Roewer, 1943 – Colombia
Meridanatus Roewer, 1943 – Venezuela
Stygnicranainae – Colombia, Ecuador
Stygnicranaus Roewer, 1913 – Colombia
Tryferos Roewer, 1931 – Ecuador
Fam. Escadabiidae – Brazil (six spp., endemic family)
Baculigerus H. E. M. Soares, 1979; Escadabius Roewer, 1949; Jim H. E.
M. Soares, 1979; Recifesius H. E. M. Soares, 1978 – Brazil
Fam. Fissiphalliidae – Colombia (endemic family)
Fissiphallius Martens, 1988 – Colombia
Fam. Gonyleptidae
Ampycella Roewer, 1931; Hexabunus Roewer, 1913; Neopachyloides Roewer,
1913; Napostygnus Roewer, 1929 – Ecuador
Thaumatopachylus Roewer, 1929; Ampycus Simon, 1879; Asarcus C.L. Koch,
1839; Bogdana Mello-Leitão, 1940; Bourgnyia Mello-Leitão, 1923; Caldasius
Roewer, 1930; Cnemoleptes Mello-Leitão, 1941; Opisthoplites Sørensen,
1884; Styloleptes Piza, 1943; Stylopisthos Roewer, 1930; Ampheres
C.L. Koch, 1839; Arthrodes C.L. Koch, 1839; Caelopygus C.L. Koch, 1839;
Garatiba Mello-Leitão, 1940; Metampheres Roewer, 1913; Metarthrodes

Roewer, 1913; Pristocnemis C.L. Koch, 1839; Proampheres Roewer, 1913;
Thereza Roewer, 1943; Adhynastes Roewer, 1930; Clarinus Roewer, 1929;
Centroleptes Roewer, 1943; Collonidium Bertkau, 1880; Currala Roewer,
1927; Deltaspidium Roewer, 1927; Friburgoia Mello-Leitão, 1926;
Geraecormobius Holmberg, 1887; Gonazula Roewer, 1930; Gonyleptellus
Roewer, 1930; Gonyleptes Kirby, 1818; Guatubesia H. Soares; Gyndesops
Roewer, 1943; Holoversia Mello-Leitão, 1940; Inhuma Piza, 1938;
Liogonyloptoides Mello-Leitão, 1925; Megapachylus Roewer, 1913;
Metagonyleptes Roewer, 1913; Mischonyx Bertkau, 1880; Multumbo Roewer,
1927; Neosadocus Mello-Leitão, 1926; Piassagera Roewer, 1928;
Proctobunoides Mello-Leitão, 1944; Progonyleptoides Roewer, 1917;
Pseudotrogulus Roewer, 1932; Sphaerobunus Roewer, 1917; Stefanasia
Soares et Soares, 1988; Tupacarana Mello-Leitão, 1939; Uracantholeptes
Mello-Leitão, 1926; Urodiabunus Mello-Leitão, 1935; Aerogonyleptes
Roewer, 1917; Ariaeus Sørensen, 1932; Despirus Roewer, 1929;
Discocyrtoides Mello-Leitão, 1923; Encheiridium Kury, 2003; Ischnotherus
Kury, 1991; Longiperna Roewer, 1929; Metanitobates Roewer, 1913;
Mitobates Sundevall, 1833; Mitobatula Roewer, 1931; Neoancistrotus Mello-
Leitão, 1929; Promitobates Roewer, 1913; Ruschia
Mello-Leitão, 1940 – Brazil
Parampherus Roewer, 1913 – Bolivia, Brazil
Chacoikeontus Roewer, 1929; Metasarcus Roewer, 1913; Nemastygnus Roewer,
1929 – Bolivia
Sibillus Roewer, 1929; Oxapampeus Roewer, 1963; Ayacucho Roewer, 1949;
Cajacayba Roewer, 1957; Cajamarca Roewer, 1952; Cargaruaya Roewer,
1956; Incasarcus Kury et Maury, 1998; Palcares Roewer; Tapacochana
Roewer, 1957; Taurisa Roewer, 1956; Tschaidicancha Roewer, 1957 – Peru
Heteropachylinae – endemic subfamily for Brazil
Aesotrinoma H. Soares, 1977; Chavesincola Soares et Soares, 1946;
Heteropachylus Roewer, 1913; Mangaratiba Mello-Leitão, 1940; Melloa
Roewer, 1930; Pseudopulcrolia Roewer, 1912; Thaumatoleptes Roewer,
1930; Tribunosoma Roewer, 1943 – Brazil
Cobaniinae – endemic subfamily for Brazil
Cobania Roewer, 1913 – Brazil
Gonyassamiinae – endemic subfamily for Brazil
Acutisoma Roewer, 1913; Goniosoma Perty, 1833; Goniosomoides Mello-
Leitão, 1932; Lyogoniosoma Mello-Leitão, 1926; Metalyogoniosoma Soares
et Soares, 1946; Phalangochilus Mello-Leitão, 1938; Trichominua Mello-
Leitão, 1938 – Brazil
Pachylinae
Acanthopachylopsis Soares et Soares, 1949; Acrographinotus Roewer, 1929;
Allogonyleptes Roewer, 1917; Anoplogynopsis H. Soares, 1966; Antetriceras
Roewer, 1949; Beckeresia H. Soares, 1970; Berlaia Mello-Leitão, 1940;
Biconisoma Roewer, 1936; Bristoweia Mello-Leitão, 1924; Bunoplus Roewer,
1927; Caldanatus Roewer, 1943; Camposicola Mello-Leitão, 1924;
Camposicoloides B. Soares, 1944; Capichabesia B. Soares, 1944; Carlotta
Roewer, 1943; Ceratoleptes Soares et Soares, 1979; Discocyrtulus Roewer, 1927;
Discocyrtus Holmberg, 1878; Eopachyloides H. Soares, 1970; Eopachylus Mello-
Leitão, 1931; Ergastria Mello-Leitão, 1941; Eugyndes Roewer, 1923;
Eugyndopsiella H. Soares, 1972; Goodnightiella Soares et Soares, 1945;
Graphinotus C.L. Koch, 1839; Guaraniticus Mello-Leitão, 1933; Gyndesoides
Mello-Leitão, 1933; Gyndoides Mello-Leitão, 1927; Harpachylus Roewer, 1943;
Huralvioides H. Soares, 1970; Hypophyllonomus Giltay, 1928; Iandumoema
Pinto-da-Rocha, 1996; Iguassua Mello-Leitão, 1935; Iguassuoides Soares et
Soares, 1954; Itatiaincola Soares et Soares, 1948; Izecksohnopilio H. Soares,
1977; Juticus Roewer, 1943; Lacronia Strand, 1942; Maromba Soares et Soares,
1954; Metadiscocyrtus Roewer, 1929; Metagraphinotus Mello-Leitão, 1927;
Metagyndoides Mello- Leitão, 1931; Metalycomedes Mello-Leitão, 1927;
Metapachyloides Roewer, 1917; Meteusarcoides Mello-Leitão, 1922;
Meteusarcus Roewer, 1913; Neopachylus Roewer, 1913; Pachylibunus Roewer,
1913; Pachylusius Mello-Leitão, 1949; Paradiscocyrtus Mello-Leitão, 1927;
Paraluederwaldtia Mello-Leitão, 1927; Paranaleptes Soares et Soares, 1947;
Paraprosontes Soares et Soares, 1947; Pareusarcus Roewer, 1929; Passosa
Roewer, 1927; Pherania Strand, 1942; Planiphalangodus Roewer, 1929;
Pseudoacrographinotus H. Soares, 1966; Pseudogyndesoides B. Soares, 1944;
Rhioxyna Soares et Bauab-Vianna, 1970; Roeweria Mello-Leitão, 1943;
Schubartesia B. Soares, 1944; Singram Mello-Leitão, 1937; Triglochinura Mello-
Leitão, 1924; Ubatubesia B. Soares, 1945; Uropachylus Mello-Leitão, 1922;
Victoriaincola Soares et Soares, 1946; Yraguara Mello-Leitão, 1937 – Brazil
Paraphalangodus Roewer, 1915 – Colombia
Bullaepus Roewer, 1930; Gerdesius Roewer, 1952; Huadquina Roewer, 1930;
Huasampillia Roewer, 1913; Hyperpachylus Roewer, 1957; Junicus Goodnight
et Goodnight, 1947; Palcapachylus Roewer, 1952; Pichitus Roewer, 1959;
Pirunipygus Roewer, 1936; Platygyndes Roewer, 1943; Polyacanthoprocta
Mello-Leitão, 1927; Punagraphinotus Soares et Bauab-Vianna, 1972;
Tarmapachylus Roewer, 1956; Tingomaria Mello-Leitão, 1948 – Peru
Punrunata Roewer, 1952 – Brazil, Peru
Acrographinotus Roewer, 1929 – Peru, Bolivia
Unduavius Roewer, 1929 – Bolivia
Ceropachylinus Mello-Leitão, 1943; Ceropachylus Mello-Leitão, 1942; Ibarra
Roewer, 1925 – Ecuador
Pachylospeleinae – endemic subfamily for Brazil
Pachylospeleus Šilhavý, 1974 – Brazil
Progonyleptoidellinae
Cadeadoius Mello-Leitão, 1936; Detigalus Roewer, 1931; Gonyleptoides
Roewer, 1913; Heliella B. Soares, 1945; Iguapeia Mello-Leitão, 1935;
Iporangia Mello-Leitão, 1935; Leptocnema C.L. Koch, 1839; Mitopernoides
B. Soares, 1945; Moreiranula Roewer, 1930; Progonyleptoidellus Piza,
1940 – Brazil
Sodreaninae – endemic subfamily for Brazil
Gertia Soares et Soares, 1946; Sodreana Mello-Leitão, 1922; Stygnobates
Mello-Leitão, 1927; Zortalia Mello-Leitão, 1942 – Brazil
Tricommatinae
Arminda Roewer, 1949; Arruda Mello-Leitão, 1940; Bebedoura Roewer, 1949;
Bissulla Roewer, 1929; Bresslauius Mello-Leitão, 1935; Bunostigma Mello-
Leitão, 1935; Camarana Mello-Leitão, 1935; Cryptogeobius Mello-Leitão,
1942; Heteromeloleptes Mello-Leitão, 1931; Lanesoares Roewer, 1949; Liops
Mello-Leitão, 1940; Lyssanvira Mello-Leitão, 1935; Pararezendesius H. Soares,
1972; Poecilosophus Mello-Leitão, 1948; Pseudopachylus Roewer, 1912;
Pseudophalangodes Roewer, 1912; Rezendezius H. Soares, 1945; Segundolus
Roewer, 1949; Spinopilar Mello-Leitão, 1940; Tachusina Strand, 1942;
Tibangara Mello-Leitão, 1940; Tricommatus Roewer, 1912; Zalanodicus Mello-
Leitão, 1936 – Brazil
Otuquisa Roewer, 1927 – Bolivia
Paratricommatus Piza, 1943 – Brazil, Paraguay
Fam. Guasiniidae – Venezuela, Brazil
Guaiquinimia González-Sponga, 1997 – Venezuela
Guasinia González-Sponga, 1997 – Venezuela, Brazil
Fam. Icaleptidae – Colombia, Ecuador
Icaleptes Kury et Pérez, 2002 – Colombia
Zalmopsylla Kury et Pérez, 2002 – Ecuador
Fam. Manaosbiidae
Azulamus Roewer, 1957 – Peru
Belemnodes Strand, 1942; Belemulus Roewer, 1932; Manaosbia Roewer, 1943;
Paramicrocranaus H. Soares, 1970; Syncranaus Roewer, 1913 – Brazil
Camelianus Roewer, 1912; Gonogotus Roewer, 1943; Cucutacola Mello-Leitão,
1940; Semostrus Roewer, 1943 – Colombia
Clavicranaus Roewer, 1915; Saramacia Roewer, 1913 – Suriname, Brazil
Dibunostra Roewer, 1943; Poecilocranaus Roewer, 1943 – Venezuela
Mazarunius Roewer, 1943 – Guyana
Meridia Roewer, 1913 – Venezuela, Suriname
Pentacranaus Roewer, 1963; Tegyra Sørensen, 1932 – Peru
Rhopalocranellus Roewer, 1925 – Ecuador
unknown genus calcar (Roewer, 1943) – Venezuela
unknown genus albituberculatus (Roewer, 1943) – Guyana

unknown genus strinatii (Šilhavý, 1979) – Venezuela
Fam. Kimulidae (= Minuidae)
Acanthominua Sørensen, 1932; Euminua Sørensen, 1932; Euminuoides Mello-
Leitão, 1935; Fudeci González-Sponga, 1997; Microminua Sørensen, 1932;
Minua Sørensen, 1932 – Venezuela
Tegipiolus Roewer, 1949 – Brazil
Pseudominua Mello-Leitão, 1935 – Venezuela, Peru
Fam. Samoidae
Cornigera González-Sponga, 1987 – Venezuela
Fam. Stygnidae
Heterostygninae
Eutimesius Roewer, 1913 – Colombia, Venezuela, Ecuador
Innoxius Pinto-da-Rocha, 1997; Minax Pinto-da-Rocha, 1997; Stenostygnellus
Roewer, 1913; Yapacana Pinto-da-Rocha, 1997 – Venezuela
Stygnidius Simon, 1879 – Brazil, French Guiana
Timesius Simon, 1879 – Colombia
Nomoclastinae – Colombia (endemic subfamily)
Nomoclastes Sørensen, 1932 – Colombia
Stygninae
Actinostygnoides Goodnight et Goodnight; Auranus Mello-Leitão, 1941
Iguarassua Roewer, 1943; Kaapora Pinto-da-Rocha, 1997; Paraphareus Goodnight
et Goodnight, 1943; Pickeliana Mello-Leitão, 1932; Planophareus Goodnight et
Goodnight, 1943; Sickesia H.E.M. Soares, 1979; Stenophareus Goodnight et
Goodnight, 1943; Verrucastygnus Pinto-da-Rocha, 1997 – Brazil
Metaphareus Roewer, 1912; Phareus Simon, 1879 – Colombia, Venezuela
Niceforoiellus Mello-Leitão, 1941 – Colombia
Ortonia Wood, 1869 – Ecuador
Protimesius Roewer, 1913 – Peru, Brazil, Ecuador, French Guiana, Suriname
Stenostygnoides Roewer, 1913 – Suriname
Stygnus Perty, 1833 – Venezuela, Brazil, Peru, Colombia, Suriname
Subfam. uncertain – Brazil
Gaibulus Roewer, 1943 – Brazil
Fam. Zalmoxidae
Absonus González-Sponga, 1987; Araguita González-Sponga, 1987;
Avilaia González-Sponga, 1998; Azulitaia González-Sponga, 1987; Buruquelia
González-Sponga, 1999; Cea González-Sponga, 1987; Chamaia González-
Sponga, 1987; Cubiria González-Sponga, 1987; Curimaguanus González-
Sponga, in Kury 2003; Granulaia González-Sponga, 1997; Guagonia
González-Sponga, 1987; Haitonia González-Sponga, 1987; Jajinia González-

Sponga, 1987; Lara M. A. González-Sponga, 1987; Micro González-Sponga,
1987; Orituco González-Sponga, 1987; Panaquire M. A. González-Sponga,
1987; Paraminuella Caporiacco, 1951; Paramo González-Sponga, 1987;
Phalangodinella Caporiacco, 1951; Retropedis González-Sponga, 1987;
Soledadiella González-Sponga, 1987; Sphoeroforma González-Sponga,
1987; Spiniella González-Sponga, 1987; Taguaza González-Sponga, 1998;
Tiara González-Sponga, 1987; Traiania H. E. M. Soares et S. Avram, 1981;
Unare González-Sponga, 1987; Unicornia González-Sponga, 1987;
Urachiche González-Sponga, 1987; Yacambuia González-Sponga, 1987;
Carayaca González-Sponga, 1998; Chirimena González-Sponga, 1999;
Guayania González-Sponga, 1999; Junquito González-Sponga, 1999;
Niquitaia González-Sponga, 1999; Ovalia González-Sponga, 1999; Pijiguaia
González-Sponga, 1998; Pilosa González-Sponga, 1999 – Venezuela
Cochirapha Roewer, 1949; Exlineia Mello-Leitão, 1942; Galanomma Juberthie,
1970 – Ecuador
Garanhunsa Roewer, 1949; Tegipiolus Roewer, 1949 – Brazil
Viacha Roewer, 1949 – Bolivia
Weyrauchiana Roewer, 1952; Sivianus Roewer, 1949 – Peru
Pirassunungoleptes H. Soares, 1966 – Brazil, Bolivia
Amblypygi From South America are recorded ca. 60 spp. of Amblypygi, belong-
ing to the genera Charinus Simon, Trichodamon Mello-Leitão, Paraphrynus
Moreno, Heterophrynus Pocock, and Phrynus Lamarck and the families Charinidae,
Phrynichidae, and Phrynidae (Pocock 1894, Baptista and Giupponi 2002ª, 2002b,
2003, Caporiacco 1947, Giupponi and Kury 2002, 2013, Gonzalez-Sponga 1998,
Goodnight and Goodnight 1946, Harvey 2013c, Jocqué and Giupponi 2012, Mello-
Leitão 1931, Pinto-da-Rocha et al. 2002, Miranda and Giupponi, 2002, Ravelo
1975a, 1975b, Quintero 1981, 1983, Vasconcelos et al. 2013, Weygoldt 1972a,
1972b).
The genus Charinus is the most widespread, including on islands like Galapagos.
From continental South America (Guyana, Brazil, Suriname, Venezuela, Peru) are
known more than 20 species, including 12 from Brazil.
Trichodamon Mello-Leitão (two spp. from Brazil) is the only genus in the family
Phrynichidae living in the Western Hemisphere (the other six genera are repre-
sented in Africa and the outlying islands). This is the only one endemic ambly-
pygid genus in South America.
Paraphrynus Moreno (18 spp. from Colombia to the Southern USA) is typical
mostly for Central America. In South America has been registered one sp.
(Colombia).
Phrynus Lamarck (five spp. in continental South America – Colombia, Ecuador,
Guyana, Venezuela, Brazil) is widespread from Brazil to the southern states of
USA (Texas, Florida), mostly in Mexico, Central America, and the Antilles.
Recently an aberrant species was described from a cave on the Flores Island in
Indonesia (Phrynus exsul Harvey, 2002), the only representative of the family
Phrynidae in the Old World.
Heterophrynus Pocock, 1894 (the only genus in the subfamily Heterophryninae) –
[Mexico] Brazil, Peru, Colombia, French Guiana, Guyana, Venezuela, Ecuador,
Suriname (14 spp.)
Map 7.25 Amblypygi in South America

Charinus – ■
Heterophrynus – ◄
Trichodamon – ●
Paraphrynus – ▼
Phrynus – ▲
Thelyphonida (Uropygi) The whip scorpions in tropical South America belong to

the subfamily Hypoctoninae and to the genera Thelyphonellus Pocock, 1894 (Brazil,
Guyana, Suriname, Venezuela, two spp.) and Mastigoproctus Pocock, 1894 (=
Amauromastigon Mello-Leitão, 1931) – widespread and reaches the Antilles,
Central America, Mexico, and the Southern USA (Haupt 2009b, Mello-Leitão
1931, Pocock 1894, Rowland 2002, Villarreal Manzanilla and Giupponi 2009).
In the subfamily Hypoctoninae, only the genus Thelyphonellus is South
American; Ravilops lives in the Dominican Republic, Valeriophonus in Costa Rica,
and the other three genera in West Africa (Etienneus), India, and Sri Lanka
(Labochirus) or Southeast Asia (Hypoctonus). In the subfamily Uroproctinae, two
monotypic genera live in Asia; all the other species belong to Mastigoproctus and
are widespread. One enigmatic species was described from Hong Kong
(Mastigoproctus transoceanicus Lazell, 2000) but is clearly introduced.
Thelyphonellus, Valeriophonus, Ravilops, and Mastigoproctus could be termed as
Neotropical.
Map 7.26 Thelyphonida in South America, Central America, and the Caribbean
Thelyphonellus – ■
Mastigoproctus – ▼
Ravilops – ●
Mayacentrum –
Valeriophonus – ▲
Mimoscorpius– ◄
Schizomida Armas (2010) made a review of Schizomida known from South

America. With some supplements, there are species of the genera Adisomus, Calima,
Surazomus, Rowlandius, Stenochrus, Tayos, Hansenochrus, Stenoschizomus, and
Wayuuzomus in seven countries: Colombia, Venezuela, Ecuador, Peru, Brazil,
Guiana, and Suriname. The southernmost finding of Schizomida on the continent is
Stenochrus portoricensis Chamberlin in Rio de Janeiro (probably introduced).
Otherwise the order does not go more southern than the Amazonia – Armas et al.
(2009), Bonaldo and Pinto-da-Rocha (2007), Giupponi et al. (2016), Kraus (1957),
Kraus and Beck (1967), Pinto-da-Rocha (1996, 2007), Reddell and Cokendolpher
(1995), Rowland and Reddell (1979), Villarreal Manzanilla et al. (2008), Moreno-
González and Manzanilla (2012), Santos and Pinto-da-Rocha (2009), Santos et al.
(2013), Moreno-González et al. (2014) (Table 7.9).
Endemic genera of Schizomida for South America are:
Fam. Hubbardiidae
Adisomus Cokendolpher et Reddell, 2000 – Brazil (one sp.)
Calima Moreno-González et Manzanilla, 2012 – Colombia (two spp.)
Colombiazomus Armas et Delgado-Santa, 2012 – Colombia (one sp.)
Stenoschizomus Gonzalez-Sponga, 1997 – Venezuela (one sp.)
Tayos Reddell et Cokendolpher, 1995 – Ecuador (one sp.)
Wayuuzomus Armas et Colmenares, 2006 – Venezuela (one sp.)
Table 7.9 Distribution of Schizomida in South America

Country Brazil Colombia Venezuela Guiana Suriname Bolivia Ecuador Peru
Taxa 11 5 8 2 3 1 3 1
Fam. + + + + + + + +
Hubbardiidae
Adisomus Cok. et 1 – – – – – – –
Reddell
Calima – 2 – – – – – –
Mor.-Gon. et
Manz.
Colombiazomus – 2 – – – – – –
Armas et Del.
Hansenochrus – – 1 2 – – – –
Red.et Cok.
Piaroa Manz. – 1 1 – – – – –
et al.
Rowlandius Red. 2 – – – – – –
et Cok.
Stenochrus 1 1 1 – – – – –
Chamberlin
Stenoschizomus – – 1 – – – – –
Gonz.-Sponga
Surazomus Red. 7 3 – – – 1 1 1
et Cok.
Tayos Red. et – – – – – – 1 –
Cokendolpher
Wayuuzomus – – 1 – – – – –
Armas et Cok.
Map 7.27 Distribution of Schizomida in South America

■ – Colombiazomus
X – Piaroa
● – Stenochrus
▼ – Surazomus
▲ – Wayuuzomus
◊ – Hansenochrus
□ – Rowlandius
► – Adisomus
▽– Stenoschizomus
◄ – Calima
– Tayos
Araneae From tropical South America (without the southernmost end) are recorded
spiders of at least 70 families. In the south (Patagonia, Chile) occur 7 more families,
or spiders of 77 families are known south of Panama (72 % of all spider families).
In the entire territory of Chile are recorded 55 families (Aguilera and Casanueva
2005). No family is endemic for the South American continent. This low level of
endemism is remarkable for the Neotropic (Huber 2000). On the small islands
Trinidad and Tobago have been recorded 51 families of spiders (plus 12 presumed
by Sewlal and Cutler 2003).
Endemic suprageneric taxa of Laniatores for the Neotropical Kingdom are:
Opiliones, Laniatores
Fam. Stygnidae – tropical South America, Lesser Antilles
Subfam. Nomoclastinae – Colombia
Fam. Agoristenidae – South America and the Caribbean
Fam. Cosmetidae – South and Central America and Mexico
Fam. Cranaidae – South and Central America and the Caribbean
Fam. Escadabiidae – Brazil
Fam. Fissiphaliidae – Colombia, Brazil
Fam. Gonyleptidae – from Patagonia to Guatemala
Fam. Guasiniidae – Brazil, Venezuela
Fam. Icaleptidae – Colombia, Ecuador
Fam. Manaosbiidae – tropical South and Central America, West Indies
Fam. Kimulidae (Minuidae) – tropical South America, West Indies
The wealth of Opiliones and Laniatores in the Neotropics is amazing and much
bigger than in any other kingdom. We have also to take into account that large areas
in South America are still almost unexplored (Tourinho and Pérez González 2006).
It is also to notice that there is no such situation in any other order in South America.
Opilioacarida From tropical South America was described Neocarus ojastii
Lehtinen, 1980 (Venezuela). N. platensis (Silvestri) is recorded also from Brazil.
Acariformes
Fam. Erythraeidae: Iguatonia Haitlinger, 2004 – Brazil (larval)
Parasitiformes (Mesostigmata and Ixodida) The very peculiar family of
Spelaeorhynchidae is confined to the New World (six spp., parasites on bats of the
families Phyllostomatidae and Mormoopidae). These mites have been found so far
in Belize, Brazil, Colombia, Cuba, Dominican Republic, Guatemala, Jamaica,
Panama, Peru, Puerto Rico, and Venezuela (Beron 2014).
Holothyrida From South America have been recorded only two species of
Holothyrida, belonging to two endemic genera of Neothyridae – one from Brazilian
Amazonas (Diplothyrus schubarti Lehtinen, 1999) and the other from the region of
Iquitos (Neothyrus ana Lehtinen, 1999), again from Amazonia, but in Peru.
Neothyrus sp. is known from Venezuela (Beron 2014).
7.34 Galapagos
7.34.1 G
and Paleogeography
The Galapagos Islands are an archipelago in the Pacific Ocean, part of Ecuador,
about 966 km far from the continent. The total area of the archipelago is 7845 km2,
the 13 main islands being Isabela (5827 km2), Fernandina (642 km2), Santa Cruz
(986 km2), Floreana (173 km2), Santiago (585 km2), Española (Hood) (60 km2), San
Cristóbal (Chatham) (558 km2), and some smaller islands. There are also six smaller
islands and 107 rocks. The highest point is Volcan Wolf in Isabela (1707 m).
The archipelago is situated on Nazca Plate which is moving with a speed of 6.4
cm/year and is being subducted under the South American Plate. Underneath is
formed the so-called Galapagos hot spot, where the core is melting and forming
volcanoes. The first islands are formed at least 8 million years ago and had no con-
nection with the mainland. At present the oldest (and most southern) island is
Fernandina, 3.5 million years ago. The colonization of the islands could have com-
menced earlier on lands, having been dry land 14 Ìa, and now are under water. The
line divergence of the terrestrial and the sea iguanas occurred 10 Ìa ago, so it hap-
pened on the older, now submerged islands (Chubb 1933, Colantoni et al. 1979,
Kuschel 1961, Lanteri 2001, Parent et al. 2008, Simkin 1984, Snell et al. 1995).
The volcanic oceanic archipelago has 749 species of higher plants, including 216
endemic. There are two species of bats, and it seems a good question why more bats
from the rich South American fauna did not fly over that distance of 966 km. On the
archipelago have radiated also 10 spp. of rice rats (Oryzomys, Nesoryzomys,
Megaoryzomys), but now only three remained. There are no amphibians, but there
are 37 species of reptiles (three snakes, geckos, tortoises, terrestrial and marine
iguanas). From (only) 30 species of birds, 25 are endemic, including the 14 spp. of
the famous Darwin’s finches (Geospizinae). About 1500 species of 1000 genera of
native insects form the entomofauna of the islands. Among the other invertebrate,
very impressive is the number of oribatid mites (202 spp., according to Schatz
1989).
Morrone (1999, 2001) includes the Galapagos as a province in the Caribbean
Subregion of the Neotropical Region.
The interesting cave fauna, found on Galapagos (Peck and Kukalova-Peck 1986,
Peck and Finstone 1993), contributes to the formulating of new ideas and biocli-
matic model of the troglomorphosis.
7.34 Galapagos 503
On the archipelago have been recorded representatives of Solifugae, Scorpiones,

Pseudoscorpiones, Opiliones (Laniatores), Amblypygi, Schizomida, Araneae,
Acariformes, and Parasitiformes (Baert 2011, 2014, Baert et al. 1995, Coolidge
1909, Roth and Craig 1970).
After Baert (version 2014), on Galapagos are recorded 434 species of Arachnida.
Schatz (1998) analyzed in details the possible ways of dispersal, colonization,
and speciation of the Oribatida on these islands. The main way is the hydrochorous
(directly and on/in floating logs or plant debris). Other ways are zoochory (birds)
and anthropochory. Some of the other Arachnida could have followed the same pat-
tern of dispersal; for some others different explanation is required. Peck and
Finstone (1993) summarized the amazing cave fauna of Galapagos and proposed an
explanation of its origin by parapatric speciation.
Solifugae Endemic genus and species is Neocleobis solitarius Roewer, 1934
(Ammotrechidae) (Baert et al. 1995, Banks 1902, Roewer 1934).
Scorpiones From Galapagos have been registered the species Centruroides exsul
Meise, 1933, and Hadruroides galapagoensis Maury, 1974 (Baert et al. 1995, Banks
1902, Kinzelbach 1973, Maury 1974).
Pseudoscorpiones From Galapagos are known 25 spp. of 9 genera

(Tyrannochthonius, Lechytia, Apolpium, Stenolpium, Serianus, Galapagodinus,
Ideobisium, Parachernes, Rhopalochernes) and 10 families (Chthoniidae,
Lechytiidae, Syarinidae, Olpiidae, Garypinidae, Garypidae, Cheiridiidae,
Atemnidae, Chernetidae, Withiidae) (Baert et al. 1995, Beier 1940, 1976d, 1978,
Mahnert 2014).
Endemic genus for Galapagos:
Fam. Garypinidae: Galapagodinus Beier, 1978
The species are almost all endemic.
Opiliones
Cyphophthalmi, Eupnoi, and Dyspnoi – not recorded
Laniatores (Baert et al. 1995, Juberthie 1970c)
Endemic genus and species is Galanomma microphthalma Juberthie, 1970
(Gonyleptidae).
Amblypygi From Galapagos is described only the endemic species Charinus
insularis Banks, 1902 (Charinidae). The genus Charinus is widespread.
Schizomida One widespread sp., Stenochrus portoricensis (Chamberlin, 1922),

known from Central America and West Indies (Baert et al. 1995)
Araneae According to Baert (2011, 2014), on Galapagos so far have been recorded
149 spp. of spiders. “Of the 124 identified species, 64 (almost 50%) are known only
from the archipelago and hence may be endemic, 35 species have a New World
distribution, 16 are cosmopolitan species, six pantropical, and three are cosmotropi-
cal species” (Baert, loc. cit.; Baert 1987, 1990, 2013, Baert and Maelfait 1986,
1997, Baert et al. 1989, 2008a, 2008b, Banks 1902, Levi 2009, Peck and Shear
1987a, b). A blind Lygromma (Gnaphosidae) and a new eyeless stridulating
Theridion have been described from the lava tubes.
Endemic genera:
Fam. Mysmenidae: Calomyspoena Baert et Maelfait, 1983
Fam. Pholcidae: Galapa Huber, 2000
Fam. Araneidae: Galaporella Levi, 2009 (one sp.)
Fam. Salticidae: Darwinneon Cutler, 1971
Acariformes
Sarcoptiformes
Oribatida According to Schatz (1998), on the archipelago have been registered
202 spp. of oribatid mites belonging to 64 families (during the 10-year study, 81
new species have been described).
The species richness of an island depends on the altitude and number of available habitats
rather than the area of the island. Many oribatid species on the Galápagos Islands have a
wide biogeographical distribution. The majority originate from the Central and South
American mainland, but several Pacific and even Holarctic elements were also found. In
comparison with the species composition of the adjacent mainland, the oribatid mite fauna
of the Galápagos Islands can be regarded as disharmonic. Sea surface transportation has
been proved at least between the islands, which also applies to oribatid species living at
higher elevations. Long distance dispersal can be assumed to be mainly hydrochorous.
Approximately 40% of all oribatid species from the Galápagos Islands have a known distri-
bution restricted to the islands and can be considered to be endemic. (Schatz 1998)
Endemic genus and species is Galapagacarus schatzi Balogh, 1985.

Parasitiformes
Ixodida
Fam. Ixodidae – several species (endemic) are described from lizards and
tortoises.
Ixodes galapagoensis Clifford et Hoogstraal, 1980; Amblyomma boulengeri
Hirst et Hirst, 1910; A. darwini Hirst et Hirst, 1910; A. usingeri Keirans,
Hoogstraal, et Clifford, 1973; A. williamsi Banks, 1924
7.35 Juan Fernandez Islands 505
7.35 Juan Fernandez Islands
7.35.1 G
and Paleogeography
The volcanic archipelago Juan Fernandez, 674 km west of South America, has an
area of 99.6 km2, of which 47.9 km2 is of Robinson Crusoe Island, or Isla Más a
Tierra (together with Santa Clara) and 49.5 km2 of Alejandro Selkirk Island (Isla
Más Afuera). The two islands are 181 km far from each other. The altitudes are El
Yunque (916 m on Robinson Crusoe) and Cerro de Los Inocentes (1319 m on
Alejandro Selkirk). The islot Santa Clara (2.2 km2) has 350 m height and is the old-
est island of the archipelago (5.8 million years old), next is Robinson Crusoe (3.8–
4.2 million years old) and Alexander Selkirk (1.0–2.4 million years old) (Kuschel
1961, Scottsberg 1954). The climate of the islands is Mediterranean.
According to UNESCO World Netwrok of Biosphere Reserves, since 1977 these
islands have been considered of scientific importance as an ecoregion, because of
the endemic species of flora and fauna There are 126 endemic species of plants
(62%), with 12 endemic genera and 1 endemic family, Lactoridaceae.
According to Morrone (2000), biogeographically Juan Fernandez Islands are a
province within the Subantarctic Subregion of the Andean region.
From the islands are known representatives (with low-level endemism) of the orders
Pseudoscorpiones (15 spp., almost all endemic, with 2 endemic genera), Opiliones,
and Araneae (2 endemic genera) and some mites.
Pseudoscorpiones The pseudoscorpions recorded on Juan Fernandez Islands
belong to the families Lechytiidae, Geogarypidae, Cheiridiidae, Chernetidae, and
Withiidae; the genera Lechytia, Geogarypus, Neocheiridium, Parazaona,
Asterochernes, Chelanops, Pseudopilanus, Asterochernes, Selachochernes, and
Protowithius; and 15 species, 14 of which endemic and 1 known from Argentina
(Beier 1955, 1957b, Mahnert 1993, 1997, 2011).
Endemic genera:
Fam. Chernetidae: Selachochernes Mahnert, 2011 (one sp.)
Fam. Withiidae: Protowithius Beier, 1955 (two spp.).
Opiliones
Eupnoi
Fam. Acropsopilionidae
Acropsopilio chilensis Silvestri, 1904 – the first Opilion discovered on the archi-
pelago, known also from Chile, including Tierra del Fuego (Pérez-González
et al. 2014)
Araneae Bäckstrom from the expedition of Skottsberg in 1916–1917 has collected
spiders, which were studied by Berland (1924 and 1935). The list of Berland, 1924b
(including the spiders recorded by F.O.P. –Cambridge and E. Simon), contains 24
spp., among them are 9 new spp. and the new genus Selkirkiella (Theridiidae).
Endemic genera are:
Fam. Linyphiidae
Schistogyna Millidge, 1991 (one sp.)
Malkinola Miller, 2007
Juanfernandezia Koçak et Kemal, 2008
7.36 North America, North of Rio Grande (incl. Greenland)
7.36.1 G
and Paleogeography
North America occupies the northern portion of the landmass generally referred to
as the New World. North America’s only land connection to South America is at the
Isthmus of Panama. The continent is delimited on the southeast by most geogra-
phers at the Darién watershed along the Colombia-Panama border, placing all of
Panama within North America. The Caribbean Islands, or West Indies, are consid-
ered part of North America.
Before the Central American isthmus was raised, the region had been underwa-
ter. The islands of the West Indies delineate a submerged former land bridge, which
had connected North America and South America via what are now Florida and
Venezuela (Kaiser et al.1972 Martin and Harrell 1957, Rogers et al. 1999, Williams
et al. 2000).
Greenland, a self-governing Danish island, and the world’s largest, is on the
same tectonic plate (the North American Plate) and is part of North America geo-
graphically. In a geologic sense, Bermuda is not part of the Americas, but an oceanic
island which was formed on the fissure of the Mid-Atlantic Ridge over 100 million
years ago.
The vast majority of North America is on the North American Plate. Parts of
western Mexico, including Baja California, and of California, including the cities of
San Diego, Los Angeles and part of San Francisco, lie on the eastern edge of the
Pacific Plate, with the two plates meeting along the San Andreas fault. The south-
ernmost portion of the continent and much of the West Indies lie on the Caribbean
Plate.
7.36 North America, North of Rio Grande (incl. Greenland) 507
The western mountains are split in the middle into the main range of the Rockies
and the coast ranges in California, Oregon, Washington, and British Columbia, with
the Great Basin – a lower area containing smaller ranges and low-lying deserts – in
between. The highest peak is Denali in Alaska (6168 m).
North America covers an area of about 24,709,000 square kilometers, about
4.8% of the planet’s surface or about 16.5% of its land area.
Laurentia is an ancient craton which forms the geologic core of North America;
it formed between 1.5 to 1.0 billion years ago during the Proterozoic eon. From the
Late Paleozoic to Early Mesozoic eras, North America was joined with the other
modern-day continents as part of the supercontinent Pangaea. One of the results of
the formation of Pangaea was the Appalachian Mountains, which formed some 480
million years ago, making it among the oldest mountain ranges in the world. When
Pangaea began to rift around 200 million years ago, North America became part of
Laurasia, before it separated from Eurasia as its own continent during the mid-
Cretaceous period. The Rockies and other western mountain ranges began forming
around this time from a period of mountain building called the Laramide orogeny,
between 80 and 55 million years ago. The formation of the Isthmus of Panama con-
nected the continent to South America about three million years ago.
In North America, north of Mexico, are found 13 orders of Arachnida, but the
“southern” orders (Amblypygi, Ricinulei, Schizomida, Uropygi, Opilioacarida, and
to big extent Solifugae and Scorpiones) are confined mostly to Texas, Arizona,
California, Florida, Louisiana, and New Mexico, some of them being on the edge of
their areals (Dice 1943, Harvey, online, Rowland and Reddell 1976, Sørensen
1898).
In Canada the southern groups are either lacking (Ricinulei, Schizomida) or
present with one cold resisting species (Scorpiones). One family of spiders
(Trogloraptoridae) is endemic to the caves of the USA.
In Greenland have been recorded 64 spp. of spiders, 1 opilion (Mitopus morio),
and 127 spp. of Acari (Jackson 1930; Sørensen 1898, Jensen and Christensen, Eds.
2003).
A remark by Shelley and Golovatch (2011) concerning Diplopoda is to be
noticed:
“All South American taxa that extend northward to North America/US through Central
America also inhabit the Antillean Arc with the exception of Epinannolenidea, which is
primarily Antillean and only ranges to central Costa Rica. Conversely, no North American
taxon spreads southward completely to South America, and those that go even part way do
so via Central America, as the Antillean paleogeographic origin excludes them as a north-
south pathway. No primarily North American taxon inhabits even part of the Antilles”.
Palpigradi According to Condé (1996) and suppl., in the USA (Texas, Louisiana)
are recorded Eukoenenia florenciae (Rucker, 1903), E. hanseni (Silvestri, 1913),
Prokoenenia californica Silvestri, 1913 (California), P. wheeleri (Rucker, 1901)
(Texas), and many unidentified (Rucker 1901, 1903, Silvestri 1813, Wheeler 1900).
In North America the order is known from the area up to 44oN (Condé 1996).
Ricinulei The only recent species of Ricinulei from the USA (Pseudocellus doro-
theae) has been described from Edinburg, Texas, Hidalgo County, and on the border
with Mexico (Gertsch and Mulaik 1939). In this country are registered also nine
fossil species of Ricinulei (Curculioididae and Poliocheridae).
Solifugae According to the checklist of Brookhart and Brookhart (2006), in conti-

nental North America have been recorded 196 species of Solifugae, from
Eremobatidae (genera Eremothera Muma, Eremorhax Roewer, Eremobates Banks,
Eremocosta Roewer, Horribates Muma, Chanbria Muma, Eremochelis Roewer,
Hemerotrecha Banks, 174 spp.) and Ammotrechidae (Branchia Muma,
Ammotrechella Roewer, Ammotrechula Roewer, Ammotrecha Banks, 22 spp.).
From these 196 spp., 77 are known from Mexico; for the area north of Rio Grande
remain ca. 119 spp. (Brookhart 1972, Muma 1951, 1962, 1970, 1974, 1976, 1986).
The deserts Chihuahuan and Sonoran contain the majority of solifugid species.
From the southern provinces of Canada are known two species: Eremobates docol-
ora Brookhart et Muma, 1981, and Hemerotrecha denticulata Muma, 1951. They
mark the northern limit of Solifugae in the Western Hemisphere.
Scorpiones In North America, north of Mexico (30 states of the USA), are distrib-
uted 111 spp. of scorpions of 20 genera and 6 families: Buthidae, Caraboctonidae,
Chactidae, Scorpionidae (incl. Diplocentridae), Superstitionidae, and Vaejovidae
(see Table 8.1). Paruroctonus boreus (Vaejovidae) is the only species that occurs in
Canada (southern Saskatchewan, Alberta, and British Columbia) (Ewing 1928,
Gertsch and Soleglad 1966, Stahnke 1956, Soleglad and Fet 2005, 2006, 2008,
2010, Soleglad et al. 2014).
The states richest in scorpions are California (51 spp.), Arizona (41 spp.), New
Mexico (20 spp.), Texas (20 spp.), Nevada (16 spp.), and Utah (12 spp.).
Pseudoscorpiones Many data have been published by Beier (1931, 1932),
Chamberlin (1925, 1929, 1930, 1931, 1949), Hoff (1946, 1956), and others.
Longtime ago Hoff (1958) made a list of the pseudoscorpions of North America,
north of Mexico. By this time the list contained 87 genera of 18 modern families.
Meanwhile, many new data were published (to mention only a few): Benedict and
Malcolm (1978), Buddle (2005), Ćurčić (1978, 1981, 1982, 1984), Harvey (online),
and Muchmore (1967, 1974). Rowland and Reddell (1976) made a list of the (then)
30 species in Texas. Now (2017) we may assume that in North America, north of
Rio Grande, are recorded about 400 species of pseudoscorpions, belonging to 105
genera and 19 families (Harvey, 2013f). From these families 13 contain 1–2 genera
and six, 1–2 species each.
Genera living only in this (quite large) area (Nearctic) are:

Fam. Chthoniidae
Neochthonius Chamberlin, 1929 – California (four spp.)
Verrucaditha J.C. Chamberlin, 1929 – USA (one sp.)
Fam. Pseudogarypidae
Pseudogarypus Ellingsen, 1909 – Canada, USA (six spp.)
Fam. Neobisiidae
Alabamocreagris Ćurčić, 1984, Alabama (two spp.); Americocreagris Ćurčić,
1982, Oregon, Washington (one sp.); Australinocreagris Ćurčić, 1984,
California (one sp.); Cryptocreagris Ćurčić, 1982, New Mexico, California
(two spp.); Fissilicreagris Ćurčić, 1984, California (four spp.); Globocreagris
Ćurčić, 1984, California (two spp.); Lissocreagris Ćurčić, 1981, USA (seven
spp.); Minicreagris Ćurčić, 1981, USA (one sp.); Novobisium Muchmore,
1967, USA (three spp.); Saetigerocreagris Ćurčić, 1984, USA (two spp.);
Tartarocreagris Ćurčić, 1984, Texas, Arkansas (six spp.); Trisetobisium
Ćurčić, 1982, USA (one sp.); Tuberocreagris Ćurčić, 1978 – USA (two spp.)
Fam. Syarinidae
Aglaochitra J.C. Chamberlin, 1952 – USA (California) (one sp.)
Fam. Cheliferidae
Aspurochelifer Benedict et Malcolm, 1979 (one sp.); Florichelifer Hoff, 1964
(Florida) (one sp.); Haplochelifer J.C. Chamberlin, 1932 (one sp.); Idiochelifer
J.C. Chamberlin, 1932 (one sp.); Juxtachelifer Hoff, 1956 (New Mexico)(one
sp.); Levichelifer Hoff, 1946, Mexico, USA (two spp.); Pachychelifer Beier,
1962, Georgia (one sp.); Paisochelifer Hoff, 1946, USA (one sp.); Phorochelifer
Hoff, 1956, USA (one sp.); Pugnochelifer Hoff, 1964, USA (one sp.);
Xenochelifer J.C. Chamberlin, 1949, California (one sp.)
Fam. Chernetidae
Acuminochernes Hoff, 1949 (two spp.); Chrysochernes Hoff, 1956 (New
Mexico) (one sp.); Illinichernes Hoff, 1949 (one sp.); Mirochernes Beier,
1930 (one sp.); Mucrochernes Muchmore, 1973 (one sp.)
In Alaska are registered three species (Neobisiidae, Globocreagris, Halobisium;
Syarinidae, Syarinus).
From Canada are registered 24 species of 19 genera and 7 families (Buddle
2005).
Opiliones The catalogue of Cokendolpher and Lee (1993) facilitated the overview
of Cyphophthalmi and “Palpatores” of Greenland, Canada, the USA, and Mexico,
together with bibliography on the Opiliones of this area. Kury (2003) analyzed the
American Laniatores. Concerning Canada and the USA, we find (with supplements)
the following supraspecific taxa, based on Briggs (1968, 1969, 1971a, 1971b, 1974),
Briggs and Hom (1966, 1967), Briggs and Ubick (1981, 1989), Ewing (1923), Giribet
and Kury (2003), Giribet and Shear (2010), Goodnight and Goodnight (1942), Hedin
and Thomas (2010), Newell (1943), Richart and Hedin (2013), Shear (1975b, 1977,
1980, 2010a), Ubick and Briggs (1989, 1992, 2002, 2004), and others.
Cyphophthalmi
Fam. Sironidae
Siro Latreille, 1804 – western USA (Oregon, California, Washington), Maryland
(nine spp., endemic)
Fam. Neogoveidae
Metasiro Juberthie, 1960 – USA (Florida, Georgia, South Carolina, endemic)
(three spp.)
Eupnoi
Fam. Phalangiidae – North America (Oligolophinae, Phalangiinae, Opilioninae)
Fam. Sclerosomatidae – from the four subfamilies, only Leiobuninae lives in
North America.
Cosmobunus Simon, 1879; Eumesosoma Cokendolpher, 1980;
HadrobunusBanks, 1900; Paranelima Caporiacco, 1938; Leuronychus Banks,
1900; Leiobunum C.L. Koch, 1839; Schenkeliobunum Staręga, 1964; Togwoteeus
Roewer, 1952
Fam. Protolophidae (endemic family to the USA and northern Mexico)
Protolophus Banks, 1893 – USA (California, Arizona, Texas, Oregon,
Washington, Utah), Mexico (eight spp.)
All families are shared with the Palearctic region, except for Protolophidae.
Many genera are endemic to more narrow areas in North America.
Dyspnoi
Acropsopilio Silvestri – USA (New York, Connecticut, Indiana, Maine, Michigan,
Ohio, Wisconsin), Canada (Ontario, Quebec) (one sp.)
Fam. Caddidae
Caddo Banks – USA (New York, Connecticut, DC, Michigan, New Brunswick,
New Jersey, etc.) (two spp.)
Fam. Ischyropsalididae
Subfam. Ceratolasmatinae
Acuclavella Shear – Idaho, Washington (seven spp.)
Ceratolasma Goodnight et Goodnight – Oregon, California (one sp.)
Fam. Taracidae
Crosbycus Roewer – Missouri, Georgia (one sp.)

Hesperonemastoma Gruber – North Carolina, California, Washington (five spp.)
Taracus Simon – Oregon, California, Colorado, Washington (seven spp.)
Fam. Sabaconidae
Sabacon Simon – Oregon, Alaska, Washington, North Carolina, California,
Kentucky (eight spp.)
Fam. Nemastomatidae
Subfam. Ortholasmatinae
Dendrolasma Banks – USA (Washington, California) (two spp.)
Ortholasma Banks – Mexico, USA (California) (five spp.)
Trilasma Goodnight et Goodnight – Mexico, Honduras (nine spp.)
Laniatores In Canada only three species of Triaenonychidae are known from one
province: British Columbia (Paranonychus brunneus Banks, 1893; Sclerobunus
nondimorphicus Briggs, 1971; S. parvus Roewer, 1931). From Alaska is known
only Paranonychus brunneus. All the other Laniatores, north of Rio Grande (29
genera and 8 families), live in the “uninterrupted” USA. Most states has one to five
spp. of Laniatores (in bold the endemics for USA and/or Canada). Exceptions are
California (63 species), Texas (18 spp.), and Oregon (15 spp.). The West Coast is
particularly rich in Laniatores, and some are considered relicts (Briggs 1971, 1973,
1974).
Subfam. Paranonychinae
Metanonychus Briggs – Idaho, California, Oregon (three spp.)
Paranonychus Briggs – Canada (British Columbia), USA (Alaska, Oregon,
Washington)
Subfam. Sclerobuninae
Sclerobunus Banks (= Cyptobunus Banks) – Canada (British Columbia), USA
(California, Colorado, New Mexico, Arizona, Montana, Nevada, Utah,
Washington, Idaho, Oregon) (three spp.)
Zuma Goodnight et Goodnight – California (two spp.)
Subfam. Triaenonychinae
Fumontana Shear – North Carolina, Tennessee (one sp.)
Fam. Cladonychiidae
Cryptomaster Briggs – Oregon (one sp.) (end.)
Erebomaster Cope – DC, Maryland, North Carolina, Virginia, Indiana, Kentucky,
West Virginia, Ohio (two spp.)
Speleomaster Briggs – Idaho (two spp.)
Theromaster Briggs – Alabama, Georgia, North Carolina, Tennessee (two spp.)
Fam. Pentanychidae
Isolachus Briggs – Oregon (one sp.)
Pentanychus Briggs – Oregon, Washington (five spp.)
Fam. Cosmetidae
Calicynorta Goodnight et Goodnight – California (one sp.)
Denticynorta Roewer – Ohio (one sp.)
Vonones Simon – Florida, Georgia, Louisiana, Oklahoma, Alabama, Illinois,
Indiana, Kansas, Texas, North Carolina, Ohio, Tennessee (two spp.)
Fam. Phalangodidae
Bishopella Roewer – Alabama, North Carolina, Florida, Georgia, South Carolina
(two spp.)
Banksula Roewer – California (11 spp.)
Calicina Ubick et Briggs – California (25 spp.)
Microcina Briggs et Ubick – California (six spp.)
Phalangodes Tellkampf – Kentucky, Tennessee (one sp.)
Phalangomma Roewer – Virginia (one sp.) (endemic)
Sitalcina Banks – California, Oregon (ten spp.)
Texella Goodnight et Goodnight – California, New Mexico, West Virginia, Texas
(21 spp.)
Tolus Goodnight et Goodnight – Tennessee (one sp.) (endemic)
Undulus Goodnight et Goodnight – Alabama (one sp.) (endemic)
Wespus Goodnight et Goodnight – Arkansas (one sp.) (endemic)
Fam. Travuniidae
Speleonychia Briggs – Washington (one sp.) (endemic)
Fam. Stygnommatidae
Stygnomma Roewer – Florida, Ohio (one sp.)
Fam. Stygnopsidae
Hoplobunus Banks – Texas (two spp.)
Family uncertain (Phalangodidae)
Crosbyella Roewer – Alabama, Arkansas, Florida, Georgia, South Carolina (five
spp.)
Amblypygi This order is recorded only from the Southern USA, with six species
of Phrynidae (Mullinex 1975, Quintero 1981, Harvey 2013):
Acanthophrynus Kraepelin, 1899 – USA (A. coronatus Butler, 1873, Arizona,

California)
Paraphrynus Moreno, 1940 – USA (Arizona, Florida) (P. mexicanus Bilimek, 1867;
P. raptator Pocock, 1902)
Phrynus Lamarck, 1801 – USA (Ph. marginemaculatus C.L. Koch, 1841, Florida;
Ph. operculatus Pocock 1902, Texas; Ph. fuscimanus C.L. Koch, 1847, Texas)
These species live also south of these states (in Mexico, etc.). They mark also the
northern limit of Amblypygi in North America.
Thelyphonida (Uropygi) In the area North of Rio Grande is known only one spe-
cies of Thelyphonida (Uropygi): Mastigoproctus giganteus (Lucas, 1835) in Texas
(Thelyphonidae). This is the northernmost species of Uropygi in the New World
(Rowland and Reddell 1976).
Schizomida North of Rio Grande Schizomida are recorded only from four states
of the USA: Arizona, California, Florida, and Texas (not counting the American
territories Hawaii and Puerto Rico) (Briggs and Hom, 1966, 1972, 1988, Chamberlin
1939, Cook 1899, Gertsch 1940, Reddell and Cokendolpher 1995, Rowland 1971,
1972, 1975). Most species live in California (eight, all of the genus Hubbardia
Cook, 1899). Another three species belong to the genera Bamazomus and Stenochrus.
From this 11 spp., 9 are endemic: Stenochrus mulaiki (Texas) and all Hubbardia
(Arizona and California). Stenochrus portoricensis is known also from Florida,
Puerto Rico, Mexico, and many other countries in Central and South America and
West Indies and is introduced also in the Canary Islands and England. Endemic
extant genus is Hubbardia Cook (Hubbardiidae) (+ the fossil Calcitro,
Calcoschizomus, and Onychothelyphonus). The family Protoschizomidae (gen. sp.
indet.) is also recorded from Texas.
Araneae In Canada and Alaska have been recorded 1413 spp. of 43 families of
spiders (highest number in Linyphiidae – 39.5%). The richest territories are Ontario
(746 spp.), British Columbia (700 spp.), and Quebec (677 spp.) (Brignoli 1973,
Griswold et al. 2012, Paquin et al. 2001, Ubick et al. (eds.) (2005), Paquin et al.
2010). These authors affirm that the 1413 spp. might represent 80% of the actual
total of spiders on this huge territory. According to Bennett (1999), the number of
spiders in Canada is ca. 1400 spp. Except for five spp. of Mygalomorphae, all the
others belong to Araneomorphae. As in most northern countries, dominant are the
Linyphiidae (>500 spp.), followed by Salticidae (110 spp.), Gnaphosidae (100
spp.), and Theridiidae (100 spp.). These four families form more than half of all
Canadian spiders.
In Greenland are found 70 spp. of spiders of nine families: Araneidae, Dictynidae,
Gnaphosidae, Hahniidae, Linyphiidae (45 spp.), Philodromidae, Tetragnathidae,
Theridiidae, and Thomisidae (Larsen and Scharff 2003, updated 2007).
Endemic family in the USA is the recently described Trogloraptoridae (Griswold,
Audisio & Ledford, 2012) from caves in Oregon and California, with one genus and
species Trogloraptor marchingtoni. Further are compared the families of spiders in
the Nearctic (56), the Palearctic (55), and the Holarctic (68), including 44 families
common between the two regions. The highest level of difference is the suborder
Mesothelae, unknown in the Nearctic. The endemism on family level is insignificant.
It is to notice that only Cuba, immensely smaller than the Nearctic, are known
almost the same number of families (52).
Opilioacarida The only representative of Opilioacarida in the USA (Texas,
Arizona) is Neocarus texanus Chamberlin et Mulaik (= Neocarus arizonicus
Chamberlin et Mulaik). It marks the northern limit of the order Opilioacarida
(Chamberlin and Mulaik 1942, Vásquez and Klompen 2002, 2009).
Parasitiformes
Ixodida Gregson (1956) published from Canada 29 spp. of Ixodida, including 5
Argasidae and 24 Ixodidae. The list of localities includes some of the northernmost
findings of Ixodida in the Western Hemisphere.
Distribution of Arachnida on the Islands of the Pacific
7.37 Polynesia and Micronesia
7.37.1 G
and Paleogeography
In the western part of the largest ocean “...the Pacific plate preserves the oldest
ocean floor on Earth at 167 Ma, of Jurassic age” (Koppers et al. 2003). The age and
the origin of the Pacific islands have been explained by Neall and Trewick (2008).
According to these authors, the paleocontinent of Zealandia has been displaced
eastward from Gondwana between 83 and 54 Ma by mid-ocean spreading of the
Tasman Sea. Remains of Zealandia are New Zealand, the Chatham Islands, and
New Caledonia. Other islands are likely to have existed in the past (between 38 and
21 Ma) (Karig 1970, 1971, 1974, Menard and Hamilton 1963, Stoddart 1992).
7.37 Polynesia and Micronesia 515
On the islands of Polynesia and Micronesia are recorded Pseudoscorpiones,

Opiliones, Araneae, Schizomida, Amblypygi, very few Scorpions, Araneae, and
Acari (Mesostigmata, Ixodida, Prostigmata). Some sources: Berland (1942), Kami
and Miller (1998), Mayr (1944), Gressitt (1961)
After a long series of publications on the spiders of the islands of the Pacific,
Berland (1934b) subdivided the Pacific into six provinces:
“La province australo-canaque” – Australia, New Zealand, New Caledonia, Loyalty
Islands, the islands Chatham, Norfolk, and Lord Howe
“La province papouasienne” – New Guinea, the islands around it, the Solomon
Island, and even Vanuatu (the New Hebrides)
“La province polynésienne” – Samoa, Fiji, Tonga, Marquesas, Society, Rapa Nui,
and even Hawaii (? subprovince)
“La province micronésienne”
“La province neotropicale” (Galapagos and Juan Fernandez Islands)
“La province antarctique” – the islands Campbell, Auckland, Bounty, Snares,
Macquarie, attached from one side to Tierra del Fuego, on the other side to
Kerguelin
Lehtinen (1980)
Palpigradi – No records
Scorpiones On many islands (Palau, Samoa, etc.) is widespread Liocheles austral-
asiae (Hormuridae). From Palau is recorded also Hormurus waigiensis (Gervais)
from the same family (Chapin 1957, Vachon 1972).
Schizomida Only Hubbardiidae indet are mentioned from Samoa, Belau, Marshall
Islands, and FS of Micronesia. From Guam and North and South Mariana Islands
(Asuncion, Saipan, Sarigan), Reddell and Cokendolpher (1995) notice the presence
of the genus Orientzomus. Cokendolpher and Reddell (2000) recorded from the
Marshall Islands the first two (new) species of Schizomida: Apozomus brignolii and
Orientzomus ralik, Cokendolpher and Tsurusaki (1994). Villarreal Manzanilla
(2010) announced the presence of genus Zomus on Samoa.
The list of pseudoscorpions of Polynesia and Micronesia counts eight families:
Chthoniidae, Tridenchthoniidae, Geogarypidae, Olpiidae, Cheiridiidae, Atemnidae,
Chernetidae, and Withiidae (Beier 1940, 1957, Chamberlin 1934, 1938, 1939a,
1939b, 1947).
Endemic genera for Polynesia and Micronesia are:
Fam. Chernetidae
Meiochernes Beier, 1957 – Caroline Islands (one sp.)
Opiliones – seven genera of four families (all Laniatores)

Cyphophthalmi are not known in Polynesia and Micronesia.
Dyspnoi are not known in Polynesia and Micronesia.
Laniatores – Roewer (1912), Goodnight and Goodnight (1957), Suzuki (1941)
Fam. Epedanidae – Dibunus Loman, 1906
Fam. Podoctidae – Metibalonius Roewer, 1915, Lomanius Roewer, 1923
Fam. Samoidea – Parasamoa C.J. Goodnight et M.L. Goodnight, 1957; Feretrius
Simon, 1879 (one sp.); Samoa Sørensen in L. Koch, 1886
Fam. Zalmoxidae – Zalmoxis Sørensen, in L. Koch 1886
Amblypygi Fam. Charontidae: On Palau has been found Charon grayi (Gervais,
1842) (Esaki 1936, Takashima 1950).
Araneae Berland (1934b) subdivided the Pacific islands into four groups.
According to him:
All information seems to indicate that the settlement of Pacific was done by migrations
coming from the Indo-Malaysian area, migrations which would probably have been multi-
ple and following several distinct currents. One of these currents would have been popu-
lated from Australia; this is what I call the “australo-canaque” province; a quite distinct
current of migration, but of same origin, would have populated Polynesia, whose current
archipelagoes constitute probably only the subdivision of a previously more extended
continent, with a branch being detached towards Hawaii; another towards Micronesia.
There is no visible relation between America and the Pacific; but on the other hand
Galapagos as well as Fernandez received their fauna from the south of America, by the
austral lands. In addition the islands of Pacific almost always present a very marked level of
endemism, which testifies to an extremely old isolation, and should be fix their separation
farther back, at least to about the middle of the Tertiary epoch, and perhaps farther. I will
add that the study of several zoological groups, as one can see it in this work, arrive, inde-
pendently from each other, at conclusions very close to mine, and so appear to me to receive
a solid confirmation from it.
7.37 Polynesia and Micronesia 517
Map 7.28 Zonation of Micronesia and Polynesia (After Berland 1934)
Berland supported the view that the islands of Polynesia were formed as a single
large continent, which became subdivided to form the islands.
Recent geological evidence has dismissed this view. The archipelagoes of Hawaii,
Marquesas and Societies are all volcanic in origin, and formed as volcanic hot spots. The
Marquesas range from Nuku Hiva, the oldest in the north at 3.7myrs, to Fatu Hiva the
youngest, in the south, at 1.4 myrs. The Society Islands range from Bora Bora, the oldest in
the north at 3.3 myrs, to Tahiti the youngest, in the south, at 1.0 myrs. (Gillespie et al. 2000)
Map 7.29 Spreading of Araneae to the islands of the Pacific (Berland, 1934)

Fam. Linyphiidae
Paro Berland, 1942 – Rapa (one sp.)
Uahuka Berland, 1935 – Marquesas Islands (two spp.)
Uapou Berland, 1935 – Marquesas Islands (one sp.)
Fam. Dictynidae
Banaidja Lehtinen, 1967 – Samoa (one sp.)
Fam. Pisauridae
Nukuhiva Berland, 1935 – Marquesas (one sp.)
Fam. Tetrablemmidae
Rhinoblemma Lehtinen, 1981 – Caroline Islands (one sp.)
7.38 Hawaii 519
Fam. Anyphaenidae
Australaena Berland, 1942 – Polynesia (two spp.)
Fam. Salticidae
Iona Peckham et Peckham, 1886 – Tonga
Rarahu Berland, 1929 – Samoa
Other sources: Beatty et al. (2008), Berland (1924, 1935a, 1935b, 1935c, 1939,
1942), Evenhuis (2006), Garb (2003), Garb and Gillespie (2006), Gillespie (2003b,
c), Gillespie et al.(2002), Lehtinen (1993), Marples (1955), Proszyński (1996)
7.38 Hawaii
7.38.1 G
and Paleogeography
Hawaiian islands are the exposed peaks of a great undersea mountain range known
as the Hawaiian-Emperor seamount chain, formed by volcanic activity over a hot
spot in the Earth’s mantle. The Hawaiian islands are about 3,000 km from the near-
est continent. The archipelago consists of eight greater and many smaller islands.
The main islands are Hawaii (10,432 km2, with the highest point Mauna Kea, 4205
m), Maui (1883 km2), Oahu (1545 km2), Kauai (1430 km2), Molokai (673 km2),
Lanai (364 km2), Niihau (180 km2), and Kahoolawe (116 km2). The total area of the
archipelago is 16,636.5 km2.
The islands range from Kauai, the oldest in the north at 5.1myrs, to Hawaii the
youngest, in the south, at up to 0.4 myrs old.
It is considered that the entire fauna of these islands, originally azoic, is a result of
the three W (wind, waves, wings) – the ancestors either arrived by rafting or were
windblown or have been brought with birds or bats (Wagner and Funk 1995,
Zimmerman 1948).
Map 7.30 The Hawaiian Islands
These islands, completely azoic in the beginning, are now housing several orders of
Arachnida: Palpigradi, Scorpiones, Schizomida, Pseudoscorpiones, and Araneae
and different mites. Even more, there are endemic genera of pseudoscorpions and
spiders and amazing cave fauna (troglobites in recently formed lava tubes). Some
other arachnids are clearly adventive from other lands, including Thailand and
North America (Berland 1934, Nishida 1994, 1997, 2002, Simon 1900).
Palpigradi On Hawaii are known two species of Palpigradi, both from the family
Eukoeneniidae – Eukoenenia florenciae (Ruckner, 1903) and E. hanseni (Silvestri,
1913). Both species are known also from the continental USA (Louisiana, Texas)
(Harvey 2013e, Rémy 1961a).
Scorpiones Isometrus maculatus (De Geer) is distributed in Oahu, Maui, and

Hawaii (Yates 1993).
Schizomida Known is only Bamazomus siamensis (Hansen) (Oahu, also from

Thailand, Hong Kong, and the Ryukyu Islands) (Reddell and Cokendolpher 1995).
Pseudoscorpiones In the first checklist of Pseudoscorpiones of Oceania

(Chamberlin 1934) are listed six spp. from Hawaii. Important contribution to the
record on the Pseudoscorpiones of Hawaiian Islands was made by Muchmore
7.38 Hawaii 521
(1979, 1983, 1989, 1996, 2000). Now (2017) from the islands are known the fami-
lies Chthoniidae, Lechytiidae, Garypinidae, Chernetidae, Cheliferidae, Cheiridiidae,
Geogarypidae, Olpiidae, and Withiidae (Beier 1932, 1940, Muchmore 2000, Harvey
1990, Harvey 2013f, Simon 1900, With 1905), according to Nishida (2002), 17 spp.
Endemic genera for Hawaiian Islands are:
Fam. Chthoniidae: Vulcanochthonius Muchmore, 2000 (three spp.)
Fam. Chernetidae: Eumecochernes Beier, 1932 (three spp.)
Of special interest are the troglobitic species in the lava tubes (Muchmore, 1979,
1983, 1989). Three species have been described: Tyrannochthonius howarthi, T.
stonei, and T. pupukeanus. T. howarthi has been transferred to the new genus
Vulcanochthonius Muchmore, 2000, including V. pohakuloae Muchmore, 2000.
In his description of Tyrannochthonius howarthi as the first cave pseudoscorpion
in Hawaii, Muchmore (1982) expressed some doubt concerning the real place of
this species in the system. According to him, “...it appears quite possible that it actu-
ally belongs to a presently unrecognized group (genus or subgenus) with a wide
distribution in the Pacific.”
Opiliones
In the checklist of terrestrial arthropods of Nishida (2002) are not mentioned any
Opiliones inhabiting Hawaii.
Araneae After Simon’s “Fauna Hawaiiensis” (1900), in the catalogue of Suman
(1964) have found place 149 spp. of 21 fam. and 66 genera. This number includes
82 endemic species (55%) of 10 fam. and 27 genera. By 2005 on the Hawaiian
Islands were known 132+ native spp. of spiders of 36 native genera and 16 native
families. Particularly interesting are the blind spiders, found by Howarth (1980) in
the relatively recent lava caves (Adelocosa anops Gertsch). At least 128 spp. are
endemic species. “Eighty percent of the known Hawaiian native spider species
belong to ten genera in seven families Cyclosa (Araneidae), Orsonwelles
(Linyphiidae), Pagiopalus and Pedinopistha (Philodromidae), Havaika (Salticidae),
Tetragnatha (Tetragnathidae), Argyrodes and Theridion (Theridiidae), and
Mecaphasa and Misumenops (Thomisidae)” (Hawai’i Comprehensive Wildlife
Conservation Strategy, 2005). Gertsch (1973) identified the collection of 20 spp. of
spiders, found in the lava tubes. Most were introduced, but six were eyeless or with
eyes reduced to vestiges (troglobites). According to Gertsch (loc. cit.), “Most impor-
tant are the two new wolf spiders (Lycosidae), one with vestigial eyes and the other
a completely eyeless species of this family of big-eyed hunting spiders.” Other
sources are: Garb (1999), Garb and Gillespie (2009), Gillespie (1991), Gillespie
et al. (1998), Gillespie et al. (2000), Gillespie et al. (2003), Simon (1899 – 1900),
Suman (1964, 1965, 1967, 1970).
Some endemisms:
Fam. Linyphiidae
Orsonwelles Hormiga, 2002
Priperia Simon, 1904 (one sp.)
Fam. Lycosidae
Adelocosa Gertsch, 1973 – Kaua’i, caves (blind)
Syroloma Simon, 1900
Fam. Tetragnathidae
Doryonychus Simon, 1900 (one sp.)
Fam. Philodromidae
Subfam. Pedinopisthinae
Pagiopalus Simon, 1900
Pedinopistha Karsch, 1880 (five spp.)
Fam. Thomisidae
Mecaphesa Simon, 1900
Fam. Salticidae
Havaika Prószyński, 2002 – 23 spp. on Hawaii, three spp. on Marquesas Islands
Also more than 30 local genera
Acari According to Garrett and Haramoto (1967), until this time on Hawaiian
Islands have been recorded 210 species of mites and ticks of 67 families and 138
genera. According to Nishida (2002), on the islands are recorded 928 spp. of Acari,
including 296 endemic (165 families, 505 genera).
Acariformes Garrett and Haramoto (1967) indicate on the archipelago 21 families

of Prostigmata and 18 families of Oribatida.
Parasitiformes (Ixodida and Mesostigmata)

Ixodida – until 1967 in Hawaii have been registered four spp. of Argasidae and
eight spp. of Ixodidae (mostly quarantine finds). Ixodes laysanensis Wilson,
1964, is described on birds.
Mesostigmata – Garrett and Haramoto (1967) indicate 15 families on the islands.
7.39 Antarctic and Subantarctic Arachnofauna
7.39.1 G
and Paleogeography
Subantarctic are termed the islands around Antarctica situated north of and adjacent
to the Antarctic Convergence (between the 48th and 61st parallel of south latitude).
The Falkland Islands, Prince Edward Islands, Crozet Islands, Amsterdam Island, St
Paul Island, Tierra del Fuego, and Macquarie Island lie north of the Antarctic
7.39 Antarctic and Subantarctic Arachnofauna 523
Convergence. Other bigger Subantarctic islands of island groups are Antipodes

Islands, Auckland Islands, Bounty Islands, Macquarie Island, Prince Edward
Islands, The Snares, and Tierra del Fuego. Some of the Antarctic islands also lie
north of 60oS: Bouvet Island, Heard Island, McDonald Islands, Kerguelen Islands,
South Georgia Islands, South Sandwich Islands, and others (Gressitt 1964, 1965,
1970, Gressitt and Weber 1959, Morrone 2000b).
According to Morrone (2000), biogeographically the Subantarctic Subregion
belongs to the Andean region of the Austral Kingdom and comprises the Austral
Andes from 37oS to Cabo de Hornos, the archipelago of southern Chile and
Argentina, and the Malvinas, South Georgia, and Juan Fernandez Islands (six prov-
inces: Maule; Valdivian; Magallanic forest; Magallanic moorland; Malvinas islands,
including South Georgia; and Juan Fernandez Islands). For each province are listed,
among the other animals, the typical elements of Arachnida.
On the Antarctic continent live only some groups of mites. From the other groups
on the Subantarctic islands are known some pseudoscorpions, opilions, and spiders
(Pugh 1993, 1997, 2003). If the most southern South America is included in the
term “Subantarctic,” we find many more groups of Arachnida, but the orders listed
above still prevail.
The ice-covered Antarctica with harsh climate and almost no vegetation and the
islands inside the Antarctic Convergence (N) have very limited arachnofauna, com-
posed mostly of Araneae and Acari. This huge area has not been always ice-covered,
and there are hypotheses for transantarctic ways of dispersion for many groups of
Arachnida, inhabiting the southern continents (Patagonia, Southern Africa,
Australia, New Zealand, New Caledonia).
Some sources: Dalenius and Wilson (1958), Fleming (1987), Forster (1964,
1970, 1971), Gressitt (1965, 1971), Hill (2009), Marshall and Pugh (1996), Morrone
(2000), Pugh (1994, 1997, 2004), Wallwork (1963, 1965, 1966, 1967, 1969, 1973,
1979)
Presently “0.34% of its area is currently ever free of snow or ice in the form of
terrestrial ecosystems including exposed nunataks, cliffs and seasonally snow and
ice-free areas” (Convey 2010, after British Antarctic Survey, 2004). Udvardy (1987)
proposed the creation of “biogeographical realm Antarctica” including four prov-
inces: Maudlandia, Marielandia, Insulantarctica, and Neozealandia.
After Hill (2009), the Antarctic land bridge, associated with a tropical to temper-
ate climate (Francis et al., 2008), was available to support the dispersal of plants and
animals for about 75–95 million years after the separation of Africa.
The study of the few arthropods in Antarctica and Subantarctic is important in
many ways – to understand the ways of dispersal and the abilities of different groups
to exist in harsh and extreme conditions of cold and devoid of vegetation environ-
ment. The only groups represented there are Acari (Acaridida, Oribatida,
Trombidiformes) and Araneae.
Some parasitic ticks and mites are brought to these lands by birds (Ixodida).
Pseudoscorpiones From the Subantarctic islands outside the ones belonging to
New Zealand and Australia (see New Zealand), Falkland Islands, and Tierra del
Fuego (see Patagonia) are recorded the following pseudoscorpions (Chamberlin
1933, Vitali-di Castri 1968):
Ile de la Possession, île Crozet – Austrochthonius insularis Vitali-di Castri

St Paul – Diplotemnus insularis Chamberlin
Opiliones. Forster (1955a)
Opiliones reported from Subantarctic and extreme southern cold temperate
islands (after Cokendolpher and Lanfranco 1985):
Neonuncia campbelli Forster – Campbell Islands (52o33′), cold temperate
(Forster 1954, 1964)
Neonuncia eastoni Forster – Auckland Islands (50o 32′), cold temperate (Forster
1954, 1964)
Neonuncia enderbyi (Hogg) – Auckland and Campbell Islands (50°32”, 50°
33”), cold temperate (Forster 1954, 1964)
Nuncia unifalculata (Enderlein) – Crozet Islands (46o30”), Subantarctic (Tambs-
Lyche 1954, Hickman 1939)
Fam. Gonyleptidae
Lycomedicus planiceps (Guerin) – Isla Hoste (55o 10”), cold temperate (Soares
and Soares 1954)
Haversia defensa (Butler) – Islas Malvinas (52o), cold temperate (Soares and
Soares 1949)
Haversia [=Hoggellula] vallentini (Hogg) – Islas Malvinas (52o), cold temperate
(Soares and Soares 1949)
Fam. Megalopsalididae
Pantopsalis distincta Forster – Auckland Islands (50° 32′), cold temperate
(Forster 1964)
Pantopsalis johnsi Forster – Auckland Islands (50o 32′), cold temperate (Forster
1964)
Pantopsalis mila Forster – Auckland Islands (50o32′), cold temperate (Forster
1964)
Pantopsalis rennelli Forster – Campbell Island (52o33′), cold temperate (Forster
1964, Gressitt et al. 1964)
Pantopsalis snaresensis Forster – Snares Island (48o), cold temperate (Forster
1964)
Fam. Neopilionidae
Thrasychirus dentichelis Simon – Isla de los Estados, Isla Hoste (54o50′, 55o30′),
cold temperate (Ringuelet 1959)
Cape Horn Archipelago 55o37′–49′ cold temperate
Thrasychirus gulosus Simon – Isla de los Estados, Isla Hoste (54o50′, 55o30′),
cold temperate (Ringuelet 1959)
Thrasychirus modestus Simon – Isla Hoste, Isla Navarino (55o 10′–30), cold
temperate (Cekalovic 1976), Isla Deceit (55o49′) cold temperate
Araneae In his paper on the spiders of islands Chatham, Bounty, Snares, Auckland,
Campbell, Macquarie, and Kerguelen, Berland (1930a) pays special attention to the
genus Myro P.-C. (Desidae), known from Kerguelen, Macquarie, South Africa,
Tasmania, and the extreme south of South America. His conclusion is that “il jalonne
donc admirablement un continent qui auraît réuni l’Amérique à l’Australie par les
subantarctiques et la Tasmanie, à Kerguelen et même au sud de l’Afrique.” Now the
distribution of this genus is restricted to Crozet, Kerguelen, Macquarie, New
Zealand, Marion, and Tasmania. There are other genera of intertidal spiders living
in South America (Porteria in Chile and even Desis galapagoensis Hirst on
Galapagos Islands), but they make no connection with Australasia or Kerguelen.
Berland also affirm that the spiders found on the small islands satelites “n’ont à
peu près aucune affinité avec la Nouvelle-Zélande.” Actually, now we know that
very few of the numerous genera of Desidae of New Zealand are represented also
on the Subantarctic islands (Gasparia Marples, Gohia Dalmas, Myro P.-Cambridge,
Laestrygones Urquhart).
Forster (1955, 1962, 1970, 1971) completed vastly the knowledge on the spider
fauna of Subantarctic islands.
Pugh (2004) analyzed the biogeography of spiders on the islands of the Southern
Ocean.
Araneae In his special paper on the possibility of mites to colonize Antarctica and
the islands of the Southern Ocean via air currents, Pugh (2003) indicates that among
the spiders “most long-range aerial colonists are minute adult Araneidae, Lycosidae,
Theridiidae, and especially Linyphiidae, representing 81% of verified specimens
collected over the Southern Ocean and 46% of Southern Ocean species.” The juve-
niles have a low colonization success. In a biogeographic analysis of the spiders on
the islands of the Southern Ocean, Pugh (2004) indicates that only 115 verified spe-
cies from 26 families are reported from these islands. On the continent the spiders
are anthropogenic immigrants.
A young spider in good condition was trapped at Marble Point on the Victoria Land coast
opposite Ross Island (Gressitt 1964).
Endemic genera:
Fam. Linyphiidae: Ringina Tambs-Lyche, 1954 – Crozet
Fam. Theridiidae: Icona Forster, 1955 – Auckland, Campbell Islands (two spp.)
Acari According to the prominent specialist of Antarctic ecology and biogeogra-

phy Prof. J.L. Gressitt, “The mites and ticks may well prove to be, in the number of
species, the richest terrestrial animal group in Antarctica. The mites seem to be
particularly tolerant of low temperatures. They have been found equally as far south
as Collembola (83°55’S)” (Gressitt 1964).
What concerns the dispersal of Acari (and other Arachnida) in Antarctica,
Gressitt (1964) says: “If the quaternary climates on the continent was too severe at
its coldest for the survival of remnants of the preceding temperate fauna, than it is
likely that all or much of the present fauna has resulted from immigration in fairly
recent time. This suggests transport in air currents, or by the agency of birds.” He
points that a mite has been trapped 3000 meters above sea level at a point halfway
between New Zealand and Antarctica.
A comprehensive checklist of the Antarctic Acari is due to Pugh (1993). It con-
tains 528 spp.
After Pugh (1994), about 70 spp. are nonindigenous.
About mites Pugh (2003) concludes that some are probably phoretic (Ascidae,
Parasitidae, Uropodidae and Veigaiidae) and that “…all Bdellidae, Eupodidae,
Acaridae, Tyroglyphidae, and Oribatida (= Cryptostigmata) relate to local edaphic/
hemiedaphic mites that were locally uplifted by storm and/or imported as shipborne
(anthropogenic) contaminants.”
Concerning the Oribatida (Cryptostigmata), Wallwork (1969) has adopted the
scheme proposed by Holdgate (1964):
1. Subantarctic zone
Kerguélin I., Prince Edward I., Crozet I., Macquarie I., South Georgia
2. Maritime Antarctic zone
Bouvet Island, South Orkney Islands, South Sandwich Islands, South Shetland
Islands, Antarctic Peninsula
3. Continental zone
Eastern Antarctic Continent
From his analysis Wallwork (1969) has drawn the following (tentative)
conclusions:
1 . The Continental zone has an impoverished, highly endemic fauna.
2. Endemism in the Maritime zone is low compared with that of the Subantarctic
zone, the fauna of the former being clearly derived from that of the latter.
3. The Subantarctic zone supports a much richer fauna than the other two zones,
and the relatively high endemicity (probably estimates) of its fauna suggests that
this zone may be recognized as a distinct faunal region, as far as the Cryptostigmata
are concerned.
Stary and Block (1998) consider 105 spp. of 20 fam. of Oribatida to inhabit the
Antarctic and Subantarctic territories. Predominant are Oppiidae and
Ameronothridae, with one endemic family (Maudheimiidae). The endemism on the

continent is 60%, in the Subantarctic 63%, against 83% in New Zealand. According
to Schatz (2004), in these territories until 2004 have been registered 118 spp. of
Oribatida. The increase of number is not so big.
Acariformes Ref.: Gressitt and Weber (1959), Marshall and Pugh (1996), Pugh
(1993, 1997, 2003, 2004), Wallwork (1965, 1969, 1973)
Parasitiformes Ref.: Gressitt and Weber (1959)
Ixodida
Fam. Ixodidae. Some ixodids have been described from marine birds (Ixodes uriae
Witte, 1852; I. kerguelenensis André et Colas-Belcour, 1942).
Map 7.31 The three widely recognized terrestrial biogeographical zones in Antarctica
(After Convey 2010)
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Chapter 8
Some Peculiarities of the Distribution
of Arachnida
Abstract Special attention is payed to the comparison between the arachnofaunas

of close areas: Mediterranean and Central European; Mexico and the USA; Central
America and the Caribbean Islands; South Africa, Madagascar, Seychelles, and
Mascarene; and special areas like Patagonia and New Guinea, Australia, New
Zealand, New Caledonia, and Lord Howe Island.
8.1 Mediterranean and Central European Arachnofauna
Both the classic Mediterranean and European-Siberian (Eurosiberian) areas are in the
Palearctic and usually are considered subregions. Malicky et al. (1983) presented argu-
ments for the existence and the horology of special Middle European (Extramediterran-
European) elements in the fauna of Europe. In Middle Europe are missing many orders
of Arachnida (Amblypygi, Uropygi, Schizomida, Ricinulei, Opilioacarida,
Holothyrida); groups like Palpigradi, Cyphophthalmi, Laniatores, and Scorpiones are
represented by symbolic number of species. Well represented are Araneae, Opiliones
(Eupnoi, Dyspnoi), and Pseudoscorpiones. Cave Arachnida are much less represented
than in the Mediterranean, and troglobites North of the Alps are almost missing.
According to Lopatin (1980), the European-Siberian Region within the Palearctic
Subkingdom is subdivided into European-Obian subregion (from Great Britain to
Kamchatka and Chukotka) and Angaran subregion (Siberia East of Yenisei – the
Johansen’s Line – as far as Sakhalin). Both subregions are subdivided into land-
scape zones: tundra, taiga, steppe, and high mountains. The Asiatic part of European-
Siberian area is with much harsher climate than in Middle Europe. Spiders are
prevailing; the part of Linyphiidae is bigger in the Nordic areas.
8.2 Arachnofauna of the Ancient Mediterranean
According to Lopatin (1980), Palearctic is a subkingdom, including three regions:

European-Siberian, Region of the Ancient Mediterranean, and East Asiatic Region.
The Region of the Ancient Mediterranean consists of two subregions – Mediterranean

https://doi.org/10.1007/978-3-319-74418-6_8
626 8 Some Peculiarities of the Distribution of Arachnida
and Saharo-Gobian. These are territories around the Mediterranean Sea and a large
band from North Africa to Central Asia and Gobi. According to the map (Lopatin
1980, p. 125), the southern border of this region follows the northern limit of the
Afrotropical Region (Paleotropical Kingdom) and also runs on the northern slope of
the Himalaya. The northern limit is following the Pyrenees, Alps, Danube, and the
mountains of Central Asia. The Mediterranean Subregion is divided into provinces:
Macaronesian, Tyrrhenian or West Mediterranean, and East Mediterranean. The
vast territory of the Saharo-Gobian Subregion includes three superprovinces:
Saharo-Sindian, Irano-Turanian, and Central Asiatic. In the respective parts of this
book are enumerated in details the Arachnida of each of these territories. Some
orders are found only in the Mediterranean Subregion, much warmer than the other
parts of the Palearctic, or are barely represented. Such groups are the Amblypygi
(Charinus, Musicodamon), the majority of Solifugae and Scorpiones, many fami-
lies of Opiliones and Araneae, most of the Palpigradi, and most of the relicts (scor-
pions, opilions, pseudoscorpions). Mediterranean or submediterranean elements are
found even in Romania and southern Slovakia, but actually the full Mediterranean
fauna stops about the mountains on the border between Bulgaria and Greece.
Submediterranean elements (Galeodes, scorpions, and other groups) enter the
present-day Bulgaria only along the valleys of Struma, Mesta, and Maritza.
Otherwise most of the Bulgarian territory is with Central European elements.
8.3 The Boundary Neotropica-Nearctic
In North America, north of Mexico, are found 13 orders of Arachnida, but the
“southern” orders (Amblypygi, Ricinulei, Schizomida, Uropygi, Opilioacarida, and
largely extending to Solifugae and Scorpiones) are confined mostly to Texas,
Arizona, California, Florida, Louisiana, and New Mexico, some of them being on
the edge of their areals (Dice 1943, Harvey, on line, Rowland and Reddell 1976;
Sørensen 1898).
8.3.1 Palpigradi
Mexico – Eukoenenia hanseni (Silvestri), E. corozalensis Montaño et Francke, and

E. tetraplumata Montaño (Eukoeneniidae).
USA – In Texas and Louisiana are recorded Eukoenenia florenciae (Rucker,
1903), E. hanseni (Silvestri, 1913), Prokoenenia californica (Silvestri, 1913)
(California), and P. wheeleri (Rucker, 1901) (Texas). In North America, the order is
known from the area up to 44oN.
8.3 The Boundary Neotropica-Nearctic 627
8.3.2 Ricinulei
Mexico – Nine species of the genus Pseudocellus Platnick.

USA – The only recent species of Ricinulei from the USA (Pseudocellus doro-
theae (Gertsch et Mulaik)) has been described from Edinburg, Texas, and Hidalgo
County, on the border of Mexico.
Solifugae According to the checklist of Brookhart and Brookhart (2006), in conti-
nental North America have been recorded 196 species of Solifugae, from
Eremobatidae (174 sp.) and Ammotrechidae (22 sp.).
Scorpiones Mexico – 272 spp. of 37 genera and 8 families. The biggest genus is

Vaejovis C.L. Koch (61 sp.). Remarkable is the presence of cave species of the fam-
ily Typhlochactidae. This family and many genera (Typhlochactas, Stygochactas,
Sotanochactas, Troglocormus, Alacran, Hoffmannichadrurus, Chaneke, Koloti,
Balsateres, Konetontli, Kuarapu, and others) are endemic for Mexico. USA – In
North America, north of Mexico (30 states of the USA), are distributed 92 spp. of
scorpions of 13 genera and 5 families. Paruroctonus boreus (Vaejovidae) is the only
species that occurs in Canada (southern Saskatchewan, Alberta, and British
Columbia). Altogether in the USA are registered 111 spp. of 20 genera and 6 fami-
lies. The states richest in scorpions are California (51 sp.), Arizona (41 spp.), New
Mexico (20 spp.), Texas (20 spp.), Nevada (16 spp.), and Utah (12 spp.).
8.3.3 Pseudoscorpiones
Mexico – Recorded are 161 spp. of Pseudoscorpiones of 63 genera and 18 families.

Six genera are endemic to Mexico.
USA – Now (2017) we may assume that in North America, north of Rio Grande
are recorded about 400 species of pseudoscorpions, belonging to 105 genera and 19
families.
8.3.4 Opiliones
Mexico – 283 spp. (1 sp. of Cyphophthalmi, 143 Laniatores, and 139 “Palpatores”).

Cyphophthalmi – Shearogovea mexasca – is a Gondwanan Neotropical element;
the laniatorid genera all have closest relations with the Neotropical fauna, including
the 16 endemic genera. About half of the “Palpatores” have close relations with the
Nearctic and half with the Neotropical fauna. Acropsopilio is considered to be a
relictual genus (Kury and Cokendolpher 2000).
USA – The catalogue of Cokendolpher and Lee (1993) facilitates the overview
of Cyphophthalmi and “Palpatores” of Greenland, Canada, the USA, and Mexico,
together with a bibliography on the Opiliones of this area. Kury (2003) analyzed the
American Laniatores.
In the USA are represented all subgenera of Opiliones: Cyphophthalmi
(Sironidae).
8.3.5 Amblypygi
In Mexico and the southern states of the USA (Arizona, California, Florida, Texas)
are found the same genera of Amblypygi (all Phrynidae): Acanthophrynus Kraepelin,
Paraphrynus Moreno, and Phrynus Lamarck. They mark also the northern limit of
Amblypygi in North America. The family Phrynidae is clearly Neotropical, despite
the finding of one species in Indonesia (Flores) (according to Armas 2006, “pudiera
ser errónea”).
8.3.6 Uropygi (Thelyphonida)
Mexico – Recorded is only one member of Mastigoproctus Pocock, 1894, M. gigan-

teus (Lucas, 1835) (Uroproctinae). The genus is known from South and Central
America, the West Indies, and the Southern USA (Texas).
USA – One species of Uropygi: Mastigoproctus giganteus (Lucas, 1835) in
Texas (Thelyphonidae), the northernmost species of the order in the New World
8.3.7 Schizomida
Mexico – Thirty-six species from the genera Mayazomus, Pacal, Stenochrus,

Protoschizomus, Agastoschizomus, Sotanostenochrus (all genera, except
Stenochrus, are endemic to Mexico.)
USA (Arizona, California, Florida, and Texas) – Most species live in California
(eight, all of the genus Hubbardia Cook, 1899). Also the genera Bamazomus and
Stenochrus. All Hubbardia (Arizona and California) are endemic in the USA.
The family Protoschizomidae (gen. sp. indet.) is also recorded from Texas.
8.3.8 Araneae
Mexico – The Mexican spider fauna consists of 2295 spp. in 423 genera and 64
families of Araneomorphae (Orthognatha), no endemic families.
USA – Endemic family in the USA is the recently described Trogloraptoridae
(Griswold, Audisio, and Ledford 2012) from caves in Oregon and California, with
one genus and species Trogloraptor marchingtoni.
8.4 Mexico: The Great American Interchange of Fauna 629
8.3.9 Opilioacarida
Mexico – At least five species, all of genus Neocarus Chamberlin et Mulaik

USA (Texas, Arizona) – One species (Neocarus texanus Chamberlin et Mulaik),
marking the northern limit of Opilioacarida
8.4 Mexico: The Great American Interchange of Fauna
South of Rio Grande is Mexico – 1,972,550 km2 of lands with exceptional biodiver-

sity. High mountains form the Cordillera Volcanica Transversal. We had the chance
to visit the highest summits and to collect some arachnids and other high mountain
animals: Îrisaba (5700 m.), Popocatepetl (5492 m.), and Nevado de Toluca (4680 m.).
Between Mexico and Colombia is stretching one of the richest in biodiversity
territories on Earth – Central America. Geographically part of North America, this
territory has nevertheless many distinctive patterns and is often considered a sub-
continent (like India). There are two main definitions of the notion of Central
America. The first comprises the seven states between Mexico and Colombia:
Belize, Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica, and Panama
(total surface of 524 000 km2). According to the other concept, Central America
starts with the Tehuantepec land bridge in Mexico – the 200 km. shortest distance
between the Atlantic and the Pacific Oceans. This concept includes in Central
America the very rich in biodiversity Yucatan Peninsula – I find this concept more
natural, as Yucatan is narrowly attached to the land south of it.
Along both seashores of Mexico, the Neotropical elements penetrate north. In
the central parts extends dry plateau with coniferous forests and mostly of Nearctic
caractere (Freeland and Dietz 1971; Martin and Harrell 1957; Barrera 1962;
Escalante et al. 2007; Espinosa et al. 2000; Ferrusquía-Villafranca 1990; Halffter
1978, 1987, 2003; Marshall and Liebherr 2000; Morrone et al. 2002).
The Mexican entomologist Prof. Gonzalo Halffter introduced (Halffter 1977) a
particular term within Mexican Transition Zone, called in Spanish “Mesoamericano
de Montaña” (the mountain rain forests, populated with animals, having evolved in
the “Central American Nucleus”). Halffter (1976) describes in details the distribu-
tion of insects in Mexican Transition Zone.
Halffter’s Mexican Transition Zone is a complex area in which the Neotropical and Nearctic
regions overlap. The generalized tracks obtained correspond broadly to Halffter’s Nearctic
and Mesoamerican patterns, respectively; however, in contrast to Halffter’s Mesoamerican
pattern, the southern generalized track extends further north through the lowlands of the
Pacific and Mexican Gulf coasts (Morrone and Márquez 2001).
The Mesoamerican distributional pattern includes Neotropical taxa that evolved in situ in
humid montane habitats (Morrone 2005).
According to Morrone (2005), Mexican Transition Zone between Nearctic and

Neotropic includes five provinces: Western Sierra Madre, Eastern Sierra Madre,
Trans-Mexican Volcanic Belt, the basins Balsas, and Sierra Madre del Sur. Morrone
(2006) also includes in the biogeographical subdivision of “Latin America” the
notion of “Mexican Transition Zone.”
The development of the Mexican Transition Zone is summarized by Morrone
(2015) into five stages: (a) Jurassic-Cretaceous, four Paleoamerican cenocrons
extend in Mexico; (b) Late Cretaceous-Paleocene, dispersal from South America of
the Plateau cenocron; (c) Oligocene-Miocene, dispersal from Central America of
the Mountain Mesoamerican cenocron; (d) Miocene-Pliocene, dispersal from North
America of the Nearctic cenocron; and (e) Pleistocene, dispersal from South
America of the typical Neotropical cenocron.
Arachnogeography
The arachnofauna of Mexico is very rich and interesting as a transition zone between
two kingdoms – Holarctic (the Nearctic Region) and Neotropica (Vázquez 1995,
1996).
In this country are lacking only the suborders Palaeoamblypygi and Mesothelae.
All orders of Arachnida are represented in Mexico, some of them with many taxa.
Especially interesting are the many cave scorpions of the family Typhlochactidae
and the 272 species of Opiliones in this transitional country.
Map 8.1 Mexico and the Southern USA

Ricinulei: Pseudocellus – ◇
Amblypygi: Phrynus – ●
Paraphrynus – ▶
Acanthophrynus – ▼
Heterophrynus – ◑
Charinus – ◻
Cyphophthalmi: Shearogovea – ♣
Metasiro – ♠
Opilioacaridae: Neocarus – ☒
Caribeacarus – △
Holothyrida: Caribothyrus – ◓
Palpigradi From Mexico are known Eukoenenia hanseni Silvestri, 1913; E. coro-
zalensis Montaño et Francke, 2006; and E. tetraplumata Montaño, 2007
(Eukoeneniidae) (Condé 1986; Harvey 2013e).
Ricinulei From Mexico are known nine species of the genus Pseudocellus Platnick,
1980 (Bolivar y Pieltain 1941, 1946; Brignoli 1974; Chamberlin and Ivie 1938;
Coronado-Gutiérrez 1970; Gertsch 1971; Goodnight and Goodnight 1952; Marquez
and Conconi 1974; Valdez-Mondragón and Francke 2011).
Solifugae In Mexico are known 77 spp. of Solifugae from 12 genera and 2
families:
Eremobatidae – Eremorhax Roewer, Eremobates Banks, Eremocosta Roewer,

Eremothera Muma, Eremochelis Roewer, Chanbria Muma, Hemerotrecha
Banks
Ammotrechidae – Ammotrechella Roewer, Ammotrechula Roewer, Ammotrecha
Banks, Branchia Muma, Ammotrechesta Roewer
Most of these genera are shared with the adjacent states of the USA (all
Eremobatidae) or with Florida, Central America, South America, or the Antilleans
(Ammotrechidae) (Muma 1976; Vazquez Rojas 1981, 1995; Brookhart and Muma
1987; Brookhart and Brookhart 2006).
Scorpiones One of the richest countries with scorpions, According to Díaz Nájera
(1975), in Mexico live 134 spp. of Scorpiones, belonging to 15 genera and 4 fami-
lies – Buthidae, Chactidae, Scorpionidae (incl. Diplocentridae), and Vaejovidae.
Since this publication many changes occurred, especially with the cave scorpions
(Francke 1977, 1981, 1982; Lourenço 1994; Lourenço and Sissom 2000; Mitchell
1968, 1971; Mitchell and Peck 1977; Prendini et al. 2010; Soleglad and Fet 2005,
2006, 2008; Williams 1970).
Now (2017) the scorpions recorded from Mexico belong to 272 spp. of 37 genera
and 8 families. The biggest genus is Vaejovis C.L. Koch (61 spp.). Remarkable is the
presence of cave species of the genera Typhlochactas Mitchell, Stygochactas Vinoli
et Prendini, Sotanochactas Francke, Troglocormus Francke, and Alacran Francke
(Typhlochactidae). According to Lourenço (1994), in Mexico live 11 out of the 13
troglobite scorpions in the world. Since then, some more troglobitic genera and spe-
cies have been published (Francke 2009). The family Typhlochactidae and many
genera (Typhlochactas, Stygochactas, Sotanochactas, Troglocormus, Alacran,
Hoffmannichadrurus, Chaneke, Koloti, Balsateres, Konetontli, Kuarapu, and
others) are endemic to Mexico (Table 8.1).
Pseudoscorpiones From Mexico have been recorded 161 spp. of Pseudoscorpiones
of 63 genera and 18 families (Beier 1933; Ceballos 2004; Chamberlin 1947;
Hentschel and Muchmore 1989; Hoff 1945; Muchmore 1973b, 1975, 1977, 1982,
1986).
Genera endemic for Mexico are:

Mexichthonius Muchmore, 1975 – Campeche (one sp.)
Apohya Muchmore, 1973b (one sp.)
Vachonium J.C. Chamberlin, 1947 – Mexico, Belize (eight spp.)
Mexichelifer Muchmore, 1973b (one sp.)
Ancalochernes Beier, 1932 (one sp.)
Epichernes Muchmore, 1982 (one sp.)
Opiliones The outline of Kury and Cokendolpher (2000) presents the overall pic-
ture of the opilionid fauna of Mexico and a review of the biogeographic affinities of
the genera (we are relying on them in our analysis). In this vast country, situated in
the “zona de transition” between the Nearctic and the Palearctic, have been recorded
283 spp. (1 sp. of Cyphophthalmi, 143 Laniatores, and 139 “Palpatores”), of which
56 remained in 2000 unnamed. Further data could be found in the papers of
Cokendolpher and Lee (1993); Goodnight and Goodnight (1942, 1945, 1951, 1953,
1971, 1977); Giribet (2011), Kury (2003, 2013), Morales Soto (1980), Roewer
(1912, 1947), Shear (1977, 1980, 2006, 2010); Šilhavý (1974, 1977), and others.
Cyphophthalmi The only Cyphophthalmid recorded from Mexico is Neogovea

mexasca Shear, 1977 (endemic, troglobitic), with Neotropical affinities, now trans-
ferred to a new genus Shearogovea Giribet, 2011, with uncertain position.
Table 8.1 Scorpions in

Country Меxico USA
Mexico and the USA
Number of species 272 111
Taxa
Fam. Buthidae + +
Centruroides Marx 40 −
Chaneke Francke et al. 2 −
Tityus C.L. Koch − 1
Fam. Caraboctonidae + +
Hadrurus Thorell 6 4
Hoffmannihadrurus Fet et Soleglad 2 −
Fam. Chactidae − +
Anuroctonus Pocock 2 −
Nullibroteas Williams 1 −
Uroctonus Thorell − 3
Fam. Euscorpiidae + −
Megacormus Karsch 4 −
Plesiochactas Pocock 2 −
Troglocormus Francke 2 1
Fam. Scorpionidae + +
Bioculus Stahnke 5 −
Diplocentrus Peters 48 5
Koloti Santiban-Lopez et al. 2 −
Fam. Superstitioniidae + +
Superstitionia Stahnke 1 1
Fam. Typhlochactidae + −
Alacran Francke 3 −
Sotanochactas Francke 1 −
Stygochactas Vignoli et Prendini 1 −
Typhlochactas Mitchell 6 −
Fam. Vaejovidae + +
Balsateres G.-Sant. et Prendini 1 −
Chihuahuanus G.-Sant. et Prendini 8 4
Franckeus Soleglad et Fen 6 −
Gerschius Graham et Soleglad 2 1
Kochius Soleglad et Fen 9 2
Konetontli G.-Sant. et Prendini 5 −
Kovarikia Soleglad, Fen et Graham − 3
Kuarapu Francke et P.S. 1 −
Maaykuyak G.-Sant. et Prendini 2 1
Mesomexovis G.-Sant. et Prendini 6 1
Paravaejovis Williams 12 4
Paruroctonus Werner 15 22
Pseudouroctonus Stahnke 9 9
Serradigitus Stahnke 14 6
(continued)
Country Меxico USA
Smeringurus Haradon 2 −
Stahnkeus Soleglad et Fen 4 4
Syntropis Kraepelin 3 −
Thorellius Soleglad et Fen 3 −
Uroctonites Williams et Savary − 4
Vaejovis C.L. Koch 39 20
Vejovoidus Stahnke 1 −
Viscaino G.-Sant. et Prendini 1 −
Wernerius Soleglad et Fen − 3
The 143 spp. of Mexican “Palpatores” are distributed as follows:

Eupnoi
Subfam. Gagrellinae
Geaya Roewer, 1910 – Ten endemic spp., the genus with Neotropical
affinities
Holcobunus Roewer, 1910 – One endemic species, genus with Nearctic affin-
ities (Eastern USA)
Krusa G. et G., 1947 – Eleven endemic spp. (partly undescribed), Neotropical
affinities
Parageaya Mello-Leitão – Three endemic spp. (genus known from Uruguay
and Argentina)
Prionostemma Pocock, 1903 – Fifteen spp. (endemic and known from Central
America)
Romerella G. et G., 1943 – Two endemic spp. (Neotropical affinities)
Trachyrhinus Weed, 1892 – Two spp. with Nearctic affinities (known also
from the USA)
Subfam. Leiobuninae
Cosmobunus G.et G., 1946 – One sp. (endemic, Neotropical affinities, mis-
identified, acc. to Kury and Cokendolpher 2000)
Hadrobunus Banks, 1900 – Two endemic spp. (Nearctic affinities, Eastern
USA)
Leiobunum C.L. Koch, 1839 – Thirty-one described and 7 undescribed spp.,
including 29 endemics (Holarctic, mostly temperate)
Leuronychus Banks, 1900 – Two spp. (one endemic) (Nearctic affinities)
Nelima Roewer, 1910 – Three endemic spp. (Holarctic, mostly temperate)
Paranelima di Caporiacco, 1938 – Five endemic spp. (Neotropical affinities,
Central America)
Undescribed subfamily
Dalquestia Cokendolpher, 1984 – Four spp. (three endemic) (Nearctic
affinities)
Diguetinus Roewer, 1912 – Two endemic spp., endemic genus
Eurybunus Banks, 1893 – One described and five undescribed spp. (Nearctic
affinities)
Globipes Banks, 1893 – Two endemic spp. (Nearctic affinities)
Metopilio Roewer, 1911 – Fourteen described and six undescribed spp.
(Mixed affinities: Southwestern USA, Central America)
Fam. Protolophidae
Protolophus Banks, 1893 – Two endemic spp. (genus with Nearctic affinities,
Western USA)
Dyspnoi
Acropsopilio Silvestri, 1904 – A. chomulae Goodnight et Goodnight, 1948
(endemic)
Fam. Nemastomatidae
Subfam. Ortholasmatinae
Martensolasma Shear, 2006 (endemic genus)
Ortholasma Banks, 1894 – Several spp. from Mexico, partly undescribed
(USA, Mexico, Honduras)
Laniatores
The 143 spp. of Mexican Laniatores are distributed as follows:
Fam. Cosmetidae
Acromares G. et G., 1942 – One endemic sp. (Neotropical affinities)
Boneta G. et G., 1944 – One endemic sp. (Neotropical affinities)
Colima G. et G., 1945 – One sp. (endemic genus)
Cynorta C.L. Koch, 1839 – Nineteen end. species (Neotropical affinities)
Cynortoides G. et G., 1945 – One end. sp. (Neotropical affinities)
Cynortula Roewer, 1912 – Two end. spp. (Neotropical affinities)
Erginiperna Roewer, 1947 – One end. sp. (endemic genus)
Erginulus Roewer, 1912 – Twelve spp., including five end. (Neotropical
affinities)
Eucynorta Roewer, 1912 – Two end. spp. (Neotropical affinities)
Eucynortoides Roewer, 1912 – One end. sp. (Neotropical affinities)
Eucynortula Roewer, 1912 – One end. sp. (Neotropical affinities)
Gueroma G. et G., 1942 – One end. sp. (Neotropical affinities)
Heterovonones Roewer, 1912 – One end. sp. (Neotropical affinities, Cuba)

Holovonones Roewer, 1912 – One sp. (Neotropical affinities)
Kevonones Chamberlin, 1925 – One end. sp. (Neotropical affinities)
Metaconomma P.-C., 1905 – One end. sp. (endemic genus)
Metacynorta P.-C., 1904 – Two spp. (one end.) (Neotropical affinities)
Metavonones P.-C., 1904 – One end. sp. (Neotropical affinities)
Meterginus P.-C., 1905 – Three spp.(one end.) (Neotropical affinities)
Michella G. et G., 1942 – One end. sp. (Neotropical affinities)
Paecilaema C.L. Koch, 1839 – Three spp., two end. (Neotropical affinities)
?Palpinus – One sp. recorded, unrecognizable
Paravonones P.-C., 1904 – Three end. spp. (Neotropical affinities)
Poala G. et G., 1942 – Two end. spp. (Neotropical affinities)
Prosontes G. et G., 1945 – One end. sp. (endemic genus)
Vononana Roewer, 1927 – One end. sp. (Neotropical affinities)
Vonones Simon, 1879 – Seven spp. (six end.) (mixed affinities, Caribbean,
Ecuador, USA)
Fam. Phalangodidae – Three spp.
Guerrobunus Goodnight et Goodnight, 1945 – Three spp. (? misplaced)
Fam. Samoidae – Three spp.
Akdalima Šilhavý, 1977 – One end. sp. (Neotropical affinities, Haiti)
Arganotus Šilhavý, 1977 – One end. sp. (Neotropical affinities, Guatemala,
Haiti)
Pellobunus Banks, 1905 – One end. sp. (Neotropical affinities)
Fam. Stygnommatidae – Eight spp.
Stygnomma Roewer, 1912 – Eight spp. (seven end., one in Belize) (Neotropical
affinities)
Fam. Stygnopsidae – Forty-nine spp.
Hoplobunus Banks, 1900 – Eleven described spp., 9 undescribed spp. (19 end.)
(Nearctic affinities, Texas)
Karos Goodnight et Goodnight, 1944 – Eleven described spp., seven undescribed
spp. (endemic genus)
Mexotroglinus Šilhavý, 1977 – One sp. (endemic genus)
Paramitraceras O.P.-Cambridge, 1905 – Three spp., one endemic sp.
(Neotropical affinities)
Sbordonia Šilhavý, 1977 – Two spp. (endemic genus)
Stygnopsis Sørensen, 1902 – Two end. spp. (endemic genus)
Tampiconus Roewer, 1949 – One sp. (endemic genus)
Troglostygnopsis Šilhavý, 1974 – Two spp. (endemic genus)
Fam. Zalmoxidae – Four spp.
Ethobunus Chamberlin, 1925 – Two spp. (one endemic) (Neotropical affinities)
Metapachylus O. Pickard-Cambridge, 1905 – One sp. (endemic genus)

Pachylicus Roewer, 1923 – One sp. (Neotropical affinities)
The transitional position of Mexico reflects in its fauna of Opiliones. The analy-
sis was made by Kury and Cokendolpher (2000), and we shall borrow from it.
For Cyphophthalmi the endemic Shearogovea mexasca (Shear 1977) is clearly a
Gondwanan Neotropical element, member of the Afro-Neotropical family
Neogoveidae.
“The occurrence of the endemic genus Guerrobunus in Mexico represents the
southernmost limit of occurrence of true Phalangodidae in the New World. This
family is typically distributed in the Holarctic Region and has 79 species recorded
in the USA. This is the single example of penetration of typical Nearctic laniatorid
fauna into Mexico. The other two genera of Laniatores which reach the Southern
USA are Vonones and Hoplobunus, both typical members of the Neotropical opilio-
fauna, their maximum northern extension being here. Of the other 40 laniatorid
genera, all have closest relations with the Neotropical fauna, including the 16
endemic genera.” About half of the “Palpatores” have close relations with the
Nearctic and half with the Neotropical fauna. Acropsopilio is considered to be a
relictual “Palpatores” genus.
Since this analysis, were published the catalogue of Kury (2003) and the papers
of Shear (2006) and other authors, and the number of identified opilionid species in
Mexico was augmented.
Actually, in seven of the northern states of Mexico are known only two to five
species of Opiliones, and these states are surely understudied. A different situation
is observed with Nuevo Leon (17 spp.), Tamaulipas (34 spp.), Guerrero (30 spp.),
San Luis Potosi (26 spp.), and Veracruz (50 spp.), the highest number being in
Chiapas (53 spp.).
Amblypygi In Mexico are represented 3 genera with 19 spp. of 1 family (Armas
1994, 1995, 2006; Armas and Gadar 2004; Bilimek 1867; Butler 1873; Gervais
1842; Mullinex 1975, 1979; Pocock 1894, 1902; Quintero 1981; Rowland 1973;
Wood 1893).
Fam. Phrynidae
Subfam. Phryninae
Acanthophrynus Kraepelin, 1899 – Mexico, USA (Arizona, California) (one sp.)
Paraphrynus Moreno, 1940 – (16 spp., 11 Mexican, all endemics) Five known from
the USA, Central America, and West Indies
Phrynus Lamarck, 1801 – Twenty-seven American spp., in Mexico are known
(seven spp., four endemic); the others known also from South America, USA
(Texas, Florida), Central America, and West Indies.
The family Phrynidae is clearly Neotropical, despite the finding of one species in
Indonesia (Flores).
Thelyphonida (Uropygi) From Mexico has been recorded only one member of
Mastigoproctus Pocock, 1894 – M. giganteus (Lucas, 1835) (Thelyphonidae,
Uroproctinae). The genus is known from South and Central America, the West
Indies, and the Southern USA (Texas) (Harvey 2003). The localities in Mexico and
Texas mark the northern limit of Uropygi in the New World.
Schizomida Mexico is very rich in Schizomida, especially in caves (Cokendolpher

and Reddell 1988; Rowland 1971a, b, 1973a, 1973b; Rowland and Reddell 1977;
Reddell and Cokendolpher 1991, 1995). So far 41 species have been recorded, from
the genera Mayazomus, Pacal, Stenochrus, Protoschizomus, Agastoschizomus,
and Sotanostenochrus (all genera, except Stenochrus, are endemic to Mexico).
Practically endemic is the family Protoschizomidae (known also in southern Texas).
This fauna is different from the fauna of the neighboring regions (Central America
and the Southern USA), largely because of the cave endemism. Actually, Mayazomus
with its seven species from Chiapas and Tabasco, is also more Central American
than connected to the rest of Mexico, so are two of the three species of Pacal Reddell
et Cokendolpher. As a whole, Mexico has the third most numerous list of Schizomida
in the world (after Australia and Cuba).
Araneae According to Hoffmann (1976), the rich spider fauna of Mexico consists
of three suborders, Orthognata, Labidognatha, and Cribellatae, and 50 families.
Now (2017) this number is greatly increased. According to Francke (2011) and
Aguayo-Morales et al. (2012), the Mexican spider fauna consists of 2295 spp. in
423 genera and 64 families. They all belong to contemporary Araneomorphae
(Orthognatha).
Opilioacarida The modern research on Mexican Opilioacarida by Vásquez and

Klompen (2002, 2004, 2009) revealed the existence in this country of at least five
species, all of the genus Neocarus Chamberlin et Mulaik. They have been collected
in Baja California Sur (N. bajacalifornicus), Jalisco (N. bajacalifornicus chame-
laensis), in Veracruz (N. veracruzensis), and in Campeche and Quintana Roo (N.
calakmulensis, N. nohbecanus, N. siankaanensis). The other members of Neocarus
are known from the neighboring USA (Texas, Arizona), Nicaragua, Cuba, and
South America.
Acariformes
Trombidiformes – Prostigmata
Fam. Erythraeidae
Endemic genus: Tepotztlana Hoffman et Mendez, 1973
Parasitiformes
Ixodida
Fam. Argasidae – Nothoaspis reddelli Keirans et Clifford, 1975 (endemic genus
and species, Yucatan).
8.5 Arachnida of the Antilleans and Mesoamerica 639
8.5 Arachnida of the Antilleans and Mesoamerica
8.5.1 Central America

Central America is extending from the Isthmus of Tehuantepec in southern Mexico

southeastward to the Isthmus of Panama where it connects to the Colombian Pacific
Lowlands in northwestern South America. Alternatively, the Trans-Mexican
Volcanic Belt delimits the region on the north. Central America has an area of some
592,000 square kilometers, shared between seven states, Belize, Guatemala,
Honduras, Costa Rica, Nicaragua, El Salvador, and Panama, but the Mexican
Yucatan is also part of it. Most of Central America rests atop the Caribbean Plate
(Halffter 1976, 1987; Morrone 2001; Khudoley and Meyerhoff 1971; Ryan). Central
America has been assigned to different zoogeographical units (Kraus 1960, 1962)
but is usually considered part of the Neotropical Region. Using the comparative
chorological method (De Lattin 1957), Müller (1972) outlined in Central America
(Yucatan included) seven centers of dispersal: Central American Rainforest, Central
American Montane Forest, Yucatan, Central American Pacific, Coco, Costa Rica,
and Talamanca-Paramo.
According to Morrone (2005), Central America is included in the Caribbean
Subregion of the Neotropical Region (Mesoamerican Dominion with five
provinces).
Central America is part of the Mesoamerican Biodiversity hot spot, with 7% of
the world’s biodiversity. As a bridge between North and South America, Central
America has many species from the Nearctic and the Neotropic ecozones.
Halffter (1978) says that his “Patron Mesoamericano de Montaña” (Mountain
Mesoamerican Pattern) is composed of elements having evolved in the Central
American Nucleus (determined by Halffter 1978, as “La mesa Central y las dos
cordilleras de Chiapas, asi como las tierras altas y sus declives de Guatemala,
Honduras, El Salvador y norte de Nicaragua”).
The Entomologist Krzhanovskiy (2002) outlined the fauna of Central American
Subregion and included in it the “Galapagos Province.”
In Central America (from the Isthmus of Tehuantepec to Panama), we can distin-
guish clearly two mountain massifs: north of the Central American Nucleus, includ-
ing 80% of the high grounds (above 600 m.) of Central America and in the south la
Sierra de Talamanca in Costa Rica and the west of Panama (Halffter 1978).
Arachnogeography
Central America, bridge between North and South America and facing the Caribbean,
is zoogeographically important and is with rich and varied nature (Armas 2004). In
the seven countries between Mexico and Colombia are represented 12 orders:
Ricinulei (18 spp. of 2 genera), Solifugae (12 spp. of 5 genera, all Ammotrechidae),
Scorpiones (6 fam., 11 genera – none is endemic), Pseudoscorpiones (13 fam., 33
gen., 1 endemic), Opiliones (1 genus of Eupnoi, 56 genera of 8 families of Laniatores,
mostly Cosmetidae), Amblypygi (three genera of the families Phrynidae and
Charinidae, four endemic spp.), Uropygi (three spp. of two genera, one is endemic),
Araneae (many), Opilioacarida (two genera), plus the superorders Acariformes and
Parasitiformes (Banks 1909a; Kury 2003; Roewer 1954).
Not recorded in Central America are Palpigradi and Holothyrida and the subor-
ders Palaeoamblypygi, Cyphophthalmi, Dyspnoi, and Mesothelae. If we consider
the entire area of Central America (incl. Yucatan) and the Caribbean, we may say
that this is the most varied area in Arachnida in the world – all orders and almost all
suborders are present (except for Palaeoamblypygi and Mesothelae).
Palpigradi – Not recorded from Central America
Ricinulei In the 7 countries of Central America (without South Mexico) are regis-
tered 18 species of Ricinulei (Cooke and Shadab 1973; Fage 1921, 1938; Merret
1960; Platnick and Pass 1982; Platnick and Shadab 1981; Chamberlin and Ivie 1938).
Cryptocellus Westwood, 1874 – Panama, Costa Rica, Honduras, Nicaragua (12

spp.)
Pseudocellus Platnick, 1980 – Guatemala, Honduras, El Salvador, Panama (six
spp.)
These are the two genera of Ricinulei (Ricinoididae), known in the New World.
They are represented also in Mexico and the Caribbean (Cuba). In Yucatan, which
is geographically also part of Central America, are known five more Ricinuleid
species.
Solifugae From the 7 countries of Central America (without South Mexico) are
known 12 spp. of Solifugae of 5 genera, all belonging to Ammotrechidae family
(Armas 1993, 1996, 2000, 2004, with supplement; Muma 1970, 1976, 1986; Pocock
1902; Roewer 1934). The highest diversity has been recorded in Nicaragua (six
spp.) and Guatemala (five spp.). According to Armas (1996), this is only half of the
presumed number of species in Central America (23).
Ammotrechinae
Ammotrecha Banks, 1900
Ammotrecha limbata (Lucas, 1835) – Guatemala
A. nigrescens Roewer, 1934 – Costa Rica, Guatemala
A. picta Pocock, 1902 – Guatemala
A. stollii (Pocock, 1895) – Costa Rica, Guatemala, USA
Ammotrechella Roewer, 1934
Ammotrechella pseustes (Chamberlin, 1925) – Panama, California, Puerto Rico

tabogana Chamberlin, 1919 – Panama
Ammotrechesta Roewer, 1934
Ammotrechesta garcetei Armas, 1993 – Nicaragua
maesi Armas, 1993 – Nicaragua
schlueteri Roewer, 1934 – Honduras
brunnea Roewer, 1934 – Costa Rica
tuzi Armas, 2000 – Mexico
Innesa Roewer, 1934
Innesa vittata (Pocock, 1902) – Guatemala
Scorpiones The list of scorpions in Central America (Mexico excluded) of Armas
and Maes (1998) contains data on 38 species of 11 genera and 5 families. From
them 24 species are Central American endemics. Meanwhile, some changes
occurred in the names of families, the place of the genera, etc. Now there are 11
genera but of 6 families (Francke 1978; Francke and Stockwell 1987; Stockwell
1988; Lourenço 1996c; Lourenço and Méndez 1984; Teruel and Stockwell 2002;
Viquez 1999) (Table 8.2).
Pseudoscorpiones In the 7 countries of Central America are recorded pseudoscor-

pions of 33 genera and 13 families (Chthoniidae, Tridenchthoniidae, Bochicidae,
Neobisiidae, Syarinidae, Ideoroncidae, Sternophoridae, Garypinidae, Atemnidae,
Olpiidae, Cheliferidae, Chernetidae, Withiidae) (Beier 1931, 1932, 1953, 1955,
1976; Heurtault 1998; Hoff 1944; Mahnert 1987; Muchmore 1973).
In the six countries of Central America (no data for Honduras), the number of the
pseudoscorpion species is as follows (Harvey 2003): Belize, 10; Guatemala, 17;
Costa Rica, 23; El Salvador, 10; Nicaragua, 1; and Panama, 15.
Endemic genus for Central America:
Fam. Chernetidae
Coprochernes Beier – Costa Rica
Opiliones
Many authors, Cokendolpher and Cokendolpher (1984), Goodnight and Goodnight
(1983), Kury (2003), Cruz-López et al. (2016), Pickard-Cambridge (1904–1905),
Roewer (1949a), Shear (2010), Šilhavý (1979), Townsend et al. (2010), and others,
have explored the rich opilionid fauna of Central America.
Cyphophthami – Unknown
Eupnoi
Geaya Roewer, 1910 – Belize, Costa Rica
Prionostemma Pocock, 1903 – Guatemala, El Salvador, Panama, Costa Rica
Dyspnoi
Fam. Nemastomatidae
642
Table 8.2 Scorpiones in Central America (Without Mexico)

Country Belize Guat. Honduras Salvador Nic. Costa Rica Panama Central America Congo DR Congo Gabon Centr. AR Angola
Number of species 4 16 9 2 4 15 17 8 18 14 9 4 24
Fam. Buthidae + + + + + + + + + + + +
Ananteris Thorell − − − − − 1 1 1
Centruroides Marx 1 6 5 1 3 4 4 10 10
Isometrus Ehrenberg − − − − − 1 1 1
Tityus C.L. Koch − − − − − 6 10 8
Fam. Chactidae − − − − − + + +
Neochactas Sol. et Fet − − − − − - 1 1
Chactas Gervais − − − − − 1 2 2
Fam. Euscorpiidae − + + − − − − +
Plesiochactas Pocock − − − − − − −
Fam. Hormuridae − + − − − + + +
Opisthacanthus Peters − 1 − − − 1 1 2
Fam. Scorpionidae + + + + + + − +
Diplocentrus Peters 4 6 3 − − − − 8
Didymocentrus Kraep. − − 1 1 2 1 −
Fam. Vaejovidae − + − − − − + +
Vaejovis C.L. Koch − 1 − − − − 1 1
8 Some Peculiarities of the Distribution of Arachnida
Ortholasmatinae
Trilasma Goodnight et Goodnight – Honduras (one sp.)
Laniatores
The catalogue of Kury (2003) enumerates 198 spp. of Laniatores from the 7 coun-
tries of Central America of 56 genera and 8 families as follows:
Fam. Cosmetidae
Acromares Goodnight et Goodnight – Belize (one sp.)
Bokwina G.et G. – Belize (one sp.)
Boneta G. et G. – Guatemala (one sp.)
Cosmetus Perty – Panama (one sp.)
Cynorta C.L. Koch – Belize, Costa Rica, Panama, Honduras, Guatemala, El
Salvador (20 spp.)
Cynortellana Roewer – Costa Rica (one sp.)
Cynortoperna Roewer – Costa Rica (one sp.)
Cynortula Roewer – Costa Rica, El Salvador, Nicaragua, Guatemala (nine spp.)
Erginoides Pickard – Cambridge-Panama, Costa Rica (two spp.)
Erginulus Roewer – Guatemala, Honduras, Belize, Costa Rica, El Salvador (21
spp.)
Eucynorta Roewer – Costa Rica, Panama, El Salvador, Guatemala (24 spp.)
Eucynortella Roewer – Guatemala, Panama (five spp.)
Eucynortoides Roewer – Costa Rica (one sp.)
Eucynortula Roewer – Costa Rica, Nicaragua, Guatemala, Belize, Panama
(seven spp.)
Eugnidia Roewer – Costa Rica (one sp.)
Eupoecilaema Roewer – Costa Rica, Panama (two spp.)
Flirtea C.L. Koch – Honduras, Panama, Costa Rica (two spp.)
Holovonones Roewer – Belize, Costa Rica, Guatemala (one sp.)
Kevonones Chamberlin – Costa Rica (one sp.)
Metacynorta Pickard – Cambridge-Guatemala (one sp.)
Metarhaucus Pickard – Cambridge-Costa Rica (one sp.)
Metavonones Pickard – Cambridge-Costa Rica (two spp.)
Meterginus Pickard – Cambridge-Guatemala, Costa Rica, El Salvador (six spp.)
Paecilaema C.L. Koch – Costa Rica, Panama, Guatemala, Honduras, Belize (12
spp.)
Paecilaemana Roewer – Panama, Costa Rica (two spp.)
Paracynorta G. et G. – Panama (one sp.)
Paravonones Pickard – Cambridge-El Salvador (one sp.)
Poecilaemula Roewer – Costa Rica (one sp.)
Reimoserius Roewer – Costa Rica (one sp.)
Tajumulcia G. et G. – Guatemala (one sp.)
Vonones Roewer – Belize (one sp.)
Vononesta Roewer – Guatemala (one sp.)
Vononula Roewer – Guatemala (one sp.)
According to Townsend et al. (2010), the family Cosmetidae in Central America

registered 133 spp. of 33 genera.
Fam. Cranaidae
Comboyus Roewer – Panama (one sp.)
Phareicranaus Roewer – Costa Rica, Panama (two spp.)
Fam. Gonyleptidae
Glysterus Roewer – Costa Rica (six spp.)
Hernandaria Soerensen – Costa Rica (two spp.)
Hernandarioides Picard – Cambridge-Panama (one sp.)
Nesopachylus Chamberlin – Panama (two spp.)
Fam. Manaosbiidae
Barrona Goodnight C. J. et M. L. Goodnight – Panama (one sp.)
Bugabitia Roewer – Panama (one sp.)
Poassa Roewer – Costa Rica (one sp.)
Zygopachylus Chamberlin – Panama (one sp.)
Fam. Samoidae
Arganotus Šilhavý – Guatemala (one sp.)
Neocynortina Goodnight C. J. et M. L. Goodnight – Costa Rica (one sp.)
Pellobunus Banks – Panama (two spp.)
Fam. Stygnommatidae
Stygnomma Roewer – Belize, Costa Rica (six spp.)
Fam. Pyramidopidae
Jarmilana Cruz-López et al. – Belize (one sp., troglobite)
Fam. Stygnopsidae
Paramitraceras P. – Cambridge-El Salvador, Guatemala, Costa Rica (two spp.)
Fam. Zalmoxidae
Ethobunus Chamberlin – Panama, Guatemala, El Salvador, Costa Rica (15 spp.)
Pachylicus Roewer – Panama, Costa Rica (eight spp.)
Panopiliops Roewer – Costa Rica (two spp.)
Phalangoduna Roewer – Costa Rica, Panama (one sp.)
Stygnoleptes Banks – Costa Rica, Panama, El Salvador (three spp.)
Family uncertain
Costabrimma Goodnight et Goodnight – Costa Rica (three spp.)
Isaeolus Roewer – El Salvador (one sp.)
Amblypygi The amblypygids are represented in Central America by the family
Phrynidae and the genera Phrynus Lamarck (four spp. in Nicaragua, Guatemala,
Honduras, Belize, Costa Rica, Panama) and Paraphrynus Moreno (four spp. in
Belize, Guatemala, Costa Rica, El Salvador, Nicaragua, Panama, Honduras). Both
genera live also in South America, Mexico, and the Antilleans. The genus Charinus
(fam. Charinidae) is found in Panama (Armas 2004; Armas de and Gonzalez 2001;
Armas de and Maes 2000; Armas de and Víquez 2001; Avila Calvo and Armas 1997;
Gervais 1842; Mullinex 1975, 1979; Pocock 1893, 1894, 1902; Roewer 1954;
Quintero 1981; Víquez et al. 2012).
There are four species endemic to Central America (between Mexico and
Colombia):
Phrynus Lamarck – Ph. maesi Armas, 1996 (Nicaragua), Ph. parvulus Pocock,
1902 (Guatemala, Belize)
Paraphrynus Moreno – P. emaciatus Mullinex, 1975 (Guatemala), P. leptus
Mullinex, 1975 (Guatemala)
Thelyphonida (Uropygi) In Central America are found five species of Uropygi
and three endemic genera Valeriophonus (Viquez et de Armas, 2005, Mayacentrum
Viquez et Armas, 2006, and Mimoscorpius Butler, 1872). They live in Costa Rica
(Valeriophonus nara Valerio), El Salvador (Mayacentrum tantalus Roewer),
Honduras (M. pijol Viquez et de Armas), Belize (M. guatemalae Viquez et de
Armas), and Guatemala (Mimoscorpius pugnator Butler) (Roewer 1954; Valerio
1981; Viquez and de Armas 2005, 2006; Armas and Viquez 2007).
Schizomida In Central America (between Mexico and Colombia) are recorded

nine species, but there is also unidentified material from many places (Armas et al.
2010; Armas and Viquez 2010; Brignoli 1973; Rowland and Reddell 1977; Reddell
and Cokendolpher 1995). Besides the largely distributed Stenochrus portoricensis
(Guatemala, Honduras, Nicaragua, Belize), from Central America are known the
genera Heteronochrus (Guatemala), Hansenochrus (Costa Rica, Panama),
Rowlandius, Piaroa and Surazomus (Costa Rica), and Belicenochrus from Belize.
Surazomus Reddell et Cokendolpher is known also from many South American
countries. Rowlandius Reddell et Cokendolpher is widespread in the Caribbean
Islands. Hansenochrus Reddell et Cokendolpher is found also in South America and
the Caribbean.
Map 8.2 Schizomida in Central America and the Caribbean

Belicenochrus – ◊
Stenochrus – ■
Hansenochrus – ▲
Rowlandius – ▼
Surazomus – ●`
Stewardpeckius – ♣
Antillostenochrus –
Piaroa –
Luisarmasius –
Cokendolpherius –
Cubacanthozomus –
Heterocubazomus –
Caribezomus – ►
Heteronochrus – ◄
Cubazomus – X
Guanazomus – □
Reddellzomus – ♦
Troglocubanus – ○
Araneae In Central America (without Mexico) are known at leat 59 families of

spiders: Microstigmatidae, Dipluridae, Nemesiidae, Theraphosidae, Paratropidae,
Barychelidae, Cyrtaucheniidae, Idiopidae, Ctenizidae, and Actinopodidae from
infraorder Mygalomorphae and Filistatidae, Drymusidae, Scytodidae, Sicariidae,
Leptonetidae, Ochyroceratidae, Telemidae, Pholcidae, Caponiidae, Tetrablemmidae,

Oonopidae, Segestriidae, Hersiliidae, Oecobiidae, Mimetidae, Deinopidae,
Uloboridae, Anapidae, Araneidae, Linyphiidae, Synphytognatidae, Tetragnathidae,
Theridiidae, Theridiosomatidae, Ctenidae, Lycosidae, Oxyopidae, Pisauridae,
Senoculidae, Trechaleidae, Udubidae, Agelenidae, Amaurobiidae, Anyphaenidae,
Dictynidae, Hahniidae, Sparassidae, Selenopidae, Zodariidae, Zoropsidae,
Clubionidae, Miturgidae, Titanoecidae, Gnaphosidae, Prodidomidae, Philodromidae,
Thomisidae, Salticidae, and Corinnidae from infraorder Araneomorphae. Most of
the endemic genera have been described from Panama by Chikering (1946, Panama).
No endemic families were noted.
Some endemic spider genera in Central America and the Caribbean:
Fam. Microstigmatidae
Micromygale Platnick et Forster, 1982 – Panama
Fam. Theraphosidae
Aenigmarachne Schmidt, 2005 – Costa Rica
Antillena Bertani, Huff et Fukushima, 2017 – Dominican Republic
Barropelma Chamberlin, 1940 – Panama (one sp.)
Caribena Fukushima et Bertani, 2017 – Puerto Rico, Cuba, US Virgin Islands,
Martinique
Crassicrus Reichling et West, 1996 – Belize
Longilyra Gabriel, 2014 – El Salvador
Mygalarachne Ausserer, 1871 – Honduras
Reichlingeria Rudloff, 2001 – Belize
Sphaerobothria Karsch, 1879 – Costa Rica, Panama
Stichoplastoris Rudloff, 1997 – El Salvador, Costa Rica, Panama
Fam. Ochyroceratidae
Fageicera Dumitrescu et Georgescu, 1992 – Cuba
Fam. Barychelidae
Fam. Agelenidae
Neowadotes Alayón, 1995 – Hispaniola
Fam. Cyatholipidae
Pokennips Griswold, 2001 – Jamaica
Fam. Tetragnathidae
Ancinosphenus Simon, 1895 – West Indies
Antillognatha Bryant, 1945 – Hispaniola
Hispanognatha Bryant, 1945 – Hispaniola
Fam. Linyphiidae
Primerigonina Wunderlich, 1995 – Panama
Sthelota Simon, 1894 – Panama, Guatemala
Fam. Filistatidae
Antilloides Breskovit et al., 2016 – Cuba, Dominican Republic, Virgin Islands,
Puerto Rico
Fam. Paratropididae
Anisaspis Simon, 1891 – Saint Vincent
Fam. Amaurobiidae
Tugana Chamberlin, 1948 – Cuba, Hispaniola
Fam. Anyphaenidae
Thaloe Brescovit, 1993 – Cuba, Hispaniola
Fam. Theridiidae
Jamaitidion Wunderlich, 1995 – Jamaica
Fam. Liocranidae
Laudetia Gertsch, 1941 – Dominican Republic, Puerto Rico
Mesobria Simon, 1897 – Saint Vincent (one sp.)
Fam. Ctenidae
Ciba Bloom et al., 2014 – Cuba, Dominican Republic
Ohvida Polotow et Brescovit, 2009 – Cuba, Puerto Rico, Bahamas
Trujillina Bryant, 1948 – Hispaniola, Puerto Rico
Fam. Pholcidae
Bryantina Brignoli, 1985 – Cuba
Ciboneya Perez, 2001 – Cuba
Platnicknia Özdikmen et Demir, 2009 – Cuba
Tainonia Huber, 2000 – Hispaniola
Fam. Tetrablemmidae
Micromatta Lehtinen, 1981 – Belize
Fam. Miturgidae
Hoedillus Simon, 1898 – Guatemala, Nicaragua
Fam. Prodidomidae
Caudalia Alayón, 1980 – Cuba
Cubanopillus Alayón et Platnick, 1993 – Cuba
Fam. Sparassidae
Decaphora Franganillo, 1931 – Cuba
Defectrix Petrunkevitch, 1925 – Panama
Fam. Thomisidae
Rejanellus Lise, 2005 – Cuba, Puerto Rico, Hispaniola
Fam. Salticidae
Albionella Chikering, 1946; Banksetosa Chickering, 1946; Carabella
Chickering, 1946
Gorgasella Chickering, 1946; Micalula Strand, 1932; Monaga Chickering,1946
Orvilleus Chickering, 1946; Toloella Chickering, 1946; Udalmella Galiano,
1994
Uluella Chickering, 1946 – All from Panama
Allodecta Bruyant, 1950 – Jamaica
Antillattus Bryant, 1943 – Hispaniola, Cuba
Bythocrotus Simon, 1903 – Hispaniola
Caribattus Bryant, 1950 – Jamaica
Cerionesta Simon, 1901 – Saint Vincent
Commoris Simon, 1902 – Guadeloupe, Dominica
Corticettus Zhang et Maddison, 2012 – Puerto Rico, Hispaniola
Paraplexippus Franganillo, 1930 – Cuba
Parasaitis Bruyant, 1950 – Jamaica
Parathiodina Bruyant, 1943 – Hispaniola
Popcornella Zhang et Maddison, 2012 – Hispaniola, Puerto Rico
Truncattus Zhang et Maddison, 2012 – Hispaniola
Opilioacarida From Central America (between Mexico and Colombia) are known
two spp. of Opilioacarida, Caribeacarus panamensis Vásquez et Klompen (Panama)
and Neocarus nicaraguensis (Vásquez et Klompen) (Nicaragua), but we can add
three species of Neocarus Chamberlin et Mulaik, 1942, from Yucatan Peninsula in
Mexico. Both Neocarus and Caribeacarus live also on the Antilleans. The species
are (so far) endemic (Vásquez and Klompen 2002, 2009).
Parasitiformes Ref.: Beron (2014), Chant and Baker (1965), Hoffmann (1962).
The purely Neotropical family of Spelaeorhynchidae (parasites on bats of the

families Phyllostomatidae and Mormoopidae) is found, together with these bats, in
Cuba, Jamaica, Puerto Rico, and Dominican Republic but also in Central America.
Holothyrida – Unknown
8.6 Antillean (Caribbean) Islands

“The West Indies and Central America, the question of the relationships between Atlantic
and Pacific Ocean, between North and South America, as well as the question of possible
land-bridges between the islands on one hand and the continent on the other, have been
dealt with in an almost interminable procession of biogeographical and geological publica-
tions.” Rutten (1935)
Antillean islands form the greater part of the West Indies in the Caribbean. The
Antilleans are divided into two major groups: the “Greater Antilles” to the north and
west, including the larger islands of Cuba, Jamaica, Hispaniola (Haiti and the
Dominican Republic), and Puerto Rico, and the smaller “Lesser Antilles” on the
southeast – comprising the northerly Leeward Islands, the southeasterly Windward
Islands, and the Leeward Antilles just north of Venezuela. The Bahamas, though
part of the West Indies, are generally not included among the Antillean islands.
Geologically, the Greater Antilles are made up of continental rock, as distinct from
the Lesser Antilles, which are mostly young volcanic or coral islands.
Cuba – area of 109,886 km2, highest point Pico Turquino (1974 m.) (Isla de la
Juventud, 2199 km2). Remain only ca. 15% of the original forest cover.
Hispaniola – Area 76 480 km2, highest point Pico Duarte (3087 m.)
Jamaica – Area 10 990 km2, highest point Blue Mountain Peak (2256 m.)
Puerto Rico – Area 9104 km2, highest point Cerro de Punta (1338 m.)
The Lesser Antilles, also known as the Caribbees, are part of the Antilles, which
together with the Bahamas, the Cayman Islands, the Turks and Caicos Islands, and
Greater Antilleans form the West Indies. The islands are a long partly volcanic
island arc, most of which wrap around the Eastern end of the Caribbean Sea on the
Western boundary with the Atlantic Ocean.
The two main groups of the Lesser Antilleans are the Windward Islands in the
south and the Leeward Islands in the north. The Leeward Antilleans in the west form
the third group; they consist of the Dutch islands just off the coast of Venezuela, plus
a group of Venezuelan islands. The main Lesser Antilles are (from north to south to
west):
Leeward Islands: Virgin Islands, Anguilla (91 km2, Saint Martin (87 km2, 414 m –
Pic Paradis), Saint-Barthélemy, Saba, Saint Eustatius, Saint Kitts, Nevis, Antigua,
Redonda, Montserrat, Guadeloupe, Dominica
8.6 Antillean (Caribbean) Islands 651
Windward Islands: Martinique, Saint Lucia, Saint Vincent, Grenadines

Barbados (a continental island found 160 km east of the Windward chain) – 430 km2,
336 m (Mount Hillaby)
Grenada – 344 km2, 840 m (Mount Saint Catherine)
Tobago – 300 km2
Trinidad – 4768 km2
Trinidad and Tobago are situated on the shelf of South America, 11 km far from the
mainland.
Dutch Antilles – Islands north of the Venezuelan coast (from west to east)
Aruba – 193 km2, 188 m (Jamanota)
Curaçao – 444 km2, 375 m (Christoffelberg)
Bonaire – 294 km2, 240 m (Brandaris)
Many papers deal with the paleogeography of the Caribbean. Among them are
Blair Hedges (2001, 2006), Bonatti and Gardner (1973), Burke et al. (1984), Curtis
et al. (2001), Freeland and Dietz (1971), Hedges (1982, 2001), Hedge et al. (1994),
Iturralde-Vinent and MacPhee (1999), Jeannel (1939), Khudoley and Meyerhoff
(1971), Koopman (1959), Meschede and Frish (1998, 2001), Meyerhoff and
Meyerhoff (1972), Morrone (2001), Page and Lydeard (1994), Schuchert (1935),
Weyl (1964), Woodring (1954), Wyatt Durham (1985), and others.
“A major deformation began in the Miocene, with folding of the Lower Tertiary
sediments and uplifting of the cores of the present-day islands. During the Late
Miocene and Pliocene, extensive peneplains were formed, which, as a result of the
youngest crustal movements in the Antilles, now stand at elevations as great as 2000
m” (Weyl 1966).
“Sykes et al. have demonstrated that the Caribbean plate has moved east-
northeast about 1400 km since late Eocene time (38 Ma). This movement changes
or affects the interpretation of many biogeographic problems of that region” (Wyatt
Durham 1985).
“In the Middle Eocene ends entirely the movement of the plate and Caribbean
Sea and its surroundings acquired approximately their actual configuration”
(Iturralde-Vinent 1975).
The Antilles and the West Indies in broader sense (including the Bahamas Bank)
have been interesting to biogeographers even since Wallace (1881). The Greater
Antilles (Cuba, Hispaniola, Jamaica, and Puerto Rico) are large enough to allow
radiative speciation within the islands and between them and even to disperse some
lines to the mainland (some examples among the vertebrates have been quoted by
Blair Hedges (2006)). This author made an analysis of the paleogeography of the
Antilles and the identity of the 1319 native terrestrial vertebrates on the islands. The
problem of how and when these animals arrived to the islands has been discussed
(Darlington 1938; Weyl 1973; Morrone 2005, and many others).
Among the non-flying terrestrial vertebrates, the endemicity is high: nearly all
173 spp. of amphibians, 96% of the 499 spp. of reptiles, 96% of the freshwater
fishes, but only 35% of birds and 50% of bats. It would be interesting to compare
these figures with the various orders of Arachnida (although with the vertebrates, we
have to deal with classes). Thanks to many local and foreign researchers, the general
picture of the Arachnida on the Antilles is more or less complete and a sound basis
for discussions.
Actually, the main argument is centered on the importance (and feasibility) of the
three ways of “arrival” of the animals: dispersal, vicariance, and land bridges. The
age of the various groups is also important. There are few fossils of Arachnida on
the islands, mostly from Dominican amber (Miocene 15–20 million years to Upper
Eocene ca. 30–45 million years), so hypotheses should be based on speculations.
Morrone (2001) includes the Antilleans as 7 of the 29 provinces of the Caribbean
Subregion of Neotropical Region (together with Central America, most of Mexico,
Ecuador, Galapagos, and other areas).
According to Morrone (2005), the Antillean Dominion is divided into eight prov-
inces: the Yucatan Peninsula province (the states of Yucatan, Campeche, and
Quintana Roo), Bahamas province, Cuba province, the Cayman Islands province,
Jamaica province, Hispaniola province, Puerto Rico province, and Lesser Antilles
province. These provinces reflect the distribution of some orders of insects in the
area.
Morrone (2017) provided a detailed regionalization of the Neotropics, dividing
the area in three subregions: Antillean, Brazilian, and Chacoan.
Based on insects (mostly Coleoptera), Krzhanovskiy (2002) considered the
Antillean Subregion part of the Neotropical Region. For him the explanation is sim-
ple: volcanic archipelago raised in the late Tertiary and populated by casual migrants
from Central and South America. Some speciation took place after that.
Arachnogeography
The arachnofauna of the islands is varied and with many endemics, well studied by
Cuban, Romanian, and other specialists (Armas 1982, 2004; Banks 1909b; Perez-
Gelabert 2008; Teruel and de Armas 2005; Armas and Avila 2015). Represented are
all orders. Remarkable are the findings of an endemic genus and species of
Holothyrida, many endemic genera of Schizomida, and very rich fauna of scorpions
(130 spp., only in Cuba there are 55 spp.). According to these authors, there is a
zoogeographical barrier between Trinidad and Tobago on one side and Grenada on
the other, “the bond line.” Lourenço (loc. cit.) indicates that the number of scorpion
species doesn’t seem to be correlated negatively with the distance from South
America but rather with the surface of the islands – five on Martinique, three on
Saint Lucie, and one on each of the smaller islands.
Lourenço (1999b) made an important analysis of the origins and the affinities of
the scorpions of the Greater Antilleans. He indicates the Northern and Central
American origins of some elements (Centruroides) and the “undoubtedly” Southern
American origin of others (Ischnuridae = Hemiscorpiidae and other Buthidae). The
explanation is the formation of a “cluster of islands,” known as Proto-Antilleans,
which drifted to South America, allowing the passage of some elements from this
continent. We may indicate that, studying the cave Isopoda of Cuba, Vandel (1973)
stressed on their exclusively South American origin.
Some attempts have been made to explain the connections of the present Greater
Antilleans with North or Central America by the lowering by 100–150 m during the
Pleistocene glaciations (Donn et al. 1962 and others) and the supposed “stepping
stones” as a consequence. However, we find hard to explain why all this rich arach-
nofauna used these “stones” and many other taxa did not. In the analysis of Weyl
(1964), no connection between the islands and any continental ground is shown
after the probable “bridge” between Jamaica-Hispaniola and Central America (the
so-called Sapperland) in the Paleocene (65–55 Ma ago).
On the islands are lacking the suborders Cyphophthalmi, Dyspnoi,
Palaeoamblypygi, and Mesothelae.
Palpigradi The only troglobite Palpigradi in the New World (Eukoenenia orghi-
dani Condé et Juberthie) was described from Cuba. Some non-described Palpigradi
have been recorded from Guadeloupe and Haiti, and the subspecies Eukoenenia
berlesei virginea Condé (1984) was described from the Virgin Islands (Condé 1984,
1986; Condé and Juberthie 1982; Remy 1948).
Solifugae From the Antilleans are known three genera of Solifugae, two of them
endemic, all belonging to Ammotrechidae (Armas 1994, 2004; Armas and Teruel
2005; Muma 1970, 1976, 1986; Muma and Nezario 1971), as follows:
Ammotrechella apejii Muma, 1981 – Jamaica (endemic)

bahamica Muma, 1986 – Bahamas (endemic)
bonariensis (Werner, 1925) – Bonaire (endemic)
[Ammotrechona] cubae (Lucas, 1835) – Cuba (endemic)
geniculata (C.L. Koch, 1842) – Bahamas, Curacao, Saint Vincent, Guadeloupe, also
USA, Mexico, and Panama
jutisi Armas et Teruel, 2005 – Cuba (endemic)
hispaniolana Armas et Alegre, 2001 – Dominican Republic (endemic)
maguirei Muma, 1986 – Caicos (endemic)
pallida Muma et Nezario, 1971 – Puerto Rico (endemic)
pseustes (Chamberlin, 1925) – Puerto Rico, Panama
Ammotrechinus gryllipes (Gervais, 1842) – Jamaica, Haiti (endemic genus and
species)
Antillotrecha iviei Armas, 2002 – Leeward Islands (endemic genus for the
Caribbean)
fraterna Armas, 1994 – Dominican Republic (endemic sp. for Dominican Rep.)
disjunctodens Armas et Teruel, 2005 – Cuba (endemic sp.)
Ammotrechella apejii Muma, 1981 – Jamaica (endemic)
bahamica Muma, 1986 – Bahamas (endemic)
bonariensis (Werner, 1925) – Bonaire (endemic)
[Ammotrechona] cubae (Lucas, 1835) – Cuba (endemic)
geniculata (C.L. Koch, 1842) – Bahamas, Curacao, Saint Vincent, Guadeloupe, also
USA, Mexico, and Panama
jutisi Armas et Teruel, 2005 – Cuba (endemic)
hispaniolana Armas et Alegre, 2001 – Dominican Republic (endemic)

maguirei Muma, 1986 – Caicos (endemic)
pallida Muma et Nezario, 1971 – Puerto Rico (endemic)
pseustes (Chamberlin, 1925) – Puerto Rico, Panama
Ammotrechinus gryllipes (Gervais, 1842) – Jamaica, Haiti (endemic genus and
species)
Antillotrecha iviei Armas, 2002 – Leeward Islands (endemic genus for the
Caribbean)
fraterna Armas, 1994 – Dominican Republic (endemic sp. for Dominican Rep.)
disjunctodens Armas et Teruel, 2005 – Cuba (endemic sp.)
Prevails the genus Ammotrechella, found also in Panama, Mexico, and the USA
(Texas and Florida). In Cuba there are eight spp. of Solifugae and in Jamaica – two
in Dominican Republic – three spp. (Armas 2004, with catalogue and analysis of the
distribution of Solifugae in the Caribbean and Central America). Since only few
taxa have been added (Armas and Teruel 2005 and others).
Ricinulei The only Ricinuleids known in the Greater Antilleans were found in
Cuba (one cave inhabiting Pseudocellus, described almost simultaneously by Cooke
(1972) and by Dumitresco and Juvara-Bals (1973), and Pseudocellus silvai (Armas,
1977)). Both species seem endemic for Cuba. Judson and Hardy (2001) described
a protonymph of Cryptocellus sp. from Tobago Island.
Scorpiones The information on the scorpions in the Antillean fauna was general-
ized by Armas (2001, 2009b) and Teruel and Kovařik (2012), who described many
of the existing taxa in the islands. Zoogeographical analysis of the origin and the
affinities of the Antillean scorpions is by Lourenço (1999a). On the Antilleans have
been recorded 4 families (Buthidae, Chactidae, Hormuridae, and Scorpionidae), 17
genera, and 130 species. Four genera with 21 species are endemic in the Antilleans
(Armas 1973, 1974, 1982, 1983, 1999, 2001, 2009b; Armas and Maes 1998; Armas
and Marcano 1987; Francke 1978; Francke and Sissom 1980; Kjellesvig-Waering
1966; Lourenço 1984, 1986, 1987; Lourenço et al. 1991; Lourenço and Huber 1999;
Lourenço and Vachon 1996; Kovařik and Teruel 2014; Moreno 1940; Teruel 2005,
and many others).
The scorpions of the Antilleans belong to 17 genera of 4 families, but Chactidae
(with 3 genera) is known only on the islands of Trinidad and Tobago near the coast
of Venezuela (also the genus Ananteris). From the other 13 genera by far, the richest
is the fauna of Cuba (55 spp.), followed of Hispaniola (Dominican Republic and
Haiti), with a total of 41 spp.
The list of the scorpions in the Lesser Antilleans (16 spp.) is provided by Lescure
et al. (1991), the one for Pinos (Isla de la Juventud, three spp. Centruroides,
Rhopalurus, no endemic) by Armas (1983).
Family Buthidae
Alayotityus Armas, 1973 (a Cuban endemism, seven species)
Tityopsis Armas, 1974 (a Cuban endemism, two species)

Family Scorpionidae (incl. Diplocentridae)
Cazierius Francke, 1978 (a Greater Antillean endemism, ten species)
Oiclus Simon, 1880 (a Lesser Antillean endemism, two species from the Leeward
Islands)
The other 11 genera are distributed as follows:
Ananteris Borelli, 1910 (one species from Trinidad and Tobago)
Centruroides Marx, 1890 (28 species on the Antilles)
Isometrus Ehrenberg, 1828 (one introduced species)
Microtityus Kjellesvig-Waering, 1966 (15 species, incl. 6 recent spp. from
Hispaniola)
Rhopalurus Thorell, 1876 (eight species from Cuba and Hispaniola)
Tityus C. L. Koch, 1836 (20 species on the Antilles, incl. 13 recent spp. from
Hispaniola)
Broteochactas Pocock, 1893 (two species from Trinidad and Tobago)
Chactas Gervais, 1844 (one species from Trinidad and Tobago)
Opisthacanthus Peters, 1861 (one species from Hispaniola)
Didymocentrus Kraepelin, 1905 (nine species from Lesser Antilles and central
Cuba)
Heteronebo Pocock, 1899 (14 species from Greater Antilles) (Table 8.3).
Pseudoscorpiones The fauna of Pseudoscorpiones in the Caribbean belongs to 16

families as follows (indicated are only the species endemic for West Indies) (Barba
Diaz and Pérez 2001; Barba Diaz and Barroso 2013; Beier 1976; Cosgrove et al.
2016; Dumitresco and Orghidan 1977, 1981; Harvey 2013b; Heurtault and Rebière
1983; Hoff 1945, 1946, 1959, 1963, 1964, 1976; Muchmore 1967, 1979, 1982,
1984, 1992, 1998; Tullgren 1907; Vitali-di-Castri 1984). According to the table in
Cosgrove (2016), the genera are 74 and the species 147.
Fam. Chthoniidae
Caribchthonius butleri Muchmore, 1976 – US Virgin Islands (endemic)
Pseudochthonius thibaudi Vitali-di-Castri, 1984 – Guadeloupe (endemic)
Tyrannochthonius guadeloupensis Vitali-di-Castri, 1984 – Guadeloupe
(endemic)
T. imitatus Hoff, 1959 – Jamaica, Dominican Republic
T. proximus Hoff, 1959 – Jamaica, Dominican Republic
Fam. Lechythiidae
Lechytia delamarei Vitali-di-Castri, 1984 – Guadeloupe (endemic)
L. trinitatis Beier, 1970 – Dominican Republic, Trinidad
Table 8.3 Scorpions in the Antilleans

Puerto Lesser Trinidad and
Country Cuba Hispaniola Jamaica Rico Bahamas Antilles Tobago
(40 + 6)
Number of species 55 41 6 17 2 16 11
Fam. Buthidae + + + + + +
Alayotityus Armas 8 − − − − − −
Ananteris Thorell − − − − − − 1
Centruroides Marx 14 5 4 5 2 8 1
Isometrus
Ehrenberg
Microtityus 9 12 − 2 − − 2
K. – W.
Rhopalurus 7 3 − 1 − − −
Thorell
Tityopsis Pocock 2 − − − − − −
Tityus C.L. Koch 1 13 1 7 − 4 4
Fam. Chactidae − − − − − − +
Neochactas Sol. et − − − − − − 1
Fet
Chactas Gervais − − − − − − 1
Broteochactas − − − − − − 1
Pocock
Fam. Hormuridae − + − − − − −
Opisthacanthus − 1 − − − − −
Peters
Fam. + + + + − + +
Scorpionidae
Casierus Francke 7 2 − 1 − 1 −
Didymocentrus 4 − − − − 3 −
Kraepelin
Cryptoiclus Ter. et 1 − − − − − −
Kov.
Heteronebo 2 5 1 1 − 5 −
Pocock
Oiclus Simon − − − − − 3 −
Tridenchthonius cubanus (Chamberlin, 1929) – Cuba
Fam. Syarinidae
Ideobisium balzanii With, 1905 – Guadeloupe, Dominica, Saint Vincent
puertoricense Muchmore, 1982 – Dominican Republic, Puerto Rico
puertoricense cavicola Muchmore, 1982 – Puerto Rico
yunquense Muchmore, 1982 – Puerto Rico
Ideoblothrus carinatus (Hoff, 1964) – Jamaica
insularum (Hoff, 1945) – Jamaica, Puerto Rico

pygmaeus (Hoff, 1964) – Jamaica, Martinique
truncatus (Hoff, 1964) – Jamaica
Fam. Bochicidae
Antillobisium mitchelli Dumitresco et Orghidan, 1977 – Cuba (endemic genus)
vachoni Dumitresco et Orghidan, 1977 – Cuba
Troglobochica Muchmore, 1984 – Jamaica (endemic genus)
T. jamaicensis Muchmore, 1984 – Jamaica
T. pecki Muchmore, 1984 – Jamaica
Mexobisium armasi Muchmore, 1980 – Cuba
M. cubanum Muchmore, 1973 – Cuba
M. sierramaestrae Muchmore, 1980 – Cuba
Nannobisium mollis (Hoff, 1964) – Jamaica
Fam. Ideoroncidae
Typhloroncus coralensis Muchmore, 1979 – US Virgin Islands (endemic) (the
other four Typhloroncus are from Mexico)
Fam. Sternophoridae
Fam. Olpiidae
Aphelolpium thibaudi Heurtault et Rebière, 1893 – Guadeloupe, Martinique
Novohorus subfuscus Hoff, 1945 – Jamaica, Puerto Rico
Olpiolum monae (Hoff, 1964) – Mona, Jamaica, Dominican Republic, Florida
O. amplum Hoff, 1964 – Jamaica
O. aureum (Hoff, 1945) – Mona, Puerto Rico
O. confundens (Hoff, 1945) – Puerto Rico
O. puertoricensis (Hoff, 1945) – Puerto Rico
Fam. Garypidae
Planctolpium arboreum Hoff, 1964 – Jamaica, Dominican Republic, Mexico
Fam. Garypinidae
Fam. Cheiridiidae
Cheiridium insulare Heurtault et Rebière, 1893 – Guadeloupe (endemic)
Fam. Pseudochiridiidae
Pseudochiridium insulae Hoff, 1964 – Cuba, Dominican Republic
Fam. Atemnidae
Paratemnoides elongatus (Banks, 1895) – Cuba, Dominican Republic, US Virgin
Islands, Central America, Mexico, USA
P. nidificator (Balzan, 1888) – Haiti, Saint Vincent, Central and South America
Fam. Cheliferidae
Cubachelifer stator Hoff, 1946 – Dominican Republic, Cuba
Parachelifer dominicanus Beier, 1976 – Dominican Republic (endemic)
P. parvus Muchmore, 1981 – US Virgin Islands (endemic)
Tyrannochelifer cubanus Hoff, 1964 – Cuba (endemic)
T. floridanus (Banks, 1891) – Dominican Republic, Florida
T. macropalpus (Tullgren, 1907) – Haiti (endemic)
Fam. Chernetidae
Americhernes puertoricensis Muchmore, 1976 – Puerto Rico
Antillochernes cruzensis Muchmore, 1984 – US Virgin Islands (endemic)
Bituberochernes jonensis Muchmore, 1979 – US Virgin Islands (endemic)
Byrso.chernes caribicus Beier, 1976 – Dominican Republic (endemic)
Caribochernes pumilus Beier, 1976 – Dominican Republic (endemic)
Chelanops ? [Dinocheirus] altimanus (Ellingsen, 1910) – St. Thomas (endemic)
Chernes hispaniolicus Beier, 1976 – Dominican Republic (end.)
Dinochernes chalumeaui Heurtault et Rebière, 1893 – Guadeloupe (endemic)
Hesperochernes vespertilionis Beier, 1976 – Dominican Republic (endemic)
Gomphochernes communis (Balzan, 1888) – South America, Trinidad,
Dominican Republic
Lustrochernes mauriesi Heurtault et Rebière, 1893 – Guadeloupe (endemic)
Parachernes (Scapanochernes) compressus (Tullgren, 1907) – Dominican
Republic, Haiti, Florida
P. dominicanus Beier, 1976 – Dominican Republic, Haiti (endemic)
Parazaona klapperichi Beier, 1976 – Dominican Republic (endemic)
Fam. Withiidae
Cacodemonius segmentidentatus [in Beier, 1976 – serratidentatus] (Balzan,
1891): Dominican Republic, South America
Dolichowithius canestrinii (Balzan, 1887) – Virgin Islands, St. Thomas, South
America
D. simplex Beier, 1932 – Dominican Republic, Puerto Rico
Endemic genera for the Antilleans are:
Antillobisium Dumitresco et Orghidan, 1977 – Cuba (two spp.)
Caribochernes Beier, 1976 – Dominican Republic (one sp.)
Cubachelifer Hoff, 1946 – Cuba, Dominican Republic (one sp.)
Troglobochica Muchmore, 1984 – Jamaica (two spp.)
The pseudoscorpions are distributed between the main countries as follows:
Cuba, 27; Jamaica, 31; Puerto Rico, 18; Haiti, 3; Dominican Republic, 26; Barbados,
1; US Virgin Island, 12; Dominica, 2; Martinique, 7; Guadeloupe, 8.
Antilleans
The fauna of Pseudoscorpiones in the Antilleans consists of 39 genera and 16
families.
Genera and families in both areas:

----------------------------------------------------------------------------------------------------
Central America Antilleans
Chthoniidae
Caribchthonius====================== Caribchthonius Muchmore
Mundochthonius Chamberlin
Pseudochthonius==========================Pseudochthonius Balzan
Tyrannochthonius =====================Tyrannochthonius Chamberlin
Lechythiidae
– Lechytia Balzan
Tridenchthoniidae
– Tridenchthonius Balzan
Bochicidae
– Antillobisium Dumitresco et Orghidan (end. genus, Cuba)
Mexobisium========================== Mexobisium Muchmore
– Troglobochica Muchmore (end. genus, Jamaica)
Neobisiidae
Microbisium Chamberlin –
Syarinidae
– Ideobisium Balzan
Ideoblothrus Balzan –
Ideoroncidae
– Typhloroncus Muchmore
Albiorix Chamberlin –
Pseudalbiorix Barba et Pérez –
Sternophoridae
Garyops ===================================== Garyops Banks
– Idiogaryops Hoff
Cheiridiidae
– Cheiridium Menge
– Cryptocheiridium Chamberlin
Pseudochiridiidae
– Pseudochiridium With
Garypinidae
Serianus ==================================Serianus Chamberlin
Atemnidae
– Atemnus Canestrini
–– Oratemnus Beier
Paratemnoides ======================== Paratemnoides Harvey
Olpiidae
– Aphelolpium Hoff

Olpiolum ====================================== Olpiolum Beier
Garypidae
– Planctolpium Hoff
Cheliferidae
– Cubachelifer Hoff (Dominican Rep., Cuba – end. genus)
Parachelifer ==============================Parachelifer Chamberlin
– Tyrannochelifer Chamberlin
Chernetidae
– Antillochernes Muchmore
– Bituberochernes Muchmore
– Byrsochernes Beier
– Caribochernes Beier (Dominican Rep., endemic genus)
– Chelanops Gervais
– Chernes Menge
– Dinochernes Beier
– Hesperochernes Chamberlin
Lustrochernes ==== =========== ================ Lustrochernes Beier
Parachernes (Parachernes) Chamberlin -
Parachernes (Scapanochernes)========Parachernes (Scapanochernes Beier)
– Parazaona Beier
Withiidae
Cacodemonius Chamberlin===================Cacodemonius Chamberlin
Dolichowithius (D.) Chamberlin============Dolichowithius (D.) Chamberlin
=========================================================
Opiliones
Cyphophthalmi – Not recorded on the Antilleans
Eupnoi (Avram 1970, 1973a, b, 1977a, b, 1981; Banks 1909; Cokendolpher and
Camilo – Rivera, 1989, Goodnight and Goodnight 1942; Rambla 1969; Roewer
1910, 1947; Šilhavý 1971, 1973, 1976, 1979; Staręga 1970)
Fam. Sclerosomatidae – Geaya Roewer, 1910 (Haiti)
Dyspnoi – Not recorded
Laniatores
According to Kury (2003), in the Caribbean area are known 117 species of Opiliones
Laniatores, belonging to 65 genera and 11 families: Cosmetidae, Agoristenidae,
Stygnidae, Stygnommatidae, Kimulidae (Minuidae), Biantidae, Samoidae,
Podoctidae, Manaosbiidae, Phalangodidae, and Zalmoxidae. The subfamily
Stenostygninae of Biantidae consists almost entirely of taxa from the Caribbean
(eight of the nine genera).
By far the best studied island is Cuba, not only because of its size but mainly as
a result of the studies of the Cubano-Romanian Expeditions (S. Avram and V.
Šilhavý described from Cuba and other islands 58 new species and many new
genera).
Here are listed the endemic taxa in the Antilleans:
Fam. Cosmetidae
Arucillus hispaniolicus Šilhavý, 1971 – Dominican Republic
Cynortula garna Goodnight et Goodnight, 1942 – Bahamas (Andros)
C. sinensis Goodnight et Goodnight, 1942 – Bahamas (New Providence)
C. juncta (Gervais, 1844) – Cuba
C. fraterna Banks, 1909 – Cuba
C. quinquesignata Franganillo Balboa, 1926 – Cuba
Cynorta sextuberculata Franganillo Balboa, 1926 – Cuba
C. hassleri Goodnight et Goodnight, 1942 – Haiti
C. lithoclasica Avram 1981 – Cuba
C. poaensis Avram 1981 – Cuba
C. quibijana Avram 1981 – Cuba
Cynortoides caraibicus (Sørensen, 1932) – US Virgin Islands, St. Thomas
C. cubanus cubanus (Banks, 1909) – Cuba
C. cubanus signatus Roewer, 1912 – Cuba
C. lateralis Roewer, 1947 – Jamaica
C. roeweri (Henriksen, 1932) – Puerto Rico
C. quadrispinosus Goodnight et Goodnight, 1942 – Jamaica
C. roeweri (Henriksen, 1932) – Cuba, Puerto Rico
C. caraibicus (Sørensen, 1932) – Dominican Republic
C. marginatus Goodnight et Goodnight, 1942 – Dominican Republic
C. v-album (Simon, 1879) – Dominican Republic, Haiti, Tortuga Island
Cynortesta laevis Roewer, 1947 – Windward Islands
C. granulata Roewer, 1947 – Windward Islands: Saint Vincent and the Grenadines
Eucynortoides antillarum Roewer, 1947 – Windward Islands: Saint Vincent and
the Grenadines
Cynortellana quadrimaculata (Gervais, 1844) – Cuba
C. bisignata (Banks, 1909) – Cuba
Heterovonones insularis Roewer, 1947 – Cuba
Erginulus castaneus (Banks, 1906) – Bahamas (Andros, New Providence)
E. quadricostatus (Franganillo Balboa, 1926) – Cuba
Metacynortoides bilineatus Goodnight et Goodnight, 1942 – Dominican
Republic
M. obscurus dorsalis Roewer, 1916 – US Virgin Islands: St. Croix, St. Thomas
M. obscurus obscurus (Banks, 1901) – Haiti, Jamaica, Puerto Rico, US Virgin
Islands, St. John
M. romanus Goodnight et Goodnight, 1942 – Dominican Republic
M. transversalis Goodnight et Goodnight, 1942 – Dominican Republic
M. scabrosus (Banks, 1909) – Cuba
Paecilaema luquillense H. Soares, 1990 – Puerto Rico
P. conspicillatus Simon, 1879 – Windward Islands, Martinique

Platycynorta secunda Roewer, 1947 – Cuba
Prasiana fallax (Sørensen, 1932) – West Indies
Proerginus lineatus Roewer, 1917 – Dominican Republic
Trinimontius darlingtoni Šilhavý, 1970 – Cuba
Vonones sayi (Simon, 1879) – Cuba
V. granulatus Roewer, 1947 – Leeward Islands, Antigua and Barbuda
V. planus Goodnight et Goodnight, 1942 – Leeward Islands, Dominica
Fam. Samoidae
Akdalima jamaicana Šilhavý, 1979 – Jamaica
Arganotus robustus Šilhavý, 1979 – Haiti
Hummelinckiolus parvus Šilhavý, 1979 – Leeward Islands: Guadeloupe,
Montserrat, St. Kitts and Nevis
Maracaynatum cubanum Šilhavý, 1979 – Cuba
M. stridulans Šilhavý 1979 – Cuba
Orsa daphne Šilhavý, 1979 – Haiti
Pellobunus haitiensis (Šilhavý, 1979) – Haiti
Reventula amabilis Šilhavý, 1979 – Jamaica
Fam. Stygnidae Heterostygninae
Stygnoplus antiguanus (Roewer, 1943) – Leeward Islands, Antigua and Barbuda
S. flavitarsis (Simon, 1879) – Leeward Islands, Guadeloupe
S. tuberculatus (Goodnight et Goodnight, 1942) – Leeward Islands, Dominica
Fam. Stygnommatidae
Stygnomma spiniferum bolivari (Goodnight et Goodnight, 1945) – Cuba
S. spiniferum spiniferum (Packard 1888) – Florida Keys (USA): Tortugas,
Jamaica
S. spirulina (Goodnight et Goodnight, 1942) – Puerto Rico
S. fiskei Rambla, 1969 – Jamaica
Fam. Agoristenidae
Agoristeninae
Agoristenus cubanus Šilhavý, 1973 – Cuba
haitiensis Šilhavý, 1973 – Dominican Republic
Ahotta hispaniolica Šilhavý, 1973 – Haiti
Calmotrinus turquinensis Šilhavý, 1973 – Cuba
Dumitrescuella ornata Avram, 1977 – Cuba
Haitimera paeninsularis Šilhavý, 1973 – Haiti
Lichirtes hexapodoides Šilhavý, 1973 – Cuba
Meriosfera gertschi Šilhavý, 1973 – Haiti
M. lineata Šilhavý, 1973 – Haiti
Orghidaniella granpiedrae Avram, 1977 – Cuba
Piratrinus calcaratus Šilhavý, 1973 – Cuba
Torreana poeyi Avram, 1977 – Cuba

T. spinata Avram, 1977 – Cuba
Vampyrostenus kratochvili Šilhavý, 1976 – Puerto Rico
Yunquenus portoricanus Šilhavý, 1973 – Puerto Rico
Fam. Phalangodidae
Phalangodes flavipes (Banks, 1908) – Cuba
Fam. Biantidae
Stenostygninae
Caribbiantes cubanus Šilhavý, 1973 – Cuba
Galibrotus carlotanus Šilhavý, 1973 – Cuba
G. matiasis Avram, 1977 – Cuba
G. riedeli Šilhavý, 1973 – Cuba
Bidoma indivisa Šilhavý, 1973 – Haiti
Decuella cubaorientalis Avram, 1977 – Cuba
Negreaella fundorai Avram, 1977 – Cuba
N. palenquensis Avram, 1977 – Cuba
N. rioindiocubanicola Avram, 1977 – Cuba
N. vinai Avram, 1977 – Cuba
N. yumuriensis Avram, 1977 – Cuba
Manahunca bielawskii Šilhavý, 1973 – Cuba
M. cuevajibarae Avram, 1977 – Cuba
M. silhavyi Avram, 1977 – Cuba
Martibianta virginsulana Šilhavý, 1973 – US Virgin Islands, St. John
Vestitecola haitensis Šilhavý, 1973 – Haiti
Fam. Podoctidae Ibaloniinae
[Santobius cubanus = Ibantila cubana (Šilhavý, 1969) – Cuba (probably intro-
duced from Melanesia, see Kury and Machado (2009))]
Fam. Kimulidae (Minuidae)
Kimula levii Šilhavý, 1969 – Cuba
K. banksi Šilhavý, 1969 – Cuba
K. goodnightiorum Šilhavý, 1969 – Cuba
K. cokendolpheri Pérez et Armas, 2000 – Dominican Republic
K. elongata Goodnight et Goodnight, 1942 – Puerto Rico
K. tuberculata Goodnight et Goodnight, 1943 – Cuba
K. turquinensis Šilhavý, 1969 – Cuba
Metakimula botosaneanui Avram, 1973 – Cuba
Minuides milleri Šilhavý 1978 – Cuba
Fam. Manaosbiidae
Cranellus balthazar Roewer, 1932 – Windward Islands
Sanvincentia tarsalis Roewer, 1943 – Windward Islands: Saint Vincent and the
Grenadines
Fam. Zalmoxidae
Cersa kratochvili Šilhavý 1979 – Cuba
Ethobunus cubensis (Šilhavý 1979) – Cuba
E. goodnighti (Rambla, 1969) – Jamaica
E. pecki (Rambla, 1969) – Jamaica
E. zebroides (Šilhavý 1979) – Cuba
Pachylicus castaneus (Šilhavý 1979) – Cuba
Family uncertain
Anamota custodiens Šilhavý 1979 – Cuba
Caribula longimana Šilhavý 1979 – Cuba
Jimeneziella negreai Avram, 1970 – Cuba (end. genus)
J. decui Avram, 1970 – Cuba
Metapellobunus unicolor (Roewer, 1912) – US Virgin Islands, St. Thomas
Mirda insulanus (Banks, 1901) – Haiti, Puerto Rico
Neoscotolemon pictipes (Banks, 1908) – Cuba
N. lutzi Goodnight et Goodnight, 1942 – Leeward Islands, Dominica
Paraconomma ovala Goodnight et Goodnight, 1942 – Puerto Rico
P. spinooculorum Goodnight et Goodnight, 1942 – Puerto Rico
Pseudomitraceras minutus Goodnight et Goodnight, 1942 – Puerto Rico
Turquinia montana Šilhavý 1979 – Cuba
Valifema blanda Šilhavý 1979 – Cuba
Endemic genera of Opiliones in the Caribbean are:
Fam. Minuidae
Jimeneziella Avram, 1970 – Cuba (two spp.)
Kimula Goodnight et Goodnight, 1942 – Cuba, Dominican Republic, Puerto
Rico (eight spp.)
Fam. Biantidae (Stenostygninae)
Bidoma Šilhavý, 1973 – Haiti (one sp.)
Caribbiantes Šilhavý, 1973 – Cuba (one sp.)
Decuella Avram, 1977 – Cuba (one sp.)
Galibrotus Šilhavý, 1973 – Cuba (three spp.)
Manahunca Šilhavý, 1973 – Cuba (three spp.)
Martibianta Šilhavý, 1973 – US Virgin Islands (one sp.)
Negreaella Avram, 1977 – Cuba (five spp.)
Vestitecola Šilhavý, 1973 – Haiti (one sp.)
Fam. Samoidae
Neoorsa Ozdikmen, 2006 (Orsa Šilhavý, 1979) – Haiti (one sp.)
Reventula Šilhavý, 1979 – Jamaica (one sp.)
Vlachiolus Šilhavý, 1979 – Cuba (one sp.)

Amblypygi
On the Antillean Archipelago have been recorded 29 species of Amblypygi from 4
genera (Charinus, Heterophrynus, Paraphrynus, Phrynus) and 2 families
(Charinidae and Phrynidae). The endemism is only on species level (25 species),
most species being confined to only 1 island (Armas 2004, 2006, 2007, 2009a,
2010, 2013; Armas and Avila Calvo 2000; Armas and Pérez Gonzalez 1994, 2001,
2002; Armas and Teruel 1997; Avila Calvo and Armas 1997; Franganillo 1926;
Mullinex 1975; Pocock 1893; Quintero 1981, 1983, 1986; Teruel 2011; Teruel and
Questel 2011; Teruel et al. 2009).
Four species (three Phrynus and one Heterophrynus) are known also from South
America (or Florida). This is in contrast with the order of scorpions, where 4 of the
15 genera are endemic to the Antilleans.
As Armas (2009b) points it, Cuba has the most diverse amblypygid fauna (2
fam., 3 genera, 16 species, incl. endemics). Follows Hispaniola with two families,
two genera, and seven species (incl. endemics). Now (2017) the distribution of the
Amblypygi in the Antillean is:
Fam. Charinidae – Charinus Simon (12 spp., Cuba, Jamaica, Dominican Rep.,
Puerto Rico, Lesser Antilleans)
Fam. Phrynidae – Heterophrynus Pocock (one sp., Trinidad and Tobago),
Paraphrynus Moreno (three spp., Bahamas and Cuba), Phrynus Lamarck (14
spp.)
According to the list of Rowland and Cooke (1973), on the Antillean islands live
only two spp. of Uropygi: Mastigoproctus baracoensis Franganillo, 1931 (Cuba),
and the second described species in Uropygi M. proscorpio (Latreille, 1806)
(Dominican Rep., Haiti, and Martinique). Another Mastigoproctus (M. pelegrini
Armas, 2000) was described later from Cuba. Armas (2002) described a new
Uropygid from the Dominican Republic, first as Telyphonellus wetherbeei and then
raised in a new genus Ravilops Viquez et Armas, 2005. So far the two Cuban
Mastigoproctus and Ravilops wetherbeei are considered endemic species; Ravilops
is also an endemic genus for the Dominican Republic (Armas 2000, 2004;
Franganillo 1931; Viquez and de Armas 2005).
Schizomida
Identified Schizomida are known (Ávila Calvo and Armas 1997; Armas 1977, 1989,
2004, 2011; Armas and Abud Antun 1990, 2002; Armas and Teruel 2002; Camilo
and Cokendolpher 1988; Dumitresco 1973, 1977; Hilton 1933; Quintero 1983;
Reddell and Cokendolpher 1995; Rowland and Reddell 1977; Teruel 2003, 2004,
2007) from:
Cuba – Antillostenochrus alejandroi, A. alticola, A. cokendolpheri, A. gibarensis, A.
holguin, A. planicauda, Cubazomus armasi, C. montanus, C. orghidani, C. row-
landi, Guanazomus armatus, Reddellzomus cubensis, Rowlandius abeli, R. alay-
oni, R. baracoae, R. biconouros, R. cubanacan, R. cupeyalensis, R. decui, R.

digitiger, R. falcifemus, R. florentiae, R. gladiger, R. gracilis, R. guantanamero,
R. labarcae, R. littoralis, R. marianae, R. melici, R. mixtus, R. monticola, R.
negreai, R. ramosi, R. recuerdo, R. reyesi, R. serrano, R. siboney, R. teruel, R.
toldo, R. vinai, Stenochrus alejandroi, S. portoricensis, Troglocubazomus orghi-
dani, T. rowlandi
Pinos (Isla de la Juventud) – Luisarmasius insulaepinorum
Hispaniola (Dominican Rep. and Haiti) – Antillostenochrus brevipatellatus, A. sub-
cerdoso, Cokendolpherius ramosi, Rowlandius anasilviae, R. casabito, R. ducou-
drayi, R. engombe, R. isabel, R. jarmillae, R. lantiguai, R. longipalpus, R.
naranjo, R. virginiae, Stenochrus subcerdoso, S. portoricensis
Dominica – Stenochrus portoricensis
Puerto Rico (incl. Isla Desecheo, Mona Island) – Antillostenochrus cerdoso,
Luisarmasius yunquensis, Rowlandius desecheo, R. monensis
Jamaica – Caribezomus laurae, Rowlandius cousinensis, R. peckorum, R. primibi-
conourus, R. viridis, Stenochrus portoricensis, Stewardpeckius troglobius
Navassa – Rowlandius steineri
Virgin Islands – Stenochrus portoricensis
Martinique – Hansenochrus dispar, Rowlandius insignis
There is also unidentified material from Barbados and Cayman Islands.
Except Stenochrus portoricensis, all species are endemic to the respective
islands. Endemic to the Caribbean are also the genera Cokendolpherius (Cuba),
Luisarmasius (Cuba, Pinos, Puerto Rico), Reddellzomus (Cuba), Stewardpeckius
(Jamaica), Cubazomus (Cuba), Troglocubazomus, and Heterocubazomus (Cuba).
Araneae
Simon (1888b); Alayón (1994, 1995, 2005), Bruyant (1940, 1947, 1948), Dumitrescu
(1973), Dumitrescu, and Georgescu (1992)
According to Alayón (2000), the spiders recorded from Cuba are 567 spp. of 52
families, including 247 endemic to the Antilleans (43.56%) with 7 endemic genera
(Troglothele Fage, Bryantina Brignoli, Fageicera Dumitresco et Georgesco,
Caudalia Alayón, Cubanopillus Alayón et Platnick, Decaphora Franganillo,
Paraplexippus Franganillo).
From the Antilleans are known some endemic genera of spiders, but there are
no endemic families:
Fam. Theraphosidae
Antillena Bertani, Huff et Fukushima, 2017 – Dominican Republic
Caribena Fukushima et Bertani, 2017 – Puerto Rico, Cuba, US Virgin Islands,
Martinique
Cubanana Ortiz, 2008 – Cuba
Nesipelma Schmidt et Kovarik, 1996 – Nevis
Fageicera Dumitrescu et Georgescu, 1992 – Cuba
Fam. Barychelidae
Fam. Agelenidae
Neowadotes Alayón, 1995 – Hispaniola
Fam. Cyatholipidae
Pokennips Griswold, 2001 – Jamaica
Fam. Tetragnathidae
Ancinosphenus Simon, 1895 – West Indies
Antillognatha Bryant, 1945 – Hispaniola
Hispanognatha Bryant, 1945 – Hispaniola
Fam. Linyphiidae
Fam. Filistatidae
Antilloides Breskovit et al., 2016 – Cuba, Dominican Republic, US Virgin
Islands, Puerto Rico
Fam. Paratropididae
Anisaspis Simon, 1891 – Saint Vincent
Fam. Amaurobiidae
Tugana Chamberlin, 1948 – Cuba, Hispaniola
Fam. Anyphaenidae
Thaloe Brescovit, 1993 – Cuba, Hispaniola
Fam. Theridiidae
Jamaitidion Wunderlich, 1995 – Jamaica
Fam. Liocranidae
Laudetia Gertsch, 1941 – Dominican Republic, Puerto Rico
Mesobria Simon, 1897 – Saint Vincent
Fam. Ctenidae
Ciba Bloom et al., 2014 – Cuba, Dominican Republic
Ohvida Polotow et Brescovit, 2009 – Cuba, Puerto Rico, Bahamas
Trujillina Bryant, 1948 – Hispaniola, Puerto Rico
Fam. Pholcidae
Bryantina Brignoli, 1985 – Cuba
Ciboneya Perez, 2001 – Cuba

Platnicknia Özdikmen et Demir, 2009 – Cuba
Tainonia Huber, 2000 – Hispaniola
Fam. Prodidomidae
Caudalia Alayón, 1980 – Cuba
Cubanopillus Alayón et Platnick, 1993 – Cuba
Fam. Sparassidae
Decaphora Franganillo, 1931 – Cuba
Fam. Thomisidae
Rejanellus Lise, 2005 – Cuba, Puerto Rico, Hispaniola
Fam. Salticidae
Allodecta Bruyant, 1950 – Jamaica
Antillattus Bryant, 1943 – Hispaniola, Cuba
Bythocrotus Simon, 1903 – Hispaniola
Caribattus Bryant, 1950 – Jamaica
Cerionesta Simon, 1901 – Saint Vincent
Commoris Simon, 1902 – Guadeloupe, Dominica
Corticettus Zhang et Maddison, 1902 – Porto Rico, Hispaniola
Paraplexippus Franganillo, 1930 – Cuba
Parasaitis Bruyant, 1950 – Jamaica
Parathiodina Bruyant, 1943 – Hispaniola
Popcornella Zhang et Maddison, 2012 – Hispaniola, Puerto Rico
Truncattus Zhang et Maddison, 2012 – Hispaniola
Opilioacarida
From Cuba and the Dominican Republic has been described the new genus
Caribeacarus Vásquez et Klompen, 2009, from which two species have been
described from Cuba, one also from the Dominican Republic, and one from Panama.
Another species, described from Cuba (and the Antillean Islands), was Neoacarus
orghidani (Juvara-Balş et Baltac 1977). Two of the three Cuban species and
Caribeacarus panamensis live in caves (Beron 2014; Juvara-Balş and Baltac 1977;
Vásquez and Klompen 2009).
Parasitiformes (Mesostigmata and Ixodida): Ref.: Beron (2014), Cruz (2001)
From Capromys pilorides (Rodentia, Capromyidae) has been described the
endemic Ixodes capromydis Černy, 1966 (endemic subgenus Alloixodes).
The purely Neotropical family of Spelaeorhynchidae (parasites on bats of the
families Phyllostomidae and Mormoopidae) is found, together with these bats, in
Cuba, Jamaica, Puerto Rico, and Dominican Republic but also in Central America.
Holothyrida The only holothyrid recorded from the Antilleans is the endemic
genus Carabothyrus Kontchán et Mahunka, 2004, with one species C. barbatus
Kontchán et Mahunka, 2004, described from the Dominican Republic.
8.6.2 C
omparison Between the Faunas of Central America
and the Antilleans
Palpigradi – Known only from the Antilleans

Ricinulei
Central America
In the 7 countries of Central America (without South Mexico) are registered 18 spe-
cies of Ricinulei:
Cryptocellus Westwood, 1874 – 12 spp.
Pseudocellus Platnick, 1980 – Six spp.
Antilleans
Two endemic spp. of Cryptocellus from Cuba
Solifugae – Endemic genera in bold
Central America – From the seven countries of Central America (without South
Mexico) are known ten spp. of Solifugae of five genera, all belonging to
Ammotrechidae family (Ammotrechinae): Ammotrecha Banks, 1900 (four spp.);
Ammotrechella Roewer, 1934 (two spp.); Ammotrechesta Roewer, 1934 (five spp.);
Innesa Roewer, 1934 (one sp., endemic genus); and Ammotrechula Roewer.
Antilleans – From the Antilleans are known four genera of Solifugae (all from
Ammotrechidae), three of them endemic, all belonging to Ammotrechidae:
Ammotrechinus Roewer, 1934 (one sp.); Ammotrechella Roewer, 1934 (nine spp.);
Ammotrechona Roewer, 1934 (one sp.); and Antillotrecha Roewer, 1934 (three
spp.).
All Solifugae in both areas belong to the same family. There is only one genus
that is common (Ammotrechella); the species are endemic.
Amblypygi
Central America – The amblypygids are represented in Central America by nine

spp.
Fam. Charinidae – Charinus Simon
Fam. Phrynidae – Phrynus Lamarck, Paraphrynus Moreno
Antilleans – Twenty-nine species of Amblypygi from four genera (Charinus,
Heterophrynus, Paraphrynus, Phrynus) and the same two families (Charinidae and
Phrynidae). The endemism is only on species level (25 species). Except
Heterophrynus, known from the islands Trinidad and Tobago by the Venezuelan
coast, the generic composition of Mesoamerica and the Antilleans is the same.
Central America – Five species (the endemics for Costa Rica Valeriophonus
Viques et Armas, Mayacentrum tantalus Viquez et Armas for El Salvador, M. pijol
V. et A. for Honduras, M. guatemalae V. et A. for Belize, and Mimoscorpius pug-
nator Butler for Guatemala).
Antilleans – Four species (three Mastigoproctus Pocock, Ravilops Viques et
Armas). Three species, including the genus Ravilops, are endemic for the Antilleans.
Schizomida
Central America
In Central America (between Mexico and Colombia) are recorded seven species, but
there is also unidentified material from many places (Reddell and Cokendolpher
1995). Besides the largely distributed Stenochrus portoricensis (Guatemala,
Honduras, Nicaragua, Belize), from Central America, are known the genera
Hansenochrus (Costa Rica, Panama), Rowlandius, and Surazomus (Costa Rica).
Surazomus Reddell et Cokendolpher is known also from many South American
countries. Rowlandius Reddell et Cokendolpher is widespread in the Caribbean
Islands. Hansenochrus Reddell et Cokendolpher is found also in South America and
the Caribbean.
Antilleans
From the Antilleans are known schizomids of nine genera (six endemic), all belong-
ing to one family – Hubbardiidae.

– Cubazomus Reddell et Cokendolpher
– Heterocubazomus Teruel
Rowlandius===========================Rowlandius Red. et Cok.
Stenochrus ======================== =========Stenochrus Chamberlin
Piaroa Manz. et al. –
– Luisarmasius Red. et Cok.
– Stewardpeckius Red. et Cok.
– Reddellzomus Armas
– Cokendolpherius Armas
Surazomus –
Hansenochrus========================Hansenochrus Red. et Cok.
Mayazomus Red. et Cock. (Heteronychus Armas et Viques) Guatemala) –
Belicenozomus Armas et Viques –
==========================================================
Ref.: Ávila Calvo and Armas (1997), Armas (1977, 1989, 2002a, b, 2004, 2010),
Armas and Abud Antun (1990, 2002), Armas and Teruel (2002), Camilo and
Cokendolpher (1988), Dumitresco (1973, 1977), Hilton (1933), Quintero (1983a),
Reddell and Cokendolpher (1995), Rowland and Reddell (1977)
Identified Schizomida are known from:
Cuba – Cubazomus armasi, C. orghidani, C. rowlandi, Rowlandius alayoni, R.
baracoae, R. biconouros, R. cubanacan, R. decui, R. digitiger, R. gladiger, R.
labarcae, R. negreai, R. recuerdo, Stenochrus alejandroi, S. portoricensis
Pinos (Isla de la Juventud) – Luisarmasius insulaepinorum
Hispaniola (Dominican Rep. and Haiti) – Rowlandius anasilviae, R. brevipatella-
tus, R. casabito, R. ducoudrayi, R. engombe, R. isabel, R. jarmillae, R. lantiguai,
R. longipalpus, R. naranjo, R. virginae, Stenochrus subcerdoso, S.
portoricensis
Dominica – Stenochrus portoricensis
Puerto Rico (incl. Isla Desecheo, Mona Island) – Luisarmasius yunquensis,
Rowlandius desecheo, R. monensis
Jamaica – Rowlandius cousinensis, R. peckorum, R. primibiconourus, R. viridis,
Stenochrus portoricensis, Stewardpeckius troglobius
Virgin Islands – Stenochrus portoricensis
Martinique – Hansenochrus dispar, H. insignis
There is also unidentified material from Barbados and Cayman Islands.
Except Stenochrus portoricensis, all species are endemic to the respective
islands. Endemic to the Caribbean are also the genera Cokendolpherius (Cuba),
Luisarmasius (Cuba, Pinos, Puerto Rico), Reddellzomus (Cuba), Stewardpeckius
(Jamaica), and Cubazomus (Cuba).
Pseudoscorpiones
Central America
In the 7 countries of Central America are recorded pseudoscorpions of 18 genera
and 12 families (Chthoniidae, Bochicidae, Neobisiidae, Syarinidae, Ideoroncidae,
Sternophoridae, Garypinidae, Atemnidae, Olpiidae, Cheliferidae, Chernetidae,
Withiidae).
Antilleans The fauna of Pseudoscorpiones in the Antilleans consists in 39 genera
and 16 families.
Genera and families in both areas

Chthoniidae
Caribchthonius Muchmore =============== Caribchthonius Muchmore
Pseudochthonius Balzan====================== Pseudochthonius Balzan
Tyrannochthonius Chamberlin===========-==Tyrannochthonius Chamberlin
Mexobisium Muchmore =================== Mexobisium Muchmore
Syarinidae
Ideobisium Balzan ======================== Ideobisium Balzan
Ideoblothrus Balzan ============================Ideoblothrus Balzan
Sternophoridae
Garyops Banks===================================Garyops Banks
Serianus Chamberlin============================Serianus Chamberlin
Atemnidae
Paratemnoides Harvey====================Paratemnoides Harvey
Olpiidae
Olpiolum Beier================================== Olpiolum Beier
Cheliferidae
Parachelifer Chamberlin=====================Parachelifer Chamberlin
Chernetidae
Lustrochernes Beier ===========================Lustrochernes Beier
Parachernes (Scapanochernes Beier)===Parachernes (Scapanochernes Beier)
Withiidae
Cacodemonius Chamberlin==================Cacodemonius Chamberlin
Dolichowithius (D.) Chamberlin===========Dolichowithius (D.) Chamberlin
Both areas have 8 families and 15 genera of pseudoscorpions in common. No

endemic families, no end. genera in Central America, and four end. genera in the
Antilleans.
Scorpiones
Central America Antilleans (without Trinidad and Tobago)
Buthidae
– = Alayotityus Armas
Centruroides ==================================Centruroides Marx
– Microtityus K.-W.
– Rhopalurus Thorell
– Tityopsis Pocock
Tityus ======================================== Tityus C.L. Koch
Chactidae –
Chactas Gervais –
Hormuridae
Opisthacanthus ==============================Opisthacanthus Peters
Scorpionidae
– Cazierus Francke
– Cryptoiclus Ter. et Kov.
Didymocentrus ==============================Didymocentrus Kraep.
– Heteronebo Pocock
Diplocentrus Peters –
Euscorpiidae
Plesiochactas Pocock –
Vaejovidae
Vaejovis C.L. Koch –
=========================================================
8.6.3 Conclusion
In the seven countries of Central America (without counting Yucatan) are known
eight spp. of Ricinulei of both American genera. From the Antilleans (Cuba) have
been recorded only two (endemic) species. The genus Cryptocellus Westwood is
common in both areas.
From the seven countries of Central America (without South Mexico) are known
Solifugae of five genera, all belonging to Ammotrechidae family (Ammotrechinae).
Two genera seem endemic. From the Antilleans are recorded four genera (all from
the same family Ammotrechidae), three of them endemic. There is only one genus
in common among both areas (Ammotrechella); the species are endemic.
The Amblypygi of the Antilleans are much numerous and varied (29 spp. of 4
genera), than in the 7 countries of Central America (9 spp. of 2 genera, the families
are the same). Again differences in the research?
Both areas have different fauna of Thelyphonida (Uropygi). Central America
has three endemic genera, the Caribbean one. Mastigoproctus is known from Cuba,
Martinique, and Haiti but is so far absent from Central America.
The Schizomida of both areas belong to one family, but the genera are very dif-
ferent (seven genera, one endemic in Central America, eight genera in the Antilleans,
six endemic). Both areas have three genera in common. Again, differences in the
research?
8.6.4 Opiliones
Cyphophthami – Unknown in both areas

Eupnoi: Sclerosomatidae (in both areas, the same genera Geaia and
Prionostemma); Dyspnoi – in Honduras the genus Trilasma Goodnight et Goodnight
(1942) (Nemastomatidae, Ortholasmatinae); in the Antilleans, Dyspnoi are not
recorded.
Laniatores The catalogue of Kury (2003) enumerates from the seven countries of
Central America 56 genera and 9 families of Laniatores (Cosmetidae, Cranaidae,
Gonyleptidae, Stygnommatidae, Samoidae, Pyramidopidae, Manaosbiidae,
Stygnopsidae, Zalmoxidae). From the Antilleans in the same catalogue are listed
117 species of Opiliones Laniatores, belonging to 65 genera and 11 families:
Cosmetidae, Agoristenidae, Stygnidae, Stygnommatidae, Kimulidae (Minuidae),
Biantidae, Samoidae, Podoctidae, Manaosbiidae, Phalangodidae, and Zalmoxidae.
From this number, there are five families (underlined) in common. The endemic
genera of opilions in the islands are 13 (Biantidae, Kimulidae). There are no
endemic families.
The fauna of pseudoscorpions in the Antilles seems much richer (39 genera, 16
families) than the fauna of the 7 countries of Central America (33 genera, 13 fami-
lies). Both areas have 15 genera in common.
This comparison shows that from 8 genera of scorpions in Central America and
11 genera in the Antilleans, there are 4 genera in common. Three families in Central
America (Chactidae, Euscorpiidae, and Vaejovidae) are not recorded in the
Antilleans. Fact is that the scorpions in the islands have more genera and species
than the scorpions of Central America region with land connection with such rich
areas like Mexico and northern South America. One explanation is the very active
speciation, especially in Cuba (55 spp.) and the Dominican Republic (40 spp.). The
other explanation is the presence in Cuba of such specialists as Armas and Teruel.
Opilioacarida – Two genera from the same family (Neocarus Chamberlin et
Mulaik and Caribeacarus Vásquez et Klompen) live in both areas. The species are
(so far) endemic.
Holothyrida are unknown in Central America. From the Dominican Republic
has been described the endemic species Caribothyrus barbatus Kontchán et
Mahunka, 2004.
In general, there is considerable difference (artificial) between the archaeofauna
of both areas but on low (genus, species) level. The Antilleans seem richer (or better
explored).
8.7 P
atagonia: South Chile, Argentina, Uruguay, and Tierra
del Fuego

Argentina’s continental area is between the Andes mountain range in the west and
the Atlantic Ocean in the east. Also claims the Falkland Island (Malvinas) and some
parts of Antarctida. The territory is ca. 2 767 000 km2 (the second biggest in Latin
America), stretching from the tropical forest of Iguazú Falls to Tierra del Fuego and
the Subantarctic climate. The southern part of Argentina and Chile is called
Patagonia (cool semidesert region of some 500 000 km2, integrating the
8.7 Patagonia: South Chile, Argentina, Uruguay, and Tierra del Fuego 675
southernmost section of the Andes mountains to the southwest toward the Pacific
Ocean, and from the east of the cordillera to the valleys it follows south through the
Colorado River toward Carmen de Patagones in the Atlantic Ocean). The Chilean
part includes the southern provinces and regions of Valdivia, Los Lagos Region, and
Greater Island of Chiloé and is composed of two subregions: the northern Aisen
and, to its south, Magallanes. The altitudinal difference is the biggest in the Western
Hemisphere – from Aconcagua (6962 m.) to the sea level.
Some sources: Cabrera and Willink (1973), Harrington (1962), Kuschel (1960),
Markgraf (1985), Mattoni (2007), Riccardi (1987), Roig-Juñent et al.(2006),
Spalletti and Franzese (1996), and Uliana and Biddle (1988).
Among the 40 centers of dispersal in the Neotropical Region, outlined by Müller
(1972), only 2 (Patagonian and Nothofagus centers) are situated in South Chile and
Patagonia.
According to Crisci et al. (1991), “The northern South American biota is most
closely related to that of North America, and southern South America constitutes a
monophyletic group together with Australia, Tasmania, New Guinea, New
Caledonia, and New Zealand, reflecting the existence of an ancient austral biota.
Four conflicting hypotheses of area relationships concerning southern South
America and the other austral areas are proposed, suggesting that southern South
America may be a composite area in itself.”
Among the biogeographical areas of Latin America of Morrone (2005), based on
the entomofauna, is also the Andean Region, which “…is included in the Austral
Kingdom, which corresponds to western Gondwanaland and comprises five other
regions: Antarctic (Antarctica), Cape or Afrotemperate (South Africa), Neoguinean
(New Guinea and New Caledonia), Temperate Australian (southeastern Australia),
and Neozealandic (New Zealand).” These ideas are developed also in the papers of
Brundin (1966), Rapoport (1968), and others. The Andean Subregion of Morrone
includes Central Chilean Subregion (between latitudes 30 and 34oSouth),
Subantarctic Subregion (from lat. 36 south to Cape Horn), and Patagonian Subregion
with two provinces (Subandean Patagonia and Central Patagonia).
8.7.2 A
rachnogeographical Character of South Chile
and Argentina
We consider here the area south of 30oS, as it is important to check from arachno-
logical point of view the sharp delimitation (for some groups of vertebrates and
insects) across Argentina and Chile of two Kingdoms – the Neogea (region
Neotropica) and the Notogea. Is it such sharp difference what concerns the Arachnida
and where is the limit (and/or transition zone)?
In this area are registered families and even genera common with faraway lands,
remnants from the Gondwana (South Africa, Australia, New Zealand) (Cabrera and
Willink 1980; Cokendolpher and Lanfranco 1985; Mello-Leitão 1939; Ringuelet
1957, 1959, 1961; Simon 1886). A catalogue of the Arachnida of the southernmost
Chile (Magallanes) and the “Antarctica Chilena” was due to Cecalovic (1976).
Amblypygi, Uropygi, Schizomida, and Ricinulei – No records in Argentina,
Chile, Uruguay, and Paraguay
Palpigradi From Chile has been described Prokoenenia chilensis Hansen
(Prokoeneniidae) (Cekalovic 1984; Hansen 1901).
Solifugae Muma (1971) lists for Chile 13 spp. of Solifugae in 9 genera and 2 fami-
lies – Ammotrechidae and Amacataidae, described by him. Later have been
described Chileotrecha Maury (1987) (endemic genus for Chile).
Maury (papers from 1970 to 1998) provided information on many Solifugae
inhabiting Argentina of the genera Pseudocleobis including P. alticola Pocock
(1900) up to 4700 m, Gaucha Mello-Leitao, Valdesia Maury, and the families
Ammotrechidae, Mummuciniidae, and Daesiidae (Kraus 1966; Maury 1976, 1980,
1981, 1982, 1983, 1987).
Scorpiones There is vast information on the scorpions, mostly of Argentina (see
table).
In Argentina are known 66 spp. of only 2 families – Bothriuridae (53 spp.) and
Buthidae (13 spp.) (Ojanguren-Affilastro, 2005 and suppl.). In Chile are recorded
53 spp. of 3 families – Bothriuridae (50 spp.), Buthidae (two spp.), and Caraboctonidae
(one sp.). In Patagonia is represented only the family Buthriuridae with the genera
Bothriurus Peters, Brachistosternus Pocock, and Urophonius Pocock. They live
even to the southern end of Tierra del Fuego (Provincia Fitogeografica Subantarctica).
It is clear that Bothriuridae (mainly the genera Bothriurus Peters and Brachistosternus
Pocock) is strongly prevailing in both countries (Maury (1968, 1975a, 1979), Mello-
Leitao (1934, 1943, 1945), Ringuelet (1953), Cekalovich (1983)).
The genera Centromachetes Lönnberg, Phoniocercus Pocock, Rumikiro
Ojanguren-Affilastro et al., and Tehuankea Cekalovic are endemic to Chile.
Uruguay has only five native genera (four Bothriuridae and one Buthidae).
Pseudoscorpiones After the very careful collection of G. Topál in the most south-
ern Andes (42oS), Beier (1964a) concluded that the pseudoscorpion fauna of
Patagonia is rather poor (20 species of 11 genera and 5 families – Chthoniidae,
Gymnobisiidae, Geogarypidae, Garypinidae, Chernetidae) (Beier 1962, 1967;
Mahnert et al. 2011; Vitali-di Castri 1970; Cekalovic 1976). Endemic genus is
Diplothrixochernes Beier (1962); the species are endemic to Patagonia in Argentina
and South Chile. Beier (1964b) is emphasizing the importance of the Andes as a
barrier along the meridian between the faunas of South Chile and South Argentina.
From Chile (incl. Juan Fernandez Isl.) are known 83 spp. of pseudoscorpions
(Cekalovic, 1984 and suppl.), from the whole of Argentina 70 spp. and from
Uruguay 5 spp.
In Argentina and Chile have been recorded pseudoscorpions of 55 genera and 16
families: Chthoniidae, Tridenchthoniidae, Pseudotyrranochthoniidae, Lechytiidae,
8.7 Patagonia: South Chile, Argentina, Uruguay, and Tierra del Fuego 677
Gymnobisiidae, Garypidae, Geogarypidae, Garypinidae, Ideoroncidae, Atemnidae,

Olpiidae, Menthidae, Cheliferidae, Cheiridiidae, Chernetidae, and Withiidae. From
Argentina are known 38 genera, from Chile also 38 genera, with 21 genera in
common. Most intriguing is the distribution of family Gymnobisiidae. From the 11
species, 1 (Beierobisium oppositum Vitali-di Castri) is an endemic genus and spe-
cies for the Falkland Islands, 2 (the genus Gymnobisium Beier) are endemic for
South Africa, and 8 have been described from Chile, Argentina, and Bolivia.
Opiliones From Argentina, Uruguay, and Chile of 30oS are known Opiliones of the
families Caddidae, Sclerosomatidae, Neopilionidae, Acropsopilionidae,
Triaenonychidae, Zalmozidae, Cosmetidae, and Gonyleptidae (Acosta 2002; Acosta
and Maury 1998; Canals 1936; nine papers from 1932 to 1943; Capocasale 1968;
Shear 1993b; Cokendolpher and Lanfranco 1985; Hunt and Cokendolpher 1991;
Maury 1986, 1988, 1991, 1992, 1993; Maury et al. 1996; Ringuelet 1955; 26 papers
from 1953 to 1963; Roewer 1961, 1962; Cecalovic 1968, 1976, 1985).
From 120 species and subspecies, 79 belong to the family Gonyleptidae (mostly
to Pachylinae). Zoogeographical analysis has been made several times, mostly by
Mello-Leitão (1939) and Ringuelet (1957, 1959).
Eupnoi Recorded are the families Caddidae, Neopilionidae, and Sclerosomatidae.
Endemic genera:
Fam. Neopilionidae
Ballarrinae – Chile, Australia, South Africa (ten spp.)
Americovibone Hunt et Cokendolpher, 1991 – Chile (one sp.)
Enantiobuninae – Brazil, Chile, Argentina (four spp.)
Thrasychirus Simon, 1884 – Chile, Argentina (three spp.)
Dyspnoi – Fam. Acropsopilionidae, Acropsopilio Forster
Laniatores Recorded are opilions of the families Gonyleptidae, Triaenonychidae,
Zalmoxidae, and Cosmetidae.
Endemic genera – Fam. Triaenonychidae: Picuchenops Maury, 1988, Argentina
(one sp.)
The zoogeography of Opiliones in Argentina has been treated several times
(Mello-Leitao 1939; Ringuelet 1957, 1959; Acosta and Maury 1998; Acosta 2002).
Mello-Leitão (1939) lists 79 spp. of Opiliones in Argentina and makes a subdivi-
sion of Argentina after the Arachnida into 5 zoogeographical provinces, including
“Provincia patagónica” (the area south of Rio Negro). The most important mono-
graph on Argentinian Opiliones is due to Ringuelet (1959). The highest number of
species exists in the three wetter areas: the Andino-Patagonian forests, the selva and
the yungas (Acosta and Maury 1998). Ringuelet (1959) proposed a subdivision of
Argentina based on Opiliones: several zones in the northeast and a narrow stripe
(mostly in Chile), called “zona austral,” including the “subzone malvinense” on the
Falkland Islands. Between these areas is spread a huge zone “Andina-Subandina-

Patagonica,” considered negative for opilions. Later some endemic (relic?) species
have been detected in this area.
The opiliofaunistical conclusions of this author (Ringuelet 1957, 1959) have
been altered by Acosta and Maury (1998) and Acosta (2002). After the last author,
in the “área Andino-patagónica” live 22 species/subspecies of Opiliones (2 of
Caddidae, 3 of Neopilionidae, 7 of Triaenonychidae, 10 of Gonyleptidae).
From the cave system, Cuchillo Curá Maury (1988) described the only
Argentinian troglobitic opilion Picunchenops spelaeus, belonging to a group far
from the other Triaenonychidae.
According to Acosta (2002), by this time the Argentinian fauna of Opiliones
includes120 “entidades” (species and subspecies), relatively few for the territory of
almost 2 800 000 km2. This is due to the dry conditions of most of the country.
Family Gonyleptidae is predominant with 79 “entidades” (69 of them belong to
Pachylinae, 57% of all Argentinian Opiliones).
Araneae In the austral part of South America are registered at least 62 families of
spiders, some of which are zoogeographically very interesting (Forster and Platnick
1984; Platnick and Forster 1989; Merian 1913; Richardson 2010; Zapfe 1961). The
families are Mecicobothriidae, Microstigmatidae, Hexathelidae, Dipluridae,
Nemesiidae, Cyrtaucheniidae, Idiopidae, Migidae, and Actinopodidae from
infraorder Mygalomorpha and Austrochilidae, Filistatidae, Drymusidae, Scytodidae,
Sicariidae, Diguetidae, Pholcidae, Caponiidae, Dysderidae, Oonopidae, Orsolabidae,
Segestriidae, Hersiliidae, Oecobiidae, Mecysmaucheniidae, Palpimanidae,
Malkaridae (incl. Pararchaeidae), Mimetidae, Deinopidae, Uloboridae, Anapidae
(incl. Micropholcommatidae), Araneidae (incl. Nephilidae), Linyphiidae,
Synotaxidae, Tetragnathidae, Theridiidae, Lycosidae, Oxyopidae, Pisauridae,
Senoculidae, Trechaleidae, Agelenidae, Amphinectidae, Amaurobiidae,
Anyphaenidae, Desidae, Dictynidae, Hahniidae, Sparassidae, Selenopidae,
Zodariidae, Zoropsidae, Clubionidae, Miturgidae (incl. Zoridae), Titanoecidae,
Gallienellidae, Gnaphosidae, Prodidomidae, Trochanteriidae, Philodromidae,
Thomisidae, Salticidae, and Corinnidae from suborder Arachnomorphae.
It is worth to analyze the distribution of some families.
Fam. Mecysmaucheniidae – The genera Mecysmauchenius Simon and
Mecysmauchenioides Forster et Platnick, 1984, are outlined by Grismado and
Lopardo (2003). The 17 species of this genus live in Patagonia and the islands
Malvinas (Falkland Island) and Juan Fernandez. Other genera of this family in
Chile and/or Argentina are Chilarchaea Forster et Platnick, 1984; Mesarchaea
Forster et Platnick, 1984; and Semysauenius Forster et Platnick, 1984. The last two
genera of the family are known from the other end of the world – New Zealand:
Aotearoa Forster et Platnick, 1984, and Zearchaea Wilton, 1946. This is one of the
amazing cases of faraway disjunctions.
Fam. Malkaridae – This small family (46 spp.) included the genera Perissopmerus
Butler, 1932, from the Australian continent; Malkara Davies, 1980, from
Queensland; Carathea Moran, 1986, from Tasmania; and, surprisingly, Chilenodes
8.8 Falkland Islands (Malvinas) 679
Platnick et Forster, 1987, from Chile and Argentina. Later to it was added the family
Pararchaeidae with six genera from Australia, Tasmania, and New Zealand.
Fam. Amphinectidae – From the 35 known genera of this family, 18 live in New
Zealand and 16 in Australia (incl. Tasmania), but 2 genera are known from Chile
(Calacadia Exline, 1960) and Western coast of South America (Metaltella Mello-
Leitão, 1932).
Opilioacarida Neocarus platensis (Silvestri, 1905) from “La Plata” (Argentina-
Uruguay) is the southernmost member of Opilioacarida in the New World.
Acariformes
Oribatida – Hammer (1962a, 1962b)
Parasitiformes – Ref.: Beron (2014)
From Panama, Brazil, and Uruguay is known the family Dasyponyssidae –
endemic to the Neotropics, as is composed by parasites on Armadillos (Cingulata,
Dasypodidae).
8.8 Falkland Islands (Malvinas)
The Falkland Islands (Islas Malvinas) are an archipelago in the Southern Atlantic
Ocean, on 51o45’S and about 480 km. from Argentina. The archipelago consists of
two main islands of East and West Falkland and ca. 200 small islands with total
surface area of 12 173 km2, on the Patagonian Shelf. The climate is cold marine,
with occasional snow all year, except in January and February, and with strong
winds throughout the year. The highest point is Mount Usborne (705 m). Falkland
Islands were originally part of South Africa. They have achieved their present posi-
tion during the fragmentation of Gondwana (Marshall 1994). Due to the strong
winds, they are treeless, with only dwarf shrubs, but have 163 spp. of native vascular
plants. The islands are biogeographically part of the mild Antarctic zone, with
strong connections to the flora and fauna of Patagonia (McDowall 2005). According
to Morrone (2000), biogeographically the Falkland Islands (Malvinas) are a prov-
ince within the Subantarctic subregion of the Andean Region.
Arachnogeography
Only one pseudoscorpion, two opilions, and some spiders and mites are known from
the Falkland Islands. They are among the southernmost arachnids (Ringuelet 1955).
Pseudoscorpiones Beierobisium oppositum Vitali-di-Castri, 1970, is the only
pseudoscorpion known from these islands (endemic genus and species of
Gymnobisiidae).
Opiliones Information about the few opilions on the islands is due to Soares and
Soares (1949), Hogg (1913), and Butler (1875).
Laniatores
Fam. Gonyleptidae: Haversia defensa (Butler), Haversia (Hoggellula) vallentini

(Hogg)
Araneae The spiders belong to the families Orsolobidae, Linyphiidae, Zodariidae,
and others (Forster and Platnick (1985), Hogg (1913), Lavery and Snazell (2013),
Platnick (1977), Schiapelli and Gershman de Pikelin (1974), Grismado and Platnick
(2008), Usher (1983)).
Endemic genera: Fam. Orsolobidae
Falklandia Forster et Platnick, 1985

Fam. Linyphiidae
Falklandoglenes Usher, 1983
Linga Lavery et Snazell, 2013
8.9 The Boundary Between Palearctic and Paleotropical
8.9.1 T
he Boundary Between Western Palearctic
and Afrotropical Arachnofaunas
It is a common knowledge that a biogeographical boundary is not a line but rather a
more or less wide transitional zone. We have also to take into account the historical
changes during the last several thousand years. Part of these changes is due to cli-
matic factors, others to human activities (also causing changes in the climate), like
deforestation, diverting of rivers, etc.
The traditional zoogeographic regions are delimitated mainly on the distribution
of terrestrial vertebrates. The difference between the regions and kingdoms is usu-
ally on much higher taxonomical level when we consider vertebrates, than with the
Arachnida. Differences between Afrotropical and Neotropical regions, or between
Holarctic and Paleotropical, are between orders or suborders and very much between
families. Historically it was not as it is today. In green Sahara and Mediterranean
Africa lived elephants, crocodiles, and giraffes; in Atlas Mountains, there were until
recently ostriches, lions, and bears. They were exterminated by man. We do not
speak about geological periods when these animals (plus Tubulidentata, Marsupialia,
8.10 Arachnida in the Afrotropical and Palearctic Regions 681
tapirs, and many others) lived in Europe. The present-day biogeographical subdivi-
sion should consider the picture of the last several thousand years.
We don’t know what was the distribution of many arachnids over the green
Sahara. It was not influenced by the rare human population during the millenia, as it
certainly was in Europe and North America with the disappearing of forests and their
replacement with anthropogenic landscape. Important were also new crops like maiz
and potatoes, chemization, and other kinds of human intruding into the ecosystems.
If we follow the classical subdivision (Africa south of Sahara and the southern-
most Arabia form the Afrotropical Region), north of this “line” is the Palearctic
(different subregions). This “line” (actually transitional zone) is separating two
kingdoms, so differences should be substantial. Let us check how the known distri-
bution of Arachnida fits into the classical scheme (Vachon 1951a, b, 1952; Beron
2017a).
8.10 Arachnida in the Afrotropical and Palearctic Regions
8.10.1 Analysis and Comments
The Afrotropical Region is bordering only the Western part of the huge Palearctic
Region (Saharo-Sindian Province). Many higher taxa are marked “present” for both
regions; but actually they may be distributed in tropical Africa and, say, Japan or
Korea. Sometime in between there is a gap of thousand kilometers, and the differ-
ence between the arachnofauna of tropical Africa and Europe is much bigger.
So some comments on the distributions of the various taxa are needed.
Palpigradi In tropical Africa are known eight described species of Palpigradi; one
genus (Allokoenenia Silv.) is endemic. In the Western Palearctic (Europe and
Lebanon) are recorded species, all from the genus Eukoenenia Börner
(Eukoeneniidae).
Ricinulei In the Old World this order is represented only by 11 species of 1 genus
(Ricinoides, endemic for West Africa).
Solifugae In the Afrotropical Region are registered eight families (three of which
from South Africa). One of the striking facts in zoogeography is the absence of
Solifugae in Madagascar.
Scorpiones In Africa south of 20oN live scorpions of five families with genera
(Bothriuridae, Buthidae, Hormuridae, Scorpionidae, Lisposomatidae). The under-
lined families are known only from Southern Africa.
Schizomida The northern limit of Schizomida in Africa is determined by their

thermopreferendum and the sands of Sahara – from Liberia to Kenya and Somalia.
There is one endemic subfamily in Southern Africa and five genera – all endemic for
narrow areas of Africa.
Thelyphonida (Uropygi) Only one genus (Etienneus) is represented in Africa (the

Western part). In the Palearctic the order is also almost not represented (one species
of Typopeltis in the far East of Russia).
Amblypygi In the Afrotropical Region (incl. South Africa) live members of five
genera of Euamblypygi of two families (Phrynichidae, Charinidae). Remarkable is
the finding in Guinea-Bissau of the most primitive member of the order – Paracharon
caecus Hansen, 1921 – sole living representative of a special family (Paracharontidae)
and even suborder (Palaeoamblypygi). Very few amblypygids have been recorded in
the Palearctic; none in Europe; Charinus in Turkey, the Dodecanese, Egypt, Israel,
and Pakistan; and Musicodamon in Algeria and Morocco.
Araneae We counted 81 families of spiders (out of the 112 families of spiders in

the world), related to both regions (living in one of them or in both). In both regions
occur 47 families, only in the Afrotropical Region 72 and also in the Palearctic
Region 72 families.
From the 72 families, represented in the Afrotropical Region, 10 are found only
in Southern Africa (south of Zambezi-Kunene).
Opilioacarida This peculiar order is represented in both territories by one family
but with different genera: Opilioacarus within South Europe and Algeria, with two
genera, endemic to West Africa (Panhaetes Naudo and Phalangiacarus Coineau
and Van der Hammen) and with Salfacarus Van der Hammen, known from Southern
Africa and Madagascar.
Differences in Subordinal Level The orders Ricinulei and Schizomida and the
suborder Palaeoamblypygi are found in Afrotropical Region but are absent in the
Palearctic. The suborders Mesothelae and Dyspnoi are represented in the Palearctic,
but not in tropical Africa (Table 8.4).
This boundary has been analyzed by Beron (2016) and other authors. Martens
(1984) concluded from his long experience in Himalaya that
We must take into account that the Himalayan fauna is predominantly one which invaded
the Himalayas at the time of or after the uplift of the mountain chain during the Tertiary and
that Himalayan fauna has only partially undergone an evolution of its own. Both climatic
belts and vegetation zones are largely in accordance with the areas of origin outside the
Himalayas of the various faunal elements. The Himalayan fauna is mainly an immigration
fauna. We distinguish five main centers of origin and thus five categories of Himalayan
fauna, three of which fall into the Palearctic (Central Asian, Himalayan West Asian,
Himalayan West Chinese) and two in the Oriental realm (Himalayan Indochinese,
Peninsular Indian.
When trekking along Kali Gandaki River, one have the strange feeling to cross a
boundary between two kingdoms – Holarctic and Paleotropic.
Table 8.4 The boundary between the Palearctic and the Indomalayan regions, indicated by
Arachnida
Group Region Region
Afrotropical Palearctic
Order Palpigradi Present Present
Fam. Eukoeneniidae Present Present
Order Ricinulei Present Absent
Fam. Ricinoididae Present Absent
Order Solifugae Present Present
Fam. Galeodidae Present Present
Fam. Karschiidae Absent Present
Fam. Daesiidae Present Present
Fam. Solpugidae Present Present
Fam. Ceromidae Present Absent
Fam. Melanoblossiidae Present (Southern Africa) Absent
Fam. Gylippidae Present (Southern Africa) Present
Fam. Hexisopodidae Present (Southern Africa) Absent
Fam. Rhagodidae Present Present
Order Scorpiones Present Present
Fam. Bothriuridae Present (Namibia) Indian Himalaya
Fam. Buthidae Present Present
Fam. Pseudochactidae Absent Present
Fam. Euscorpiidae Absent Present
Fam. Scorpiopidae Absent Present
Fam. Troglotayosicidae Absent ? Present
Fam. Iuridae Absent Present
Fam. Diplocentridae Absent Present
Fam. Hemiscorpiidae Absent Present
Fam. Hormuridae Present Present
Fam. Scorpionidae Present Present
Fam. Akravidae Absent Present
Fam. Lisposomidae Present (South Africa) Absent
Order Schizomida Present Absent
Fam. Hubbardiidae Present Absent
Order Uropygi Present (relict) Present
Fam. Hypoctonidae Present Present
Order Amblypygi Present Present
Suborder Neoamblypygi Present Present
Fam. Charinidae Present Present
Fam. Phrynichidae Present Present
Suborder Palaeoamblypygi Present Absent
Fam. Paracharontidae Present Absent
Order Opiliones Present Present
Suborder Cyphophthalmi Present Present
(continued)
Fam. Sironidae Present Present
Fam. Ogoveidae Present Absent
Fam. Neogoveidae Present Absent
Fam. Pettalidae Present (South Africa) Absent
Suborder Eupnoi Present Present
Fam. Caddidae Present (South Africa) Present (Japan)
Fam. Phalangiidae Present Present
Fam. Sclerosomatidae Present Present
Fam. Neopilionidae Present (South Africa) Absent
Suborder Dyspnoi Absent Present
Fam. Ischyropsalididae Absent Present
Fam. Sabaconidae Absent Present
Fam. Dicranolasmatidae Absent Present
Fam. Trogulidae Absent Present
Fam. Nemastomatidae Absent Present
Fam. Nipponopsalididae Absent Present (Japan, Korea)
Suborder Laniatores Present Present
Fam. Cladonychiidae Absent Present
(Holoscotolemon)
Fam. Travuniidae Absent Present
Fam. Triaenonychidae Present (South Africa) Present (Japan, Korea)
Fam. Assamiidae Present Absent
Fam. Biantidae Present Nepal, India
Fam. Epedanidae Absent Japan, India, Taiwan
Fam. Phalangodidae Absent Present
Fam. Podoctidae Present Present (Japan, India)
Fam. Pyramidopidae Present Absent
Fam. Samoidae Present (Mozambique) Absent
Fam. Zalmoxidae Seychelles Absent
Order Pseudoscorpiones Present Present
Suborder Epiocheirata Present Present
Fam. Chthoniidae Present Present
Fam. Tridenchthoniidae Present, India, Seychelles Japan, Bhutan, Nepal,
Fam. Absent Present
Pseudotyrannochthoniidae
Fam. Lechytiidae Present Present
Fam. Feaellidae Present India
Suborder Iocheirata Present Present
Fam. Ideoroncidae Present Present
Fam. Hyidae Absent India
Fam. Gymnobisiidae Present (South Africa) Present
Fam. Neobisiidae Present Present
Fam. Syarinidae Present Present
(continued)
Fam. Garypidae Present Present
Fam. Geogarypidae Present Present
Fam. Larcidae Absent Present
Fam. Cheiridiidae Present Present
Fam. Pseudochiridiidae Present India, Nepal
Fam. Olpiidae Present Present
Fam. Garypinidae Present (South Africa) Present
Fam. Menthidae Present (Socotra) Present (Israel)
Fam. Sternophoridae Present India
Fam. Withiidae Present Present
Fam. Cheliferidae Present Present
Fam. Atemnidae Present Present
Fam. Chernetidae Present Present
Order Araneae Present Present
Suborder Mesothelae Absent Present
Fam. Liphistiidae Absent Present
Suborder Orthothelae Present Present
Infraorder Mygalomorphae Present Present
Fam. Microstigmatidae Present (South Africa) Absent
Fam. Hexathelidae
(Macrothelinae) Present Present
Fam. Dipluridae (Euagrinae) Present Present
Fam. Nemesiidae Present (Southern Africa) Present
Fam. Theraphosidae Present Present
Fam. Atypidae Present Present
Fam. Antrodiaetidae Absent Present (Japan)
Fam. Cyrtaucheniidae Present Present
Fam. Idiopidae Present Present
Fam. Ctenizidae Present (Southern Africa) Present
Fam. Migidae Present Absent
Infraorder Araneomorphae Present Present
Fam. Archaeidae Present (Southern Africa) Absent
Fam. Hypochilidae Absent Present
Fam. Austrochilidae Present (Southern Africa) Absent
Fam. Filistatidae Present Present
Fam. Drymusidae Present (South Africa) Absent
Fam. Scytodidae Present Present
Fam. Sicariidae Present Present
Fam. Leptonetidae Absent Present
Fam. Ochyroceratidae Present ? Present (China)
Fam. Telemidae Present Present
Fam. Pholcidae Present Present
Fam. Caponiidae Present Absent
(continued)
Fam. Tetrablemmidae Present India
Fam. Dysderidae Present Absent
Fam. Oonopidae Present Present
Fam. Orsolobidae Present (Southern Africa) Absent
Fam. Segestriidae Present Present
Fam. Eresidae Present Present
Fam. Hersiliidae Present Present
Fam. Oecobiidae Present Present
Fam. Palpimanidae Present Present
Fam. Mimetidae Present Present
Fam. Deinopidae Present Present
Fam. Uloboridae Present Present
Fam. Anapidae Present Present
Fam. Araneidae Present Present
Fam. Cyatholipidae Present Present
Fam. Linyphiidae Present Present
Fam. Sinopimoidae Absent China (doubtful status)
Fam. Symphytognathidae Present Present (one sp. in Japan)
Fam. Synaphridae Absent (but present on Present
Madagascar)
Fam. Tetragnathidae Present Present
Fam. Theridiidae Present Present
Fam. Theridiosomatidae Present Present
Fam. Ctenidae Present Present
Fam. Lycosidae Present Present
Fam. Oxyopidae Present Present
Fam. Pisauridae Present Present
Fam. Psechridae Absent Present
Fam. Trechaleidae Absent Present
Fam. Udubidae Present Absent
Fam. Zoropsidae Present (South Africa) Present
Fam. Agelenidae Present Present
Fam. Amaurobiidae Present Present
Fam. Anyphaenidae Present (South Africa) Present
Fam. Cybaeidae Absent Present
Fam. Desidae Present Present
Fam. Dictynidae Present Present
Fam. Hahniidae Present Present
Fam. Sparassidae Present Present
Fam. Selenopidae Present Present
Fam. Zodariidae Present Present
Fam. Chummidae Present (South Africa, end.) Absent
Fam. Clubionidae Present Present
(continued)
Fam. Miturgidae Present Present
Fam. Phyxelididae Present Present
Fam. Titanoecidae Absent Present
Fam. Ammoxenidae Present (Southern Africa) Absent
Fam. Cithaeronidae Present Present
Fam. Gallieniellidae Present Absent
Fam. Gnaphosidae Present Present
Fam. Prodidomidae Present Present
Fam. Trochanteriidae Present Present
Fam. Philodromidae Present Present
Fam. Thomisidae Present Present
Fam. Salticidae Present Present
Fam. Corinnidae Present Present
Fam. Liocranidae Present Present
Order Opilioacarida Present Present
Fam. Opilioacaridae Present Present
Order Holothyrida Absent (only Seychelles) Absent
Order Ixodida Present Present
Fam. Argasidae Present Present
Fam. Ixodidae Present Present
Fam. Nuttalliellidae Present (Southern Africa) Absent
Order Mesostigmata Present Present
Order Sarcoptiformes Present Present
Order Trombidiformes Present Present
In his analysis of the distribution of Mammals in the Himalayan ranges, Hoffmann

(2001) concludes that, because of the strong altitudinal gradient, “Palearctic elements
dominate higher, and Indomalayan elements, lower elevations” (being in the Himalaya
almost equally represented). Further Hoffmann (2001) analyzes in details the transi-
tional zone in such complicated area as North Burma, Sichuan, and Yunnan (Map 8.2).
Concerning the insects, Krijanovskij (2002) accepted the existence of an Indo-
Malayan Region with three subregions: Malayan (Malacca, the Sunda Islands, and
the Philippines), Indian (most of Hindustan and Sri Lanka), and Indo-Chinese
(Assam, Manipur, Indo-China without Tenasserim, Malacca, and the high Northern
Burma, also the tropical South China and the islands Hainan, Taiwan, and Ryukyu).
According to the catalogue of Fet (1988) and other sources, from the present ter-
ritory of Russia (including Crimea) are known five spp. of scorpions.
The neighboring Kirghizstan has also only the same two Mesobuthus (cauca-
sicus and eupeus), but Tajikistan, situated near the scorpion-rich (29 spp.)
Afghanistan, has already 6 spp., including Hottentotta, Orthochirus, Psammobuthus,
and Pseudochactas.
Gromov (2001) outlined the northern limit of the distribution of Scorpiones in
Central Asia. From the area in the Palearctic East from Yenisei River are known
scorpions of the families Euscorpiidae, Buthidae, and Hemiscorpiidae (Russia East

of Yenisei, North China, Mongolia, Korea, and Palearctic Japan). In Mongolia live
only three species of genus Mesobuthus Vachon (M. eupeus, M. caucasicus, M.
martensii). In China north of Yangtze are recorded very few scorpions. In Korea are
recorded only two scorpions: Mesobuthus martensii and the widespread Liocheles
australasiae. Japan has also two widespread species – the same Liocheles austral-
asiae and Isometrus maculatus.
The scorpiofauna of Pakistan reflects the transitional fauna of this country: 46
species of the families Buthidae (Androctonus Ehr., Baloorthochirus Kov., Buthacus
Birula, Buthoscorpio Werner, Compsobuthus Vachon, Hemibuthus Pocock,
Hottentotta Birula, Isometrus Ehr., Lychas C.L. Koch, Mesobuthus Vachon,
Odontobuthus Vachon, Orthochirus Karsch, Plesiobuthus Pocock, Sassandiothus
Farzanpay, Vachonus Tikader et Bastawade), Euscorpiidae (Euscorpiops Vachon,
Scorpiops Vachon), Hemiscorpiidae (Hemiscorpius Peters), and Scorpionidae
(Heterometrus Ehr.).
8.11 The Boundary Across China
According to Corbet (1978), the Yangtze River was “…just beyond the southern
limit of the [Palearctic] region,” and further “…in lowland China the boundary is
taken very arbitrarily as Latitude 35oN, corresponding in part with the Hwang Ho
(Yellow River).” The broad area between these two rivers has warm temperate cli-
mate (Fan 1990) and transitional zone between the two realms. Corbet and Hill
(1992) confirm the idea that the arbitrary northern boundary between the Palearctic
and the Indomalayan region is about 35oN (Yellow River). Zhang and Zhao (1978)
placed the median line “a little south of Yangtze.” Again Corbet and Hill (1992)
defined three divisions of the transitional zone between the Hwang Ho and the
Yangtze (see Hoffmann 2001 for details).
All these subdivisions were based on vertebrates, a group with many anthropo-
genic changes during the last centuries. It would be interesting to compare this dis-
cussion with the results obtained by a detailed analysis on the distribution of all
orders of Arachnida (so far much less known in this area).
8.11.1 The Transitional Zone
We discussed already the map and the analysis on mammals (Hoffmann 2001).
Recently Morrone (2015) published a biogeographic map, showing the “Chinese
Transitional Zone.” A higher category has been assigned to this (or a similar) zone
in the subdivision of Holt et al. (2013) – an independent “Sino-Japanese Realm.”
Most families (93) of Arachnida are in common to both regions. There are no
endemic orders or suborders in any of them. What concerns the Arachnida is that
8.11 The Boundary Across China 689
their distribution does not justify the sharp difference between the two kingdoms
(Paleotropical and Holarctic) in Eastern Eurasia. The transitional zone (Sino-
Japanese Realm) of Holt et al. (2013) also does not satisfy the criteria for outlining
an area on the same footing as Palearctic and Indomalayan Realms.
Cluster analyses, based again on terrestrial vertebrates (He et al. 2017), revealed
nine major zoogeographical regions in China: South China, the Yungui Plateau,
Taiwan, North China, the Inner Mongolia Plateau, Northwest China, the Longzhong
Plateau, the Tibetan Plateau, and East Himalaya. He et al. (2017) suggest a major
south-north-oriented Palearctic-Oriental boundary in China.
Other sources: Palestrini and Zunino (1986), Palestrini et al. (1986).
8.11.2 Arachnogeography of Himalaya
Typical for the Himalaya is the giant mountain system on the limit or transition
between two regions and even kingdoms. The prominent arachnologist and explorer
of the Himalaya, Jochen Martens, analyzed in several articles the distribution of the
Palearctic and Indomalayan elements in the mountain (Martens 1973, 1983, 1984,
1987, 1993). The arachnofauna of Himalaya was analyzed by Beron (2001a, 2001b,
2008), Mani (1959), and others.
Map 8.3 The transitional zone between Palearctic and Indomalayan regions (After Hoffmann,
2001, modified)
8.11.3 E
ast Asiatic (Himalayan-Chinese) Region of Lopatin
(1980)
Although much smaller (the extratropical parts of Southeast Asia, including parts of
the Himalaya, Tibet, Northeast China, Primorye, Korea, and the three bigger islands
of Japan South of Hokkaido), this area is very interesting and contains some “south-
ern” elements like Uropygi (Typopeltis). Still missing are Amblypygi, Schizomida,
Palpigrada, Opilioacarida, and Holothyrida; there are very few scorpions
(Mesobuthus, Liocheles).
Some of the arachnids of Japan are clearly relicts (opilions like Suzukiellus,
Yuria, spiders of suborder Mesothelae); some genera and even species (Caddo agi-
lis), as well as the subfamily Paranonychinae, show affinities with the Nearctic
fauna.
The climate of this area is very different from the climate of the European-Obian
and the Angaran Subregions. The lowlands have mild, even warm climate, the prox-
imity of the ocean is causing high humidity, and the forests are broadleaf, some of
them even evergreen. Of course, the high mountains have many belts with different
climates. In the area there are many.
8.12 The Cape Arachnofauna
8.12.1 Africa South of Zambezi-Kunene

This arbitrary-outlined area includes South Africa, Lesotho, Swaziland, Botswana,

Namibia, Zimbabwe, and Mozambique (south of Zambezi). Southern Africa has a
wide diversity of ecoregions including grassland, bushveld, karoo, savannah, and
riparian zones. It has complex plateaus that create massive mountain structures
along the South African border. The highest mountains are Drakensberg (Mafadi,
3450 m) in South Africa and Lesotho and Brandberg (2606 m) in Namibia. The
highest peaks in South Africa not part of the Drakensberg Escarpment are located in
the Karoo range in the southwestern part of the country. The two highest peaks in
the Karoo are Compassberg and Seweweekspoortpiek, with summit elevations of
2504 m and 2325 m, respectively (Killick 1978; Mahaney (Ed.)1989; Martin 1961;
Moreau 1952, 1963; Taylor 1978; White 1978; van Zinderen Bakker 1975).
8.12 The Cape Arachnofauna 691
Map 8.4 Map of Southern Africa
The Mediterranean climates are restricted to the extreme southwestern Cape.

The Cape flora is so distinct that it is recognized as a floral kingdom of its own
(Capensis). This is justified by the richness of species, the high degree of endemism
(especially of families), and the disjunct distribution of many of its members
(Werger 1978).
Arachnogeography
Southern Africa is very different botanically from the continent more to the North,
so different, that is, considered a kingdom. Much less are the differences in the
fauna – usually we speak about Cape Subregion of the Afrotropical Region. This
land of deserts, mountains, and an analogue of the Mediterranean climate still con-
tains many endemic groups, mostly on a level genus. There are many riddles in this
southern end of Africa – connections with other southern lands and sharp difference
with the fauna of Madagascar and the Mascarene.
The arachnofauna of southern parts of Africa is rich and relatively well studied
by people like Lawrence, Beier, Prendini, Starega, and other researchers. We now
know that in the area south of Kunene-Zambezi are present the orders Palpigradi (1
end. sp.), Solifugae (the richest part of the world that concerns this order), Scorpiones
(140 spp., 1 end. subfamily), Opiliones (3 end. genera of Cyphophthalmi, 1 end.
subfamily of Eupnoi and several endemic genera, 1 end. subfamily in Laniatores,
and 164 spp., all South African genera are endemic), Pseudoscorpiones (122 spp.,
73% endemic, 14 end. genera), Amblypygi (rich fauna with 1 endemic genus), and
Schizomida (1 endemic subfamily, all species are endemic).
Araneae (64 families, 2 of them endemic), Opilioacarida (2 spp., part of a genus
known in Madagascar), Ixodida (1 extremely interesting endemic family, known from
Tanzania, South Africa, and Namibia). There are endemic genera also in group like
Prostigmata and other Acari. Lacking from Southern Africa are the orders Ricinulei
and Holothyrida and the suborders Palaeoamblypygi, Mesothelae, and Dyspnoi
(Haddad 2004; Lawrence 1955, 1963) (Monod 1957; Newlands 1978; Werger 1978).
A total of 821 spp., 296 genera, and 69 families of non-acarine arachnids
(Araneae, Solifugi, Scorpiones, Pseudoscorpiones, Opiliones, and Amblypygi)
were known from Namibia (Griffin 1998); now (2017) they are certainly more.
Palpigradi Eukoenenia lawrencei Remy, 1957, is the only species of this order,
known from the described area.
Solifugae The area under consideration (South Africa, Lesotho, Namibia,

Swaziland, Botswana, Zimbabwe, southern part of Mozambique) is extremely rich
in solpugids. Here are represented 6 of the 12 families (incl. 17 end. genera) and
many of the 153 known genera of Solifugae, including the endemic family
Hexisopodidae and the endemic genera Hexisopus (15 spp.) and Chelypus (9 spp.)
(Dippenaar-Schoenman and González Reyes 2006; Hewitt 1919; Lawrence 1955;
Purcell 1903; Roewer 1941; Wharton 1981). With 127 spp. of Solifugae, 47 (37%)
of which are endemic and 9 genera occurring only in Namibia, this country appears
to be the richest in the world in Solifugae (Griffin 1998).
An older, but most reliable, source of information on the group is the checklist of
Lawrence (1955), listing from this area 181 species of 33 genera and 6 families.
Wharton (1981) outlined the solpugids of Namibia. According to Harvey (2013b),
in South Africa are recorded 163 spp. of Solifugae of 18 genera and 6 families.
The families Gyllipidae and Rhagodidae are shared with Asia, Solpugidae only
with the rest of Africa, Ceromidae with southern part of tropical Africa, and only
Daesiidae also with South America. Fossil Ceromid is recorded from Brazil.
Endemic genera of Solifugae south of Zambezi-Kunene:
Fam. Ceromidae – Ceromella Roewer (three spp.), Toreus Purcell (one sp.)
Fam. Gyllipidae – Bdellophaga Wharton (one sp.), Trichotoma Lawrence (three
spp.)
Fam. Hexisopodidae – Hexisopus Karsch (15 spp.), Chelypus Purcell (9 spp.)
Fam. Daesiidae – Namibesia Lawrence (one sp.)
Fam. Melanoblossiidae (Melanoblossiinae end. in Southern Africa) – Daesiella

Hewitt (one sp.), Lawrencega Roewer (seven spp.), Melanoblossia Purcell (four
spp.), Microblossia Roewer (one sp.), Unguiblossia Roewer (two spp.)
Solpugidae – Metasolpuga Roewer (one sp.), Prosolpuga Roewer (one sp.),
Solpuguna Roewer
Scorpiones The checklist of Lawrence (1955, with additions) contains 104 spp. of
scorpions from Southern Africa with many subspecies. More recent checklist was
provided by Prendini (2005). The scorpions south of Kunene-Zambezi belong to 14
genera and to the present-day families Bothriuridae (Brandbergia Prendini,
Lisposoma Lawrence) and Buthidae.
An important review of the scorpions of Namibia belongs to Lamoral (1979). In
the monograph find place two families (Buthidae and Scorpionidae) and seven gen-
era (Buthotus, Karasbergia, Parabuthus, Uroplectes, Hadogenes, Lisposoma,
Opisthophthalmus). The conclusions of Lamoral are that “The scorpion fauna of
Namibia is derived from elements that originated in Laurasia during Pangean times.
These elements migrated overland to the North African region of Gondwanaland
and ultimately to southern Africa in the wake of the southward shift of the warm
tropical belt. …No affinities could be found between the scorpion fauna of either
Namibia or southern Africa and South America.”
“Namibia has the highest species richness and endemism of scorpions in south-
ern Africa. All four families, seven genera (64%), and 57 sp. (44%) of southern
African scorpions occur within its borders, of which one genus and at least 27 spe-
cies (21%) are endemic” (Prendini, 2003a). After the description of Brandbergia
Prendini, 2003, the endemism becomes even higher but still cannot compare with
Madagascar (100%). African, Australian, and the only Indian member of
Bothriuridae could be considered as palaeoendemics (many authors, generalized by
Prendini (2003a)). Most Bothriurids live in South America, so Bothriuridae displays
a purely Gondwanan distribution, as do Scorpionoidea as a whole.
The survey of Prendini (2005) on the scorpions of Southern Africa contains data
on 13 genera and 140 spp.
Pseudoscorpiones The checklist of Dippenaar-Schoenman and Harvey (2000) enu-
merates 135 species from South Africa, belonging to 65 genera and 15 families (Beier
14 papers from 1931 to 1970, Ellingsen 1912; Lawrence 1935; Newlands 1978).
Endemic genera in South Africa (incl. Lesotho, Botswana, Namibia, and
Swaziland):
Fam. Chthoniidae
Selachochthonius Chamberlin, 1929 – South Africa, Lesotho (three spp.)
Fam. Garypidae
Ammogarypus Beier, 1962 – Namibia, South Africa (three spp.)
Eremogarypus Beier, 1955 – Namibia (four spp.)
Meiogarypus Beier, 1955 – Namibia (one sp.)
Fam. Gymnobisiidae
Gymnobisium Beier, 1931 – South Africa, Lesotho (two spp.)
Fam. Ideoroncidae
Botswanoncus Harvey et Du Preez, 2014 – Botswana (one sp.)
Fam. Cheliferidae
Aperittochelifer Beier, 1955 – South Africa (five spp.)
Litochelifer Beier, 1948 – South Africa (one sp.)
Rhopalochelifer Beier, 1964 – South Africa (one sp.)
Fam. Chernetidae
Myrmochernes Tullgren, 1907 – South Africa (one sp.)
Fam. Withidae
Afrowithius J.C. Chamberlin, 1931 – South Africa (one sp.)
Cyrtowithius Beier, 1955 – Namibia, South Africa (two spp.)
Paragoniochernes Beier, 1932 – South Africa (two spp.)
Parallowithius Beier, 1955 – Namibia, South Africa (two spp.)
Pseudatemnus Beier, 1947 – South Africa (one sp.)
Very high endemism on species level: Ninety-seven species (73%) are known
only from South Africa, 33 spp. are more widespread in the Afrotropical Region,
and 3 spp. are cosmopolitan. In the paper of Harvey and Du Preez (2014) are
described a new genus and species of cave pseudoscorpion from Botswana
(Botswanoncus ellisi, Ideoroncidae).
Opiliones A “preliminary” analysis of the zoogeography of Southern African
Opiliones was made by Starêga (1989a). According to him, south of Kunene-
Zambezi live 217 nominal species of Opiliones and “the largest part of this fauna
(63,6%) is of Gondwanan origin. Other groups represented belong to the
Paleotropical (19,8%) and Mediterrano-Afrotropical element (16%).” Since this
analysis, more species have been added (Lotz 2009; de Bivort and Giribet 2010), or
now (2017) the species are 225 (222 endemic) from 53 genera and 9 families
(Hansen and Soerensen 1904; Kauri 1961; Lawrence 1931, 1933, 1934, 1938, 1963;
Rosas Costa 1950; Starêga 1984, 1992, 2008). Starega (1989a) concludes that “the
largest part of the southern African harvestmen fauna shows clear relationship with
the faunas of other remnants of the former Gondwana.”
Cyphophthalmi In the recent paper of de Bivort and Giribet (2010) have been
described 7 new species, bringing the number of South African Cyphophthalmi to
15 spp. of 3 endemic genera (Purcellia Hansen et Soerensen, 1904 [syn. Speleosiro
Lawrence, 1931], Parapurcellia Rosas Costa, 1950) and 1 family – Pettalidae,
shared with Madagascar, Chile, Sri Lanka, Australia, and New Zealand.
Eupnoi. In the area have been recorded 46 spp. of 5 genera of Caddidae,
Phalangiidae, and Neopilionidae (Lotz 2009).
Endemic genera are:
Fam. Caddidae: Caddella Hirst, 1925 – Six spp.
Fam. Neopilionidae
Neopilioninae – Endemic subfamily (three spp.)
Neopilio Lawrence, 1931 – Two spp.
Vibone Kauri, 1961 – One sp.
Dyspnoi – Not recorded in the described area
Laniatores In the area have been recorded 164 spp. from 45 genera and 5 families:
Triaenonychidae, Assamiidae, Samoinae, Trionyxellidae, and Biantidae. Family
Triaenonychidae (from the six subfamilies, only Adaeinae is endemic in Southern
Africa) is “the most important component of the South African fauna” (Staręga
1989a) with 125 spp. (more than 50% of all Opiliones in this area). All South
African genera of Triaenonychidae are endemic.
Adaeulum Roewer, 1914 (12 spp.); Amatola Lawrence, 1931 (6 spp.);
Austromontia Lawrence, 1931 (6 spp.); Austronuntia Lawrence, 1931; Biacumontia
Lawrence, 1931 (7 spp.); Cryptadaeum Lawrence, 1931 (1 sp.); Graemontia
Lawrence, 1931 (6 spp.); Larifuga Loman, 1898 (10 spp.); Larifugella Lawrence,
1933 (5 spp.); Lawrencella Strand, 1932 (1 sp.); Lizamontia Kury, 2004 (4 spp.);
Mensamontia Lawrence, 1931 (2 spp.); Micradaeum Lawrence, 1931 (1 sp.);
Micromontia Lawrence, 1939 (1 sp.); Monomontia Lawrence, 1931 (17 spp.);
Montadaeum Lawrence, 1931 (1 sp.); Paradaeum Lawrence, 1931 (1 sp.);
Paramontia Lawrence, 1934 (2 spp.); Roeweriana Lawrence, 1934 (1 sp.);
Rostromontia Lawrence, 1931 (3 spp.); Speleomontia Lawrence, 1931 (1 sp.);
Yulella Lawrence, 1939 (1 sp.)
Fam. Assamiidae
Cryptopygoplus Lawrence, 1931 – Three spp.
Polycoryphus Loman, 1902 – Namibia, South Africa (one sp.)
Umtaliella Lawrence, 1934 – Zimbabwe (one sp.)
Fam. Biantidae
Biantessus Roewer, 1949 (two spp.)
Cryptobiantes Kauri, 1962 (one spp.)
Amblypygi In the catalogue of Lawrence (1955) are listed three species of
Amblypygi from Southern Africa: Damon variegates (Perty, 1834) (Damoninae),
Phrynichus scullyi (Purcell, 1901), and Ph. bacillifer (Gerstächer, 1873)
(Phrynichidae).
Weygoldt (1996, 1999) added the endemic genus and species Phrynichodamon
scullyi Weygoldt and the new species Damon brachialis, among others.
Prendini et al. (2005) described another new species (Damon sylviae) and pro-
vided a table of all 11 spp. in genus Damon and a key with the 5 species from
Southern Africa: Damon annulatipes, D. brachialis, D. gracilis, D. variegates, and
D. sylviae.
Schizomida
Endemic subfamily (Megaschizominae), genus (Megaschizomus Lawrence,

1969), and two species – M. mossambicus (Lawrence) and M. zuluanus (Lawrence) –
are the only Schizomida south of Kunene-Zambezi (Lawrence 1947, 1955, 1969).
Araneae
In Southern Africa are recorded spiders of 64 families, 2 of them endemic (Griffin

and Dippenaar-Shoeman 1992; Griswold and Platnick 1987; Haddat et al. 2009;
Hewitt 1916, 1917; Huber 2003; Jocqué 2001; Lawrence 1938; Purcell 1902;
Scharff 1992; Tucker 1923). South of Zambezi-Kunene have been recorded at least
105 endemic genera and 2 endemic families of spiders.
Endemic spider supraspecific taxa in Southern Africa:
Fam. Anapidae
Dippenaaria Wunderlich, 1995 – South Africa (one sp.)
Fam. Theraphosidae
Brachionopus Pocock, 1897 – South Africa (five spp.)
Ceratogyrus Pocock, 1897 – Botswana, South Africa, Zimbabwe, Malawi,
Mozambique, Namibia
Harpactira Ausserer, 1871 – South Africa, Namibia (16 spp.)
[Harpactirella Purcell, 1902 – South Africa (11 spp.), ? Morocco (one sp.)
Idiothele Hewith, 1919 – Southern Africa (two spp.)
Trichognathiella Gallon, 2004 – South Africa (one sp.)
Fam. Chummidae – Endemic family (two spp.)
Chumma Jocqué, 2001 – (two spp.)
Fam. Microstigmatidae (the other genera are known from South America)
Microstigmata Strand, 1932 – South Africa (six spp.)
Fam. Gallieniellidae
Austrachelas Lawrence, 1938 (nine spp.), Drassodella Hewitt, 1916 (seven
spp.) – South Africa
Fam. Orsolobidae
Afrilobus Griswold et Platnick, 1987 (three spp.), Azanialobus Griswold et
Platnick, 1987 (one sp.)
Fam. Amaurobiidae
Chresiona Simon, 1903 (three spp.); Obatala Lehtinen, 1967 (one sp.);
Pseudauxinum Simon, 1902 – South Africa
Fam. Dictynidae
Shango Lehtinen, 1967 – South Africa (one sp.)
Fam. Cyrtaucheniidae
Homostela Simon, 1892 – South Africa (five spp.)
Fam. Trachelidae
Afroceto Lyle et Haddad, 2010, Southern Africa (16 spp.); Fuchiba Haddad et
Lyle, 2008, South Africa, Botswana, Lesotho, Mozambique (6 spp.);
Fuchibotulus Haddad et Lyle, 2008, South Africa, Mozambique (3 spp.);
Poachelas Haddad et Lyle, 2008, South Africa, Zimbabwe (4 spp.);
Spinotrachelas Haddad, 2006, South Africa (5 spp.)
Fam. Caponiidae
Diplogena Purcell, 1904 – South Africa, Namibia, Botswana (six spp.)
Fam. Mysmenidae
Isela Griswold, 1985 – South Africa (one sp.)
Fam. Oecobiidae
Paroecobius Lamoral, 1981 – South Africa, Botswana (two spp.)
Uroecobius Kullmann et Zimmermann, 1976 – South Africa (one sp.)
Fam. Zodariidae
Caesetius Simon, 1893, South Africa, Namibia, Mozambique, Malawi (ten spp.);
Cicynethus Simon, 1910, South Africa, Namibia (three spp.); Heradida
Simon, 1893, South Africa, Namibia (six spp.); Procydrela Jocqué, 1999,
South Africa (two spp.); Psammodnon Jocqué, 1991, South Africa, Namibia
(three spp.); Psammorygma Jocqué, 1991, South Africa, Namibia; Rotundrela
Jocqué, 1999, South Africa (two spp.); Thaumastochilus Simon, 1897, South
Africa (two spp.)
Fam. Zoropsidae
Griswoldia Dippenaar-Schoeman et Jocqué, 1997 – South Africa (12 spp.)
Phanotea Simon, 1896 – South Africa (13 spp.)
Fam. Pisauridae
Tapinothelella Strand, 1909, South Africa (one sp.); Voraptipus Roewer, 1955,
Mozambique (one sp.); Walrencea Blandin, 1979, South Africa (five spp.)
Fam. Oonopidae
Australoonops Hewitt, 1915, South Africa (one sp.); Calculus Purcell, 1910,
South Africa (one sp.); Dalmasula Platnick et al., 2012, South Africa, Namibia
(five spp.); Nephrochirus Simon, 1910, Namibia; Pseudoscaphiella Simon,
1907, South Africa (one sp.)
Fam. Eresidae
Paradonea Lawrence, 1968 – South Africa, Zimbabwe, Botswana, Namibia
(five spp.)
Seothyra Purcell, 1903 – Namibia, Botswana, South Africa, Angola (13 spp.)
Fam. Araneidae
Ideocaira Simon, 1903 (three spp.), Nemospiza Simon, 1903 (one sp.) – South
Africa
Fam. Ammoxenidae
Ammoxenus Simon, 1893 – South Africa, Namibia, Botswana (six spp.)
Rastellus Platnick et Griffin, 1990 – South Africa, Namibia, Botswana,
Zimbabwe (seven spp.)
Fam. Palpimanidae
Ikuma Lawrence, 1938 – Namibia (two spp.)
Fam. Penestomidae – Endemic family!
Penestomus Simon, 1902 – South Africa, Lesotho (three spp.)
Fam. Theridiidae
Chorizopella Lawrence, 1947 (one sp.), Histagonia Simon, 1895 (one sp.) –
South Africa
Fam. Liocranidae
Coryssiphus Simon, 1903 – South Africa (three spp.)
Fam. Clubionidae
Carteroniella Strand, 1907 – South Africa (one sp.)
Fam. Prodidomidae
Austrodomus Lawrence, 1947 (two spp.); Eleleis Simon, 1893 (one sp.);
Purcelliana Cooke, 1964 (one sp.) – South Africa
Namundra Platnick et Bird, 2007 – Namibia, Angola (four spp.)
Theuma Simon, 1893 – South Africa, Namibia (25 spp.)

Fam. Idiopidae
Ctenolophus Purcell, 1904, Segregara Tucker, 1917 – South Africa
Galeosoma Purcell, 1903, Gorgyrella Purcell, 1902 – Southern Africa
Fam. Miturgidae
Parapostenus Lessert, 1923 – South Africa, Lesotho (one sp.)
Fam. Phyxelididae
Lamaika Griswold, 1990 (one sp.); Malaika Lehtinen, 1967 (two spp.);
Matundua Lehtinen, 1967 (one sp.); Namaquarachne Griswold, 1990 (five
spp.); Pongolania Griswold, 1990 (two spp.); Themacrys Simon, 1906 (five
spp.); Vidole Lehtinen, 1967 (five spp.) – South Africa
Xevioso Lehtinen, 1967 – South Africa, Malawi, Zimbabwe, Swaziland (nine
spp.)
Fam. Hersiliidae
Tyrotama Foord et Dippenaar-Shoeman, 2005 – South Africa, Namibia, Angola
(eight spp.)
Fam. Eutrichuridae
Lessertina Lawrence, 1942 – South Africa (two spp.)
Fam. Microstigmatidae
Microstigmata Strand, 1932 – South Africa (six spp.)
Fam. Cyatholipidae
Ilisoa Griswold, 1987 (three spp.), Ubacisi Griswold, 2001 – South Africa
Fam. Nemesiidae
Lepthercus Purcell, 1902; Pionothele Purcell, 1902; Spiroctenus Simon, 1889 –
South Africa
Fam. Linyphiidae
Afribactrus Wunderlich, 1995 (one sp.); Lucrinus O.P.-Cambridge, 1904;
Erigonops Scharff, 1990 (one sp.) – South Africa
Fam. Sparassidae
Arandisa Lawrence, 1938 (one sp.); Carparachne Lawrence, 1962 (two spp.);
Mierorchestris Lawrence, 1962 (two spp.); Orchestrella Lawrence, 1965
(two spp.); Palystella Lawrence, 1928 (four spp.) – Namibia
Panaretella Lawrence, 1937 (five spp.), Parapalystes Croeser, 1996 (five spp.) –
South Africa
Fam. Lycosidae
Minicosa Alderweireldt et Jocqué, 2007 – South Africa
Fam. Gnaphosidae
Smionia Dalmas, 1920 – South Africa
Titus O.P.-Cambridge, 1901 – Zimbabwe
Trephopada Tucker, 1923, Trichothyse Tucker, 1923 – South Africa, Namibia
Fam. Philodromidae
Hirriusa Strand, 1932 – South Africa, Namibia (three spp.)
Fam. Thomisidae
Avelis Simon, 1895 – South Africa (one sp.)
Stiphropella Lawrence, 1952 – South Africa (one sp.)
Fam. Salticidae
Araegeus Simon, 1901 – South Africa, Mozambique
Ureta Wesolowska et Haddad, 2013 – South Africa
Zulunigma Weselowska et Cumming, 2011 – South Africa (one sp.)
Fam. Corinnidae
Austrophaea Lawrence, 1952; Pronophaea Simon, 1887 (three spp.);
Vendaphaea Haddad, 2009 (one sp.) – South Africa
Opilioacarida Two species have been described, both in the genus Salfacarus Van
der Hammen, common with Tanzania and Madagascar (S. dispar V.d. Hammen and
S. lawrencei V.d. Hammen) (Van der Hammen 1977).
Acariformes
Fam. Erythraeidae
Endemic genera: Kakamasia Lawrence, 1944 – South Africa
Oribatida. Niedbala (2006) presents an outline of the ptyctimous mites in South
Africa.
Parasitiformes
Ixodida From Tanzania to Namibia and South Africa lives one of the strangest
ticks in the world, strange enough to form a family of its own, Nuttalliellidae, on the
same level with Ixodidae and Argasidae. This is Nuttalliella namaqua Bedford,
1931, described from the Little Namaqualand.
8.14 Madagascar 701
Conclusion – Very high endemism on species level: Ninety-seven species

(73%) are known only from South Africa, 33 spp. are more widespread in the
Afrotropical Region, and 3 spp. are cosmopolitan (in the paper of Harvey and Du
Preez (2014)).
8.13 Arachnofauna of Madagascar and the Adjacent Islands
Here is outlined and compared the arachnofauna of Madagascar, the Seychelles, and
the island groups Comoro, Aldabra, and Mascarene (Benoit 1977, 1979; Fage 1929,
1954; Heurtault 1986; Hoogstraal 1953; Lawrence 1959; Lourenço 1995, 1996a, b,
1999, 2000a; Lourenço et al. 2006, Millot 1948, Vachon 1979b, Vasquez and
Klompen 2010).
8.14 Madagascar
8.14.1 G
and Paleogeography
The Big Island (587, 041 km2) is 419 km far from the coast of Mozambique. The
highest point of Madagascar is Maromokotro (2876 m.), in the TsaratananaMassif;
other higher massifs are Ankaratra (2642 m.), Andringitra (2658 m.), and others. In
the last century, 95% of the native forests have been destroyed by logging and slash-
and-burn cultivation. The population (over 20 million, ten times more than in 1900)
is growing very fast, and the unique nature of the island is under severe threat
(Badyukov et al. 1989; Embleton and McElhinny 1975; Flores 1970; Furon 1949;
Katz and Premoli 1979; McElhinny et al. 1976; Paulian 1951; Rabinowitz et al.
1983; Smith and Hallam 1970).
The prehistoric breakup of the Gondwana supercontinent separated the
Madagascar-Antarctica-India landmass from the Africa-South America landmass
around 135 million years ago. Madagascar later split from India around 88 million
years ago, allowing plants and animals on the island to evolve in complete isolation.
In the mid-Eocene to the early Miocene, the channel may have been spanned by a
land bridge allowing terrestrial mammals to move between Madagascar and the
mainland. Now the Mozambique Channel is 3292 m. deep.
According to Warren et al. (2010): “Corollary of island biogeographical theory
is that islands are largely colonized from their nearest mainland source. Despite
Madagascar’s extreme isolation from India and proximity to Africa, a high propor-
tion of the biota of the Madagascar region has Asian affinities. This pattern has
rarely been viewed as surprising, as it is consistent with Gondwanan vicariance.

Molecular phylogenetic data provide strong support for such Asian affinities, but
often not for their vicariant origin; most divergences between lineages in Asia and
the Madagascar region post-date the separation of India and Madagascar consider-
ably (up to 87 Myr), implying a high frequency of dispersal that mirrors coloniza-
tion of the Hawaiian archipelago in distance. Indian Ocean bathymetry and the
magnitude of recent sea-level lowstands support the repeated existence of sizeable
islands across the western Indian Ocean, greatly reducing the isolation of Madagascar
from Asia.”
The general information about Madagascar has been summarized mainly by Paulian
(1951); Battistini and Richard-Vindard (1972); and Lourenço (Ed.) (1996).
Map 8.5 Madagascar, Comoro Islands, and the Mascarene
Arachnogeography
The fauna of Madagascar has puzzled the biologists since the beginning of the zoo-
geography. The endemism of many groups of vertebrate and invertebrate fauna has
been described many times (Dippenaar-Schoenman and González Reyes 2006;
Hewitt 1919; Lawrence 1955; Purcell 1903; Roewer 1941; Wharton 1981; Millot
1948a, b). No need to repeat it here, but the recent achievement of the knowledge
concerning the Arachnofauna of Madagascar is worth analyzing.
8.14 Madagascar 703
In the books on zoogeography or biogeography of the world, Madagascar is

treated differently. From phytogeographic point of view, with 9 endemic families
and over 300 endemic genera of plants and 85% of endemic species, the island is
a region (subkingdom) on equal footing with the Afrotropical or Indomalayan
regions (Takhtadjian 1970). The different treatment of the island by zoogeographers
is explained further, but now is prevailing the idea of independent region
(Krijanovskij 1980).
What concerns the insects, Krijanovskiy (2002) accepts a special Madagascar
Region, with two subregions: Madagascar Subregion and Mascarene Subregion. He
writes that “The fauna of the Madagascar Region is completely comparable in the
richness with the faunas of the other regions of Paleotropical Kingdom and is highly
peculiar. The level of the species endemism in the majority of the insect groups
under consideration is overpassing 90%, the level of generic endemism oscillates
between 35 and 90%. There are several endemic taxa of higher rank – tribes and
subfamilies. Not less are remarkable also the negative particularities of the fauna
under consideration. The distinctions of this fauna with the faunas of the Afrotropical
and the Indomalayan regions are not less than the distinction of these regions among
themselves. The history of the fauna of Madagascar developed at least since the
early Miocene (more probably since the Oligocene) independently from the faunas
of the other regions of the Paleotropical Kingdom” (we translated these important
lines, written in Russian).
On the Big Island are known Palpigradi, Scorpiones, Amblypygi,
Pseudoscorpiones, Opiliones, Araneae, Opilioacarida, and many other Acari.
Lacking are Ricinulei, Uropygi, Solifugae (most intriguing absence!), and
Holothyrida (another puzzling absence). Lacking are also the suborders
Palaeoamblypygi, Mesothelae, Eupnoi (another puzzle!), and Dyspnoi. The fauna
of Palpigradi (19 spp.) is the richest in the world, and the scorpions are almost 100%
endemic.
Palpigradi The papers of Rémy (1950, 1960a) are the base and were longtime the
only source of information about Palpigradi from Madagascar (13 Eukoenenia
Börner and 3 Koeneniodes Silvestri, all Eukoeneniidae). Some are endemic in the
“Grande Terre,” so is Triadokoenenia Condé (1991) (Prokoeneniidae, the only
endemic genus). Others have been found also on Réunion, Mauritius, and even as far
as Sri Lanka (Eukoenenia chartoni, Koeneniodes madecassus) (Condé 1991, 1996).
Scorpiones The immense progress in the study of scorpions of Madagascar could

be assessed by comparing the three “faunas” – of Fage (1929) and Millot (1948),
listing 10 species each, and of Lourenço (1996) with 19 species of 9 genera and 3
families (Buthidae, Scorpionidae, and Hemiscorpiidae – Ischnuridae).
Both Fage (1929) and Millot (1948) came to the conclusion that, because of their
large size, scorpions are easily collected and “we know all the populations of the
island as well as their relative density and distribution” (Fage 1929). Since this opti-
mistic (but premature) statement of the prominent French arachnologist, however,
the number of species known from Madagascar has more than doubled and is far
from being exhaustive (Lourenço 1995, 1996, 2001, 2003, 2012, etc.).
Microcharminae became a family and later has been downgraded again.
Vachon (1969/1979) added two more species. Since the beginning of the studies
of Lourenço in 1995 have been described many new species of smaller dimensions,
unknown to the former researchers. Also important analysis took place, due to
Lourenço (1996b, 1996e, 2000a). According to Lourenço (1996b), “The most
remarkable characteristic of the scorpion fauna of Madagascar is the impressive
level of endemicity, both in species and in genera. This supports the hypothesis of
the very early isolation of the island from other land masses.”
Fet, Gantenbein et al. (2003) concluded from molecular analysis that the endemic
genus Grosphus is cladistically close to the South American genera Centruroides
and Rhopalurus.
The present-day (2017) list of the scorpions of Madagascar contains 52 sp., 13
genera, and 3 families (Gervais 1844; Kraepelin 1895, 1901; Fage 1929, 1946; Vachon
1969, 1979; Lourenço 1995, 1996b, 1996e, 1997, 1999, 2000a, c, 2001, 2003c, 2012c;
Lourenço et al. 2004, 2006; Lourenço and Goodman 2002, 2004, 2009; Lourenço and
Ythier 2010; Pocock 1889, 1890, 1893, 1894, 2002, and others).
Fam. Buthidae Grosphus Simon, 1880

G. ankarafantsika Lourenço, 2003 T. guillaumeti Lourenço, 1995
G. ankarana Lourenço et T. ivohibe Lourenço et Goodman, 1999
Goodman, 2003
G. annulatus Fage, 1929 T. judsoni Lourenço, 1996
G. bistriatus Kraepelin, 1901 T. lucileae Lourenço, 1995
G. darainensis Lourenço, Goodman et T. manonae Lourenço, 2000
Ramilijaona, 2004
G. feti Lourenço, 1996 T. monodi Lourenço, 2000
G. flavopiceus Kraepelin, 1901 T. parriloi Lourenço, 1996
G. garciai Lourenço, 2001 T. petrae Lourenço, 1996
G. grandidieri Kraepelin, 1901 T. pococki Lourenço, 1995
G. hirtus Kraepelin, 1901 T. racotondravonyi Lourenço et
Goodman, 2003
G. intertidalis Lourenço, 1999 Troglotityobuthus Lourenço, 2000
G. limbatus Pocock, 1889 T. gracilis (Fage, 1946)
G. madagascariensis (Gervais, 1844) [Fam. Microcharmidae]
G. mahafaliensis Lourenço, Goodman et Microcharmus Lourenço, 1995
Ramilijaona, 2004
G. olgae Lourenço, 2004 M. cloudsleythompsoni Lourenço, 1995
G. simoni Lourenço, Goodman et M. hauseri Lourenço, 1996
Ramilijaona, 2004
Neogrosphus Lourenço, 1995 M. jussarae Lourenço, 1996
N. blanci Lourenço, 1996 M. sabineae Lourenço, 1996
N. griveaudi (Vachon, 1969) Neoprotobuthus Lourenço, 2000
Isometrus Hemprich et Ehrenberg, 1829 N. intermedius Lourenço, 2000
I. maculatus (DeGeer, 1778) – Alien Fam. Heteroscorpiidae
8.14 Madagascar 705
Pseudouroplectus Lourenço, 1995 Heteroscorpion Birula, 1903

P. betschi Lourenço, 1995 H. goodmani Lourenço, 1996
P. lalyae Lourenço et Ythier, 2010 H. kaii Lourenço et Goodman, 2009
P. pidgeoni Lourenço et Goodman, 1999 H. magnus Lourenço et
Goodman, 2002
Palaeogrosphus Lourenço, 2000 H. opistacanthoides (Kraepelin, 1895)
P. copalensis (Lourenço, 1996) H. raselimananai Lourenço et
Goodman, 2004
Tityobuthus Pocock, 1893 Fam. Hormuridae (Liochelidae)
T. baroni (Pocock, 1890) Opisthacanthus Peters, 1861
T. darainensis Lourenço et O. madagascariensis Kraepelin, 1894
Goodman, 2002
T. dastychi Lourenço, 1997 Iomachus Pocock, 1893
T. griswoldi Lourenço, 2000 I. punctulatus Pocock, 1897
Paleocheloctonus Lourenco, 1996
P. pauliani Lourenço, 1996
Endemism: Except the alien Isometrus maculatus, all 12 native genera (Grosphus,
Neogrosphus, Pseudouroplectus, Tiyobuthus, Palaeogrosphus, Microcharmus,
Neoprotobuthus, Pseudouroplectes, Heteroscorpion, Opisthacanthus,
Paleocheloctonus, Troglotityobuthus) and 100% of the species are endemic in
Madagascar, partly with the Comoros. After Goodman and Benstead (2005), the
scorpions known from Madagascar are 40 spp. The family Microcharmidae, consid-
ered endemic, was synonymized with Buthidae by Volschenk et al. (2008). Despite
the high endemism, several affinities can be detected. They have been hinted by the
older authors (Pocock, Vachon) but analyzed in details by Lourenço (1996b, 1996e,
1999, 2000a, 2000d) and Lourenço et al. (2006).
Pseudoscorpiones According to Millot (1948), in Madagascar are known 11 spp.
of Pseudoscorpions of 5 families. Vachon (1951, 1960) added also the presence of
Chthoniidae, Tridenchthoniidae (Dithidae), and Faella sp. (Faellidae).
Now (2017, partial information from M. Harvey) from Madagascar are known
14 spp. of Pseudoscorpions of 10 genera and the families Hyidae, Feaellidae,
Garypidae, Olpiidae, Pseudochiridiidae, Atemnidae, Chernetidae, and Withiidae.
Endemic genera of Pseudoscorpiones of Madagascar:
Fam. Garypidae
Paragarypus Vachon, 1937 – One sp.
Paracheiridium Vachon, 1938 – Two spp. (also Iles Glorieuses)
The endemism is relatively low, compared to other groups.
Opiliones According to the Fauna of Lawrence (1959) and the supplements, since
this time in Madagascar have been recorded Opiliones of 17 genera and 5 families:
Pettalidae, Triaenonychidae, Biantidae, Samoidae, and Zalmoxidae (Fage 1945;
Loman 1898; Mendes and Kury 2012; Pocock 1902, 1903; Roewer 1914, 1949;
Shear and Gruber 1996; Starêga 1992).
Remarkable is the lack of families like Assamiidae, widely distributed in Africa
and other tropical countries of the Eastern Hemisphere (Lawrence 1959). Even
more intriguing is the lack of Dyspnoi and, even more, of Eupnoi, so widespread in
continental Africa.
Cyphophthalmi From Madagascar have been described two (endemic and mono-
typic) genera:
Fam. Pettalidae.
Manangotria Shear et Gruber, 1996 – One sp.
Incertae sedis:
? Ankaratra Shear et Gruber, 1996 – One sp.
Eupnoi and Dyspnoi
No native “Palpatores” are known in Madagascar.

Laniatores From Madagascar are known Laniatores of 15 endemic genera and the
families Biantidae, Triaenonychidae, Samoidae, Podoctidae, and Zalmoxidae.
Endemic genera of Opiliones Laniatores in Madagascar are:
Acumontia Loman, 1898 (syn. Triaenomontia Roewer, 1914; syn. Tanalaius
Roewer, 1914; syn. Triacumontia Roewer, 1915; syn. Spinimontia Roewer, 1915;
syn. Bezavonia Roewer, 1949) (22 spp.); Ankaratrix Lawrence, 1959 (two spp.);
Antongila Roewer, 1931 (one sp.); Decarynella Fage, 1945 (one sp.);
Hovanuncia Lawrence, 1959 (syn. Flavonuncia Lawrence, 1959) (three spp.);
Ivohibea Lawrence, 1959 (one sp.); Millomontia Lawrence, 1959 (two spp.);
Millotonyx Lawrence, 1959 (one sp.); Paulianyx Lawrence, 1959 (two spp.)
Fam. Biantidae: Anaceros Lawrence, 1959 (four spp.), Fageibiantes Roewer, 1949
(two spp.)
Fam. Samoidae: Hovanoceros Lawrence, 1959, Malgaceros Lawrence, 1959
Fam. Zalmoxidae: Bunofagea Lawrence, 1959, Remyus Roewer, 1949
Schizomida Lawrence (1969). Three (endemic?) species of Schizomida have been
recorded from Madagascar (Lawrence, 1931), referred as genus Bamazomus by
Harvey (2001).
8.14 Madagascar 707
Amblypygi
Fam. Charinidae
Charinus madagascariensis Fage, 1954 (endemic species), is the only representa-
tive of Amblypygi from the Big Island.
Araneae On Madagascar are known about 400 spp. of spiders of 46 families
(Griswold 1997; Blandin 1979; Emerit 1974; Griswold and Ledford 2001; Haddad
et al. 2009; Huber 2000, 2003; Jocqué 1994; Lawrence 1938; Lehtinen 1967; Millot
1946, 1947; Platnick 1984, 1995; Pocock 1895; Simon 1889, 1902, 1903), but
according to Alderweireldt and Jocqué (1994), this figure is only 10% of the real
Malagasy spider fauna. According to Goodman and Benstead (2005), in Madagascar
are known 459 spp. of spiders, 390 of which are endemic (85%). The opinion of
Emerit (1978) is the same – that the “Prétendues lacunes faunistiques de la Région
Malgache” are only due to the insufficient collection. We may say that Madagascar
is very rich in endemics (seen well from the vertebrates), but one should be careful
to judge the endemism in the invertebrates, taking into account the intensity of col-
lecting in a given area.
Emerit (1997) recorded for Madagascar the Asian subfamily Chorizopinae
(Araneidae); in 1980 he recorded the family Mimetidae.
Some endemic genera of spiders for Madagascar:
Fam. Barychelidae (found on Madagascar, but not in Southern Africa; the genus
Tigidia Simon is known from Madagascar, seven spp.; Mauritius, one; and India,
four)
Zophoryctes Simon, 1902 (one species)
Fam. Zoropsidae (the other genera are known from America and New Zealand)
Calamistrula Dahl, 1901 (one sp.)
Fam. Gallieniellidae
Gallieniella Millot, 1947 (three spp. in Madagascar, one sp. in Comoro Islands)
Legendrena Platnick, 1984 (seven spp.)
Fam. Ctenidae
Mahafelytenus Silva-Dávila, 2007 (seven spp.)
Fam. Hersiliidae
Prima Foord, 2008 (one sp.)
Fam. Theraphosidae
Encyocrates Simon, 1892 (one sp.)
Fam. Ubudidae
Ubuda Simon, 1880 (three spp.)
Zorodictyna Strand, 1907 (two spp.)
Fam. Cyatholipidae
Alaranea Griswold, 1997 (four spp.)
Vazaha Griswold, 1997 (two spp.)
Fam. Pisauridae (incl. Halidae Jocqué, 1994, and the genera Hala Jocqué, 1994,
two spp., and Tolma Jocqué, 1994, one sp.)
Caripetella Strand, 1928 – Madagascar, Comoro Islands (one sp.)
Paracladycnis Blandin, 1979 (one sp.)
Ransonia Blandin, 1979 (one sp.)
Tallonia Simon, 1889 (one sp.)
Thalassiopsis Roewer, 1955 (one sp.)
Fam. Synaphridae
Africepheia Miller, 2007 (one sp.)
Fam. Araneidae
Augusta O.P.-Cambridge, 1877 (one sp.); Exechocentrus Simon, 1889 (two
spp.); Parmatergus Emerit, 1994 (three spp.)
Fam. Phyxelididae
Ambohima Griswold, 1990 (ten spp.)
Manampoka Griswold et al., 2012 (one sp.)
Rahavavy Griswold et al., 2012 (three spp.)
Fam. Theridiidae
Asygyna Agnarsson, 2006 (two spp.)
Fam. Idiopidae
Genysinae
Genysa Simon, 1889 (two spp.), Hiboka Fage, 1922 (one sp.)
Fam. Migidae
Paramiginae
Micromesomma Pocock, 1895 (1 sp.); Madacantha Emerit, 1970 (1 sp.);
Paramigas Pocock, 1895 (11 spp.); Thyropoeus Pocock, 1895 (2 sp.)
Fam. Filistatidae
Andoharano Lehtinen, 1967
8.14 Madagascar 709
Fam. Selenopidae
Garcorops Corronca, 2003 – Madagascar, Comoro Islands (three spp.)
Hovops Benoit, 1968 (nine spp.)
Fam. Liocranidae
Donuea Strand, 1932 (two spp.)
Fam. Lycosidae
Katableps Jocqué, Russell-Smith et Alderweireldt, 2011 (three spp.)
Fam. Linyphiidae
Thapsagus Simon, 1894 (one sp.), Thyreobaeus Simon, 1889 (one sp.)
Tmeticides Strand, 1907 (one sp.)
Fam. Oxyopidae
Hostus Simon, 1898 (one sp.)
Fam. Pholcidae
Paramicromerys Millot, 1946 (14 spp.), Zatavua Huber, 2003a (17 spp.)
Fam. Sparassidae
Chrosioderma Simon, 1897 (nine spp.); Megaloremmius Simon, 1903 (one
sp.); Staianus Simon, 1889 (one sp.)
Fam. Zodariidae
Aschema Jocque, 1991 (two spp.), Omucukia Kocak et Kemal, 2008 (two spp.)
Omucutia Kocak et Kemal, 2008 (two spp.)
Fam. Thomisidae
Apyretina Strand, 1929 (five spp.)
Cyriogonus Simon, 1886 (six spp.)
Diplotychus Simon, 1903 (one sp.)
Emplesiogonus Simon, 1903 (two spp.)
Herbessus Simon, 1903 (one sp.)
Iphoctesis Simon, 1903 (one sp.)
Lampertia Strand, 1907 (one sp.)
Plastonomus Simon, 1903 (one sp.)
Pseudoporrhopis Simon, 1886 (one sp.)
Soelteria Dahl, 1907 (one sp.)
Fam. Salticidae
Echinussa Simon, 1901 (three spp.)
Goleta Peckham et Peckham, 1894
Macopaeus Simon, 1900

Myrmanu Prósz., 2016
Myrmele Prósz., 2016
Phaulostyleus Simon, 1902 (four spp.)
Poessa Simon, 1902
Tomobella Szüts et Scharff, 2009 (two spp.)
Tomocyrba Simon, 1900 (six spp.)
Opilioacarida Vásquez and Klompen (2010) contributed to the study of
Opilioacarids of Madagascar with another four species to the two already described
by Van der Hammen (1977), all in his genus Salfacarus. Other members of this
genus live on the African continent (South Africa and Tanzania). The six species are
all endemic.
Acariformes
Oribatida In the fauna of Oribatida of Madagascar, 80% of the species are

endemic. The genera are distributed evenly between the Afrotropical and the
“Oriental” Regions (Schatz, 2004). Other sources: Mahunca (2002) and nine other
papers; Niedbala 2004.
Fauna of ptyctimous mites from Madagascar and neighbouring islands Seychelles,

Mauritius, Comoros and Reunion (MAG) comprises 111 species for which morphological
diagnoses have been made. …The similarity of the fauna of MAG (111 species) to the fauna
of Afrotropical Region (200 species) is much higher (15 % common species) than to the
fauna of Oriental Region (262 species) (4 % common species). The fauna of MAG shows
no similarity to that of India. In the years 2010–2015 new soil samples (171) were collected
on Madagascar and they contained 78 species. The high number of endemic species among
the dominants and high-frequency species confirm the unique character of the fauna of
Madagascar. (Niedbala 2017)
Parasitiformes
Ixodida The base of our knowledge is the paper of Hoogstraal (1953), in which
find place 11 endemic or presumably endemic species of Ixodida (9 Haemaphysalis,
1 Ixodes, and 1 Argasidae indet). Here we quat the conclusions of Hoogstraal con-
cerning the origin of the Malagasy tick fauna:
Millot (1948, 1952) stated that Malagasy ticks are certainly all of African origin. This is
entirely or almost entirely true for introduced species. The endemic species show some
affinities with African fauna, but many more with that of southern Asia. …As in Madagascar,
haemaphysalid species form the bulk of the southern Asiatic ixodid tick fauna….The very
important African genus Rhipicephalus, consisting of over forty species, is entirely unrep-
resented by endemic species in the Malagasy Faunal Region. Amblyomma ticks, repre-
8.15 Mascarene Islands 711
sented in Africa by many more species than in Asia, are lacking in endemic species in the
Malagasy Region except in the Seychelles.
According to Klompen (2003) and Goodman and Benstead (2005), there are 27 spp.
of Ixodida, among them 25 are endemic.
8.15 Mascarene Islands
8.15.1 G
and Paleogeography
The Mascarene Islands (or Mascarenhas Archipelago) is a group of islands in the
Indian Ocean east of Madagascar comprising Mauritius, Réunion, Rodrigues,
Cargados Carajos shoals, etc. The islands share a common geologic origin in the
volcanism of the Réunion hot spot beneath the Mascarene Plateau and form a dis-
tinct ecoregion with a unique flora and fauna. The youngest islands to form were
Mauritius (7–10 ma), the oldest of the existing islands, created along with the
undersea Rodrigues ridge. The islands of Rodrigues and Réunion were created in
the last 2 million years. The Mascarene Plateau is an undersea plateau that extends
approximately 2000 km, from the Seychelles to Réunion. The plateau covers an
area of over 115,000 km² of shallow water, with depths ranging from 8 to 150 m.
The islands have never been connected to the mainland, so the flora and fauna of the
Mascarenes arrived from over the sea. Prehistoric islands of the Mascarene Plateau,
now disappeared under the sea, may have served as “stepping stones” which allowed
species to island-hop from the Seychelles or Madagascar.
Réunion – A French island, 200 km southwest of Mauritius, the nearest island. Area
2512 km2, 63 km long, 45 km wide. Highest point is Piton des Neiges (3070 m),
also the shield volcano Piton de la Fournaise, more than 2631 m.
Mauritius – An island 900 km. east of Madagascar, 570 km. from Rodrigues.
Area – 2040 km2, highest point 828 m
Rodrigues – An island, part of the Republic of Mauritius, 109 km2, highest point
355 m
Arachnogeography
These small strange islands with fantastic (already gone) vertebrate fauna have sev-
eral orders of Arachnida, but no endemisms higher than genus. They are Palpigradi,
Scorpiones (one endemic species), Pseudoscorpiones (six endemic species, two
endemic genera), Opiliones (seven end. species of Laniatores, one endemic
genus), Araneae (one endemic genus), and different Acari, of which most interest-
ing is the presence of Holothyrida (three sp., incl. one endemic genus on Mauritius).
Nothing similar to the fauna of Africa or Madagascar. From the Mascarene are
missing the orders Ricinulei, Thelyphonida, Solifugae, and Opilioacarida and the
suborders Palaeoamblypygi, Mesothelae, Cyphophthalmi, and Dyspnoi
(Maureemootoo, 2000).
Palpigradi The information about the Palpigradi on these islands comes from
Remy (1952b – Réunion, 1958 - Mauritius), in total three spp. (one of Eukoenenia,
two of Koeneniodes), all of them belonging to Eukoeneniidae. On Réunion have
been collected Eukoenenia hanseni (Silvestri), Koeneniodes frondiger Remy, and K.
madecassus Remy, all found also on Madagascar and Mauritius and in Indonesia.
Scorpiones
Endemic sp. on Mauritius is Lychas serratus (Buthidae). Chiromachus ochropus

(Hormuridae) is shared with the Seychelles and Zanzibar (Probst 1973).
Pseudoscorpiones On the islands (Réunion, Mauritius) have been recorded eight
species of Pseudoscorpions (Chthoniidae, Cheliferidae, Chernetidae, and Withiidae),
six of them are endemic in the Archipelago, and two endemic genera (Beier 1935,
1940; Mahnert 1975).
Thaumatowithius Beier, 1940 – Réunion, Mauritius (two spp.)
Pilochelifer Beier, 1935 – Réunion, Mauritius (one sp.)
Opiliones
From Mascarene Islands are known seven genera of Opiliones (Roewer 1927, 1936,
1949, 1954; Simon 1879; Staręga 1989b).
Cyphophthalmi – None
Eupnoi
Fam. Sclerosomatidae – Gagrella Stoliczka
Dyspnoi – None
Laniatores
Fam. Assamiidae – Maccabeesa Roewer (one sp.)
Fam. Biantidae – Hinzuanius Karsch (one sp.)
Fam. Podoctidae – Erecanana Strand (one sp.), Sitalcicus Roewer (one sp.),
Ibalonius Karsch (one sp.)
Fam. Zalmoxidae – Zalmoxis Sørensen (Zalmoxomma Roewer) (one sp.)
Endemic Opiliones on the islands are:
Fam. Biantidae
Hinzuanius mauriticus Roewer, 1927 – Mauritius
Fam. Assamiidae
Maccabeesa lawrencei Roewer, 1936 – Mauritius
8.16 Seychelles (incl. Aldabra), Comores 713
Fam. Podoctidae
Erecanana insulana Roewer, 1949 – Réunion
Sitalcicus novemtuberculatus Simon, 1879 – Réunion
Ibalonius breoni Simon, 1879 – Réunion
Fam. Zalmoxidae
Zalmoxis [Zalmoxomma] occidentalis (Roewer, 1949) – Mauritius
Gagrella reunionis (Roewer, 1954) – Réunion
Amblypygi From Mauritius is known Phrynichus scaber (Gervais, 1844)
(Phrynichidae), known also from the Seychelles.
Holothyrida From Mauritius have been described two endemic species:

Holothyrus coccinella Gervais (1842), the first holothyrid ever described, and H.
legendrei Van der Hammen (1983). According to the modern system of Holothyrida
(Lehtinen 1995, Beron 2014), the genus Holothyrus remains endemic in Mauritius.
The order Holothyrida is known also from Rodrigues.
8.16 Seychelles (incl. Aldabra), Comores
8.16.1 G
and Paleogeography
The Republic of Seychelles consists of four archipelagoes (Seychelles, Amirantes,

Farquhar, and Aldabra), with some 155 islands of total surface area of 451 km2. The
most interesting for arachnology are the 42 granite islands, remnants of a much big-
ger landmass. They lay to the northeast of Madagascar and c. 1600 km. east of
Kenya (the African coast). Some scientists claim that 18 000 years ago, the sea level
was 120 m lower and today’s Seychelles was part of a landmass covering c.
32 000 km2. The granite islands are very old (the oldest islands of any ocean), and
careful research is being done on their flora and fauna (Benoit 1978; Badyukov
et al. 1989; Gerlach et al. 2013; Scott 1958).
The granite islands are situated in a radius of 56 km from the main island Mae
(142 km2, highest point 914 m.). They are part of the so-called Inner islands, on
which live 82 500 inhabitants. Some other bigger
The Outer Islands (211 km2, 2% of the population) are coral (Amirante, Aldabra,
Farcouar, and others) but are also interesting.
These granitic islands are part of the Mascarene Plateau, detached from the
Indian Plate 65 million years ago.
The system of national parks covers 42% of the land and 260 km2 of the sea around
it. The flora of the Seychelles includes 1170 species of higher plants (75 endemic spe-
cies); among the animals 15 bird species, 3 mammal species, 30 species of amphibi-
ans and reptiles, and hundreds of species of invertebrates are not found elsewhere.
Starting with the words of Jeannel (1942), “Dans tous les groupes d’Insects, les
monographes s’accordent pour affirmer que la faune des Seychelles a un caractère
nettement indo-malais, les éléments malgaches s’y trouvant en majorité, tout élé-
ment africain faisant défaut,” we should check the rightness of this conclusion on
what concerns the arachnids.
The very important contribution to the fauna of the Seychelles (Séchelles) was
the series of papers as a result of the mission of P.L.G. Benoit and J.J. Van Mol in
1972. The Arachnida were treated by Benoit (1978, series of papers on spiders),
Mahnert (1978, pseudoscorpions), and Benoit (1979, scorpions). The results were
published in Belgium under the editorship of P. Benoit, who wrote the introduction
of this important series. Many other specialists have explored the strange islands
and their non-insect inhabitants (see Gerlach, J. & J. Marusik, Eds, 2010).
Arachnogeography
The amazing fauna of the granitic islands, combining African, Indian, and endemic
elements, received attention by very qualified arachnologists, and now we know
that:
The Seychelles are home of the orders Palpigradi, Scorpiones, Pseudoscorpiones,
Opiliones (four endemic genera), Amblypygi, Schizomida, Araneae, and some
groups of Acari (Gerlach 2003; Gerlach and Marusik (Eds) 2010; Hirst 1913;
Harvey 2001).
Absent are the orders Ricinulei, Solifugae, and Opilioacarida and the suborders
Palaeoamblypygi, Mesothelae, Cyphophthalmi, Dyspnoi, and Eupnoi. To be noticed
is the presence of Holothyrida (three endemic genera of the family Holothyridae),
the big poisonous creatures, absent in Madagascar and in the nearby Africa.
Schizomida are also very well represented by seven genera (two endemic) and seven
species (five endemic). Among the 228 spp. of spiders of 40 families (Saaristo
2003), there are at least 12 endemic genera. Despite this interesting fauna, the level
of endemism is only to genus, and in some orders, there are only endemic species.
Palpigradi Koeneniodes madecassus is recorded from La Digue (known also from
Madagascar and Indonesia) (Harvey 2001).
Scorpiones From the Seychelles have been reported the species Isometrus macu-
latus, Lychas braueri (Scorpionidae), and Chiromachus [Ischnurus] ochropus
(Hormuridae, known from Zanzibar and Mauritius). Lychas braueri (Kraepelin,
1896) is endemic in the Seychelles (Benoit 1979; Harvey 2001; Kraepelin 1896;
Lourenço and Goodman 2009; Prendini 2010).
Pseudoscorpiones The 9 species of Pseudoscorpiones, mentioned by Beier (1940)

from the Seychelles, have been added up in 1978 to 4 more, so the figures, quoted
by Mahnert (1978) and Harvey (2001, 2009, 2011), are 23 species of 18 genera and
12 families: Chthoniidae, Tridenchthoniidae, Syarinidae, Garypidae, Garypinidae,

Geogarypidae, Pseudochiridiidae, Olpiidae, Atemnidae, Feaellidae, Chernetidae,
and Withiidae (Beier 1974; Chamberlin 1930; Hirst 1911).
No endemic genera, seven endemic species:
Compsaditha seychellensis Beier, 1974 – Mahé
Fam. Feaellidae
Feaella affinis Hirst, 1911 – Praslin, Silhouette
Fam. Geogarypidae
Afrogarypus seychellensis (Beier, 1940) – Praslin, Mahé
Geogarypus ocellatus Mahnert, 1978 – Praslin
Fam. Olpiidae
Beierolpium benoiti Mahnert, 1978 – Curieuse, Silhouette (one endemic sp.)
Fam. Syarinidae
Ideoblothrus seychellesensis (Chamberlin, 1930) – Silhouette
Fam. Atemnidae
Oratemnus brevidigitatus Beier, 1940 – Mahé, Silhouette, Aride
The fauna has affinities with both the Afrotropical and Indomalayan faunas.
From the other ten spp. in the family Feaellidae (the only genus Feaella), eight live
on the African continent (incl. four in Southern Africa), one in India, and one in
Australia. Compsaditha has 12 spp. (5 from Africa, 5 from Asia and Oceania, 1
from the Seychelles, and 1 from South America). From 14 spp. of Afrogarypus, 1 is
from the Seychelles and 13 from the African continent (A. impressus is known from
Aldabra, Mozambique, and South Africa). Beierolpium has also 14 known species,
from Africa (8), Australia, Oceania, Malaysia, and Venezuela. Oratemnus is typi-
cally Oriental – 14 spp. from Asia, 4 from Australia, 1 from the Seychelles, and 1
from Oceania and the Caribbean. Geogarypus and Ideoblothrus are widespread.
Opiliones
Cyphophthalmi, Eupnoi, and Dyspnoi – Not known on the Seychelles

Laniatores The paper of Rambla (1984) summarizes the knowledge on the
Opiliones of the Seychelles – 17 spp., all of Laniatores of 9 genera and the families
Phalangodidae, Biantidae, Podoctidae, Samoidae, and Zalmoxidae (Loman 1902;
Hirst 1911; Roewer 1912, 1949).
Fam. Podoctidae
Centrobunus braueri Loman, 1902; Holozoster ovalis Loman, 1902; Ibalonius
inscriptus Loman, 1902; I. lornani Hirst, 1911; l. bimaculatus Loman, 1902;
I. karschi Loman, 1902; l. flavopictus Hirst, 1911; Sitalcicus gardineri Hirst,
1911; S. incertus Rambla, 1984
Fam. Samoidae
Benoitinus elegans Rambla, 1984; Mitraceras crassipalpum Loman, 1902; M.
pulchra Rambla, 1984; Samoa sechellana Rambla, 1984
Fam. Biantidae
Biantes parvulus (Hirst, 1911), B. albimanus (Loman, 1902), B. minimus
(Rambla, 1984)
Fam. Zalmoxidae
Metazalmoxis ferruginea Roewer, 1912
Endemic genera for the Seychelles are:
Fam. Podoctidae
Centrobunus Loman, 1902 (one sp.), Holozoster Loman, 1902 (one sp.)
Peromona Roewer, 1949 (one sp.)
Fam. Samoidae
Benoitinus Rambla, 1984 (one sp.), Mitraceras Loman, 1902 (one sp.)
Fam. Zalmoxidae
Metazalmoxis Roewer, 1912 (one sp.)
Amblypygi. From the Seychelles are known Phrynichus scaber (Gervais, 1844)
(Phrynichidae), known also from Mauritius, and Charinus seychellarum
Kraepelin, 1898 (Benoit 1979, Harvey 2001, Kraepelin 1898) (Charinidae)
(endemic).
Schizomida. On the Seychelles are found seven species of Schizomida. Harvey
(2001) describes four new monotypic and partly endemic genera:
Fam. Hubbardiidae
Mahezomus apicoporus Harvey, 2001 – Endemic genus and species
Secozomus latipes (Hansen, 1905) – Endemic species
Zomus bagnalii (Jackson, 1908) – Known from Southeast Asia
Anepsiozomus sobrinus Harvey, 2001 – Endemic genus and species
Ovozomus lunatus (Gravely, 1911) – Known from the Seychelles, Cook Islands,
Mayotte and other islands, Ceylon, India
Bamazomus aviculus Harvey, 2001 – Endemic species
Apozomus gerlahi Harvey, 2001 – Endemic species
Three species of Schizomida, described from Madagascar, are transferred to

Bamazomus Harvey: B. madagassus (Lawrence), B. milloti (Lawrence), and B.
vadoni (Lawrence). As a result, on the Seychelles are known seven spp. of
Schizomida of seven genera, including four endemic genera and five end. species.
Araneae Many specialists (Simon, Benoit, Brignoli, and others) contributed to the
study of the spiders of the fascinating islands. Gerlach (2003) quoted the figure of
198 spp. on the islands. After a series of papers (1978, 1998, 2001, 2002) on the
spiders of the Seychelles, Saaristo (2003) published online a list of 228 spp. from 40
families, living on the islands (198 spp. from the Granitic Islands and 30 more spe-
cies on the reefs and atolls) (Benoit 1978 – several papers, 1979; Grasshoff 1980;
Wanless 1984).
Endemic genera of spiders:
Fam. Theraphosidae
Nesiergus Simon, 1903 (three spp.)
Fam. Tetrablemmidae
Mariblemma Lehtinen, 1981
Euso Saaristo, 2001, Roche Saaristo, 1998
Fam. Oonopidae – Gamasomorphinae
Diblemma O. P.- Cambridge, 1908 (one sp.); Ischnothyrella Saaristo, 2001 (one
sp.); Lionneta Benoit, 1979 (eight spp.); Lisna Saaristo, 2001 (one sp.);
Matyotia Saaristo, 2001 (one sp.); Patri Saaristo, 2001 (one sp.); Prida
Saaristo, 2001 (one sp.); Aridella Saaristo, 2002 (one sp.); Cousinea Saaristo,
2001 (one sp.); Farqua Saaristo, 2001 – Farquhar Islands – (one sp.)
Fam. Gnaphosidae
Microdrassus Dalmas, 1919
Fam. Tetragnathidae
Pholcipes Schmidt et Krause, 1993 – Comoro Islands (one sp.)
Fam. Theridiidae
Argyrodella Saaristo, 2006
Bardala Saaristo, 2006 – Aldabra (one sp.)
Nanume Saaristo, 2006 – Aldabra (one sp.)
Sesato Saaristo, 2006 (one sp.)
Seycellesa Kocak et Kemal, 2008 (one sp.)
Spinembolia Saaristo, 2006 (one sp.)
Stoda Saaristo, 2006 (one sp.)
Fam. Ctenidae
Apolania Simon, 1898 (one sp.)
Fam. Scytodidae
Soeuria Saaristo, 1997
Fam. Sparassidae
Pleurotus Simon, 1898 (one sp.)
Rhacocnemis Simon, 1897 (one sp.)
Stipax Simon, 1898 (one sp.)
Thomasettia Hirst, 1911 (one sp.)
Fam. Salticidae
Baviola Simon, 1896
Holothyrida There are three endemic genera (Sternothyrus Lehtinen, 1995;
Michaelothyrus Gerlah, Lehtinen et Madl, 2010; and Dicrognatus Gerlah, Lehtinen
et Madl, 2010) in the Seychelles, one genus (Hammenius Lehtinen, 1981) is shared
with New Guinea, and one more species is incertae sedis. The presence of these
peculiar mites on the Seychelles and the Mascarene Islands is as curious as their
absence in Africa and Madagascar (Beron 2014; Gerlah 1995; Gerlah et al. 2010;
Hammen 1983; Lehtinen 1981, 1995; Thon 1906).
Discussion concerning the affinities and differences in the Arachnid faunas
of Madagascar and the adjacent islands:
We may start with the words of Millot (1952):
“Parmi les Invertébrés, les Arachnides fournissent des documents particulierement signifi-
catifs. Les espèces de la Réunion aussi bien que celles de Maurice ou de Mahé sont tout à
fait différentes de celles de Madagascar: Araignées, Opilions, Scorpions, Pseudoscorpions
sont presque aussi dissemblables que s’il s’agissait de régions fort separées – et alors que
les formes malgaches sont d’affinités nettement africaines, les espèces des Séchelles et des
Mascaraignes sont de parenté non moins nettement indo-malaise”.
Palpigradi Four genera are known in the Big Island, including the endemic genus
and species Triadokoenenia Condé, 1991 (Prokoeneniidae). The other genera
(Eukoenenia, Prokoenenia, and Koeneniodes) are represented also on Mauritius,
Réunion, and even as far as Sri Lanka, with the same species. On the Seychelles (La
Digue) has been recorded Koeneniodes madecassus.
Solifugae – Absent on all these islands, but numerous in East and South Africa
Scorpiones Several attempts to analyze the origins and affinities of the scorpions in
Madagascar and the surrounding islands (Fage 1929; Millot 1948; Legendre 1972;
Vachon 1979b) were crowned by the contribution and analysis of Lourenço (1996).
After this paper, many more new species have been published by Lourenço and his
coauthors, but the main conclusions remain valid. To quote Lourenço (1996b):
“…the present representatives of the Madagascar scorpion fauna are derived from protoele-
ments of both the families Buthidae and Ischnuridae which were already present in
Gondwanaland, previous to fragmentation and continental drift” The endemicity of scorpi-
ons in Madagascar is the highest in the world: 100% of the species known and in the native
genera. All 12 native genera are endemic, Grosphus also on the Comores. There is also an
endemic (contested) family (Microcharmidae). This high endemicity (and not only of scor-
pions) “supports the hypothesis of the very early isolation of the island of other land masses
(Lourenço 1996).
Meanwhile, in their subdivision of the world, based on scorpions, Nenilin and

Fet (1992) include Madagascar, the Mascarene, and the Seychelles in the Afrotropical
Region as Madagascar Subregion, divided into Madagascar Rayon and Mascarene –
Seychelles Rayon.
If we compare the scorpion fauna of Madagascar with Southern Africa, we may
find similarities only on family level.
Pseudoscorpiones From the total area under study (Madagascar, Seychelles,
Aldabra, Comores, Mascarene Islands), there are relatively few species of pseudo-
scorpions recorded (24 genera and 14 families). No endemic families, only two
endemic genera (Mauritius, Réunion). If we compare the generic lists of the vari-
ous island groups, we get the following distribution:
Fam. Chthoniidae
Tyrannochthonius Chamberlin, 1929 – Seychelles
Chthonius C.L. Koch, 1843 – Seychelles
Compsaditha Chamberlin, 1929 – Seychelles (endemic sp.)
Fam. Syarinidae
Ideoblothrus Balzan, 1872 – Seychelles (one endemic sp.)
Fam. Hyidae
Indohya Beier, 1974 – Madagascar, India
Fam. Garypidae
Paragarypus Vachon, 1937 – Madagascar (one sp., endemic)
Fam. Geogarypidae
Afrogarypus Beier, 1931 – Aldabra, Seychelles (endemic sp.)
Geogarypus Chamberlin, 1930 – Seychelles (endemic sp.)
Fam. Garypinidae
Aldabrinus Chamberlin, 1930 – Aldabra (one endemic sp.)
Fam. Feaellidae – Madagascar
Feaella Ellingsen, 1906 – Seychelles (one endemic sp.)
Fam. Olpiidae
Beierolpium Heurtault, 1976 – Seychelles
Xenolpium Chamberlin, 1930 – Madagascar, Seychelles, Aldabra
Paracheiridium Vachon, 1938 – Madagascar (two spp., endemic, but also Iles
Glorieuses)
Pseudochiridium With, 1906 – Seychelles
Fam. Atemnidae
Anatemnus Beier, 1932 – Madagascar, Seychelles
Oratemnus Beier, 1932 – Seychelles
Paratemnoides Harvey, 1990 – Madagascar
Fam. Chernetidae
Haplochernes Beier, 1932 – Madagascar (one endemic sp.)
Lamprochernes Tomosvary, 1882 – Seychelles
Metagoniochernes Vachon, 1939 – Madagascar (endemic sp.)
Fam. Withiidae
Nesowithius Beier, 1940 – Seychelles
Thaumatowithius Beier, 1940 – Réunion, Mauritius (two spp.)
Withius Kew, 1911 – Madagascar, Seychelles
Fam. Cheliferidae
Pilochelifer Beier, 1935 – Réunion, Mauritius (one sp.)
Heurtault (1986) writes that on Madagascar live members of 12 endemic genera
of Pseudoscorpiones but did not provide a list of these genera.
According to the information available, there is a striking difference between the
huge endemicity in Madagascar by the scorpions (and many other groups) and the
very low level by the pseudoscorpions.
Opiliones According to Lawrence (1959), the most striking feature in the fauna of
Opiliones is its total absence in Madagascar and the quasi total on the adjacent
islands of native “Palpatores.” If we complete the table in the monograph of Lawrence
(1959) with the data of Rambla (1984) and others, it will take shape as follows:
Area Cyphophthalmi Triaenonychidae Phalangodidae Assamiidae

rMadagascar + + − −
Seychelles − − + −
Mauritius − − − +
Réunion − − − −
Area Biantidae Podoctidae Samoidae Zalmoxidae
Madagascar + − + +
Seychelles − + + +
Mauritius + + − +
Réunion − + − −
Genera in common:
Madagascar – Seychelles: none
Seychelles – Mauritius: none
Seychelles – Réunion: Ibalonius, Sitalcicus (Podoctidae)
Madagascar – the Mascarene: none
Réunion – Mauritius: none
Schizomida
Madagascar: Three endemic Bamazomus species

Seychelles: Seven spp. known, five of them endemic, three endemic genera
(Mahezomus, Anepsiozomus, Secozomus). Zomus bagnalii (Jackson) is known
from Malaysia (incl. Sarawak), Singapore, and England (Kew Gardens).
Mauritius: Zomus bagnallii (Jackson), Schizomus vinsoni Lawrence
Rodrigues: Schizomus tenuipes Lawrence, Schizomus virescens Lawrence
Réunion: Ovozomus lunatus (Gravely)
Comoro Islands: Ovozomus lunatus (Gravely) (Mayotte)
Amblypygi
Madagascar: Charinus madagascariensis Fage, 1954 (endemic species)

Mauritius: Phrynichus scaber (Gervais, 1844)
Seychelles: Charinus seychellarum (Kraepelin, 1898), Phrynichus scaber (Gervais,
1844)
Araneae The only family considered endemic in Madagascar (Halidae) has been
merged with Pisauridae in 2006.
Opilioacarida Known only from Madagascar (six endemic species of genus

Salfacarus Hammen, living also in Southern Africa)
Holothyrida Four species on the Seychelles (incl. endemic genus Sternothyrus

Lehtinen), two endemic species of Holothyrus Gervais, endemic genus for
Mauritius. The order is not known from Madagascar and Africa but is found in Sri
Lanka, India, New Guinea, etc. (Table 8.5)
Table 8.5 Distribution of the arachnids in the areas concerned

Afrotropical Mascarene
Group (Southern Africa) Madagascar Seychelles Islands
Order Palpigradi Present Present Present Present
Fam. Eukoeneniidae Present Present Present Present
Order Ricinulei Absent in South Absent Absent Absent
Africa
Fam. Ricinuididae Absent in South Absent Absent Absent
Africa
Order Solifugae Present Absent Absent Absent
Fam. Galeodidae Present Absent Absent Absent
Fam. Daesiidae Present Absent Absent Absent
Fam. Solpugidae Present Absent Absent Absent
Fam. Ceromidae Present Absent Absent Absent
Fam. Melanoblosiidae Present Absent Absent Absent
Fam. Gylippidae Present Absent Absent Absent
Fam. Hexisopodidae Present Absent Absent Absent
Order Scorpiones Present Present Present Present
Fam. Bothriuridae Present Absent Absent
Fam. Buthidae Present Present Present Present
Fam. [Microcharmidae] Absent Present Absent Absent
Fam. Heteroscorpionidae Absent Present Absent Absent
Fam. Hormuridae Present Present Present Present
Fam. Scorpionidae Present Absent Absent Present
Fam. Lisposomidae Present Absent Absent Absent
Order Schizomida Present Present Present Present
Fam. Hubbardiidae Present Present Present Present
Order Uropygi Absent in Southern Absent Absent Absent
Africa
Fam. Hypoctonidae Absent in Southern Absent Absent Absent
Africa
Order Amblypygi Present Present Present Present
Suborder Neoamblypygi Present Present Present Present
Fam. Charinidae Present Present Present Absent
Fam. Phrynichidae Present Present Present Present
Suborder Palaeoamblypygi Absent in Southern Absent Absent Absent
Africa
Order Opiliones Present Present Present Present
Suborder Cyphophthalmi Present Present Present Absent
Fam. Ogoveidae Absent in Southern Absent Absent Absent
Africa
Fam. Neogoveidae Absent in Southern Absent Absent Absent
Africa
Fam. Pettalidae Present Present Absent Absent
Suborder Eupnoi Present Present Present Present
(continued)
Fam. Caddidae Present Absent Absent Absent
Fam. Phalangiidae Present Absent Present Absent
Fam. Sclerosomatidae Present Absent Present Present
Fam. Neopilionidae Present Absent Absent Absent
Suborder Dyspnoi Absent Absent Absent Absent
Suborder Laniatores Present Present Present Present
Fam. Triaenonychidae Present Present Absent Absent
Fam. Assamiidae Present Absent Absent Present
Fam. Biantidae Present Present Present Present
Fam. Podoctidae Present Present Present Present
Fam. Pyramidopidae Present Absent Absent Absent
Fam. Samoidae Present Absent Present Absent
Fam. Zalmoxidae Absent Absent Present Present
Order Pseudoscorpiones Present Present Present Present
Suborder Epiocheirata Present Present Present Present
Fam. Chthoniidae Present Present Present Present
Fam. Tridenchthoniidae Present Absent Present Absent
Fam. Present Absent Absent Absent
Pseudotyrannochthoniidae
Fam. Lechytiidae Present Absent Absent Absent
Fam. Feaellidae Present Present Absent Absent
Suborder Iocheirata Present Present Present Present
Fam. Ideoroncidae Present Absent Absent Absent
Fam. Hyidae Absent Present Absent Absent
Fam. Neobisiidae Present Absent Absent Absent
Fam. Syarinidae Present Absent Present Absent
Fam. Garypidae Present Absent Present Absent
Fam. Geogarypidae Present Absent Present Absent
Fam. Cheiridiidae Present Absent Present Absent
Fam. Pseudochiridiidae Present Present Absent Absent
Fam. Olpiidae Present Present Present Absent
Fam. Garypinidae Present Absent Absent Absent
Fam. Menthidae Absent in Southern Absent Absent Absent
Africa
Fam. Sternophoridae Present Absent Absent Absent
Fam. Withiidae Present Present Present Present
Fam. Cheliferidae Present Absent Present Present
Fam. Atemnidae Present Present Present Absent
Fam. Chernetidae Present Present Present Present
Order Araneae Present Present Present Present
Suborder Mesothelae Absent Absent Absent Absent
(continued)
Suborder Orthothelae Present Present Present Present
Infraorder Mygalomorphae Present Present Present Present
Fam. Microstigmatidae Present Absent Absent Absent
Fam. Hexathelidae Present Absent Present Absent
Fam. Dipluridae Present Present Present Absent
Fam. Nemesiidae Present Present Present Absent
Fam. Atypidae Present Absent Present Absent
Fam. Cyrtaucheniidae Present Absent Present Absent
Fam. Idiopidae Present Present Absent Absent
Fam. Ctenizidae Present Absent Present Absent
Fam. Migidae Present Absent Absent Absent
Infraorder Araneomorphae Present Present Present Present
Fam. Archaeidae Present Present Absent Absent
Fam. Hypochilidae Absent Absent Present Absent
am. Austrochilidae Present Absent Absent Absent
Fam. Filistatidae Present Present Present Absent
Fam. Drymusidae Present Absent Absent Absent
Fam. Scytodidae Present Present Present Absent
Fam. Sicariidae Present Absent Present Absent
Fam. Leptonetidae Absent Absent Present Absent
Fam. Ochyroceratidae Present Absent Present Absent
Fam. Telemidae Present Absent Present Absent
Fam. Pholcidae Present Present Present Present
Fam. Caponiidae Present Absent Absent Absent
Fam. Tetrablemmidae Present Present Present Absent
Fam. Dysderidae Present Absent Absent Absent
Fam. Oonopidae Present Absent Present Present
Fam. Orsolobidae Present Absent Absent Present
Fam. Segestriidae Present Present Present Absent
Fam. Eresidae Present Present Present Absent
Fam. Hersiliidae Present Present Present Absent
Fam. Oecobiidae Present Absent Present Absent
Fam. Palpimanidae Present Absent Present Absent
Fam. Mimetidae Present Present Present Absent
Fam. Deinopidae Present Present Present Absent
Fam. Uloboridae Present Present Present Absent
Fam. Udubidae Absent Present Absent Absent
Fam. Caponiidae Present Absent Absent Absent
Fam. Dysderidae Present Absent Absent Absent
Fam. Oonopidae Present Absent Present Absent
Fam. Segestriidae Present Present Present Absent
(continued)
Fam. Eresidae Present Present Present Absent
Fam. Anapidae Present Absent Present Absent
Fam. Araneidae Present Present Present Present
Fam. Cyatholipidae Present Present Present Present
Fam. Linyphiidae Present Present Present Present
Fam. Symphytognathidae Present Present Present Absent
Fam. Synaphridae Absent Present Present Absent
Fam. Tetragnathidae Present Present Present Absent
Fam. Theridiidae Present Present Present Present
Fam. Theraphosidae Present Absent Present Present
Fam. Theridiosomatidae Present Absent Present Absent
Fam. Ctenidae Present Present Present Present
Fam. Lycosidae Present Present Present Present
Fam. Oxyopidae Present Present Present Present
Fam. Pisauridae Present Present Present Present
Fam. Psechridae Absent Absent Present Absent
Fam. Trechaleidae Absent Absent Present Absent
Fam. Zoropsidae Present Absent Present Absent
Fam. Agelenidae Present Absent Present Present
Fam. Amaurobiidae Present Absent Present Absent
Fam. Anyphaenidae Present Absent Present Absent
Fam. Cybaeidae Absent Absent Present Absent
Fam. Desidae Present Present Present Absent
Fam. Dictynidae Present Absent Present Absent
Fam. Hahniidae Present Absent Present Absent
Fam. Sparassidae Present Present Present Absent
Fam. Selenopidae Present Present Present Present
Fam. Zodariidae Present Present Present Absent
Fam. Chummidae Present Absent Absent Absent
Fam. Clubionidae Present Present Present Present
Fam. Miturgidae Present Present Present Absent
Fam. Phyxelididae Present Present Present
Fam. Titanoecidae Absent Absent Present Absent
Fam. Ammoxenidae Present Absent Absent Absent
Fam. Cithaeronidae Present Absent Present Absent
Fam. Gallielienillidae Present Present Absent
Fam. Gnaphosidae Present Present Present Present
Fam. Prodidomidae Present Absent Present Absent
Fam. Trochanteriidae Present Present Present Present
Fam. Philodromidae Present Present Present Absent
Fam. Thomisidae Present Present Present Present
(continued)
Fam. Salticidae Present Present Present Present
Fam. Corinnidae Present Present Present Absent
Fam. Liocranidae Present Present Present Absent
Order Opilioacarida Present Present Absent Absent
Fam. Opilioacaridae Present Present Absent Absent
Order Holothyrida Absent Absent Present Present
Fam. Holothyridae Absent Absent Present Present
Order Ixodida Present Present Present Present
Fam. Ixodidae Present Present Present
Fam. Argasidae Present Present Present
Fam. Nuttalliellidae Present Absent Absent Absent
Order Mesostigmata Present Present Present Present
Order Sarcoptiformes Present Present Present Present
Order Trombidiformes Present Present Present Present
Oribatida On the island of Madagascar and five neighbouring islands: Aldabra,

the Comoros, Mauritius, Réunion and the Seychelles, the number of hitherto found
Ptyctima species is 57, including 37 Phthiracaroidea, 18 Euphthiracaroidea, and
two Mesoplophoroidea. Of those 57 species, 30 are endemic (53%), ten Ethiopian
species (17%), 12 pantropical species (21%) three oriental (5%) and two semicos-
mopolitan (3%). Endemism of Phthiracaroidea (62%) is similar to Euphthiracaroidea
(59%). Similarity between the ptyctimous fauna of Madagascar and the neighbour-
ing islands is very small - only one or two species are common. Moreover, the fauna
of Mauritius is completely different from that of Madagascar. The ptyctimous fauna
of Madagascar and the near islands must have been shaped before separation of
Madagascar from Africa (Niedbala 2004).
8.16.2 Remarkable Distributions and Absences
Southern Africa – Missing orders or suborders: Orders Holothyrida, Ricinulei,

Thelyphonida (Uropygi); suborders Dyspnoi (Opiliones), Mesothelae (Araneae),
Palaeoamblypygi (Amblypygi); present orders Opilioacarida (1 genus in common
with Madagascar), Schizomida (endemic genus and subfamily), Cyphophthalmi (3
endemic genera of a family shared with Madagascar and other countries), Eupnoi
(3 endemic genera, 1 endemic subfamily), Laniatores (1 endemic subfamily),
Pseudoscorpiones (14 endemic genera, no endemic families), Scorpiones, Solifugae
(endemic families), Araneae (48 endemic genera, 2 endemic families), Acari
Ixodida (1 endemic family)
8.17 Evaluation of Wallace’s Line and the Other Lines in Southeast Asia… 727
Madagascar – Missing orders or suborders: Holothyrida, Solifugae, Ricinulei,

Uropygi; suborders Eupnoi and Dyspnoi (Opiliones), Mesothelae (Araneae),
Palaeoamblypygi (Amblypygi); recent orders Palpigradi (1 endemic genus),
Opilioacarida (1 genus in common with Southern Africa), Scorpiones (12 endemic
native genera, 100% native species, 1 endemic family, sharp difference with
Southern Africa or the other islands), Pseudoscorpiones (no endemic families),
Araneae (no endemic families)
Seychelles – Missing orders or suborders: Orders Opilioacarida, Ricinulei,
Uropygi; suborders Cyphophthalmi, Eupnoi and Dyspnoi (Opiliones), Mesothelae
(Araneae), Palaeoamblypygi (Amblypygi); present orders Holothyrida (three
endemic genera), Schizomida (four endemic genera)
Mascarene – Missing orders or suborders: Orders Opilioacarida, Ricinulei,
Uropygi; suborders Cyphophthalmi and Dyspnoi (Opiliones), Mesothelae
(Araneae), Palaeoamblypygi (Amblypygi); present orders Holothyrida (endemic
family), Pseudoscorpiones (two endemic genera), Eupnoi (Opiliones)
On all smaller islands is missing the Opilion family Triaenonychidae, having in
Madagascar 15 endemic genera. There are almost no genera of Opiliones in com-
mon between Seychelles and Mascarene and no genera in common between
Madagascar and the other island groups.
Groups of no special zoogeographic interest (no endemisms above species):
Southern Africa – Palpigradi
Madagascar – Amblypygi, Opilioacarida, Schizomida, Pseudoscorpiones
Seychelles – Amblypygi
Mascarene - Amblypygi, Eupnoi (Opiliones), Scorpiones
Conclusion: Arachnologically Madagascar, the Seychelles, and the Mascarene
are so different from the African continent that they deserve the rank of region pair-
ing with Africa. There is also considerable difference between Madagascar and the
smaller islands, which are often called subregions. Important is the absence of many
groups (orders and families) abundant in Africa but lacking in the islands.
8.17 E
valuation of Wallace’s Line and the Other Lines
in Southeast Asia According to the Arachnida.
“La frontière, en biogéographie, n’est pas, en

général, une ligne, mais une zone complexe don’t
l’emplacement et la topographie ont subi de
multiples variations au cours des temps
géologiques... On peut cependant affirmer que la
ligne Wallace – pour certaines formes de Scorpions
est une frontière réellement existante.” Max Vachon (1953)
8.17.1 Wallace’s Line
In result of his studies of the nature of Australasia, Wallace (1876) concluded that
“We may consider it established that the Strait of Lombok [between Bali and
Lombok] (only 15 miles wide) marks the limit and abruptly separates two of the
great zoological regions of the globe.” This famous line, called by Huxley (1868)
“Wallace’s line”, runs through Makassar Strait between Borneo and Sulawesi and
then the line was based mostly on mammals, but Mayr (1944b) made the remark
that “An equally pronounced faunal difference exists among birds, insects, and
other groups of animals in the two regions.”
When crossing the famous line in 1994 and 1995, I asked myself: “Is this state-
ment true also for such zoogeographically interesting animals as Arachnida?”
Meanwhile, a lot of new information was accumulated on the taxonomy of such
groups as Schizomida, Opiliones, Scorpiones, and others, and time has come to
check how this zoogeographical “rift” (the term belongs to Prof. Vachon) reflects
the past and present distribution of the various Arachnida. Some orders (Solifugae,
Opilioacarida, Ricinulei) are not known in this area.
8.17.2 Lydekker’s Line
The line proposed in 1895 by the British naturalist Richard Lydekker (1849–1915)
is separating Wallacea from the shelf of Australia – New Guinea.
8.17.3 Weber’s Line
Max Weber (1852–1937) proposed a line passing between Sulawesi and Halmahera
and marks the balance of the Indomalayan and Australian elements in the fauna. It
is a line based on mammals and should be tested what concerns the invertebrates
(Arachnida and others).
Wallacea Observing longtime the strange transition in the animal world of the
islands of Malay Archipelago, Wallace presumed, “such facts could be explained
only by major changes in the Earth’s surface.” Now we know that in the Pleistocene,
the major continental islands Borneo, Java, Sumatra, and Bali have been connected
with the Asian mainland. The rainforest existed by this time, and the sea level was
lower by 180 m. That is why on these islands, we find the same animals like in
Malaya (sometimes other subspecies) – elephants, rhinos, tapirs, tigers, leopards,
and primates. Some of them meanwhile have disappeared on the continent but still
live on the islands (the orangutans).
Crossing the narrow (only 25 km) Lombok Chanel between Bali and Lombok,
all these animals disappear. Appear cockatoo parrots, some marsupials, and differ-
ent reptiles.
During his research between 1854 and 1862, Wallace found that the birds on Bali
and Java are almost the same (97%), and as soon as we cross the 25 km straight, the
picture changes abruptly – Bali and Lombok have only 50% of their bird species in
common; one may think that the others were not capable to fly over this distance.
Starts Wallacea, most interesting territory of a mix of fauna.
The ancestors of the present-day animals and plants of Wallacea as a whole origi-
nate either from Asia or from Australia – New Guinea – but on the bigger islands
occurred an active autochthonous speciation, and there are many endemics. Some of
the emblematic Indonesian endemics are actually inhabiting Wallacea (anoa, baby-
russa, “Comodo dragon”). Most islands have not been interconnected, they are
separated by deep straights, and this was important to explain the high endemism.
According to Conservation International, Wallacea has more than 10 000 species
of plants, including ca. 1500 (15%) endemic. Among the terrestrial vertebrates, the
endemism is even higher: from 1142 species, almost half (529) are endemics. In
Wallacea live 223 species of native mammals, 126 of them endemics. Only bats
count 124 species – almost one tenth of the world’s bats.
The birds of Wallacea are extremely diverse (650 species, including 265
еndemic). More than half of them live in Sulawesi alone (356 species, 96 endemic).
In Wallace’s time, the area has been covered by lush tropical forests. Only small
fraction of them remains – 45% of the surface is covered by some sort of forest, but
only in 52,017 km², or 15% – the forest is in its pristine state. From the total surface
of Wallacea (347,000 km²), some 20,000 km² is protected (at least legally). The
remaining forest is most often under concessions of powerful Western or Japanese
companies, which destroy not only the irreplaceable rainforest but also hundreds of
its described, semidescribed, and undescribed inhabitants.
Map 8.6 Location map of Torres Strait in relation to the regional Quaternary shoreline changes
(Adapted from Rowe 2007)
8.17.4 D
evelopment of Wallacea and the Surrounding Lands
of Southeast Asia and the Malayan Archipelago
The analysis of Moss and Wilson (1998) concerning the biogeographic implications
of the Tertiary paleogeographic evolution of Sulawesi and Borneo is worth quoting.
Wallacea, according to them, is a biogeographic region, situated between the areas
with Asiatic and Australian floras and faunas, where organisms show a high degree of
endemism. A land connection between Borneo and mainland Southeast Asia may
have existed throughout much of the Tertiary and would have allowed migration of
terrestrial biota. Western Sulawesi had been connected to Eastern Borneo by the Late
Cretaceous and by the Early Eocene with possibilities of dispersal of fauna between
Borneo and Western Sulawesi. The East Sulawesi ophiolite was accreted onto Sulawesi
during or after the late Oligocene and resulted in the formation of more extensive land
areas of Sulawesi. “Microcontinental fragments accreted onto eastern Sulawesi in the
Miocene to Pleistocene may have been emergent as they drifted towards Sulawesi and
allowed island hopping or rafting for biota of Australian affinity. Island hopping routes
for the dispersal of organisms between Borneo-Sulawesi and the Philippines may have
existed along volcanic arcs, such as the long-lived North Sulawesi arc, the Sulu and
Sangihe arcs, and the Cagayan arc” (Moss and Wilson 1998).
Remarks Concerning Araneae From the three suborders of spider, one

(Mesothelae) does not live east of Wallace’s Line but is known from Southeast Asia.
From 112 families of spiders in the world, 19 are known from Australasia east of
this line, but not from Southeast Asia. Some are endemic for Australia, New Zealand,
or New Caledonia; others live also in faraway parts of the world, mostly in South
Africa, Chile, or Argentina. Typical are Austrochilidae, Orsolobidae, Malkaridae,
Cyatholipidae, Amphinectidae, Zoropsidae, and Ammoxenidae, also subfam.
Prithinae of fam. Filistatidae (one genus in New Guinea, two in Australia, seven in
Africa, North and South America). Some subfamilies of widespread families are
endemic for the area east of Wallace’s Line: Hexathelidae (Hexathelinae), Dipluridae
(Masteriinae), Cyrtaucheniidae (Aporoptychinae), Idiopidae (Arbanitinae),
Miturgidae (Diaprograptinae), Gallieniellidae (Meedoinae), and others.
8.17.5 The “Lines” and Some Other Groups
In their Atlas of Diplopoda, Shelley and Golovatch (2011) conclude that

From present records, the hypothetical borders separating Asian and Australian faunas in
Indonesia (Wallace’s, Weber’s, and Lydekker’s ‘lines’) hold little relevance for diplopods.
Of the four exclusively Laurasian orders, only Glomerida and Platydesmida (…) expand
into the East Indies/Indonesia, as Julida and Callipodida (…) terminate on the southeast
Asian continent. In the north, Glomerida spread eastward to Weber’s line, between Sulawesi
and Halmahera, while the southern edge conforms roughly to Wallace’s line. Platydesmida,
however, extend to Wallace’s line between Borneo and Sulawesi while the southern border
passes between Sumatra/Java and corresponds to nothing. We would not expect these
‘lines’ to be operative for the Gondwanan representatives, which were carried passively to
southeast Asia by the terranes, and this is indeed the case. The only taxon conforming pre-
cisely to one of these hypothetical boundaries is Spirostreptidea, derived from Gondwana I,
whose eastern border matches Lydekker’s line completely. While future discoveries may
alter East Indian taxon borders, hypotheses like these lines may not apply to low vagility
organisms like millipeds, whose geographies are so dependent on geological events; con-
formity of Spirostreptidea may therefore be coincidence.
In a recent analysis of the caddisflies (Trichoptera) of Lombok, Bali, and Java,

Malicky et al. (2014) conclude that “The caddisfly fauna of the three islands is of
Asiatic origin, no Australian influence was noted. The well-known Wallace’s line
does not act as a faunistic border between Bali and Lombok for Trichoptera.”
It is interesting to compare this analysis with our data concerning the Arachnida –
other non-insect group of relatively small animals with (mostly) low vagility. Of
course, there are differences – almost all Arachnida are predators, not living so
much in rotten logs, some are ballooning, and others are small and relatively easy to
be dispersed by air or otherwise (at least, it seems so). According to Lehtinen (1980),
“Various lines limiting and dividing the Wallacea seem to have no equivalents in the
arachnological zoogeography.” It seems that Gressitt (1974) arrived to a similar
conclusion on what concerns the zoogeography of insects.
Map 8.7 Australasia and the lines (After Hall 2002, modified)
8.17.6 Arachnida and the Lines
Palpigradi
Two families (Eukoeneniidae and Prokoeneniidae) are represented in the area.
West of Wallace’s Line: Fam. Prokoeneniidae and the genus Prokoenenia (one sp.
on Sulawesi) – Thailand, Java, Sulawesi
East of Wallace’s Line: None
Both sides: Fam. Eukoeneniidae (genera Eukoenenia and Koeneniodes)
Amblypygi
Two families are represented in the area, Charinidae and Charontidae, both of the
two sides of the Wallace’s Line. Special case is the only species of Phrynus (from
the American family Phrynidae) from a cave on Flores (Indonesia).
Only West of Wallace’s Line:
Catageus Thorell, 1889 – Burma
Only East of Wallace’s Line – None
Both sides:
Charinus Simon, 1892 (Charinides Gravely) – Australia, Andaman Islands, India,
W. Samoa, Vanuatu, Indonesia (Java, Borneo), Singapore, Malaysia, Papua New
Guinea, New Caledonia, Solomon Islands
Sarax Simon, 1892 (Phrynichosarax Gravely) – Solomon Islands, Malaysia,
Singapore, the Philippines (Luzon), Indonesia (Java, Kalimantan), India, Andaman
Islands, Papua New Guinea, New Britain, Vietnam, Laos, Cambodia, Borneo
Charon Karsch, 1879 – Australia (Northern Territory, Queensland, Christmas
Island), Indonesia (Java, Maluku, Sumbawa), Malaysia (incl. Borneo), Palau, Papua
New Guinea, the Philippines, Singapore, Solomon Islands
Stygophrynus Kraepelin, 1895 – Solomon Islands (Guadalcanal), Burma,
Thailand, Vietnam, Malaysia, Indonesia (Java, Sumatra, Kalimantan)
Uropygi (Thelyphonidae, incl. Hypoctoninae) are known from the area (from both
sides of Wallace’s Line).
West of Wallace’s Line:
Ginosigma Speijer, 1936 – Sunda Islands, Thailand
Hypoctonus Thorell, 1889 – Burma (Myanmar), South China, Malaysia, Thailand,
Bangladesh, Java, India
East of Wallace’s Line:
Chajnus Speijer, 1936 – Sunda Islands (Lombok)
Glyptogluteus Rowland, 1973 – Philippines (Panay)
Thelyphonoides Krehenwinkel et al., 2009 – Philippines (Panay)
Both sides:
Thelyphonus Latreille, 1802 (syn. Abaliella Strand, 1928, syn. Minbosius Speijer,
1936, syn. Tetrabalius Thorell, 1889, fide Haupt, 2009a) – Indonesia, Singapore,
Philippines, Vanuatu (New Hebrides), Fiji, Rennell, Solomon Islands, New
Guinea, New Britain, Burma, Sri Lanka, India, Thailand, W. Samoa, Borneo,
Moluccas
Typopeltis Pocock, 1894 (syn. Teltus Speijer, 1936) – China, ? Russia, Taiwan,
Hainan, Japan, Thailand, Vietnam
Schizomida
One family (Hubbardiidae) is represented in the area.
Schizomus Packard-Cambridge, 1972
Clavizomus Reddell et Cokendolpher, 1995 – Java, West Malaysia, Singapore
Trithyreus Kraepelin, 1899 – Burma (Myanmar)
Zomus Reddell et Cokendolpher, 1995 – Malaysia (incl. Sarawak), Singapore,
England (Kew Gardens)

Attenuizomus Harvey, 2000 – Australia, NT
Brignolizomus Harvey, 2000 – Australia
Draculoides Harvey, 1992 – Western Australia
Julattenius Harvey, 1992 – Queensland (Australia)
Notozomus Harvey, 1992 - Queensland (Australia)
Orientzomus Cokendopher et Tsurusaki, 1994 – Philippines (Luzon), Japan, Bonin
Island
Paradraculoides Harvey et al., 2008 – Western Australia
Hubbardiidae indet. – New Guinea, New Britain, New Ireland; Schizomus
modestus
Both sides:
Apozomus Harvey, 1992 – Australia, Japan, Papua New Guinea, Taiwan, Sabah
(Borneo), W. Malaysia, Vietnam
Bamazomus Harvey, 1992 – Thailand, Hong Kong, W. Malaysia, Ryukyu Islands,
Papua New Guinea, Australia, Hawaii
Ovozomus Harvey, 2001 – Seychelles, Comoro Islands, Cook Islands, Ceylon,
India, Christmas Island, Réunion
Scorpiones
Seven families of scorpions are known from the area.
Fam. Chaerilidae – Bangladesh, India, Indonesia, Malaysia, Nepal, Singapore, Sri
Lanka, Vietnam
Fam. Pseudochactidae – Caves of Laos and Vietnam
Fam. Scorpiopsidae – Southeast Asia, India, Indonesia, Malaysia
Fam. Scorpionidae – South and Southeast Asia, Indonesia
Fam. Urodacidae – Australia
Both sides:
Fam. Buthidae – All continents, tropical and subtropical countries and in some
countries with moderate climate
Fam. Hormuridae – Australia, Southeast Asia, Pacific islands, Indian Ocean
Islands
Pseudoscorpiones
In the area have been registered representatives of 20 families. Most of them (18)
are represented from both sides of Wallace’s Line and usually also on other conti-
nents or worldwide.
Fam. Ideoroncidae – Malaysia, India, Cambodia, Thailand, Vietnam, Indonesia
(Sumatra)

Fam. Pseudogarypidae – Tasmania (one extant species)
Both sides:
Fam. Chthoniidae – Cosmopolitic
Fam. Tridenchthoniidae (Dithidae) – Australia, New Caledonia, Lord Howe
Island, Caroline Islands, India, Vietnam, Philippines, Indonesia (Moluccas,
Sumatra, Java), New Guinea, Japan, Palau Island, Bhutan, Nepal, Laos, Marcus
Island, Malaysia
Fam. Lechytiidae – Vietnam, Marshall Islands, Marcus Island
Fam. Hyidae – Philippine Islands, Indonesia (Java, Sumatra), Australia
Fam. Neobisiidae – Philippines, Vietnam, Thailand, Japan, Burma, China, Taiwan
Fam. Syarinidae – Bhutan, India, Sri Lanka, Nepal, Tuvalu, Philippines, Solomon
Island, New Zealand, New Caledonia, New Guinea
Fam. Parahyidae – Caroline Islands, Singapore
Fam. Garypidae (Synsphyronidae) – India, Japan, Indonesia (Krakatau, Nicobar
Island), Maldives Island, Sri Lanka-Marshall Islands, Australia (incl. Tasmania),
New Zealand
Fam. Menthidae – Socotra, Australia
Fam. Cheiridiidae – China, Taiwan, Nepal, Philippines, Japan, Hawaii-Papua New
Guinea, Salomon Islands, Mariana Islands, Australia, New Zealand
Fam. Olpiidae – Australia, New Zealand, Norfolk Island, Indonesia, India, Burma,
Japan, Cambodia, Vietnam, Papua New Guinea, New Caledonia, Salomon
Islands, Tuvalu, Samoa, Caroline Islands, Mariana Islands, Marshall Islands
Fam. Garypinidae – Solomon Islands, Australia, New Guinea, Hawaii, Indonesia,
Thailand, Cambodia, Vietnam
Fam. Sternophoridae – Australia, Papua New Guinea, India, Sri Lanka, Laos,
Cambodia, Vietnam
Fam. Withiidae – New Zealand, Lord Howe Island, Vietnam, Indonesia, Christmas
Island, India, Cambodia, Malaysia, Thailand, Philippines, Solomon Islands,
Nepal, China, Taiwan, Mariana Islands, Marshall Islands, Papua New Guinea
Fam. Cheliferidae – Indochina, India, China, Indonesia-Australia, New Zealand,
Papua New Guinea
Fam. Atemnidae (Miratemnidae) – South East Asia, Philippines, Bhutan, Nepal,
India, Nicobar Islands, Solomon Islands, Sri Lanka, Indonesia, China-New
Caledonia, Lord Howe Island, Caroline Islands, Mariana Islands, Christmas
Island, Marquesas Islands, Marshall Islands, Papua New Guinea
Fam. Chernetidae (Myrmochernetidae) – New Guinea, Solomon Islands, etc.
Opiliones
All suborders are represented in the area.
Cyphophthalmi (three fam.)
Only west of Wallace’s Line: None
Only east of Wallace’s Line:
Fam. Troglosironidae – New Caledonia

Both sides:
Fam. Stylocellidae – Southeast Asia (from India to New Guinea)
Fam. Pettalidae – New Zealand, Australia-Sri Lanka
Eupnoi
Fam. Phalangiidae – Mitopiella Banks, Borneo
Fam. Caddidae – Australia (incl. Tasmania), New Zealand (but also in North
America, South America, and South Africa, not found in Southeast Asia)
Fam. Sclerosomatidae – Europe, Asia, Africa, South, Central and North
America, Antilleans, Australia, New Guinea, Solomon Island (not found in
Southeast Asia)
Fam. Neopilionidae – Australia, New Zealand (but also in South America and
South Africa, not found in Southeast Asia)
Both sides: None
Dyspnoi
Fam. Nemastomatidae (Ortholasmatinae, Cladolasma Suzuki) – Thailand,
China, Japan (three spp.)
East of Wallace’s Line: None
Both sides: None
Laniatores
Fam. Stygnommatidae – ? Neotropical family with two doubtful spp. from
Malaya of genus Stygnomimus Roewer, 1927
Fam. Biantidae (Biantes, Probiantes) – Nepal, India, Burma, Sumatra, Malaya
Fam. Pyramidopidae – Africa, Central America
Fam. Synthetonychiidae – New Zealand (14 spp.)
Fam. Triaenonychidae – Australia, Tasmania, New Zealand, New Caledonia
(but also in Madagascar, South Africa, South America)
Both sides:
Fam. Assamiidae – Southern Asia, subfam. Dampetrinae to Indonesia, New
Guinea, and Australia
Fam. Samoidae – Australia (New South Wales), Samoa, Indonesia
Fam. Podoctidae (incl. Erecananinae and Ibaloniinae) – Palau, Taiwan, Indonesia,

Malaysia, New Guinea, India, Philippines, Vanuatu
Fam. Sandokanidae [Oncopodidae, praeoccup.] – Philippines, Sumatra, Borneo,
Burma, Nepal, India, Malaysia, Singapore, Thailand
Philippines, New Guinea, Thailand, Sulawesi, Borneo, Sumatra, Japan, Malaysia,
India, Taiwan, China, Vietnam
Fam. Zalmoxidae (Stygnoleptinae) – New Guinea, Solomon Islands, Fiji, Bismarck
Archipelago, Philippines, Java, Sulawesi, Moluccas, Australia, Marianas, New
Caledonia, Marshall Islands, Caroline Islands, Polynesia
Remarks Concerning Araneae From the three suborders of spider, one
(Mesothelae) does not live east of Wallace’s Line but is known from Southeast Asia.
From 112 families of spiders in the world, 19 are known from Australasia east of this
line, but not from Southeast Asia. Some are endemic for Australia, New Zealand, or
New Caledonia; others live also in faraway parts of the world, mostly in South Africa,
Chile, or Argentina. Typical are family Austrochilidae, Orsolobidae, Malkaridae,
Cyatholipidae, Amphinectidae, Zoropsidae, and Ammoxenidae, also subfam.
Prithinae of fam. Filistatidae (one genus in New Guinea, two in Australia, seven in
Africa, North and South America). Some subfamilies of widespread families are
endemic for the area east of Wallace’s Line: Hexathelidae (Hexathelinae), Dipluridae
(Masteriinae), Cyrtaucheniidae (Aporoptychinae), Idiopidae (Arbanitinae),
Miturgidae (Diaprograptinae), Gallieniellidae (Meedoinae), and others.
Araneae
Only West of Wallace’s Line (Especially in Southeast Asia)
Suborder Mesothelae – Southeast Asia, China, Japan
Fam. Liphistiidae – Southeast Asia, China, Japan
Suborder Orthognathi
Infraorder Mygalomorphae – Both sides
Fam. Atypidae
Fam. Hexathelidae – New Zealand, Australia, Tasmania, Asia (Macrothele)
Subfam. Hexathelinae – Australia, New Zealand
Subfam. Macrothelinae – Asia, New Zealand
Fam. Dipluridae – Australia, Oceania, Taiwan, Thailand, New Caledonia
Subfam. Euagrinae – Australia, Taiwan, New Caledonia
Subfam. Masteriinae – Oceania, Australia
Fam. Nemesiidae – Burma, India, China, Australia, New Zealand, Ryukyu Islands,
Southeast Asia
Fam. Theraphosidae – Australia, South and Southeast Asia, New Guinea
Fam. Barychelidae – Australia, New Caledonia, Southeast Asia, New Guinea, Sri
Lanka, Oceania, Fiji
Subfam. Barychelinae – Australia, New Caledonia

Subfam. Sasoninae – Australasia
Fam. Atypidae – Asia
Fam. Cyrtaucheniidae – Australia, Thailand
Subfam. Aporoptychinae – Australia
Fam. Idiopidae – Australia, New Zealand, India, Sri Lanka, South Asia, Thailand
Subfam. Arbanitinae – Australia, New Zealand
Fam. Ctenizidae – Taiwan, China, Thailand, Australian region, New Guinea
Subfam. Ctenizinae – Thailand, China, Taiwan, Australian region
Subfam. Pachylomerinae – Australian region
Fam. Migidae – Australia, New Zealand
Subfam. Calathotarsinae – Australia
Subfam. Migidae – New Zealand, Australia
Fam. Actinopodidae – Australia
Suborder Araneomorphae
Fam. Austrochilidae – Tasmania
Subfamily Hickmaniinae (as family Hickmaniidae)
Fam. Gradungulidae – New Zealand, Australia (A.C.T., Queensland, New South
Wales, Victoria)
Fam. Filistatidae – Australia, New Guinea
Subfam. Prithinae – New Guinea
Fam. Periegopidae – Australia (Queensland), New Zealand
Fam. Scytodidae – Worldwide, except of the Far North
Fam. Sicariidae – Worldwide, except of the Far North
Fam. Ochyroceratidae – South Asia, China, Pacific Islands, New Guinea
Fam. Telemidae – Sumatra, Malaysia, China, Vietnam, New Caledonia, New
Guinea
Fam. Pholcidae – Worldwide
Fam. Tetrablemmidae – Southeast Asia, Borneo, India, Indonesia (Sulawesi),
Samoa, Caroline Islands
Fam. Dysderidae – Dysdera, worldwide
Subfam. Dysderinae – Dysdera, worldwide
Fam. Oonopidae – Philippines, Southeast Asia, Australia, Tasmania, New Zealand,
New Caledonia, Hawaii, Sri Lanka
Subfam. Gamasomorphinae – Bhutan, Nepal, China, Philippines, Southeast
Asia, Australia, New Zealand, Hawaii
Subfam. Oonopidae – New Caledonia, Tasmania, Sri Lanka
Fam. Orsolobidae – Australia, New Zealand, Tasmania

Fam. Segestriidae – Asia, Australia, New Zealand
Fam. Lamponidae – Australia, New Zealand, New Caledonia (genus Centrocalia)
Subfam. Centrothelinae – Australia, New Caledonia
Subfam. Lamponinae – Australia, New Zealand
Subfam. Pseudolamponinae – Australia
Fam. Prodidomidae – Australia
Fam. Eresidae – Eurasia
Fam. Hersiliidae – Australasia, India, Sri Lanka, Australia, Borneo
Fam. Oecobiidae – Asia; Oecobius is cosmopolitan
Fam. Holarchaeidae – Tasmania, New Zealand
Fam. Mecysmaucheniidae – New Zealand
Subfam. Mecysmaucheniinae Simon, 1895 – New Zealand
Subfam. Zearchaeinae Forster et Platnick, 1984 – New Zealand
Fam. Micropholcommatidae – New Zealand, Australia, Tasmania, New Guinea
Fam. Pararchaeidae – Australia (incl. Tasmania), New Zealand, New Caledonia
Fam. Huttoniidae – New Zealand
Fam. Palpimanidae – South Asia, Sri Lanka
Subfam. Chediminae – South Asia, Sri Lanka
New Guinea, Fiji, Borneo, Vietnam, India
Fam. Malkaridae – Australia
Fam. Mimetidae – Palearctic, Africa, Principe, Central and South America, USA,
Asia, Australia: Ero and Mimetus worldwide
Fam. Deinopidae – All continents, except Europe and Antarctica
Fam. Uloboridae – Philippines, Samoa, Fiji, Vanuatu, New Guinea, Oceania,
Australia, New Zealand; Uloborus is spread worldwide; Zosis is pantropical
Fam. Anapidae – New Caledonia, Australia, Tasmania, New Zealand, New Guinea,
China, Taiwan
Fam. Araneidae – Worldwide
Fam. Cyatholipidae – Australia
Fam. Linyphiidae – Worldwide
Fam. Mysmenidae – Samoa, New Caledonia, Tasmania
Fam. Nesticidae – Worldwide
Fam. Pimoidae – North America, Asia, Europe, Japan
The Pimoidae form a relictual group along the Western coast of North America,
Europe (Alps, Apennines, and Cantabrian Mountains of northern Spain), and
the Himalaya. This makes a Holarctic predecessor probable. In 2003, a spe-
cies was found in Japan.
Fam. Sinopimoidae – (one sp.)

Fam. Symphytognathidae – In the tropics of Central and South America and the
Australian region (with Oceania), three species (Anapistula benoiti, A. caecula,
Symphytognatha imbulunga) found in Africa and one (Anapistula ishikawai) in
Japan. Anapistula jerai occurs in Southeast Asia.
Fam. Synotaxidae – Australia, Tasmania, New Zealand
Subfam. Pahorinae – Endemic to New Zealand
Subfam. Physogleninae – Australia, Tasmania, New Zealand
Subfam. Synotaxinae – Australia
Fam. Nephilidae – Pantropical
Fam. Theridiidae – Cosmopolitic
Fam. Theridiosomatidae – Australia, China, Malaysia, Sri Lanka, Philippines,
Sumatra, Samoa, Taiwan
Fam. Ctenidae – Australia, China, Sumatra, New Guinea
Fam. Lycosidae – Cosmopolitic
Fam. Oxyopidae – South East Asia
Fam. Pisauridae – Africa, South America, North America, Asia, New Caledonia,
Canary Islands, Madagascar, New Guinea, Cuba, Chatham Islands, Australia,
etc.
Fam. Psechridae – China, India, Nepal, Sri Lanka, Southeast Asia, Nicobar Islands,
Philippines, Australia
Fam. Stiphidiidae – Australia, Tasmania, New Zealand, Madagascar, Mauritius
Fam. Trechaleidae – South and Central America, Mexico, USA
Fam. Udubidae – Sri Lanka, Madagascar, Africa, USA to Panama
Fam. Zoropsidae – Australia, New Zealand, Sri Lanka, China, Korea, Japan,
Cyprus, Mediterranean, South Africa
Fam. Agelenidae – Mediterranean, Europe to Central Asia, USA to Chile, New
Zealand, Socotra, Himalaya, Cuba, Bahamas, Cyprus, China; Tegenaria,
worldwide
Fam. Amphinectidae – Australia, Tasmania, New Zealand, Chile, South America
Fam. Amaurobiidae – Russia, Korea, China, Japan, Argentina, Paraguay, Chile,
Taiwan, Australia, New Zealand, USA, Canada, Falkland Islands, India, Laos,
Thailand, Vietnam, Balkans, Greece, Crete, Germany, Switzerland, Italy,
Micronesia, Ethiopia, Libya, Nepal, Bhutan, Pakistan, Tajikistan, etc.
Fam. Anyphaenidae – Cosmopolitic
Fam. Cybaeidae – America, Europe, Japan, Korea, China, Turkmenistan, Tajikistan,
Uzbekistan, Sumatra, Venezuela, Colombia; Argyroneta, Palearctic
Fam. Desidae – Australia, Tasmania, New Zealand, New Caledonia, Oceania,
Korea, Japan, the USA, Paraguay, Chile
Fam. Dictynidae – Europe, China, Mongolia, India, Algeria, Kazakhstan, Canary
Islands, New Zealand, Australia, Sri Lanka, South Africa, USA, Canada, Mexico,
Kyrgyzstan, West Indies, Hawaii, South America, St. Helena, Andaman Islands,
Galapagos Island, New Caledonia, etc.
Fam. Hahniidae – Oceania, Australia, Tasmania, New Zealand, India, Philippines,

Sumatra
Fam. Nicodamidae – Australia, Tasmania, New Zealand, New Guinea
Fam. Sparassidae – Worldwide, except of the Far North
Fam. Selenopidae – Asia
Fam. Zodariidae – Australia, New Zealand, New Guinea, Southeast Asia
Fam. Zoropsidae – New Zealand
Fam. Clubionidae – Lord Howe Island, Asia, Australia, Oceania, Sri Lanka
Fam. Cycloctenidae – New Zealand, Australia, Indonesia (Java)
Fam. Miturgidae – Thailand, Indonesia, Australia, Tasmania, New Zealand, New
Guinea, Central and South America, Israel; Zora, Palearctic
Subfam. Eutichurinae – Australia, Thailand, Indonesia
Subfam. Miturginae – Australia, New Guinea
Subfam. Systariinae – Southeast Asia
Subfam. Diaprograptinae – Timor, Australia, New Zealand
Subfam. Zorinae – Central and South America, Australia, Tasmania, New
Zealand, Israel; Zora, Palearctic
Fam. Phyxelididae – Sumatra, Borneo
Fam. Titanoecidae – India, Sri Lanka to China, New Guinea, Marquesas Islands,
Europe, Mexico, South America; Titanoeca, Holarctic
Fam. Ammoxenidae – Australia, Tasmania, Southern Africa
Fam. Cithaeronidae – Africa, India, Malaysia, Greece, introduced to Australia and
Brazil
Fam. Gallieniellidae – Australia, Argentina, Madagascar, South Africa, Kenya
Subfam. Meedoinae – Australia (five gen.)
Fam. Gnaphosidae – Worldwide
Fam. Trochanteriidae – Australia, Sulawesi, Christmas Island, China, India, New
Caledonia
Fam. Philodromidae – India, New Guinea, Southeast Asia, Hawaii
Fam. Thomisidae – Worldwide
Fam. Salticidae – Worldwide
Fam. Corinnidae – Worldwide
Fam. Liocranidae – Burma, Thailand, Borneo, Sumatra, Australia, Sri Lanka, New
Guinea
Opilioacarida (both sides of the “Line”)
Ref.: Beron (1914), Das and Bastawade (2007), Leclerc (1989), Walter and Proctor
(1998)
Only one family (Opiliocaridae) in the world.
Only west side of Wallace’s Line:
Endemic genus Indiacarus Das et Bastawade, 2007 – India
Endemic genera Siamacarus Leclerc, 1989 and Vanderhammenacarus Leclerc,
1989 – Thailand (caves)
East side of Wallace’s Line: Indet. Opilioacarida from Australia (Walter and
Proctor 1998)
Holothyrida (both sides of the Line)
Ref.: Berlese (1923), Beron (1914), Domrow (1955), Lehtinen (1981, 1991, 1995),
Thorell (1882), Van der Hammen (1961, 1983), Womersley (1935)
Only west side of Wallace’s Line: None in Southeast Asia
East side of Wallace’s Line:
Fam. Allothyridae – Australia, New Zealand (two gen., three spp.)
Both sides:
Fam. Holothyridae – New Guinea, Seychelles, New Caledonia, Mauritius, Sri
Lanka, Lord Howe Island (23 spp.)
Remarks: It seems interesting that in New Caledonia and Lord Howe Island live
holothyrids of the genera Lindothyrus Lehtinen, 1995, and Haplothyrus Lehtinen,
1995, of fam. Holothyridae and not of Allothyridae.
Some Arachnida and the Lydekker’s Line
Palpigradi – Both sides of Lydekker’s Line (LL)
Amblypygi – Charinidae and Charontidae live on both sides (usually the same gen-
era), the only species of Phrynidae from Flores is west of LL.
Thelyphonida (Uropygi) – Only one widespread genus (Thelyphonus Latreille s.
lato) is found east of LL (in New Guinea), species of seven genera live in
Southeast Asia, Indonesia, or the Philippines.
Schizomida – The only family (Hubbardiidae) of LL, with some endemic genera on
either side.
Pseudoscorpiones – As with the Wallace’s Line, only one family (Ideoroncidae) is
known from the west side, one family (Pseudogarypidae, relict in Tasmania LL.)
only is from the east side, and 18 families of Pseudoscorpiones live on both sides
of
Opiliones Cyphophthalmi – Only the relict family Troglosironidae is known from
the east side of LL (New Caledonia), no family is known only from the Western
side, and two families (Stylocellidae and Pettalidae) have been recorded from
both sides (as with Wallace’s Line).
Opiliones Dyspnoi – Practically missing in the area (one sp. in Thailand)
Opiliones Eupnoi – Fam. Neopilionidae is found in Australia, New Zealand (but,
similarly to Triaenonychidae, also in South Africa and South America)
Opiliones Laniatores – Synthetonychiidae is endemic in New Zealand;
Triaenonychidae is known east of LL but has representatives also in
South Africa, South America, and Madagascar. Six families are distributed on both
sides of LL.
Scorpiones – As with the Wallace’s Line, only one family (Urodacidae) is typical
for the Australian side, four families are known only in the west, and two
(Buthidae and Hormuridae) live on both sides.
Araneae – Suborder Mesothelae lives only West of all “lines.” Mygalomorphae and
Araneomorphae are crossing all “lines”; many families are restricted to the land east
of Lydekker’s Line (see the comments above).
Acari – All orders are found on both sides of Lydekker’s Line. Allothyridae
(Holothyrida) is confined to Australia and New Zealand.
According to Lehtinen (1980), “Various lines limiting and dividing the Wallacea
seem to have no equivalents in the arachnological zoogeography.” It seems that
Gressitt (1974) arrived to a similar conclusion on what concerns the zoogeography
of insects.
8.17.7 Conclusions
From the orders of Arachnida, some (Ricinulei and almost all Solifugae) are miss-
ing from the analyzed area (Indomalayan and Australian regions). The situation
with the other orders is as follows.
Palpigradi – Fam. Eukoeneniidae is found on both sides (the same genera), fam.
Prokoeneniidae is found in Thailand, Java, and Sulawesi, almost not living east
of Wallace’s Line.
Amblypygi – Endemism on species level, all families and almost all genera are
found on both sides.
Thelyphonida (Uropygi) – Low level of endemism (only genera), one family on
both sides.
Schizomida – Low level of endemism (only genera), one family on both sides.
Scorpiones – Four families live only west of the “Line,” only one (Urodacidae) is
known only east of it. Two families are known on both sides.
Solifugae – Represented only by one species (Dinorhax rostrumpsittaci Simon) in
Vietnam and Maluku Islands (both sides of Wallace’s Line).
Pseudoscorpiones – One family (Ideoroncidae) only west of Wallace’s Line, 1 fam-
ily (Pseudogarypidae) only east of the Line (relict in Tasmania), 18 families on
both sides.
Opiliones – Cyphophthalmi. Two families live on both sides, one (the relict
Troglosironidae in New Caledonia) only on the east side, none on the west side.
Opiliones – Eupnoi. One widespread family (Phalangiidae) is known on the west
side (Borneo); three families live on the east side but are recorded also from
faraway countries.
Opiliones – Dyspnoi. One family (Nemastomatidae) is represented in Thailand
with one species; none is known from the east side or from both sides (practically
the suborder is absent in the whole area).
Opiliones – Laniatores. Two families are known only from the west side, and two
others are known only from the east side, but one of them (Triaenonychidae) is
recorded also from Madagascar, South Africa, and South America (but not found
in Southeast Asia). The other (Synthetonychiidae) is endemic in New Zealand.
Six families live on both sides.
Araneae
Mesothelae – Only west of the “Lines”

Mygalomorphae – Both sides of the “Lines”
Araneomorphae – Worldwide distributed suborder, living on both sides of the
“Lines”
Only on the west side of Wallace’s Line are known five families.
Only on the east side are known 20 families. Some are endemic for Australia,
New Zealand, or New Caledonia; others live also in faraway parts of the world,
mostly in South Africa, Chile, or Argentina, but not in Southeast Asia.
On both sides are recorded at least 70 families (out of the 114 families of spiders
in the world), mostly widespread.
Opilioacarida – The only family of the order Opilioacaridae is known from both
sides of the “Line.” Two endemic genera live in India and Thailand; from Australia
the order is recorded, but without further data.
Parasitiformes – We shall consider here only the zoogeographically interesting
suborder Holothyrida, living on both sides of the “Line.” The family Holothyridae
is known with endemic genera from both sides of the “Line” (not recorded in
Southeast Asia, but living in India, Seychelles, Mauritius, and Sri Lanka). Only East
of the “Line” (Australia and New Zealand) is known the family Allothyridae.
None of the higher groups of Acari (Opilioacarida, Sarcoptiformes, Prostigmata,
Acaridida, Oribatida, Parasitiformes, Mesostigmata, Ixodida, and Holothyrida) is
limited by the “Lines.”
In general, the analysis of Arachnida on both sides of Wallace’s Line confirms
the conclusion of Shelley and Golovatch (2011) that “… the hypothetical borders
separating Asian and Australian faunas in Indonesia (Wallace’s, Weber’s, and
Lydekker’s ‘lines’) hold little relevance for diplopods.” The “lines” seem to “hold
little relevance” also for Arachnida. There is no order of Arachnida living only on
one side of the “Lines.” Only the spider suborder Mesothelae seems limited to the
Western side. Some confirmation of the Vachon’s opinion (1953) (“On peut cepen-
dant affirmer que la ligne Wallace – pour certaines formes de Scorpions – est une
frontière réellement existante”) could be confirmed what concerns the scorpions.

There are families and genera, known only on one side, but the level of endemism
is relatively low. Within Arachnida we cannot detect anything similar to the endemic
orders and even subclasses in the vertebrates which are the base of the construction
of the “Lines.”
Conclusion of Lehtinen (1980): “Arachnological data suggests a rearrangement
of the classical Oriental and Australian regions into a big, fairly homogenous Indo-
Pacific region and a South-Gondwanan region, the latter extending also to southern
parts of other continents of the southern hemisphere. The limitation of their subre-
gions is largely different from the classical limitations based on the distribution of
vertebrates. Various lines limiting and dividing the Wallacea seem to have no equiv-
alents in the arachnological zoogeography.” This very experienced author thinks
also that “The spider fauna of Melanesia, Micronesia, and Polynesia, with the
exception of New Caledonia, Lord Howe Island and surrounding small archipela-
goes, is simply an impoverished Oriental fauna.”
Principal geological events important to Wallace’s Line (Partly after Audley-Charles 1981)
Million Period
Q Pleistocene
10
20 Pliocene
30 Miocene
40 Oligocene
50 Eocene
60 Palaeocene
70
Q – Quaternary
1. Australia/New Guinea splits from Antarctica (c. 53 Ma)
2. Formation of the Philippines by collision of Asian continental fragment with an island arc
(Oligocene).
3. Possible land connection(s) across Makassar Strait (mid-Miocene)
4. Collision between New Guinea and a Tertiary island arc (c.15 Ma)
5. Collision between Gondwana (Sula Peninsula) and Laurasia at or near east Sulawesi (c. 15 Ma)
but submarine
6. Island chain established between east Sulawesi and Australia (Late Miocene to Late Pliocene)
7. Collision between parts of Gondwanan Outer Banda Arc and Laurasian (volcanic) Inner Banda
Arc (Latest Miocene to Early Pliocene)
8. Probable land connection (s) across south Makassar Strait (from Late Pliocene)
8.18 The Arachnida of the Papuan Subregion
8.18.1 New Guinea and Melanesia

Melanesia consists of the island of New Guinea, the Bismarck Archipelago,

d’Entrecasteaux Islands, Louisiade Archipelago, Maluku Islands (not included
here), Fiji, Norfolk Island, Solomon Islands, Schouten Islands, Torres Strait Islands,
Trobriand Islands, Vanuatu, Woodlark Island, and some other islands. Central is the
huge island of New Guinea (829 000 km2, maximal height 4884 m.). It is actually a
small continent – only the surface of the state of Papua New Guinea is 462 840 km2.
Within this state are also the “smaller” islands – New Britain (35.600 km2, more
than Belgium), New Ireland (8.600 km2, comparable to Corsica), Bougainville
(10.500 km2), and others.
About 25% of New Guinea is situated higher than 1000 m. The giant mountain
chain, stretching along the entire island, is long more than 2000 m. and is higher
than the Alps (4884 m). The other islands are also mountainous (New Britain 2440
m, Bougainville 2740 m). The rivers of the Big Island (fed by up to 6500 mm rain
annually) are impressive. Sepik, Fly, Ramu, and Baliem are comparable to Rhine or
Don. In the Papuan phytogeographic Province (of the Malesian Region) are known
more than 9000 species and 1400 genera of higher plants, from which 8500 species
and 140 genera are endemic.
“When New Guinea, the leading edge of the Australian plate, was first elevated
to form an extensive land area in the late Oligocene, it was colonized largely from
the adjacent rich tropical lowlands of Malaysia. Biogeographic affinities with
Australia are poorly developed, but there has been a spectacular late Tertiary radia-
tion of many groups…Land connections between Australia and New Guinea are
also suggested by the mid-Pliocene appearance of the marsupials…The island as a
whole is a region of faunal and floral mixing, survival, and evolution in the middle
to late Tertiary” (Raven and Axelrod 1972).
“The consensus of contemporary zoogeographic studies is that though New
Guinea and its shelf islands were joined to Australia during several phases of the
Pleistocene, New Guinea has a basic, rich biota, much more ancient in origin as well
as evolutionary history, and independent of Australian faunal or floral influence”
(Udvardy 1975).
Some data on the glaciation and climate change in New Guinea we find in Löffler
(1982):
The Pleistocene glacial history of New Guinea may date back to about 700000 -+
100 000 years BP when lava erupted under ice on Mt Giluwe....The last glaciation lasted
until about 15 000 years BP when the ice receded rather rapidly to completely disappear by
about 9000 years BP…The snowline during the last glaciation was at about 3550 – 3600 m
altitude indicating a lowering of the snowline by about 1000 – 1100 m. This would corre-
spond to a temperature depression of about 5–6 oC assuming that precipitation was similar
to the present.
8.18 The Arachnida of the Papuan Subregion 747
On the summit of Jaya (Carstensz), the glacial ice covers 6.9 km2 (16.4 km2 in
the middle of the nineteenth century), retreating rapidly. Its thickness is ca. 40 m.
Some sources: Axelrod and Raven (1982), Chase (1971), Coleman (1980),
Doutch (1972), Gressitt (1956, 1958, 1967, 1971, 1974, 1982a, 1982b), Heads
(2002, 2009), Kemp (1984), Krijanovskiy (1980), Mitchell and Warden (1971),
Raven and Axelrod (1972), Robbins (1971), Solem (1958), Smith (1990), Taylor,
Goodliffe and Martinez (1999), Thompson (1967), Whitmore (1981).
Arachnogeography
New Guinea, Bismarck, and Salomon islands are with different positions in the
biogeographical subdivision of Australasia, view by botanists, mammalogists, orni-
thologists, and entomologists. According to the botanists and entomologists, they
are clearly part of the Indomalayan region. Krijanovskiy (1980) also includes them
into his Paleotropical dominion, together with tropical Africa, Madagascar, and the
Indomalayan region. The peculiar high-ranking vertebrates send Papuan area into
the Notogea (Australian Realm).
What concerns the Arachnida, our conclusion, is that the level of endemism is
much lower than in the mammals or birds. In the Papuan area (still very understud-
ied) are found the orders Palpigradi, Scorpiones, Pseudoscorpiones, Opiliones,
Amblypygi, Uropygi, Schizomida, Araneae, and many groups of Acari. Most inter-
esting is the order Holothyrida, represented in New Guinea by 11 species of 3
endemic genera of the family Holothyridae.
Absent from New Guinean area are the orders Ricinulei, Solifugae, and
Opilioacarida and the suborders Amblypygi, Mesothelae, and Dyspnoi.
Palpigradi Recorded are only two spp.: Eukoenenia cf. lawrencei and Koeneniodes
cf. frondiger.
(Eukoeneniidae) (Condé 1980, 1981).
Scorpiones According to the monograph of Koch (1977) and the older papers of
Takashima (1948, 1950), in New Guinea (incl. Aru and Bougainville) live only six
widespread species of the genera Isometrus, Lychas (Buthidae), and Liocheles
(Hormuridae). They are shared with Australia. From Rennell Island (the Solomon
Islands) have been found three spp., including the newly described species
(endemic) Liocheles penta Francke et Lourenço (1991) (Browning and Hyatt 1957).
Pseudoscorpiones Beier (1940, 1965b, 1967, 1968, 1982) brought the pseudo-
scorpions from Papua New Guinea (incl. Bismarck Islands to 60 species of 36 gen-
era and 12 families) (Chthoniidae, Tridenchthoniidae, Syarinidae, Cheiridiidae,
Garypinidae, Geogarypidae, Atemnidae, Olpiidae, Sternophoridae, Cheliferidae,
Chernetidae, Withiidae). Other data are provided by Beier (1940, 1982) and
Morikawa (1963).
Beier (1966b) lists 24 genera and 8 families of pseudoscorpions from the
Solomon Islands (incl. Bougainville and Buka Islands, which belong to Papua New
Guinea), mostly from Guadalcanal (26 spp.). Now the known species of
Pseudoscorpiones in the islands are 38, including some endemics. Species living on
both Solomon Islands and New Guinea and the Bismarcks are Acanthicochernes
biseriatus Beier (1964), and the common genus of both is Gelachernes Beier
(1940) – four spp. (Chernetidae).
Endemic genus for New Guinea and the Bismarck Archipelago:
Fam. Cheliferidae
Papuchelifer Beier, 1965 – Indonesia (Papua), Papua New Guinea (three spp.)
Endemic genus for the Solomon Islands:
Fam. Chernetidae
Cyclochernes Beier, 1970 – Guadalcanal (one sp.)
The known pseudoscorpions from Vanuatu (New Hebrides) are only four species
of four genera and the families Atemnidae, Cheliferidae, and Chernetidae.
Lissochelifer insularis (Beier) is common in New Guinea and Paratemnoides salo-
monis and Haplochernes hebridicus in the Solomon Islands. Only Hebridochernes
paradoxus is an endemic species to Vanuatu.
On Fiji are registered three spp. of the genera Geogarypus (Geogarypidae) and
Haplochernes (Chernetidae). Geogarypus longidigitatus (Rainbow, 1897) and
Haplochernes funafutensis (With, 1907) were described from Funafuti (Tuvalu) and
Haplochernes ellenae Chamberlin (1938) from Viti Levu (Fiji).
Opiliones From Melanesia have been recorded Opiliones of the families
Stylocellidae, Sclerosomatidae, Assamiidae, Podoctidae, Epedanidae, and
Samoidae, following Clouse and Giribet (2007), Forster (1949b), Goodnight and
Goodnight (1947), Loman (1906), Roewer (1911, 1912, 1913, 1915b, 1935–1936,
1949, 1963), and Suzuki (1982). The collection of P. Beron from New Guinea, New
Britain, and New Ireland is still under study.
Cyphophthalmi Until recently, Cyphophthalmi were not registered east of the

Lyddeker’s Line. However, two species of Stylocellus Westwood, 1874, genus
known from Malaysia, Indonesia, Singapore, and Palawan (Philippines), have been
found in Papua (Indonesian New Guinea) (Clouse and Giribet 2007). These animals
don’t cross saltwater areas, so a problem arises about their establishment in New
Guinea.
Eupnoi
Fam. Sclerosomatidae – Gagrella Stoliczka
Dyspnoi – Not recorded from Melanesia
Laniatores
In Melanesia are recorded Laniatores of the families Assamiidae, Podoctidae,
Epedanidae, Samoidae, and Zalmoxidae.
Endemic genera are:
Fam. Assamiidae
Granobunus Roewer, 1912 – New Guinea (one sp.)
Heteropygoplus Roewer, 1915 – New Guinea (one sp.)
Apygoplus Roewer, 1912 – New Guinea (three spp.)

Macrodampetrus Roewer, 1915 – New Guinea (two spp.)
Metadampetrus Roewer, 1915 – New Guinea (two spp.)
Metamosoia Roewer, 1915 – New Guinea (one sp.)
Mosoia Roewer, 1912 – New Guinea (one sp.)
Neonothippus Roewer, 1912 – New Guinea (one sp.)
Fam. Podoctidae (incl. Erecananinae and Ibaloniinae)
Asproleria Roewer, 1949 – New Guinea (one sp.)
Heteroibalonius C.J. Goodnight et M.L. Goodnight, 1947 – New Guinea (one
sp.)
Heteropodoctis Roewer, 1912 – New Guinea (one sp.)
Orobunus Goodnight et Goodnight, 1947 – (one sp.)
Paramesoceras Roewer, 1915 – New Guinea (one sp.)
Podoctinus Roewer, 1923 – New Britain (one sp.)
Proholozoster Roewer, 1915 – New Guinea (one sp.)
Santobius Roewer, 1949 (Mesoceras Soerensen, 1886, preoccup.,=Mesoceratula
Roewer, 1949) – Vanuatu, Fiji (two spp.)
Fam. Samoidae
Badessa Sørensen, in L. Koch 1886 – Fiji (one sp.)
Fijicolana Roewer, 1963 – Melanesia (one sp.)
Fam. Zalmoxidae – In New Guinea and the nearby islands live 26 spp. of the genus
Zalmoxis Sorensen (the only Zalmoxis in the Eastern Hemisphere).
Amblypygi In Melanesia are represented four genera of Amblypygi (Dunn 1949;
Gravely 1915; Rahmadi and Kojima 2010; Thorell 1888):
Fam. Charinidae
Charinus Simon – Ch. australianus (L. Koch) (Samoa, Fiji), Ch. pescotti Dunn
(Solomon Island)
Sarax Simon – S. monodenticulatus Rahmadi et Kojima (Papua, Waigeo), S.
newbritainensis Rahmadi et Kojima (New Britain, New Ireland), S.
sarawakensis (Thorell) (New Guinea, Solomon Island), S. willeyi Gravely
(New Britain), S. subterraneus Thorell (papuanus) (New Guinea, Maluku).
Fam. Charontidae
Charon Karsch – Ch. grayi (Gervais) (? = papuanus Thorell) (New Guinea)
Stygophrynus Kraepelin – S. (Neocharon) forsteri Dunn (Solomon Island,
endemic for Guadalcanal)
Thelyphonida (Uropygi) On the island of New Guinea and nearby islands of
Fergusson, New Ireland, and New Britain live three species of Abaliella Strand. One
Thelyphonus was described by Rowland (1973a) from Guadalcanal (Solomon
Islands), added to the other species from the Solomon’s Thelyphonus leucurus
Pocock (1898). The widespread Minbosius manilanus (C.L. Koch) lives on the
Moluccas, in New Guinea, and the Philippines. All of them belong to Thelyphonidae
and Thelyphoninae, typical for the Papuan area, Samoa, the Philippines, and
Southeast Asia. Haupt (2009a) synonymized Abaliella Strand, Minbosius Speijer,
and Tetrabalius Thorell with Thelyphonus Latreille.
Schizomida Schizomus modestus Hansen, 1905, was described from New Guinea
and recorded also in New Britain and W. Malaysia (unverified record of Buxton
(1917)). Hubbardiidae indet. are known from New Guinea, New Britain, and New
Ireland (Reddell and Cokendolpher, 1995). In my collection in Sofia, there are
schizomids, collected by me in New Ireland in 1975.
Araneae
The Spiders of New Guinea
In New Guinea and the Bismarck Archipelago (incl. Buka and Bougainville) have
been recorded spiders of at least 50 families (Balogh 1936; Baehr and Baehr 1993;
Baert 1979, 1980, 1982, 1984; Bourne 1980; Chrysanthus, many papers since 1958;
Lehtinen and Saaristo 1980; Main 1982; Robinson 1982; Thorell 1881; Versteirt
et al. 2010; York 1982). The collection, brought by me (43 spp. of 15 families), was
identified only partially by Brignoli (1981), but his untimely death did not allow
him to describe the many new taxa in this collection. It has noted that some species
related to groups believed “typical” of New Zealand (Stiphidiidae). The preliminary
conclusion of Brignoli (1981) is “It is still too early to appreciate the value of the
proposal of Lehtinen (1980), of abandoning the traditional Oriental and Austral
regions in favour of an Indo-Pacific and a South Gondwanan region, but I would like
to observe that, if it is probably possible to accept the limited value of Wallace’s and
Weber’s lines for spiders, I do not see every definite border between the two new
regions proposed by Lehtinen.”
Several families of spiders are known from Australia (Zoropsidae, Synotaxidae),
but not from New Guinea. Others (Telemidae, Ochyroceratidae, Stenochilidae), on
the contrary, have been recorded from New Guinea, but not from Australia, despite
the proximity of both territories. None of the many Zoropsidae described in
Queensland had ventured across the Torres Strait.
The list of Araneae in Fiji contains 122 spp. of spiders (part of them indet.), of 35
families (Evenhuis 2006). Lakarobius Berry et al. (Salticidae) is an endemic genus.
Endemic genera of spiders in the “Papuan area” are:
Fam. Telemidae
Jocquella Baert, 1980 – New Guinea
Fam. Theridiidae (Hadrotarsinae)

Yoroa Baert, 1984 – New Guinea
Fam. Barychelidae
Fijocrypta Raven, 1994 – Fiji
Fam. Anapidae
Guiniella Rix et Harvey, 2010 – New Guinea (one sp.)
Fam. Philodromidae
Pseudopsellonus Balogh, 1936 – New Guinea (one sp.)
Senoculifer Balogh, 1936 – New Guinea (three spp.)
Fam. Clubionidae
Arabellata Baert, Versteirt et Jocqué, 2010 – New Guinea
Invexillata Versteirt, Baert and Jocqué, 2010 – New Guinea
Fam. Mysmenidae
Kekenboschiella Baert, 1982 – New Guinea (four spp.)
Fam. Pisauridae
Campostichommides Strand, 1911 – New Guinea (Kei Island)
Hesydrimorpha Strand, 1911 – New Guinea
Fam. Selenopidae
Godumops Crews et Harvey, 2011 – New Guinea (one sp.)
Fam. Uloboridae
Lubinella Opell, 1984 – New Guinea
Fam. Pisauridae
Hesydrimorpha Strand, 1911 – New Guinea
Papakula Strand, 1911 – New Guinea (one sp.)
Fam. Sparassidae
Clastes Walckenaer, 1837 – Moluccas, New Guinea (one sp.)
Exopalistes Hogg, 1914 – New Guinea
Strandiellum Kolosváry, 1934 – New Guinea
Fam. Stiphidiidae
Asmea Gray et Smith, 2008 – New Guinea (four spp.)
Fam. Lycosidae
Satta Lehtinen et Hippa, 1979 – New Guinea
Fam. Thomisidae
Cetratus Kulcziński, 1911 – New Guinea
Latifrons Kulcziński, 1911 – New Guinea
Rhaebobates Thorell, 1881 – New Guinea (two spp.)
Fam. Salticidae
Allococalodes Wanless, 1982 – New Guinea
Aruana Strand, 1911 – New Guinea, Aru
Bulolia Zabka, 1996 – New Guinea
Chalcolemia Zhang et Maddison, 2012 – New Guinea
[Coccorchestes Thorell, 1881 – 38 spp. from New Guinea, 1 from New Britain,
1 from Queensland
Cucudeta Maddison, 2009 – New Guinea
Diolemus Thorell, 1870 – Sixteen spp. in New Guinea, two in Moluccas
Furculattus Balogh, 1980 – New Guinea, New Britain (one sp.)
Leptathamas Balogh, 1980 – New Guinea
Opistoncana Strand, 1913 – New Ireland
Papuaneon Maddison, 2016 – New Guinea
Paraharmochirus Szombathy, 1915 – New Guinea
Porius Thorell, 1892 – New Guinea (two spp.)
Saitissus Roewer, 1938 – New Guinea
Tabuina Maddison, 2009 – New Guinea (three spp.)
Tarodus Pocock, 1899 – New Britain
Variratina Zhang et Maddison, 2012 – New Guinea
Viribestus Zhang et Maddison, 2012 – New Guinea
Zabkattus Zhang et Maddison, 2012 – New Guinea
Fam. Corinnidae
Melanesotypus Raven, 2015 – Solomon Islands
Holothyrida With 11 out of 29 species in the order Holothyrida, the island of New
Guinea looks like the center of speciation of these strange arachnids. All species are
endemic to New Guinea: Hammenius fujuge fujuge, H. fujuge iora, H. ingii, H.
insularis, Hammenius grandjeani grandjeani, Leiothyrus armatus, L. holthuisi, L.
nitidissimus, Thonius berlesei, Th. longipes, Th. montanus, and Th. mendi. So are
the genera Hammenius Lehtinen and Thonius Lehtinen. Hammenius niger (Thon)
lives on Silhouette Island (the Seychelles), a highly interesting distribution for these
conservative animals (Thorell 1882, Beron 2014; Canestrini 1897; Lehtinen 1981,
1995; Van der Hammen 1983).
8.19 Arachnogeography of Australia and Tasmania 753
8.19 Arachnogeography of Australia and Tasmania
8.19.1 Australia
The continent of Australia has an area of 7 692 000 km2 (without Tasmania but
including the offshore islands). The highest point is Mount Kosciuszko (2228 m.).
The continent was detached from Antarctic some 95–80 Ma (Late Cretaceous)
and drifted northward, coming close to the equator and to Sundaland, the present-
day Indochina, and the Greater Sunda islands. The present-day Australia consists of
several subregions of very different landscape. Huge areas in Western and central
Australia are deserts and form the so-called Eremial.
The northernmost part of the continent, on Torres Strait, consists of dry savanna
(in the Western part) and of rainforests similar to the forests in New Guinea (in the
Eastern part). Udvardy (1975) notes that Usinger (1963) attached Cape York (the
northern peninsula of Queensland) to his Papuan subdivision of the Oriental region,
but Gressitt (1961) treated it as a clearly transitional area together with southern
New Guinea and in 1975 still opines (in a letter to Udvardy) that “the overlap of
Australian and Oriental in southern New Guinea and Northern Australia needs to be
shown as an overlap zone with dominance of Oriental elements….” Prof. Gressitt
told the same personally to the present author during a visit in Wau in the same year
(1975). However, this remark concerns mostly to the flora and the invertebrates,
especially the insects. Speaking of vertebrates, Papuan Subregion is clearly part of
the Australian region (the differences are between kingdoms!).
To quote partly Keast (1959):
New Guinea and Tasmania, lying on the Australian continental shelf, would be brought into
contact by a fall in sea-level of about 10 m and 50 m, respectively. They are known to have
been joined to Australia during part of the Tertiary and apparently twice during the
Pleistocene…This being so, it would seem surprising that the flora of New Guinea bears
little resemblance to that of Australia…and much of its invertebrate fauna is likewise Indo-
Malayan…In explanation of this it has been suggested that the geologically ancient north-
ern part of New Guinea was once, presumably in the Mesozoic, in direct land contact with
Asia … Another proposal is that Australia did not always lie in close proximity to New
Guinea but ‘drifted’ into it.
According to Keast (1959), “approximately one-third of Australian continent lies

within the tropics and the rest is either temperate or sub-temperate.” Special
adaptations to cold are little developed in Australian animals. Also after this author,
Australia has not been in direct land contact with Asia since the beginning of
Tertiary, a period of over 50 Ma. Actually, after the modern concepts, Australia has
never been in direct contact with Asia, as it drifted from the south.
If for the final phase of extinction of the giant animals in Australia (Diprotodon,
Palorchestes, Thylacoleo) could be blamed the combination of aridity and the arrival
of aboriginal man and the dingo, eventual changes in the composition of the arach-
nofauna are to be explained only with the aridization. Some sources: Brown,
Campbell and Crook (1968), Embleton (1973), Filewood (1984), Keast (Ed) (1981),
McElhinny et al. (1976), Veevers and McElhinny (1976), Walker (Ed.)(1972),
Weissel et al. (1976).
Several attempts have been made to subdivide Australia zoogeographically
(Hedley 1894; Spencer 1896; Sloane 1915; Campbell 1943; Udvardy 1975). The
history of the regionalization has been narrated by Ebach (2012). A recent subdivi-
sion is due to Ebach, Gill, Kwan, Anyong, Murphy, and Cassis (2013). On the main-
land are outlined 15 regions; Tasmania is a subregion.
The zoogeographical subdivision of Ebach et al. (2013) of the “Australian
Sub-realm”:
Sub-realm Australia
Bassian Region
Subregions: Southeastern Forest, McPherson-MacPherson-Macleay, Adelaide,
Tasmania
Torresian-Bassian Region
Subregions: Atherton Plateau, Cape York Peninsula, Arnhem Land, Kimberley
Plateau
Eyrean Region
Subregions: Northern Desert, Eastern Desert, Pilbara, Western Desert, Eyre
Peninsula, South-West Australia, Riverina
“Breakup between Australia-Antarctica and the northern part of Greater India
commenced ca. = 130 Ma, and between Australia and Antarctica around 96 Ma. At
the beginning of the Palaeogene, Australia commenced its northward drift towards
its present position. Seafloor spreading between Australia and Antarctica was at first
slow, but increased to ca. 5 cm. per year around 45 Ma. By 35 Ma, the Antarctic
Circumpolar Current became established, thereby triggering glaciation in Antarctica.
Northern Australia reached the tropics by the beginning of the Miocene, and Australia
has progressively moved northwards at 7–8 cm per year since” (Li and Powell 2001).
Arachnogeography Concerning the vertebrates, Australia is a continent with
many endemic high-ranking taxa, and many groups are lacking in the native fauna.
About Arachnida there are no endemic orders or suborders, and lacking are only a
few: the orders Ricinulei, Thelyphonida (Uropygi), and Solifugae and the suborders
Paleamblypygi, Dyspnoi, and Mesothelae.
Some of the 11 orders, represented on the continent, contain endemic taxa, but of
low rank, incomparable with the situation with the vertebrates.
Palpigradi, Amblypygi, and Opilioacarida are represented with only several spe-
cies. The scorpions are of four families (1 family is endemic, 42 spp.);
Pseudoscorpions (150 spp.) and Schizomida (53 spp., 9 genera, mostly endemic)
are well studied, thanks, besides Beier and other older authors, mainly to the ener-
getic researcher M. Harvey and his team. On the continent there are also good
specialists of opilions, spiders, and many groups of Acari. Still, very few high-
ranking endemic taxa have been found. There is 1 subendemic (shared with New
Zealand) family of Holothyrida; 5 endemic genera of Pseudoscorpiones; 20 end.
genera of Opiliones; 338 genera of Araneae, of which 88 (26%) are endemic; and
many endemic genera of Acari.
Yeates, Harvey, and Austin (2003) tried to make an actual (2003) estimate of the
number of Arachnida in Australia and to predict the real number of different orders,
living there. Here is their table (after 13 years, this should be modified somehow
(added sp. in brackets)).
Taxa Described Australian species Estimated total Australian species

Arachnida total 5705 42 195
Palpigrada 1 (native) 5
Amblypygi 4 10
Schizomida 46 (now 52) 80
Pseudoscorpiones 181 750
Opiliones 194 1200
Acarina 2871 20 000
Palpigradi Besides Eukoenenia mirabilis (imported), from Western Australia was

described an indigenous member of Palpigradi: Eukoenenia guzikae Barranco et
Harvey, 2008.
Scorpiones In his monograph, Koch (1977) enumerates for the “Australo-Papuan

Region” 29 spp. of scorpions from the genera Cercophonius (Bothriuridae), Lychas,
Isometroides, Isometrus (Buthidae), Liocheles (Hormuridae), and Urodacus
(Urodacidae). Endemic family is Urodacidae and end. genera are Urodacus Peters
with 19 species (incl. 7 newly described by L.E. Koch) and Isometroides Keyserling.
Cercophonius Peters with one sp. (Australia and Tasmania) has been recently found
in the Himalaya. As there are no strictly Papuan species (not found in Australia),
this is the final figure 32, to add the new genus and sp. Aops oncodactylus Volschenk
et Prendini, 2008, and the two new Urodacus, described by Volschenk, Smith, and
Harvey (2000) and Volschenk, Harvey, and Prendini (2012) of species of scorpions
on the Australian mainland or in Tasmania (the last has no endemic scorpion and
shares with the mainland Cercophonius squama). We should extract from the
32 spp. Lychas variatus (also in Fiji) and the widespread Isometrus maculatus, I.
melanodactylus, Liocheles australasiae, L. waigiensis, and L. karschii. Remain the
26 species, endemic to Australia (Cercophonius squama also in Tasmania). Australia
and New Guinea actually share only the six widespread species of Isometrus,
Lychas, and Liocheles. Now (2017) some of the family names are different.
To quote from the analysis of Koch (1977): “The highly speciated urodacine scor-
pionid genus Urodacus is widespread in Australia but absent from both New Guinea
and Tasmania. Because of the time required for the evolution of a genus, there are two
possibilities: either the genus Urodacus has always been confined to inland Australia,
or there have been species outside Australia and these became extinct. The exclusion
from Tasmania of the genera Urodacus and Lychas…could be due to cold. There is
however an absence of factors…which might exclude Urodacus from New Guinea. I
consider therefore that either its species in the more peripheral northern areas of the
Australian mainland have reached there relatively recently (since the last major trans-
gression of the sea) or that species in New Guinea have been eliminated.”
In his article on the zoogeography of Australian scorpions, Koch (1981) distrib-
uted the scorpion taxa in three “levels of endemism”:
1. Cosmopolitan species or non-endemic species shared with Asia: Isometrus mac-
ulatus, Liocheles australasiae, L. waigiensis, L. karschii
2. Species that have arisen in Australo-Papua from non-endemic genera: Lychas
(three spp.), Isometrus (one sp.)
3. Species that have arisen in Australo-Papua: Isometroides (one sp.), Cercophonius
(one sp.), Urodacus (20 spp.)
He also classified the genera regarding the continental drift:
1. Genera derived from an ancient stock with related genera only in South America:
Cercophonius
2. Australian genus with less certain relationships and probably ultimately derived
from Asia: Urodacus
3. Genera in common with Asia: Isometrus, Lychas, Liocheles
4. Genus clearly derived from Lychas in Australia: Isometroides
Further, the scorpions are distributed into three groups: Southern, Central, and
Northern species.
Endemic family is Urodacidae.
Pseudoscorpiones In Australia (Tasmania excluded) have been registered 181 spp.
of Pseudoscorpions of 59 genera and 20 families (Beier 1948, 1954, 1966, 1975;
Edwards and Harvey 2008; Harvey 1992, 2006, and many others, Harvey and
Edward 2007; Harvey and Mei Chen Leng 2008a, b; Harvey and Muchmore 1990;
Hoff 1947; Harvey 2013f, actualized).
Endemic genera of pseudoscorpions in Australia (without Tasmania) are:
Fam. Menthidae
Theхmus Harvey, 1990 (two spp.)
Fam. Olpiidae
Austrohorus Beier, 1966 – (one sp.)
Barbaraella Harvey, 1005 – (one sp.)
Linnaeolpium Harvey et Leng, 2008 – (one sp.)
Fam. Cheliferidae
Australochelifer Beier, 1975 – (one sp.)
Fam. Chernetidae
Conicochernes Beier, 1948 – (four spp.)

Marachernes Harvey, 1992 – (three spp.)
Opiliones In Australia (without Tasmania) are known ten indigenous families of
Opiliones.
Cyphophthalmi In Australia have been registered nine spp. of Cyphophthalmi of

two genera of the family Pettalidae (Boyer and Giribet 2007; Clouse and Giribet
2007; Giribet 2003; Juberthie 1988b). The genera are endemic in Queensland
(Austropurcellia Juberthie, 1988, six spp.) and Western Australia (Karripurcellia
Giribet, 2003, three spp.).
Eupnoi (Forster 1949a, 1952b, 1955; Hunt 1971, 1985, 1992; Hunt and
Cokendolpher 1991; Hunt and Hickman 1993; Kauri 1954; Roewer 1914–1949;
Shear 1996, 2001; Taylor 2009, 2011; Hickman 1957, 1958; Hogg 1909; Hunt
1990, 1995)
Fam. Neopilionidae – Arrallaba Hunt et Cokendolpher (one sp.), Ballarra Hunt
et Cokendolpher (six spp.), Plesioballarra Hunt et Cokendolpher (one sp.),
Hypomegalopsalis Taylor, 2011 (one sp.)
Fam. Sclerosomatidae – Gagrella Stoliczka (one sp.)
Genus Hesperopilio Shear (WA, one sp.) was nested in Phalangioidea by Groh
and Giribet (2015) but with unclear family belonging.
Dyspnoi – Fam. Acropsopilionidae-Acropsopilio Silvestri, 1905, Queensland
(one sp.) [Austropsopilio Forster, 1955 (syn. Tasmanopilio Hickman, 1957), NSW,
Queensland (two endemic spp., the genus known also from Tasmania)]
Laniatores In Australia (without Tasmania) are known Laniatores of 54 genera (39
genera in Triaenonychidae) and 6 families (Assamiidae, Phalangodidae (mis-
placed?), Podoctidae, Samoidae, Zalmoxidae, Triaenonychidae).
Endemic genera of Opiliones Laniatores for Australia (Tasmania excluded) are:
Breviacantha Kauri, 1954 (one sp.); Callihamina Roewer, 1942 (one sp.);
Callihamus Roewer, 1931 (one sp.); Cluniella Forster, 1955 (three spp.);
Heteronuncia Roewer, 1920 (one sp., Queensland); Holonuncia Forster, 1955 (13
spp., ACT, NSW, Victoria); Perthacantha Roewer, 1931 (one sp.); Yatala Roewer,
1942 (SA, one sp.); Conoculus Forster, 1949 (one sp.); Dingupa Forster, 1952 (one
sp., WA); Dipristes Roewer, 1931 (one sp.)
Amblypygi In Australia (the Northern Territory, Queensland, and the Australian
Christmas Island) are known six spp. of Amblypygi of the genera Charon Karsch
(three spp.) and Charinus Simon (three spp.) (Charinidae). They are proper only to
the tropical part of the continent (Dunn (1949), Harvey (1985), Harvey et al. (2012),
Harvey and West (1998), Monteith (1965).
All species are (so far) endemic to the areas of description. From Western
Australia has been recorded another amblypygid (Harvey et al. 2012).
Map 8.8 Distribution of Amblypygi and Schizomida in Australia

Amblypygi
Charon – ▲
Charinus – ▼
Schizomida
Draculoides – ■
Apozomus – □
Bamazomus – ◊
Notozomus – ♠
Julattenius –
Brignolizomus – ◄
Ovozomus – ►
Paradraculoides – Ω
Attenuizomus – ♦
Thelyphonida (Uropygi) – Not recorded in Australia.

Schizomida The first member of order Schizomida from Australia (a troglobitic
one) was described by Harvey (1988), despite the fact that the order was recorded
from this continent as early as 1963. In his important paper in 1992, Harvey described
5 new genera (Draculoides, Apozomus, Bamazomus, Notozomus, and Julattenius)
and 24 new species of Schizomida. From the map (p. 79), it is clear that all these spe-
cies are known from the periphery of the continent: the far west, far north, and east-
ernmost parts of Australia. Later Harvey (2000) added two more genera Brignolizomus
and Attenuizomus, with three new species. New descriptions followed (Harvey (1992,
2000a, 2000b, 2001b, 2013g); Harvey and Humphreys (1995); Harvey et al. (2008).
Now (2017) in Australia are known 56 spp. of the genera Draculoides (6),
Apozomus (11), Bamazomus (5), Notozomus (17), Julattenius (2), Brignolizomus
(3), Ovozomus (1), Paradraculoides (3), and Attenuizomus (4) (endemic genera in
bold).
Araneae
Australia (continental) Following the table of Main (1981) in Australia are regis-
tered 46 families of spiders. Without Salticidae, they contained 276 indigenous gen-
era, of which 77 (27.9%) are endemic; including the Salticidae with 338 genera of
which 88 (26%) are endemic. The family Salticidae contained 62 genera, of which
11 are endemic. Other families with many genera are Araneidae (33, 6 end.),
Lycosidae (25, 2 end.), and Thomisidae (23, 5 end.).
Since 1981 many new records of spiders in Australia have changed these
numbers.
Now (2017) in Australia are registered 73 families. The families Gradungulidae,
Periegopidae, Holarchaeidae (Tasmania), Malkaridae-Pararchaeidae (also New
Caledonia), and Nicodamidae (also New Guinea) are shared only (or almost only)
with New Zealand; the families Hexathelidae (Hexathelinae), Migidae
(Calathotarsinidae), Malkaridae, and Amphinectidae are shared with southern South
America (Argentina and Chile) (Baehr 2003, b, c, 2004a, b; Baehr and Baehr 1987;
Baehr and Churchill 2003; Baehr and Jocqué 2000; Balogh 1980; Crews and Harvey
2011; Davies 1995a, b, 1998, 2003a, b; Dunn 1951; Framenau 2006a, b; Gardziñska
1996; Gardzinska and Zabka 2010; Gray 1973; Harms and Harvey 2009, 2013;
Harvey 1995; Hawkeswood 2003; Huber 2001; Jocqué 1993; Koch 1871–1881; Main
1969, 1976, 1981a, 1981b, 1987; Patoleta 2002; Patoleta and Zabka 1999; Platnick
1991, 2000, 2002, 2004; Platnick and Baehr 2006; Platnick and Forster 1989; Raven
(1976, 1978, 1979, 1980a, 1980b, 1981a, 1981b, 1985, 1994, 2009), Raven and
Stumkat (2005); Richardson and Zabka 2004; Richardson et al. 2006; Rix 2005,
2006; Rix and Harvey 2010, 2010b, 2011, 2012a, 2012b, 2012c; Rix et al. 2010;
Thorell 1881; Wanless 1988; Zabka 1987a, b, 1988, 1989, 1990a, b 1991a, b, 1992a,
b, 1993b, 1994, 1995b, 1999, 2000, 2001, 2002, 2003, 2006a, b, 2009, 2012, 2014).
Endemic genera of spiders in Australia (Tasmania generally excluded) are:

Fam. Hexathelidae
Hexathelinae – Atrax O. P-Cambridge, 1877 (3 spp.); Bymainiella Raven, 1978
(4 spp.); Hadronyche L. Koch, 1873 (31 spp., incl. 1 from Tasmania);
Paraembolides Raven, 1980 (8 spp.); Teranodes Raven, 1985 (2 spp.)
Fam. Dipluridae
Subfam. Euagrinae – Australothele Raven, 1984 (7 spp.); Carrai Raven, 1984 (1
sp.); Cethegus Thorell, 1881 (12 spp.); Namirea Raven, 1984 (7 spp.)
incertae sedis
Troglodiplura Main, 1969 (one sp.)
Fam. Anapidae – Chasmocephalon O. P.-Cambridge, 1889 (eight spp.); Maxanapis
Platnick et Forster, 1989 (nine spp.); Nortanapis Platnick et Forster, 1989 (one
sp.); Octanapis Platnick et Forster, 1989 (two spp.); Queenslanapis Platnick et
Forster, 1989 (one sp.); Risdonius Hickman, 1939 (three spp.); Spinanapis
Platnick et Forster, 1989 (nine spp.); Victanapis Platnick et Forster, 1989 (one
sp.)
Subfam. Micropholcommatinae: Micropholcomma Crosby et Bishop, 1927 (8 spp.,
with Tasmania)
Fam. Archaeidae – Austrarchaea Forster et Platnick, 1984 (27 spp.)
Fam. Malkaridae – Malkara Davies, 1980 (1 sp.), Perissopmeros Butler, 1932 (7
spp.); syn. Pararchaeidae: Anarchaea Rix, 2006 (4 spp.); Flavarchaea Rix,
2006 (7 spp.); Nanarchaea Rix, 2006 (2 spp.); Ozarchaea (16 spp.);
Westrarchaea Rix, 2006 (3 spp.)
Fam. Nemesiidae – Aname L. Koch, 1873; Chenistonia Hogg, 1901; Ixamatus
Simon, 1887; Kwonkan Main, 1983; Merredinia Main, 1983; Namea Raven,
1984; Pseudoteyl Main, 1985; Teyl Main, 1975; Teyloides Main, 1985; Xamiatus
Raven, 1981; Yilgarnia Main, 1986
Fam. Synotaxidae – Calcarsynotaxus Wunderlich, 1995 (two spp.); Microsynotaxus
Wunderlich, 2008 (two spp.); Paratupua Platnick, 1990 (one sp.)
Fam. Cyatholipidae – Forstera Koçak et Kemal, 2008 (one sp.); Matilda Forster,
1988 (one sp.); Teemenaarus Davies, 1978 (one sp.); Tekellatus Wunderlich,
1978 (one sp.)
Fam. Theridiidae – Magnopholcomma Wunderlich, 2008 (one sp.)
Fam Ctenidae – Bengalla Gray et Thompson, 2001 (one sp.), Janusia Gray, 1973
(one sp.)
Fam. Lycosidae – Anomalosa Roewer, 1960 (two spp.); Costacosa Framenau et
Leung, 2013 (two spp.); Hoggicosa Roewer, 1960 (ten spp.); Kangarosa
Framenau, 2010 (ten spp., one in Tasmania); Knoelle Framenau, 2006 (one sp.);
Mainosa Framenau, 2006 (one sp.); Tapetosa Framenau et al., 2009 (one sp.);
Tuberculosa Framenau et Yoo, 2006 (four spp.); Venator Hogg, 1900 (one sp.)
Fam. Arkyidae – Arkys Walck., 18370, Demendians Strand, 1929
Fam. Pholcidae – Wugigarra Huber, 2001 (20 spp.); Trichocyclus Simon, 1908 (23
spp.); Micromerys Bradley, 1877(7 spp.); Panjange Deeleman-Reinhold et
Deeleman, 1983 (6 spp.)
Fam. Oxyopidae – Pseudohostus Rainbow, 1915 (one sp.)
Fam. Pisauridae – Inola Davies, 1982 (three spp.)
Fam. Psechridae – Baiami Lehtinen, 1967 (9 spp.); Barahna Davies, 2003 (8 spp.);
Borrala Gray et Smith, 2004 (4 spp.); Carbinea Davies, 1999 (4 spp.); Corasoides
Butler, 1929 (1 spp.); Couranga Gray et Smith, 2008 (2 spp.); Elleguna Gray et
Smith, 2008 (2 spp.); Jamberoo Gray et Smith, 2008 (4 spp.); Kababina Davies,
1995 (9 spp.); Karriella Gray et Smith, 2008 (2 spp.); Malarina Davies et
Lambkin, 2000 (4 spp.); Malarina Davies et Lambkin, 2000 (4 spp.);
Procambridgea Forster et Wilton, 1973 (12 spp., one of them also in NZ – trans-
ported?); Tartarus Gray, 1973 (4 spp.); Therlinya Gray et Smith, 2002 (11 spp.);
Wabua Davies, 2000 (11 spp.)
Fam. Miturgidae (Zorinae) – Elassoctenus Simon, 1909 (one sp.); Hestimodema
Simon, 1909 (two spp.); Simonus Ritsema, 1881 (one sp.); Thasyraea L. Koch,
1878 (two spp.); Thasyraea L. Koch, 1878 (three spp.); Tuxoctenus Raven,
2008 (three spp.)
Fam. Zoropsidae – Birrana Raven et Stumkat, 2005 (one sp.); Huntia Gray et
Thompson, 2001 (two spp.); Kilyana Raven et Stumkat, 2005 (ten sp.); Krukt
Raven et Stumkat, 2005 (five spp.); Megateg Raven et Stumkat, 2005 (eight
spp.)
Fam. Amphinectidae – Austmusia Gray, 1983 (three spp.); Buyina Davies, 1998
(two spp.); Cunnawarra Davies, 1998 (two spp.); Jalkaraburra Davies, 1998
(one sp.); Keera Davies, 1998 (one sp.); Magua Davies, 1998 (one sp.); Penaoola
Davies, 1998 (two spp.); Quemusia Davies, 1998
Fam. Amaurobiidae – Bakala Davies, 1990 (1 sp.); Dardurus Davies, 1976 (6
spp.); Daviesa Koçak et Kemal, 2008 (2 spp.); Jamara Davies, 1995 (1 sp.);
Manjala Davies, 1990 (3 spp.); Midgee Davies, 1995 (9 spp.); Storenosoma
Hogg, 1900 (13 spp., 1 of them in Tasmania); Wabarra Davies, 1996 (2 spp.)
Fam. Desidae – Colcarteria Gray, 1992 (three spp.); Forsterina Lehtinen, 1967
(eight spp. in mainland Australia, one in New Caledonia); Lathyarcha Simon,
1908 (three spp.); Paramatachia Dalmas, 1918 (five spp., one of them shared
with Tasmania); Phryganoporus Simon, 1908 (five spp., one of them shared
with Tasmania); Pitonga Davies, 1984 (one sp.)
Fam. Dictynidae – Callevophthalmus Simon, 1906 (two spp., one shared with Lord
Howe Island)
Fam. Nicodamidae – Ambicodamus Harvey, 1995 (11 spp., three of them shared
with Tasmania); Durodamus Harvey, 1995 (3 spp.); Litodamus Harvey, 1995 (3
spp.); Nicodamus Simon, 1887 (2 spp.); Novodamus Harvey, 1995 (2 spp.);
Oncodamus Harvey, 1995 (2 spp.)
Fam. Sparassidae – Anchognatha Thorell, 1881 (1 sp.); Holconia Thorell, 1877 (9
spp.); Irileka Hirst, 1998 (1 sp.); Keilira Hirst, 1989 (2 spp.); Neosparassus
Hogg, 1903 (19 spp., 2 of them also in Tasmania); Yiinthi Davies, 1994 (8 spp.,
one of them shared with New Guinea); Zachria L. Koch, 1875 (2 spp.)
Fam. Selenopidae – Karaops Crews et Harvey, 2011 (37 spp.)

Fam. Zodariidae – Asteron Jocqué, 1991 (8 spp., 2 also in Tasmania); Cavasteron
Baehr et Jocqué, 2000 (12 spp.); Chilumena Jocqué, 1995 (2 spp.); Euasteron
Baehr, 2003 (17 spp.); Hetaerica Rainbow, 1916 (2 spp.); Holasteron Baehr,
2004 (16 spp.); Leptasteron Baehr et Jocqué, 2001 (2 spp.); Masasteron Baehr,
2004 (21 spp.); Minasteron Baehr et Jocqué, 2000 (3 spp.); Neostorena Rainbow,
1914 (7 spp.); Nostera Jocqué, 1991 (2 spp., 1 of them shared with Lord Howe
Island); Notasteron Baehr, 2005 (2 spp.); Phenasteron Baehr et Jocqué, 2001 (2
spp.); Pseudasteron Jocqué et Baehr, 2001(1 sp.); Spinasteron Baehr, 2003 (19
spp.); Storosa Jocqué, 1991 (2 spp.); Subasteron Baehr et Jocqué, 2001(one
sp.); Tropasteron Baehr, 2003 (22 spp.); Zillimata Jocqué, 1995 (one sp.)
Fam. Zoropsidae – Austrotengella Raven, 2012 (six spp.)
Fam. Miturgidae – Diaprograpta Simon, 1909 (five spp.); Eupograpta Raven, 2009
(two spp.); Mitzoruga Raven, 2009 (three spp.); Nuliodon Raven, 2009 (one sp.)
Fam. Ammoxenidae – Austrammo Platnick, 2002 (four spp.), Barrowammo
Platnick, 2002 (one sp.)
Fam. Gallienelliidae – Meedo Main, 1987 (13 spp.); Neato Platnick, 2002 (7 spp.);
Oreo Platnick, 2002 (5 spp.); Peeto Platnick, 2002 (1 sp.); Questo Platnick, 2002
(1 sp.)
Fam. Gnaphosidae – Battalus Karsch, 1878 (one sp.); Ceryerda Simon, 1909 (one
sp.); Encoptarthria Main, 1954 (five spp.); Epicharitus Rainbow, 1916 (one
sp.); Homoeothele Simon, 1908 (one sp.); Montebello Hogg, 1914 (one sp.);
Symphanodes Rainbow, 1916 (one sp.)
Fam. Lamponidae – Asadipus Simon, 1897 (20 spp.); Bigenditia Platnick, 2000 (2
spp.); Centroina Platnick, 2002 (11 spp.); Centrothele L. Koch, 1873 (10 spp.);
Centsymplia Platnick, 2000 (1 sp.); Graycassis Platnick, 2000 (10 spp.);
Lamponata Platnick, 2000 (2 spp.); Lamponega Platnick, 2000 (3 spp.);
Lamponicta Platnick, 2000 (1 sp.); Lamponina Strand, 1913 (6 spp.);
Lamponoides Platnick, 2000 (1 sp.); Lamponusa Platnick, 2000 (1 sp.); Longepi
Platnick, 2000 (8 spp.); Notsodipus Platnick, 2000 (18 spp.); Paralampona
Platnick, 2000 (8 spp.); Platylampona Platnick, 2004 (1 sp.); Pseudolampona
Platnick, 2000 (12 spp.); Queenvic Platnick, 2000 (4 spp.)
Fam. Prodidomidae – Cryptoerithus Rainbow, 1915 (19 spp.); Molycria Simon,
1887 (36 spp.); Myandra Simon, 1887 (4 spp., 1 of them shared with Tanzania);
Nomindra Platnick et Baehr, 2006 (16 spp.); Wesmaldra Platnick et Baehr, 2006
(14 spp.); Wydundra Platnick et Baehr, 2006 (45 spp. in Australia, 1 in Malaysia,
Moluccas)
Fam. Trochanteriidae – Bolathana Platnick, 2002 (2 spp.); Desognanops Platnick,
2008 (1 sp.); Desognaphosa Platnick, 2002 (26 spp., 1 on Solomon Islands);
Fissarena Henschel, Davies et Dickman, 1995 (9 spp.); Hemicloeina Simon,
1893 (9 spp., 1 in New Guinea); Longrita Platnick, 2002 (10 spp.); Morebilus
Platnick, 2002 (13 spp.); Platorish Platnick, 2002 (5 spp.); Rebilus Simon, 1880
(17 spp.); Tinytrema Platnick, 2002 (5 spp., 1 also in Tasmania); Trachycosmus
Simon, 1893 (4 spp.); Trachyspina Platnick, 2002 (8 spp.); Trachytrema Simon,
1909 (2 spp.)
Fam. Thomisidae – Corynethrix L. Koch, 1876 (one sp.); Isala L. Koch, 1876 (one
sp.); Poecilothomisus Simon, 1895 (one sp.); Saccodomus Rainbow, 1900 (one
sp.)
Fam. Salticidae – Abracadabrella Zabka, 1991 (3 spp.); Allococalodes Wanless,
1982 (3 spp.); Ananeon Richardson, 2013 (1 sp.); Allococalodes Wanless, 1982
(3 spp.); Aruana Strand, 1911 (2 spp., 1 from Aru Island); Astia L. Koch, 1879
(1 sp.); Astilodes Zabk, 2009 (1 sp.); Barraina Richardson, 2013 (1 sp.);
Coccorchestes Thorell, 1881 (39 spp. In New Guinea, 1 sp. in Queensland);
Damoetas Peckham et Peckham, 1886 (1 sp.); Frewena Richardson, 2013 (1
sp.); Furculattus Balogh, 1980 (1 sp.); Grayenulla Zabka, 1992 (7 sp.);
Huntiglennia Zabka et Gray, 2004 (1 sp.); Jacksonoides Wanless, 1988 (7 sp.);
Judalana Rix, 1999 (1 sp.); Leptathamas Balogh, 1980 (1 sp.); Maratus Karsch,
1878 (37 spp. in Australia, 1 sp. in China); Maddisonia Zabka, 2014 (3 sp.);
Megaloastia Zabka, 1995 (1 sp.); Paraharnochirus Szombathy, 1915 (2 sp.);
Parahelpis Gardzinska et Zabka, 2010 (2 sp.); Paraphilaeus Zabka, 2003 (1
sp.); Porius Thorell, 1892 (2 sp.); Pseudosynagelides Zabka, 1991 (6 sp.);
Pungalina Richardson, 2013 (1 sp.); Rhombonotus L. Koch, 1879 (1 sp.);
Simaethula Simon, 1902 (7 spp.); Sondra Wanless, 1988 (15 spp.); Tauala
Wanless, 1988 (7 spp. in Australia, 1 in Taiwan); Udvardya Prószyński, 1992 (1
sp.); Urogelides Zabka, 2009 (1 sp.); Viroqua Peckham et Peckham, 1901 (1 sp.)
Fam. Corinnidae – Leichhardteus Raven et Baehr, 2013 (eight spp.); Poecilipta
Simon, 1897 (two spp.)
Fam. Liocranidae – Liparochrysis Simon, 1909 (one sp.)
Opilioacarida
No species has been published, but there are indications for the presence of
Opilioacarida in Australia (Walter and Proctor 1998).
Acariformes
Beron (2008), Domrow (1992), Domrow and Lester (1985), Haliday (1998)
Trombidiformes
Suborder Prostigmata
Fam. Smarididae – Endemic genus Sphaerotarsus Womersley, 1936
Fam. Erythraeidae
Endemic genera:
Erythrellus Southcott, 1946; Erythrites Southcott, 1946; Erythroides Southcott,
1946; Rainbowia Southcott, 1961; Pussardia Southcott, 1961; Mypongia
Southcott, 1961; Wartookia Southcott, 1961
Sarcoptiformes
Suborder Oribatida
According to the Catalogue of Australian Oribatida (Colloff and Halliday, 1998), in
Australia are registered 340 named species (256 endemic) and 237 unnamed spe-
cies or a total of 577 species. There are no endemic superfamilies indigenous to
Australia.

Order Ixodida (Domrow 1987, Haliday 1998, Roberts 1970)
Fam. Argasidae – Six spp. of Argas Latreille and one of Otobius Banks
Fam. Ixodidae – Sixty-eight spp., no endemic genera, some endemic species
Holothyrida Known are two species of the genus Allothyrus (fam. Allothiridae,
confined to Australia and New Zealand) (Domrow, 1955; Van der Hammen, 1961,
1983; Womersley, 1935).
8.20 Tasmania
8.20.1 G
and Paleogeography
Tasmania is an Australian island and state. It is 240 km south of the continent, sepa-
rated by Bass Strait. The state includes the island of Tasmania and some smaller
islands (state area 68,401 km2), of which the main island covers 62,409 km2. The
highest point is Mount Ossa (1614 m.).
Situated at 42oS, Tasmania has a cool temperate climate with four distinct
seasons.
The island was joined to the mainland of Australia until the end of the last glacial
period approximately 10,000 years ago. Much of Tasmania is still densely forested,
with the Southwest National Park and neighboring areas holding some of the last
temperate rain forests in the Southern Hemisphere. The Tarkine, located in the
island’s far North West, is the largest temperate rainforest area in Australia covering
approximately 3800 square kilometers.
According to Darlington (1960), the fauna of Tasmania is depauperate, both in
diversity of basic stocks and in total number of species. Many groups, both verte-
brate (Varanus monitor lizards, pythons, Typhlopidae, and many others) and inver-
tebrates (Uropygi, Amblypygi, Cyphophthalmi, Dyspnoi), are missing in the island.
Still, in Tasmania are (or were) preserved animals extinct in mainland Australia.
“Tasmania has long been regarded as a biological treasure trove, supporting an
astounding range of species found nowhere else, many of which have survived the
perturbations of climate change during the Pleistocene” (Harvey 1998).
Arachnogeography
No records for the order Palpigradi, Ricinulei, Solifugae, Amblypygi, Uropygi,
Schizomida, Opilioacarida, and Holothyrida and the suborders Paleamblypygi,
Cyphophthalmi, and Dyspnoi. Almost no scorpions, many endemic genera of
Triaenonychidae (Laniatores), 2 endemic subfamilies, and 29 endemic genera of
spiders, many endemic genera of Acari (Erythraeidae and others).
Scorpiones From Tasmania has been recorded only Cercophonius squama
(Bothriuridae), shared with mainland Australia. Also on Flinders I. and King I.
(Koch, 1977).
8.20 Tasmania 765
Pseudoscorpiones
In Tasmania are registered Pseudoscorpiones from several families (Chthoniidae,
Pseudogarypidae, Pseudotyrannochthoniidae, Garypidae, Syarinidae, Garypinidae,
etc.) (Benedict and Malcolm 1978; Dartnall 1970; Harvey 1998; Morris 1948).
Endemic genera for Tasmania are Neopseudogarypus Morris, 1948
(Pseudogarypidae), and Oreolpium Harvey et Štáhlavský (Garypinidae).
Opiliones. Ref.: Hickman (1957, 1958), Hogg (1909), Hunt (1990, 1995),
Roewer (1914)
In Tasmania have been registered the following Opiliones:
Cyphophthalmi – Not recorded in Tasmania
Eupnoi – Fam. Neopilionidae (incl. Monoscutidae) (Spinicrus Forster, 1949)
Dyspnoi – Fam. Acropsopilionidae: Austropsopilio Forster, 1955 (Tasmanopilio
Hickman, 1957)
In Tasmania are registered Laniatores of 27 endemic genera of the family
Triaenonychidae.
Endemic genera are:
Allonuncia Hickman, 1958 (one sp.); Ankylonuncia Hickman, 1958 (three spp.);
Bryonuncia Hickman, 1958 (one sp.); Calliuncus Roewer, 1931 (five spp.);
Cluniella Forster, 1955 (three spp.); Hickmanoxyomma Hunt, 1990 (seven
spp.); Leionuncia Hickman, 1958 (one sp.); Notonuncia Hickman, 1958
(three spp.); Nucina Hickman, 1958 (two spp.); Nuncioides Hickman, 1958
(two spp.); Odontonuncia Hickman, 1958 (one sp.); Parattahia Roewer, 1914
(one sp.); Rhynchobunus Hickman, 1958 (one sp.); Stylonuncia Hickman,
1958 (one sp.); Tasmanonyx Hickman, 1958 (one sp.); Pyenganella Hickman,
1958 (one sp.); Tasmanonuncia Hickman, 1958 (one sp.); Thelbunus Hickman,
1958 (one sp.); Allobunus Hickman, 1958 (one sp.); Chilobunus Hickman,
1958 (one sp.); Chrestobunus Roewer, 1914 (three spp.); Eubunus Hickman,
1958 (one sp.); Glyptobunus Roewer, 1914 (two spp.); Mestonia Hickman,
1958 (one sp.); Miobunus Roewer, 1915 (six spp., Tasmania); Phanerobunus
Roewer, 1915 (four spp.); Phoxobunus Hickman, 1958 (two spp.)
Araneae In Tasmania have been recorded 2 endemic subfamilies (Plesiothelinae
and Hickmanniinae) and 29 endemic genera of spiders (Musgrave 1947; Gertsch
1958; Hickman 1957, 1958, 1969; Platnick and Forster 1989; Raven 1978). Endemic
genera and subfamilies:
Fam. Hexathelidae
Plesiothelinae
Plesiothele Raven, 1978 – One sp.
Fam. Anapidae
Acrobleps Hickman, 1979 (one sp.); Epigastrina Rix et Harvey, 2010 (three
spp.); Hickmanapis Platnick et Forster, 1989 (two spp.); Olgania Hickman,
1979 (five spp.); Tasmanapis Platnick et Forster, 1989 (one sp.)
Fam. Austrochilidae – Chile, Argentina, Tasmania (nine spp.)
Hickmaniinae (as family Hickmaniidae)
Hickmania Gertsch, 1958 – One sp.: Hickmania troglodytes (Higgins et Petterd,
1883), Cave
Fam. Orsolobidae
Cornifalx Hickman, 1979 (one sp.); Hickmanolobus Forster et Platnick, 1985;
Olgania Hickman, 1979; Carathea Moran, 1986; Tupua Platnick in
Forster, Platnick et Coddington, 1990; Cicirra Simon, 1876; Namandia Lehtinen,
1967; Ommatauxesis Simon, 1903; Toxops Hickman, 1940
Fam. Malkaridae – Carathea Moran, 1986 (two spp.)
Fam. Anapidae (subfam. Micropholcommatinae) – Epigastrina Rix et Harvey, 2010
(three spp.)
Fam. Synotaxidae – Tupua Platnick, 1990 (four spp.)
Fam. Lycosidae – Tasmanicosa Roewer, 1959 (one sp.)
Fam. Psechridae – Tjurunga Lehtinen, 1967 (one sp.)
Fam. Miturgidae (Zorinae) – Odomasta Simon, 1909 (one sp.)
Fam. Araneidae – Collina Urquhart, 1891 (one sp.)
Fam. Amphinectidae
Tanganoides Davies, 2005 (six spp., incl. one in Victoria); Tasmabrochus
Davies, 2002 (five spp.); Tasmarubrius Davies, 1998 (five spp.); Teeatta
Davies, 2005 (three spp.)
Fam. Amaurobiidae – Only endemic species
Fam. Desidae
Cicirra Simon, 1886 (one sp.); Namandia Lehtinen, 1967 (one sp.);
Ommatauxesis Simon, 1903 (one sp.); Toxops Hickman, 1940 (one sp.)
Fam. Hahniidae – Neoaviola Butler, 1929 (one sp.)
Fam. Stiphidiidae – Tjurunga Lehtinen, 1967 (one sp.)
Notes on several families:
Actinopodidae Three genera, two in South America and one (Missulena
Walckenaer, 1805) sheared between Australia (10 spp.) and Chile (Missulena tus-
sulena Goloboff, 1994). Greater Nothogea?
Austrochilidae Small family of nine spp., illustrating well the concept of Notogea
including the southern end of South America. Two subfamilies: Austrochilinae with
8.21 Parasitiformes (Some Ixodida and Mesostigmata): Australia and New Zealand 767
two genera from Chile and Argentina and Hickmaniinae with one species from a
cave in Tasmania. The same could be said concerning the fam. Malkaridae: one
genus in Tasmania, two in mainland Australia, and one (Chilenodes Platnick et
Forster, 1987) in Chile and Argentina.
Hexathelidae Atracinae (three genera with three spp., endemic to Australia),

Plesiothelinae is endemic to Tasmania and many others in Europe, Asia, and Africa,
Macrothelinae has an isolated end. genus in New Zealand (Porrhothele Simon,
1892), and Hexathelinae is also a good example for a bigger Notogea – three genera
in Australia, two in Chile and Argentina, and one in New Zealand.
Nicodamidae Nine genera with 29 spp. Four genera endemic to Australian conti-
nent, one end. to Tasmania, one (Dimidamus Harvey, 1995) distributed in New
Guinea and Australia, and one (Ambicodamus Harvey, 1995) shared between
Australia and Tasmania.
Malkaridae Four genera, two (Malkara and Perissopmeros) endemic to mainland

Australia, one (Carathea) endemic to Tasmania, and one (Chilenodes Platnick and
Forster, 1987) known from Chile and Argentina. Again “Greater Nothogea”! To this
family was attached as junior synonym Pararchaeidae from Australia, Tasmania,
and New Zealand.
Subfam. Micropholcommatinae The distribution of the subfamily of Anapidae

comprises mainland Australia, Tasmania (22 spp.), Auckland Islands, New Guinea,
Campbell Islands, New Zealand, New Caledonia, Lord Howe Island, Chile, and
Brazil. Thanks to the work of Rix and Harvey (2010), now we have several exam-
ples of “traditional” Gondwanan distribution: Gigiella Rix et Harvey, 2010 (one sp.
in mainland Australia and Tasmania, one sp. in Chile); Eperiella Rix et Harvey,
2010 (one sp. in Tasmania, one sp. in Chile); and Normplatnicka Rix et Harvey,
2010 (two spp. in m. Australia, one sp. in Chile). They are also good examples for a
bigger Notogea.
Synotaxidae Three subfamilies: Pahorinae (endemic to New Zealand),

Physogleninae (Australia, Tasmania, New Zealand, and Chile), and Synotaxinae
(Australia, Chile, South America).
8.21 P
arasitiformes (Some Ixodida and Mesostigmata):
Australia and New Zealand
Ixodida New Zealand has 11 named species of ticks, four of which are endemic:
Aponomma sphenodonti (Dumbleton, 1953) (on tuatara); Ixodes anatis Chilton,
1904 (on kiwi and Anatidae); I. jacksoni Hoogstraal, 1967 (from nest of Stictocarbo-
Phalacrocorax punctatus); and Carios quadridentatus Heath, 2012 (from the

endemic bat Mystacina tuberculata). Six are known also from Australia (Dumbleton
1953, 1963; Heath 1977, 2012; Heath et al. 2011; Hoogstraal 1967; Spain and
Luxton 1971).
Acariformes (Some Examples)

Sarcoptiformes
Suborder Oribatida
Ref.: Australia. Colloff and Haliday (1998); New Zealand. Hammer (1966,
1967, 1968), Colloff and Cameron (2014).
According to the catalogue of Australian Oribatida (Colloff and Halliday 1998),
in Australia are registered 340 named species (256 endemic) and 237 unnamed spe-
cies or a total of 577 species. There are no superfamilies indigenous to Australia.
Thanks to the research of M. Hammer, from New Zealand have been listed at least
50 families of Oribatida (Spain and Luxton 1971).
Trombidiformes (Some examples)
Ref.: Beron (2008a), Domrow (1978), Domrow and Lester (1985), Haliday
(1998), Southcott (1961), Womersley (1936), Southcott (1966), Hirst (1926),
Southcott (1988), Zhang (2000)
Prostigmata Erythraeoidea
Endemic Genera in Australia
Fam. Smarididae – Sphaerotarsus Womersley, 1936
Fam. Erythraeidae
Erythrellus Southcott, 1946; Erythrites Southcott, 1946; Erythroides Southcott,
1946; Rainbowia Southcott, 1961; Pussardia Southcott, 1961; Mypongia
Southcott, 1961; Wartookia Southcott, 1961
End. species in New Caledonia: Charletonia rageaui Southcott, 1966
Endemic genera in New Zealand: Neosmaris Hirst, 1926; Taranakia Southcott,
1988; Ramsayella Zhang, 2000
8.22 New Zealand
8.22.1 G
and Paleogeography
The archipelago, called by the Europeans in 1645 New Zealand, is one of the amaz-
ing places on Earth, when discussing zoogeography. It is composed of two bigger
islands (North and South Islands), separated by the Cook Straight (22 km. wide),
one less big (Stewart), and many smaller islands with a total surface of 268 thousand
km2. New Zealand is 1500 km. far from Australia. The relief is very varied with
highest point Mount Cook, or Aoraki (3754 m.), and 17 other summits higher than
3000 m. The biota is very altered by humans, mostly by the white settlers and the
8.22 New Zealand 769
animals they introduced, one way or another. For the invertebrates, most important
was the change in the environment because of the land use, deforestation, etc.
To understand the history of the biota of New Caledonia, New Zealand, and Lord
Howe Island, we have to follow up the hypothetical development of Zealandia, the
New Zealand microcontinent, having existed until some 23 million years ago as a
result of the breakaway from Australia about 60–85 million years ago and from
Antarctic 85–130 Ma ago (Craw 1988, 1989; Fleming 1949, 1962, 1963a, 1963b,
1967, 1970, 1975; Grehan 1989; Griffiths 1975; Page 1989; Sharma and Wheeler
2013; Udvardy 1975).
According to Fleming (1975), the isolation of New Zealand becomes effective
for the land vertebrates (and maybe invertebrates) before the end of Cretaceous.
Cracraft (1973) thinks that “Australia separated from Antarctica in the Eocene and
with spreading rates approximately twice as fast as those for New Zealand and this
differential movement between Australia-New Zealand and Antarctica formed the
Tasman Sea” (see also Griffiths and Varne, 1972). In any case, New Zealand was
detached from Antarctida more than 80 Ma ago (Cracraft, 1973), together with the
ancestors not only of the moa, kiwi, tuatara, and other well-known cases also with
the ancestors of many insects and other invertebrates. In Cracraft (1974), we read
that “The next portion of Gondwanaland to separate was New Zealand, which was
adjacent to West Antarctica and which began drifting in the late Cretaceous at about
40–45 Ma ago.”
Arachnogeography It would be hardly possible to assess the entire endemism and
richness of New Zealand arachnofauna some 40 years ago. Forster (1949c, 1973,
1975) analyzed the particularities of this amazing fauna, but since (mostly with the
efforts of Forster himself and his collaborators), new data have been accumulated.
The orders Scorpiones, Ricinulei, Thelyphonida (Uropygi), Amblypygi, Schizomida,
and Opilioacarida are completely missing from New Zealand and the surrounding
islands (naturally). Represented are Pseudoscorpiones, with six endemic genera,
Opiliones (amazingly rich in endemics and with elements in common with faraway
countries, one endemic family), and Araneae (huge number of spiders in an island
country). Among spiders there are 93 endemic genera and 93% endemic species.
Pseudoscorpiones The pseudoscorpion fauna of New Zealand and the Subantarctic

islands in the area consists now of 70 species of 28 genera and 9 families:
Chthoniidae, Syarinidae, Garypidae, Garypinidae, Olpiidae, Cheiridiidae,
Cheliferidae, Chernetidae, Withiidae, including some species and 6 genera endemic.
One (Philomaoria) is shared with Lord Howe Island, another (Synsphyronus) with
Tasmania, and 11 with Australia (Beier 1966, 1967, 1969, 1973, 1976; Chamberlin
1925; Harvey 2009, 2013f).
Endemic genera:
Fam. Chthoniidae
Maorichthonius Chamberlin, 1925 – New Zealand (one sp.)
Sathrochthoniella Beier, 1967 – New Zealand (one sp.)
Tyrannochthoniella Beier, 1966 – New Zealand (one sp.)

Fam. Olpiidae
Nelsoninus Beier, 1967 – New Zealand (South Island) (one sp.)
Fam. Chernetidae
Apatochernes Beier, 1948 – New Zealand (Campbell Islands, Snares Islands,
Auckland Islands, Chatham Islands, Norfolk Island (17 spp.)
Heterochernes Beier, 1966 – New Zealand (one sp.)
Opiliones Ref.: Boyer and Giribet (2007, 2009), Crawford (1992), Forster (1947,
1948, 1952, 1954, 1975, 16 papers from 1943 to 1975), Groh and Giribet (2015),
Hirst (1925), Roewer (1931), Taylor (2004, 2009, 2011, 2013).
In New Zealand (s. str.) live 27 genera of Opiliones, out of them 24 endemic
(Forster, 1975, actualized), from the families Pettalidae, Acropsopilionidae,
Neopilionidae (incl. Monoscutidae), Synthetonychiidae, and Triaenonychidae.
Cyphophthalmi
In New Zealand have been recorded 23 spp. of the fam. Pettalidae:
Rakaia Hirst, 1925 – Twelve spp. (endemic genus)
Aoraki Boyer et Giribet, 2007 – Eight spp. (endemic genus)
Neopurcellia Forster, 1948 – South Island (three spp.) (endemic genus)
Eupnoi
Fam. Neopilionidae – Megalopsalidinae: Megalopsalis Forster, Forsteropsalis
Taylor (nine spp.), Mangatangi Taylor (one sp.), Pantopsalis Simon (nine sp.)
Monoscutinae (endemic subfamily): Acihasta Forster, Monoscutum Forster,
Templar Taylor (New Zealand, Auckland, Snares, Campbell Islands)
Dyspnoi: Fam. Acropsopilionidae – Acropsopilio Silvestri (one sp.)
Laniatores
In New Zealand are registered 68 spp. of Laniatores of 15 genera and 2
families.
Endemic supraspecific taxa:
Fam. Synthetonychiidae – Endemic family
Synthetonychia Forster, 1954 – Fourteen spp.
Hedwiga Roewer, 1931 (1 sp.); Hendea Roewer, 1931 (14 spp.); Hendeola
Forster, 1954 (2 spp.); Metanuncia Roewer, 1914 (Stewart Island) (1 sp.);
Neonuncia Roewer, 1914 (5 spp.); Prasma Roewer, 1931 (3 spp.); Prasmiola
Forster, 1954 (one sp.); Psalenoba Roewer, 1931 (one sp.); Triregia Forster,
1948 (3 spp.); Algidia Hogg, 1920 (8 spp.); Cenefia Roewer, 1931 (4 spp.);
Muscicola Forster, 1954 (1 sp.); Pristobunus Roewer, 1931 (2 spp.)
Subfam. Soerensenellinae – Endemic subfamily (eight spp.)
Karamea Forster, 1954 (four spp.); Soerensenella Pocock, 1903 – four species
Araneae Recently an overview of New Zealand spiders has been published by
Paquin, Vink, and Dupérré (2010). Best characterized is the spider fauna of New
Zealand by the book review of Duffey (2010, Newsl. Br. Arachnol. Soc., 119):
“Britain and New Zealand are comparable in area but the former has 658 species
while the latter has 1126 described and another 536 awaiting description, making a
present total of about 1662 species in 236 genera and 57 families. As new species
are still being found the authors think the true total could be about 2000. One can’t
help feeling this is a conservative estimate because even the well-worked fauna in
Britain, which recorded a total of 584 in 1958, has since added another 74 species.
The best known New Zealand arachnologist, R.R. Forster, though the New Zealand
total could be as much as 2500 species. Whether 2000 or 2500, the fascinating ques-
tion is why New Zealand has such a rich fauna when no country in Europe reaches
even the lower figure. France, one of the largest, is two and half times the area [of
NZ], but has a total of 1569 sp. (Le Peru 2007). The family Linyphiidae is the larg-
est in New Zealand but only 12% of the total and they are all in the subfamily
Linyphiinae. Endemic Erigoninae apparently do not exist as all known species are
introduced. In Britain about 40% of the spider fauna are Linyphiidae, of which over
70% are Erigoninae.”
The endemism of New Zealand spiders is amazing: 93% of the fauna consists of
endemic species.
From New Zealand (including the nearby islands Auckland, Campbell, Snares,
etc.) have been recorded 2 endemic families (Huttoniidae and Megadictynidae),
1 subfamily (Pahorinae), and 93 endemic genera, and at least 93% of the species
of spiders are endemic (Simon 1892; Blest 1979; Forster, 1955, 1959, 1964, 1970,
1975; Forster and Blest 1979; Forster and Platnick 1977, 1984; Forster and Wilton
1968, 1973; Parrott 1942; Platnick and Forster 1989; Rix 2006; Vink and Dupérré
2010; Zabka et al. 2002):
Fam. Hexathelidae
Hexathele Ausserer, 1871 (20 spp.); Porrhothele Simon, 1892 (5 spp.)
Fam. Anapidae
Novanapis Platnick et Forster, 1989 (one sp.); Paranapis Platnick et Forster,
1989 (two spp.); Zealanapis Platnick et Forster, 1989 (ten sp.)
Fam. Pararchaeidae – Forstrarchaea Rix, 2006 (one sp.); Pararchaea Forster, 1955
(one sp.)
Fam. Idiopidae – Cantuaria Hogg, 1902 (42 spp.)
Fam. Gradungulidae
Gradungula Forster, 1955; Pianoa Forster, 1987; Spelungula Forster, 1987 (one
sp.)
Fam. Huttoniidae: Huttonia Pickard-Cambridge, 1880
Fam. Orsolobidae: Anopsolobus Forster et Platnick, 1985; Ascuta Forster, 1956;

Bealeyia Forster et Platnick, 1985; Dugdalea Forster et Platnick, 1985; Duripelta
Forster, 1956; Maoriata Forster et Platnick, 1985; Orongia Forster et Platnick,
1985; Paralobus Forster et Platnick, 1985; Pounamuella Forster et Platnick,
1985; Subantarctia Forster, 1955; Tangata Forster et Platnick, 1985; Tautukua
Forster et Platnick, 1985; Turretia Forster et Platnick, 1985; Waiporia Forster et
Platnick, 1985; Waipoua Forster et Platnick, 1985; Wiltonia Forster et Platnick,
1985
Fam. Mecysmauchenidae (Aotearoa Forster et Platnick, 1974; Zearchaea Wilton,
1946
Fam. Anapidae (Micropholcommatinae): Pua Forster, 1959; Tinytrella Rix et
Harvey, 2010; Taliniella Rix et Harvey, 2010
Fam. Cyatholippidae: Hanea Forster, 1988; Tekella Urquhart, 1894; Tekelloides
Forster, 1988
Fam. Synotaxidae: Nomaua Forster, 1990; Pahora Forster, 1990; Pahoroides
Forster, 1990; Runga Forster, 1990; Meringa Forster, 1990; Mangua Forster,
1990; Zeatupua Fitzgerald et Sirvid, 2009
Fam. Agelenidae: Ahua Forster et Wilton, 1973; Huka Forster et Wilton, 1973;
Mahura Forster et Wilton, 1973; Neoramia Forster et Wilton, 1973; Neorepukia
Forster et Wilton, 1973; Oramia Forster et Wilton, 1973 (also in Lord Howe
Island); Oramiella Forster et Wilton, 1973; Orepukia Forster et Wilton, 1973;
Paramyro Forster et Wilton, 1973; Porotaka Forster et Wilton, 1973; Tararua
Forster et Wilton, 1973; Tuapoka Forster et Wilton, 1973
Fam. Tetragnathidae: Eryciniolia Strand, 1912 (one sp.); Nanometa Simon, 1908
(one sp.); Pinkfloydia Dimitrov et Hormiga, 2011 (one sp.)
Fam. Lycosidae: Notocosa Vink, 2002 (one sp.)
Fam. Psechridae – [Borrala Gray et Smith, 2004 (31 spp., 1 in New Caledonia)];
Nanocambridgea Forster et Wilton, 1973 (1 sp.); Pillara Gray et Smith, 2004 (4
spp.)
Fam Amaurobiidae – Auhunga Forster et Wilton, 1973 (1 sp.); Maloides Forster et
Wilton, 1989 (1 sp.); Muritaia Forster et Wilton, 1973 (5 spp.); Otira Forster et
Wilton, 1973 (6 spp.); Pakeha Forster et Wilton, 1973 (18 spp.); Paravoca
Forster et Wilton, 1973 (2 spp.); Poaka Forster et Wilton, 1973 (1 sp.); Waitetola
Forster et Wilton, 1973 (1 sp.)
Fam. Amphinectidae:
Akatorea Forster et Wilton, 1973 (2 spp.); Amphinecta Simon, 1898 (11 sp.);
Aorangia Forster et Wilton, 1973 (16 sp.); Dunstanoides Forster et Wilton, 1989
(9 spp.); Holomamoea Forster et Wilton, 1973 (1 sp.); Huara Forster, 1964 (12
spp.); Makora Forster et Wilton, 1973 (5 spp.); Mamoea Forster et Wilton, 1973
(19 spp.); Maniho Marples, 1959 (10 spp.); Marplesia Lehtinen, 1967 (2 spp.);
Neolana Forster et Wilton, 1973 (3 spp.); Neororea Forster et Wilton, 1973 (2
spp.); Oparara Forster et Wilton, 1973 (2 spp.); Paramamoea Forster et Wilton,
1973 (10 spp.); Rangitata Forster et Wilton, 1973 (1 sp.); Reinga Forster et
Wilton, 1973 (5 spp.); Rorea Forster et Wilton, 1973 (2 spp.); Waterea Forster et
Wilton, 1973 (1 sp.)
Fam. Desidae: Gasparia Marples, 1956; Gohia Dalmas, 1917; Goyenia Forster et
Wilton, 1970; Hapona Forster, 1970; Helsonia Forster, 1970; Hulua Forster et
Wilton, 1973; Lamina Forster, 1970; Mangareia Forster, 1970; Matachia
Dalmas, 1917; Mesudus Özdikmen, 2007; Neomyro Forster et Wilton, 1973;
Notomatachia Forster, 1970; Nuisiana Forster et Wilton, 1973; Otagoa Forster,
1970; Panoa Forster, 1970; Rapua Forster, 1970; Toxopsoides Forster et Wilton,
1973; Tuakana Forster, 1970
Fam. Dictynidae: Arangina Lehtinen, 1967; Paradictyna Forster, 1970; Viridictyna
Forster, 1970
Fam. Hahniidae: Kapanga Forster, 1970; Porioides Forster, 1989; Rinawa Forster,
1970
Fam. Megadictynidae: Forstertyna Harvey, 1995; Megadictyna Dahl, 1906
Fam. Zodariidae: Forsterella Jocqué, 1991
Fam. Zoropsidae: Haurokoa Forster et Wilson, 1973
Fam. Cycloctenidae: Plectophanes Bryant, 1935 (5 spp.); Toxopsiella Forster, 1964
(12 spp.); Uzakia Koçak et Kemal, 2008 (1 sp.)
Fam. Miturgidae: Pacificana Hogg, 1904; Zealoctenus Forster et Wilton, 1973
Fam. Gnaphosidae: Kaitawa Forster, 1979; Matua Forster, 1979; Nauhea Forster,
1979; Notiodrassus Bruyant, 1935; Zelanda Özdikmen, 2009
Fam. Thomisidae: Cymbachina Bryant, 1933 (one sp.)
Fam. Salticidae: Hinewaia Zabka et Pollard, 2002 (one sp.)
Acariformes
Ref.: Beron (2008a), Clark (2014), Hammer (1966, 1968), Spain and Luxton (1971)
Trombidiformes Prostigmata
Fam. Erythraeidae
Endemic genera: Neosmaris Hirst, 1926; Ramsayella Zhang, 2000; Taranakia
Southcott, 1988; Pukakia Clark, 2014
Parasitiformes
Ixodida
New Zealand has nine named species of ticks, three of which are endemic:
Aponomma sphenodonti (Dumbleton, 1953) (on tuatara); Ixodes anatis Chilton,
1904 (on kiwi and Anatidae); and I. jacksoni Hoogstraal, 1967 (from nest of
Stictocarbo-Phalacrocorax punctatus). Six are known also from Australia
(Dumbleton 1953, 1963; Heath 1977; Hoogstraal 1967; Spain and Luxton 1971).
One unnamed Argasidae is known from the endemic bat Mystacina tuberculata.
Holothyrida
Recorded (loc.?) is Allothyrus (?) australasiae (Womersley, 1935) known from
Australia.
8.23 New Caledonia
8.23.1 G
and Paleogeography
New Caledonia is located in the subregion of Melanesia in the South West Pacific.
It comprises the main island (Grande Terre), the Loyalty Islands, and several smaller
islands. It has a land area of 18,575.5 square kilometers.
The Grande Terre is by far the largest of the islands, with an area of 16,372 square
kilometers, and is elongated northwest-southeast, 350 km in length and 50–70 km
wide. A mountain range runs the length of the island, with five peaks over 1500 m.
The highest point is Mont Panié at 1628 m elevation.
New Caledonia is one of the northernmost parts of a (93%) submerged continent
called Zealandia. It sank after rifting away from Australia 60–85 million years ago
(mya) and from Antarctica between 130 and 85 mya. New Caledonia itself is sepa-
rated from Australia in the Late Cretaceous (65–66 Ma ago), and subsequently drifted
in a north-eastern direction, reaching its present position about 50 Ma ago. According
to Neall and Trewick (2008), New Caledonia separated from New Zealand 83 Ma ago.
There is also a hypothesis that the islands emerged 37 Ma ago (Oligocene) and that its
biota started developing by this time. Some geologists insist that New Caledonia has
been entirely submerged several times and repopulated after that. However, biologists
accept that parts of the land remained above water as refugia for the archaic animals
and plants. Many species from the Gondwanan flora in the Late Cretaceous and early
Tertiary probably survived in the temperate climate of New Caledonia and died out in
Australia during its strong aridisation. This opinion is contested too.
The detailed analysis of Grandcolas et al. (2008) concerning the formation of the
New Caledonian biota comes to the conclusion that it is not a continental, but rather
an oceanic island, having its biota formed not earlier than 37 Ma ago during the
Oligocene (confirmed by molecular research).
New Caledonia is extremely interesting from biogeographical point of view, and
different problems are raised. As a matter of fact, it is an archipelago of one big and
very varied island, the Loyalty Islands, the Pine Island, and many other islets and
reefs. The polulation is 258 000. The isolation of New Caledonia dates at least from
the Miocene (perhaps from the Oligocene) and thus have been preserved many relict
animal and plant, and also neoendemics have been formed. After the arrival of the
first settlers (the Canaques) ca. 3200–3300 years ago, many endemic animals disap-
peared, and the Europeans (since 1853 New Caledonia is a French territory) contrib-
uted to it mostly by mining and agrarian activities, forest destruction, planting foreign
trees, and others. New Caledonia is 1300 km. far from Australia, 1500 km. from New
Zealand, and 1200 km. from Fiji (Darlington 1957; Heads 2008a, 2008b, 2010; Lillie
and Brothers 1970; Neall and Trewick 2008; Paramonov 1958, 1960; Sarasin 1925).
Arachnogeography There are several hypotheses about the origin of the arachno-
fauna (and the remaining fauna) of the Big Island, far away from any mainland.
Besides the animals, brought on the “Three Ws” (wind, waves, and wings), the
fauna is either quite recent (after the total submersion of the island) or contains
8.23 New Caledonia 775
remains from much older fauna (Grandcolas et al., 2008). The presence, at least, of
the amazing relict family of (cave dwelling) opilions Troglosironidae speaks rather
for at least some elements surviving from older fauna. Besides, on the islands, there
are some other orders of Arachnida: Scorpiones, Amblypygi, Schizomida, and
Pseudoscorpiones, endemic genera of Laniatores, many endemic spiders, and most
interesting Holothyrida – one genus endemic and one shared with Lord Howe
Island.
Scorpiones Three species (two endemic and one widespread) are recorded
(Kraepelin 1914; Simon 1877; Vachon 1976).
Fam. Buthidae
Isometrus (Reddyanus) heimi Vachon
Fam. Hormuridae
Liocheles australasiae (Fabricius)
L. neocaledonicus Simon
Amblypygi Charinus neocaledonicus Simon, 1895 (Charinidae), endemic spe-
cies of a widespread genus, is recorded from New Caledonia. Other members of the
same genus (all of them known from caves) are three subspecies of Ch. australianus
and Charinus pecki Weigoldt, 2006.
Schizomida Only Hubbardiidae indet. are mentioned from New Caledonia

(Reddell and Cokendolpher 1995).
Pseudoscorpiones In New Caledonia and the Loyalty Islands are known 14 spe-
cies of Pseudoscorpiones from 10 genera of 7 families: Chthoniidae,
Tridenchthoniidae, Parahyidae, Syarinidae, Garypinidae, Atemnidae, and
Chernetidae (Beier (1940, 1964, 1966, 1966d, 1968, 1976, 1979; Simon 1880).
Among them are the endemic species Hebridochernes caledonicus, H. gressitti, H.
maximus, Nesidiochernes caledonicus, Paraldabrinus (end. genus) novaecale-
doniae, Amblyolpium ruficeps, Ideobisium antipodum, Anaulacodithella novacale-
donica, A. reticulata, Sathrochthonius kaltenbachi, Tyrannochthonius troglophilus,
and T. zonatus – 12 spp. or 86% endemism.
Opiliones
The most interesting is the endemic family of Cyphophthalmi Troglosironidae
with 1 genus Troglosiro Juberthie, 1979, and 13 spp. (Shear 1993; Sharma and
Giribet 2009).
Dyspnoi - Missing
Laniatores
From New Caledonia and the Loyalty Islands are known Laniatores from the
families Triaenonychidae and Zalmoxidae (eight endemic species of genus

Zalmoxis) (Simon 1881; Roewer 1912, 1914, 1949; Goodnight and Goodnight
1948; Sharma and Giribet 2012).
Endemic Laniatores in New Caledonia are the following genera:
Diaenobunus Roewer, 1914; Triconobunus Roewer, 1914
Santobius Roewer, 1949 (Mesoceras Soerensen, 1886; preoccup., Mesoceratula
Roewer, 1949) – Vanuatu, Fiji
The presence of fam. Zalmoxidae has been explained by Sharma and Giribet
(2012): “The family Zalmoxidae, similar to the Pacific iguanas, constitutes the
unusual case of a lineage of Neotropical origin that colonized the Indo-Pacific likely
by ancient transoceanic dispersal during the Late Cretaceous.”
Araneae In New Caledonia have been recorded spiders of al least 22 families:
Anapidae, Barychelidae, Dipluridae, Desidae, Pararchaeidae, Lamponidae,
Theridiidae, Pisauridae, Miturgidae (Zorinae), Salticidae, Scytodidae,
Tetrablemmidae, Telemidae, Segestriidae, Oonopidae, Mimetidae, Deinopidae,
Uloboridae, Mysmenidae, Linyphiidae, Tetragnathidae, and Araneidae (Berland
1924, 1929; Platnick and Forster 1989, 1993; Platnick 1993; Rainbow 1920; Raven
1994; Rix and Harvey 2010; Simon 1889; 1892). Until 1993 have been recorded
spiders of 112 genera and 194 spp. (Platnick 1993). Platnick and Forster (1993)
synonymized the “endemic family” Bradystichidae Simon, 1884, with Pisauridae.
Rix and Harvey (2010b) recorded from the island the first representative of
Pararchaeidae outside Australia and New Zealand.
Endemic genera:
Fam. Dipluridae: Stenygrocercus Simon, 1892 (six spp.)
Fam. Anapidae: Caledanapis Platnick et Forster, 1989 (six spp.); Mandanapis
Platnick et Forster, 1989 (one sp.); Montanapis Platnick et Forster, 1989 (one
sp.); Caledothele Raven, 1991
Fam. Barychelidae, Barychelinae: Barycheloides Raven, 1994 (5 spp.); Barychelus
Simon, 1889 (2 spp.); Encyocrypta Simon, 1889 (32 spp.); Natgeogia Raven,
1994 (1 sp.); Orstom Raven, 1994 (6 spp.); Questocrypta Raven, 1994 (1 sp.)
Fam. Desidae: Canala Gray, 1992 (three spp.)
Fam. Lamponidae: Centrocalia Platnick, 2000 (three spp.)
Fam. Theridiidae: Anatea Berland, 1927 (one sp.)
Fam. Pisauridae: Bradystichus Simon, 1884 (five sp.), Pseudohostus Rainbow,
1915 (five spp.)
Fam. Miturgidae (incl. Zoridae): Zoroides Berland, 1924 (one sp.)
Fam. Salticidae: Corambis Simon, 1901 (two spp.); Lystrocteisa Simon, 1884 (one
sp.); Rhondes Simon, 1901 (six spp.)
Acariformes - Ref.: Beron (2008)
Fam. Erythraeidae: Charletonia rageaui Southcott, 1966
8.24 Lord Howe Island 777

Holothyrida First recorded by Berlese (1923). Endemic genus is Haplothyrus
Lehtinen (two spp.). One genus (Lindothyrus Lehtinen) is shared with Lord Howe
Island (on New Caledonia endemic sp. Lindothyrus rubellus Lehtinen, 1995). Both
genera belong to the family Holothyridae and not to the Allothyridae, known from
Australia and New Zealand. The New Guinean genera also are not represented in
New Caledonia. The same observation is valid for Lord Howe Island.
8.24 Lord Howe Island
The small Lord Howe Island is called “The riddle of Pacific” (Paramonov 1958). On
a surface of 14.55 km2 (10 km long and up to 2 km wide) live many endemic spe-
cies, and the island is 600 km. far from Australia and 900 km. far from Norfolk
Island. Its inhabitants (387 permanent and up to 400 tourists) are in the “settled
area,” in the lowland. It is because since 1981 70% of the island is a reserve and,
fortunately, there are some forests left on the 875 meter high Mount Gower. The
archipelago includes also 28 uninhabited islands.
It is considered that Lord Howe is part of the island chain, having existed along
the Western edge of the shelf called Lord Howe Rise, 3000 km. long and 300 wide.
This Rise extended from New Zealand to west of New Caledonia and consists of
continental rocks, detached from Australian plate 60–80 Ma ago. The shelf is part
of Zealandia microcontinent.
Pseudoscorpiones From Lord Howe Island, Beier (1976) has published eight spp.
of pseudoscorpions (four endemics in the island), seven genera, and six families
(Chthoniidae, Dithidae, Olpiidae, Atemnidae, Chernetidae, Cheliferidae). One
endemic subgenus Pholeochthonius – troglobite. Notogean genera are
Anaulacodithella and Philomaoria.
Araneae Rix and Harvey (2010a) described the endemic genus and species of
Micropholcommatidae Patelliella adusta and another endemic micropholcommatid
species – Taphiassa magna.
Endemic genera
Fam. Micropholcommatidae
Patelliellini (end. tribe)
Patelliella Rix et Harvey, 2010
Fam. Cyatholepidae
Lordhowea Griswold, 2001
Fam. Zodariidae
Basasteron Baehr, 2003
Fam. Phrurolithidae
Dorymetaecus Rainbow, 1920 (one sp.)
8.25 Conclusion
The level of representation of Arachnida in the classical Notogea (with Papuan area
but without Patagonia) is much lower than the level in the vertebrates, with their
endemic subclasses, orders, and suborders. Even in the most isolated area (New
Zealand), there are no endemics of very high rank. Here are the endemisms above
genus:
Australia (cont.): One endemic family of scorpions (Urodacidae)
Tasmania: Only endemic subfamilies of spiders (Plesiothelinae and Hickmanniinae)
New Guinean area: No endemics above genus
New Caledonia: One endemic family of Opiliones (Troglosironidae)
New Zealand: One endemic family of spiders (Huttoniidae) and one of Opiliones
(Synthetonychiidae)
Characteristics of the Arachnida in Australia, Tasmania, Papuan Area, New
Zealand, New Caledonia, and Lord Howe Island
Kingdom Notogaea – No endemic orders or suborders among Arachnida
Region of Australia
Palpigradi – Two spp. brought from Europe, one local (?)
Solifugae - Missing
Amblypygi – Fam. Charinidae (Charinus Simon)
Thelyphonida (Uropygi) – Not found in Australia
Schizomida – Fam. Hubbardiidae (Apozomus Harvey, Attenuizomus Harvey,
Bamazomus Harvey, Brignolizomus Harvey, Draculoides Harvey, Julattenius
Harvey, Notozomus Harvey, Ovozomus Harvey, Paradraculoides Harvey
et al.)
Scorpiones – Endemic family is Urodacidae, end. genera Urodacus Peters
with 19 species, Isometroides Keyserling and Cercophonius Peters (Australia
and Tasmania), and Aops Volschenk et Prendini.
Pseudoscorpiones – No endemic families; 150 spp., 17 fam. Endemic genera
of pseudoscorpions in Australia (without Tasmania) are fam. Olpiidae,
Austrohorus Beier (one sp.), Linnaeolpium Harvey et Leng (one sp.); fam.
Cheliferidae, Australochelifer Beier (one sp.); fam. Chernetidae,
Conicochernes Beier (four spp.), Marachernes Harvey (three spp.); in
Tasmania Neopseudogarypus Morris
Opiliones – No endemic families
Cyphophthalmi – Fam. Pettalidae (two genera, endemic in Queensland)
(Austropurcellia Juberthie) and Western Australia (Karripurcellia Giribet)
8.25 Conclusion 779
Laniatores – Fam. Triaenonychidae (endemic: Breviacantha Kauri, Callihamina

Roewer, Callihamus Roewer, Cluniella Forster, Heteronuncia Roewer,
Holonuncia Forster, Perthacantha Roewer, Yatala Roewer, Conoculus
Forster, Dingupa Forster, Dipristes Roewer), Assamiidae (Dampetrus
Karsch, Metamermerus Roewer), Samoidae, Zalmoxidae (Stygnoleptinae)
Eupnoi – Fam. Neopilionidae (incl. Monoscutidae), Ballarriinae (Arrallaba
Hunt et Cokendolpher, Ballarra Hunt et Cokendolpher, Plesioballarra crinis
Hunt et Cokendolpher, Vibone Kauri). Hesperopilio Shear – Unclear family
place
Dyspnoi – Fam. Acropsopilionidae (Acropsopilio Silvestri)
Araneae – Seventy-three families with at least 237 endemic or subendemic gen-
era (without Tasmania). In Tasmania – Two endemic subfamilies and 29
endemic genera
Opilioacarida – One undeterm. recorded
Holothyrida – Fam. Allothyridae (Allothyrus van der Hammen, Australothyrus
van der Hammen)
Region New Zealand
Very unbalance fauna. Seven orders of Arachnida are missing; the remaining are
Pseudoscorpiones (67 spp.), Opiliones (111 spp.), Araneae (1662 spp.), Holothyrida
(1 sp.), Ixodida (11 spp.), Mesostigmata, Sarcoptiformes, and Trombidiformes.
Particularly well is represented the order Araneae.
Palpigradi – Missing
Solifugae – Missing
Amblypygi – Missing
Thelyphonida (Uropygi) – Missing
Schizomida – Missing
Scorpiones – Missing
Pseudoscorpiones – No endemic families; 67 spp. of 27 genera and 8 families.
Endemic genera: Maorichthonius Chamberlin, Sathrochthoniella Beier,
Tyrannochthoniella Beier, Nelsoninus Beier, Apatochernes Beier (together with
Campbell Islands, Snares Islands, Auckland Islands, Chatham Islands, Norfolk
Island), Heterochernes Beier
Opiliones – One endemic family
Cyphophthalmi – Three endemic genera (Rakaia Hirst, Aoraki Boyer et
Giribet, Neopurcellia Foster) from the family Pettalidae (in total 21 spp.)
Laniatores – Endemic family: Synthetonychiidae (Synthetonychia Forster),
Triaenonychidae: Hedwiga Roewer, Hendea Roewer, Hendeola Forster,
Metanuncia Roewer (Stewart Island), Neonuncia Roewer, Prasma Roewer,
Prasmiola Forster, Psalenoba Roewer, Triregia Forster, Algidia Hogg,
Cenefia Roewer, Muscicola Forster, Pristobunus Roewer, subfam.
Sørensenellinae (Karamea Forster, Sorensenella Pocock)
Eupnoi – Fam. Caddidae (Acropsopilio Silvestri), Neopilionidae (incl.

Monoscutidae) (Forsteropsalis Taylor (nine spp.), Mangatangi Taylor,
Pantopsalis Simon; Monoscutinae (end. subfamily): Acihasta Forster,
Monoscutum Forster, Templar Taylor) (New Zealand, Auckland, Snares,
Campbell Islands)
Dyspnoi – Missing
Araneae: One endemic family (Huttoniidae), 1 subfamily (Pahorinae), and
at least 93 endemic genera of spiders:
Fam. Huttoniidae (Huttonia Pickard-Cambridge), Hexathelidae (Hexathele
Ausserer, Porrhothele Simon), Anapidae (Novanapis Platnick et Forster,
Paranapis Platnick et Forster, Zealanapis Platnick et Forster), Pararcheidae
(Forstrarchaea Rix, Pararchaea Forster), Cycloctenidae (Plectophanes Bryant,
Toxopsiella Forster, Uzakia Koçak et Kemal, Cantuarea Hogg), Gradungulidae
(Gradungula Forster, Pianoa Forster, Spelungula Forster), Orsolobidae
(Anopsolobus Forster et Platnick, Ascuta Forster, Bealeyia Forster et Platnick,
Dugdalea Forster et Platnick, Duripelta Forster, Maoriata Forster et Platnick,
Orongia Forster et Platnick, Paralobus Forster et Platnick, Pounamuella Forster
et Platnick, Subantarctia Forster, Tangata Forster et Platnick, Tautukua Forster
et Platnick, Turretia Forster et Platnick, Waiporia Forster et Platnick, Wiltonia
Forster et Platnick), Mecysmauchenidae (Aotearoa Forster et Platnick,
Zearchaea Wilton, Parapua Forster, Pua Forster, Forstarchaea Rix, Tekelloides
Forster, Waitkerra Opell, Nomana Forster, Pahora Forster, Pahoroides Forster,
Runga Forster, Wairua Forster, Meringa Forster, Mangua Forster, Ahua Forster
et Wilton, Huka Forster et Wilton, Mahura Forster et Wilton, Neoramia Forster
et Wilton, Oramia Forster et Wilton, Oramiella Forster et Wilton, Orepukia
Forster et Wilton, Paramyro Forster et Wilton, Porotaka Forster et Wilton,
Tararua Forster et Wilton, Tuapoka Forster et Wilton, Anhunga Forster et
Wilton, Maloides Forster et Wilton, Muritaia Forster et Wilton, Pakeha Forster
et Wilton, Paravoka Forster et Wilton, Poaka Forster et Wilton, Gasparia Forster
et Wilton, Gohia Forster et Wilton, Goyenia Forster et Wilton, Hapona Forster,
Helsinia Forster, Hulua Forster et Wilton, Lamina Forster, Mangareia Forster,
Matachia Dalmas, Mesudus Özdikmen, Neomyro Forster et Wilton,
Notomatachia Forster, Nuisiana Forster et Wilton, Otagoa Forster, Panoa
Forster, Rapua Forster, Toxopsoides Forster et Wilton, Tuakana Forster),
Dictynidae (Paradictyna Forster, Viridictyna Forster, Karanga Forster, Poroides
Forster, Forstertyna Harvey, Megadictyna Dahl, Haurokoa Koçak et Kemal,
Zealoctenus Forster et Wilton, Pacificana Hogg, Kaitawa Forster, Matua
Forster, Nauhea Forster, Notiodrassus Bruyant, Zelanda Özdikmen)
Opilioacarida – Missing
Holothyrida – No endemic families or genera: Allothyrus van der Hammen
(one sp., in common with Australia)
Ixodida – No endemic genera. Fam. Ixodidae and Argasidae (11 spp., 4
endemic)
8.25 Conclusion 781
Other Acari: No endemic families

Trombidiformes
Prostigmata – Fam. Erythraeidae (Neosmaris Hirst, Taranakia Southcott,
Ramsayella Zhang)
Papuan Area (New Guinea, Bismarck, Salomon Islands)
Arachnida: No endemic families
Palpigradi – Fam. Eukoeneniidae (Eukoenenia cf. lawrencei et Koeneniodes cf.
frondiger)
Amblypygi – Fam. Charinidae (Charinus Simon, Sarax Simon, New Guinea),
Charontidae (Charon Karsch, New Guinea)
Thelyphonida (Uropygi) – Fam. Thelyphonidae (Thelyphoninae). Genera:
Abaliella, Thelyphonus, and Minbosius (New Guinea, Fergusson, New
Ireland, New Britain, Guadalcanal). Haupt (2009a) synonymized Abaliella
Strand, Minbosius Speijer, and Tetrabalius Thorell with Thelyphonus Latreille.
Only endemic species
Schizomida – Fam. Hubbardiidae (Apozomus Harvey, Bamazomus Harvey –
New Guinea)
Scorpiones – In New Guinea (incl. Aru and Bougainville) are known six wide-
spread species of the genera Isometrus Ehr., Lychas C.L. Koch (Buthidae),
and Liocheles Sundevall (Hormuridae).
Pseudoscorpiones – Fam. Chthoniidae, Tridenchthoniidae, Syarinidae,
Cheiridiidae, Garypinidae, Geogarypidae, Atemnidae, Olpiidae,
Sternophoridae, Cheliferidae, Chernetidae, Withiidae. Endemic genera:
Papuchelifer Beier (Cheliferidae), Cyclochernes Beier (Chernetidae)
Opiliones
Cyphophthalmi – Fam. Stylocellidae (two spp. of Leptopsalis on Bird’s Head of
New Guinea)
Laniatores – Fam. Assamiidae (Dampetrinae), Epedanidae, Podoctidae (incl.
Erecananinae and Ibaloniinae) (Asproleria Roewer, Heteropodoctis Roewer,
Paramesoceras Roewer, Podoctinus Roewer (New Britain), Proholozoster
Roewer), Zalmoxidae (Stygnoleptinae) (Zalmoxis Soerensen)
Eupnoi – Fam. Sclerosomatidae (Gagrella Stoliczka)
Dyspnoi – Missing
Araneae – ca. 200 spp. Endemic genera: Jocquella Baert (Telemidae), Yoroa
Baert (Theridiidae), Pseudopsellonus Balogh, Senoculifer Balogh (Philodromidae)
Holothyrida – Fam. Holothyridae (Hammenius Lehtinen, Leiothyrus van der
Hammen) (all in New Guinea)
New Caledonia
Palpigradi – Present, unidentified
Amblypygi – Fam. Charinidae (Charinus Simon)

Thelyphonida (Uropygi) – Missing
Schizomida – Hubbardiidae indet.
Scorpiones – Two species of Liocheles Sundevall (Hormuridae)
Pseudoscorpiones – Families Chthoniidae, Tridenchthoniidae, Syarinidae,
Cheiridiidae, Garypinidae, Geogarypidae, Atemnidae, Olpiidae, Sternophoridae,
Cheliferidae, Chernetidae, Withiidae
Opiliones
Cyphophthalmi – One endemic fam. Troglosironidae with genus Troglosiro Jub.
(13 spp.)
Laniatores – Fam. Assamiidae, Epedanidae, Podoctidae (incl. Erecananinae and
Ibaloniinae), Zalmoxidae (Stygnoleptinae)
Dyspnoi – Missing
Araneae – Many endemic genera
Holothyrida – Fam. Holothyridae: two genera, one of them endemic (Haplothyrus
Lehtinen, two spp.), the other (Lindothyrus Lehtinen) is shared with Lord Howe
Island.
8.25.1 Conclusion
The level of representation of Arachnida in the classical Notogea (with Papuan area
but without Patagonia) is much lower than the level in the vertebrates, with their
endemic subclasses, orders, and suborders. Even in the most isolated area (New
Zealand), there are no endemics of very high rank. Here are the endemisms above
genus:
Tasmania: Only endemic subfamilies of spiders (Plesiothelinae and Hickmanniinae)
New Zealand: One endemic family of spiders (Huttoniidae) and one of Opiliones
(Synthetonychiidae)
Tasmania: Only endemic subfamilies of spiders (Plesiothelinae and
Hickmanniinae)
New Zealand: One endemic family of spiders (Huttoniidae) and one of
Opiliones (Synthetonychiidae)
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Chapter 9
Cave Arachnida
Abstract Almost all orders of Arachnida are represented in caves, except for

Solifugae. In some orders (Pseudoscorpiones) there are many troglobitic blind spe-
cies. Such species are found also among spiders, scorpions, opilions, and others.
Further species are prospected in different groups and regions of the World, and the
ways in which Arachnida have invaded the caves are discussed here (especially the
new discoveries in tropical caves, the subterranean fauna of Hawaii, Galapagos, and
unusual caverns such as Movile and Ayalon).
9.1 Cave Life in Different Orders
Palpigradi Many species of Palpigradi are known only from caves (Eukoenenia
Börner).
Cave Palpigradi have been found in Bulgaria, Greece, France, Italy, and Brazil.
Palpigradi are known also from hyporheic environments or iron mines
(Leptokoenenia Condé).
Scorpions These arachnids are not typical cave dwellers, but there are several tro-
globites among the 1725 species in the order. They are known mostly from Mexico
(studies of Mitchell 1968, 1971, Francke, many papers), but also from Ecuador,
Venezuela, Sarawak, Laos, Vietnam, Australia, and Israel. A general survey of cave
scorpions was made by Lourenço and Francke (1985). The list of all troglobitic
scorpions in Lourenço (1994) contains only 13 species (11 from Mexico), but since
then many new investigations have added greatly to the knowledge on cave scorpi-
ons (from Israel, Laos, Vietnam, and Australia). Troglophylic scorpions are known
to reside in Cuba. Some trogloxenic scorpions (Euscorpius) have been found in
South European caves: E. deltshevi Fet et al., 2014, E. solegladi Fet et al., 2014, E.
candiota (Birula, 1903), and others.
Fam. Buthidae According to Armas (1973), the species Alayotityus delacruzi,
described by him from the cave Cueva de los Majáes (38 °C), is the first clearly
cavernicolous scorpion in Cuba. Cave Centruroides have been found in Mexico
(Reddell 1981).

https://doi.org/10.1007/978-3-319-74418-6_9
834 9 Cave Arachnida
Fam. Chactidae The enigmatic Pyrenean blind scorpion Belisarius xambeui from
the Chactidae (Broteinae) family was identified by Fet and Soleglad (2003).
Lourenço (1998) places it in the family Troglotayosicidae (also South American).
These are the affinities of the recently described Akrav genus from Israeli caves (and
new family Akravidae).
Fam. Scorpionidae (incl. Diplocentridae) Three species of cave Diplocentrus
have been described by Francke (1977 1978), of which D. anophthalmus is
eyeless.
Fam. Pseudochactidae The genus Pseudochactas Gromov (1998) was described
as being found in Central Asia (Kazachstan), and then more cave species were found
in Laos (Troglokhammouanus steineri Lourenço (2007), Vietbocap lao Lourenço
(2012)), and in Vietnam (Vietbocap canhi Lourenço and Dinh-Sac Pham (2010), V.
thienduongensis Lourenço and Ðình Sắc Pham (2012).
Fam. Akravidae In 2006 Israeli cavers found about 20 dead scorpions in the
Ayalon Cave (which was before that completely closed), described by Levy (2007)
as a new species, genus, and even family (Akrav israchanani, fam. Akravidae). Fet,
Soleglad, and Zonstein (2011) studied in detail this new enigmatic species and con-
firmed its Neotropical affinities.
Fam. Troglotayosicidae The new genus and species Troglotayosicus vachoni was
described by Lourenço (1981) after being found in the famous Cueva de los Tayos
in Ecuador, first in the Chactidae, but later (Lourenço 1998) he created a special
family for it.
Fam. Typhlochactidae Vignoli and Prendini (2009) revised the family to having
nine species of endemic troglomorphic scorpions from eastern Mexico
(Typhlochactinae – Sotanochactas, Stygochactas, and Typhlochactas;
Alacraninae – Alacran).
According to Prendini et al. (2009), troglobitism is not an evolutionary
dead-end.
Fam. Chaerilidae Chaerilus chapmani Vachon et Lourenço (1985) have been
described as being from Borneo (Sarawak) and the eyeless Ch. sabinae Lourenço
(1995) was found in Gua Atah on Sulawesi (not India, as stated in the original
description.
Pseudoscorpiones More than 3538 described species in 26 families and more than
454 genera (Harvey 2002, 2013f) have been recognized. Cave-inhabiting species
are included in several families, but only six (Chthoniidae, Pseudotyrannochthoniidae,
Neobisiidae, Syarinidae, Bochicidae, and Olpiidae) contain troglobites.
9.1 Cave Life in Different Orders 835
Fam. Chthoniidae There are around 650 species of this family, many troglobites
of the genera Chthonius, Aphrastochthonius, Apochthonius, Kleptochthonius,
Lagynochthonius, Neobalkanella, Pseudochthonius, Troglochthonius,
Tyrannochthonius, etc. from Europe, Mexico, the USA, Hawaii, Guatemala,
Sarawak, Australia, and Japan. Other cave species (troglophile, trogloxene) have
been identified from Korea, Japan, Brazil, and Kenya.
Fam. Pseudotyrannochthoniidae Five species of Pseudotyrannochthonius Beier
are considered to be troglobitic in Australia (incl. Tasmania).
Fam. Tridenchthoniidae (= Dithidae) This family has 71 species, and only one
(Tridenchthonius juxtlahuaca from Mexico) is found in a cave.
Fam. Lechytiidae This family is made up of 23 recent species, and only one
(Lechytia cavicola from Mexico) is found in a cave.
Fam. Pseudogarypidae Seven extant species in this family, two of them
(Pseudogarypus)are found in caves in California (USA).
Fam. Ideoroncidae Fifty-nine species belong to this family, ten living in caves
(Albiorix bolivari, A. mirabilis, A. reddelli from Mexico, Dhanus doveri, Dh. suma-
tranus from Malaya and Sumatra, Negroroncus aelleni from Congo, and four spe-
cies of Typhloroncus from Mexico).
Fam. Bochicidae (= Vachoniidae) This family has 44 species, 14 of them known
from caves: Antillobisium mitchelli, A. vachoni from Cuba, three sp. of Leucohya
from Mexico and Texas, Troglobochica jamaicensis, T. pecki from Jamaica, three
sp. of Mexobisium from Guatemala and Mexico, Troglohya caranzai, T. mitchelli
from Mexico, Arcanobisium from Spain, Titanobochica from Portugal. Many of
them are troglobites.
Fam. Neobisiidae 595 species in this family, many of them from caves, including
troglobites from Europe, Turkey, Caucasus (the genera Neobisium, Roncus,
Balkanoroncus, Roncobisium, etc.), and the USA (Tartarocreagris, Trisetobisium).
Fam. Syarinidae 111 species in this family, 18 from caves of USA (Chitrella),
Hadoblothrus (Greece, Italy), Ideobisium (Venezuela, Ecuador, and Puerto Rico),
Ideoblothrus (Mexico), Pseudoblothrus (Italy, Crimea, France, and Switzerland),
Troglobisium (Spain). Most of the European species are troglobites and relicts
(Beier 1970).
Fam. Larcidae This family has 15 species, three species of which are found in
caves (Italy, Spain, and California).
Fam. Cheiridiidae This family has 73 species; only one (Cryptocheiridium ele-
gans from Cuba) has been identified as being from a cave.
Fam. Olpiidae This family has 268 species, only six are known to inhabit caves
(including Amblyolpium graecum from Greece and the troglomorphic endemic
genus Linnaeolpium linnaei (Harvey et Mei Chen Leng, 2008) from Australian
caves, the only Olpiid with reduced eyes).
Fam. Garypinidae There 76 species in this family.
Fam. Sternophoridae There are 20 species in this family, one was found in a cave
by us (Afrosternophorus cavernae, Papua New Guinea).
Fam. Withiidae 158 species, only two (Parawithius schlingeri from Peru and
Pycnowithius cavernicola from Kenya) are known from caves.
Fam. Cheliferidae 273 species, five of them found in caves (Lissochelifer gracili-
pes from Kenya, Mexichelifer reddelli from Mexico, Protochelifer cavernarum, P.
naracoortensis from Australia and Stygiochelifer cavernae from Java).
Fam. Atemnidae (= Miratemnidae) 178 species, only two found in caves (Atemnus
strinatii in the Philippines and Oratemnus cavernicola in Australia).
Fam. Chernetidae (= Myrmochernetidae) 663 species, including from caves in
Macedonia, Cuba, Sicily, Azerbaijan, Japan, Thailand, Uganda, Mexico, Australia,
and many other places).
Opiliones This order contains 45 families, 500 genera, and more than 6500 spe-
cies, according to Harvey (2002), Pinto-da-Rocha and Giribet (2007) and Kury
(2013), in four suborders: Cyphophthalmi, Laniatores (mostly tropical) and
Palpatores (mostly Holarctic). All four contain cave-inhabiting species.
Cyphophthalmi The troglobite «Neogovea» mexasca Shear has been identified in
Mexican caves. It is currently the only species of the genus Shearogovea Giribet
with uncertain affinities.
Fam. Neogoveidae An endemic genus and species Canga renatae has been found
in a Brazilian cave, and Metagovea philipi in a cave in Ecuador.
Fam. Sironidae The strange genus and species Marwe coarctata Shear from
Kenya is now considered to not belong to Sironidae, but has no other family attach-
ment. According to Rambla and Juberthie (1994), the troglobite of Bulgarian
Central Stara Planina Tranteeva paradoxa Kratochvil is by far the most modified.
The genus Tranteeva was synonymized under Cyphophthalmus by Karaman (2009).
Fam. Pettalidae All are anophthalm dwellers of the southern hemisphere with one
endemic troglobite from South Africa (Purcellia [Speleosiro] argasiformes
Lawrence).
Fam. Troglosironidae This is an endemic family of troglobites in New Caledonia.
So far 13 species have been described by Juberthie (1979), Shear (1993), and
Sharma and Giribet (2005); some more awaiting description.
Dyspnoi
Fam. Nemastomatidae According to Rambla and Juberthie (1994), 11 troglo-

bitic species are included in this family from caves of from the Balkans, Crimea,
Georgia, Italy, Tunisia, and the USA. In Bulgarian caves two troglobitic species
have been reported: Paranemastoma bureschi and P. beroni, and are largely repre-
sented in caves.
Fam. Ischyropsalididae This is a European family with at least ten troglobitic
species from the cool caves in the Alps, Cantabric Mountains, and the Pyrenees.
Four species are considered troglophyles.
Fam. Sabaconidae In the Pyrenees the troglobite Sabacon picosantrum described
by Martens (1982) are found (described as being from Picos de Europa) and the
troglophyle S. paradoxum Simon.
Laniatores
Fam. Phalangodidae Some European species (Paralola, Ausobskya) are troglo-

bites, and are usually considered relicts, but opinions differ on this.
Fam. Assamiidae Tropical species of Africa and SE Asia, some of them anoph-
thalm, from leaf litter, termite nests, but also described as being from caves
(Sijucavernicus kempi and Calloristus cavernarum from India, occulated species
from Gabon, India, Thailand, and Tropical Africa).
Fam. Pyramidopidae Maiorerus randoi Rambla, 1993 from a lava cave in
Fuerteventura (Canary Isl.) is the only eyeless pyramidopid known. Another troglo-
bite (Jarmilana Cruz-López, Proud et Pérez-Gonzalez, 2016, the first member of
the family in the New World) has been identified recently from a cave in Belize.
Fam. Gonyleptidae This family is rarely modified for cave life. Many species
(troglophyles and trogloxenes) have been described from South American caves.
Fam. Agoristenidae Members of the genus Trinella Goodnight and Goodnight

have been described under different names (Vima, Phalangozea) from caves in
Venezuela (Rambla 1978; Muñoz-Cuevas 1976). Some are troglobites.
Fam. Triaenonychidae Some have been found in North American lava tubes
(Briggs 1974).
Fam. Cladonychiidae (= Erebomastridae) This family has been identified in
caves in the USA (Erebomaster).
Fam. Travuniidae A most interesting family, containing relicts of a very old pre-
Tertiary fauna (Kratochvíl 1958; Rambla and Juberthie 1994). In this latter paper
threre is a map of a very disjunct distribution in South Europe (Pyrenees, Sardinia,
Dalmatia), USA (Speleonychia), Japan (Yuria pulcra Suzuki, outside and inside
caves), and Korea (Peltonychia coreana). Kury and Mendes (2007) analyzed the
taxonomic status of European genera of Travuniidae. They concluded that this relict
family contains in Europe only five genera and 17 species, most of which are troglo-
bites: Arbasus (A. caecus from France), Buemarinoa (B. patrizii from Sardinia),
Dinaria (D. vietrenicae from Bosnia and Herzegovina), Hadziana = Peltonychia
(6–8 species from France, Spain, Italy, Switzerland, and Slovenia), Travunia =
Abasola (four sp. from Croatia, Montenegro, Bosnia, and Herzegovina. The new
genus and species Trojanella serbica Karaman (2005), described from a cave in
Stara planina (now in Serbia) was also found in Travuniidae.
Some other families (Trogulidae, Sclerosomatidae, Phalangiidae) contain spe-
cies found in caves, but only as troglophyles and trogloxenes.
Araneae About one-third (37) of the 113 families of spiders in the world are
known to be from caves. We can find a detailed list in Ribera and Juberthie (1994),
which has been updated.
With regard to the biogeographical importance of cave spiders, we have to keep
in mind the opinion of Brignoli (1979), that « …les araignées cavernicoles n’ont
aucune valeur spéciale en biogéographie [The cave species don’t have any special
value for the Biogeography]».
Liphistiomorpha Microphthalmes are known from caves in Malaysia and Thailand
(Liphistius). Five families of Mygalomorpha contain cave inhabiting species:
Ctenizidae (Guinea, Ecuador), Hexatelidae (New Zealand), Dipluridae (Philippines,
Jamaica, Australia, Mexico, New Zealand), Barychelidae (Cuba), and Theraphosidae
(Mexico). According to Ribera and Juberthie (1994), in these groups there are
around 20 troglobitic spiders living in tropical caves.
The bulk of the cave spiders are included in the Araneomorpha, and especially in
Haplogynae.
Fam. Austrochilidae Hickmania troglodytes (Higgins et Peters) – endemic troglo-
bite from the caves of Tasmania.
Fam. Leptonetidae Many troglobites and troglophiles from the caves of South
Europe, North Africa, Turkey, USA, Mexico, Japan, Korea, and China (Leptoneta,
Protoleptoneta, Paraleptoneta, Barusia, Sulcia, Leptonetela, Teloleptoneta,
Saturana, Falcileptoneta, Masirana, Archoleptoneta, Neoleptoneta, Appaleptoneta,
Callileptoneta). As a whole family, the cave Leptonetids show a disjunct distribu-
tion in the temperate regions.
Fam. Ochyroceratidae Small spiders typical in the southern hemisphere. Cave
species in Cuba (Fageicera, Speocera, Ochyrocera), Guatemala, Peru (Ochyrocera),
Mexico (Ochyrocera, Theotima), Jamaica (Theotima), Hawaii (Theotima),
Philippines (Simonicera), Burma (Psiloderces, Theotima, Althepus), Malaysia
(Psiloderces), and Okinawa (Speocera). Some species are microphthalm and could
be accepted as troglobites.
Fam. Dysderidae Many troglobites and troglophiles in the caves of South Europe
(Spain, Majorca, Portugal, Italy, Sardinia, Dalmatia, Morocco, Algeria, Creta –
Dysdera, Stalita, Folkia, Stalagtia, Minotauria, Rhodera, Rhode, Harpactea,
Speleoharpactea) and Canary Islands (Dysdera). Stalita taenaria Schiödte from
Postoina jama (Slovenia) was the first troglobitic spider ever described.
Fam. Oonopidae Many anophthalm species live in tropical caves (Mexico, Cuba,
Ecuador, New Guinea, India, Ceylon, etc.), from genera like Wanops, Oonops,
Triaeris, Oonopsoides, Gamasomorpha, Opopaea, Ischnothyreus. They live also in
the forest litter and cannot be assumed to be troglobites.
Fam. Telemidae An ancient family with troglobites in Europe (Telema tenella),
Japan (T. niponica), Guatemala (T. mayana), Usophila from the caves of USA and
New Caledonia, Apneumonella oculata (Tanzania), Cangoderces lewisi (Cango
Cave in South Africa) and others.
Fam. Tetrablemidae Like Oonopidae, these spiders live in the tropical forests and
caves in Mexico (Tetrablemma, Matta), Okinawa (Tetrablemma), and Sumatra
(Ablemma).
Fam. Loxoscelidae, Scytodidae Often found in caves, mostly troglophyles.
Loxosceles vonweredei Newlands is a troglobite from South Africa.
Fam. Pholcidae Cosmopolytic family, often found in caves. Some are considered
troglobites (Mexico, Jamaica, Venezuela, Ecuador, New Guinea, Canaries,
Tasmania, and others).
Fam. Linyphiidae This huge family has many species in caves of Europe, North
Africa, Canary Islands., etc. In Europe important species are Lepthyphantes s. lato,
Troglohyphantes, Porrhomma, Centromerus, Caviphantes dobrogica living in
Movile Cave in Romania. Other troglobites are Tunnagyna antricola (Mexico),
Anthrobia monmouthia, and Phanetta subterranea (USA), Meioneta gagnei
(Hawaii), Allomengea coreana and Jacsonella sexoculata (Korea), Thyphonychia
meridionalis and Troglodytia bouidwijni (ex -Yugoslavia), and Dunedinia

(Australia).
Fam. Pimoidae Pimoa breuli is known from Cantabric Mts. (troglophyle).
Fam. Metidae Troglophilic spiders from South Europe, North America, Far East
of Russia, Guinea, Tasmania.
Fam. Synotaxidae Two troglobitic and one troglophylic species of the endemic
genus Tupua are described from Tasmania.
Fam. Nesticidae Many species, mostly troglophylic, in caves from the USA to
Japan. Some Nesticus are troglobites, also Typhlonesticus from Dalmatia and the
blind Nesticella marapu Benjamin (2004) from Marro Cave in Sumba, Indonesia.
The anophthalm Canarionesticus from the Canaries is considered a glacial relict.
Fam. Theridiidae Found often in tropical caves, rarely in temperate caves. Some
are troglobites: Coscinidia, Stenops, and Achaearanea from New Guinea, Steatoda,
Pholeomma, and Icona from Australia and the most interesting spiders in the lava
caves of the Açores, Canaries, and Galapagos (Theridion spp.).
Fam. Theridiosomatidae Many troglophylic species live in the caves of tropical
countries. Some (Wendilgarda from Venezuela and Guadelupa, Andasta from
Ceylon) are known as only being from caves.
Fam. Anapidae Some troglophyles from American caves (Anapis, Anapisoma)
and Tasmania (Chasmocephalon, Pseudoanapis) and troglobites from South Africa
(Pseudoanapis) and Korea (Conoculus).
Fam. Mysmenidae One relict Trogloneta from Tasmania, some troglophylic
Mysmena from Mexico and the USA.
Fam. Clubionidae Berlandia tenericola from a cave in Tanzania, Brachyanillus
liocraninus (anophthalm) from Algeria, Lascona cristiani from Movile Cave in
Romania (troglobite and relict), Agroecina canariensis from lava tubes in the
Canary Islands.
Fam. Gnaphosidae Two troglobite species from lava tubes in Galapagos:
Lygromma gertschi and L. anops.
Fam. Lycosidae Living outside caves, except for two troglobite species from the
Hawaiian lava tubes: Lycosa howarthi and Adelocosa howarthi (Gertsch 1973).
Fam. Prodidomidae Found in many tropical caves. Some are considered troglo-
bites: Lygromma gertschi from Jamaica, L. anops from Galapagos.
Fam. Stiphidiidae In Australia, incl.Tasmania, Stiphidion, Baiami and Tartarus

have troglophyle and troglobitic species.
Fam. Agelenidae Often found in cave entrances, especially Tegenaria in Europe,
Blabomma in Korea, and others. Some are microphthalms.
Fam. Hahniidae Rare in caves, exceptions are the troglobites Iberina mazaredoi
from France and Spain and I. caeca in Movile Cave in Romania.
Fam. Mimetidae Mimetus strinatii from a cave in Ceylon.
Fam. Cybeidae and Amaurobiidae Microphthalm cave species of Cybaeus and
Coelotes from Japan and Korea.
Fam. Holarchaeidae Subterranean family of two species in Tasmania and New
Zealand.
Fam. Cycloctenidae Subterranean Cycloctenus and Toxopsiella in Australia and
Tasmania.
Fam. Sparassidae Giant spiders, often found in caves of Southeast Asia, but not
troglobitic.
Schizomida Altogether 33 species (three Protoschizomidae of genus
Agastoschizomus and 30 Schizomidae of genus Schizomus) have been found in
caves, 25 of them are considered troglobites or at least more or less troglomorphic
(Georgesco 1994, actualized).
Ricinulei This relict order is known only from Neotropical America (up to Northern
Mexico and Texas) and West Africa. Two of the three extant genera are represented
in caves. According to Juberthie (1994), only ten species of Ricinulei are known
from underground environments (nine Cryptocellus, eight from Mexico and one
from Cuba; one Cryptocellus (= Heteroricinoides) from Venezuela). All Ricinulei
are blind, and the cave species have relatively longer legs.
Amblypygi This order contains a small number of large and conspicuous « whip
spiders », found often in caves (according to Harvey (2003), 136 species, reorganized
(Miranda et al. 2018) into 146 species with 17 genera and five families). They are
known from caves in tropical countries. The American species (from southern USA,
Mexico, Central America, Caribbean Islands, and South America) belong to the fam-
ily Phrynidae (Phrynus and Paraphrynus), the cave species of the Old World belong
to the families Charinidae (Charinus), Phrynichidae (Phrynichus, Euphrynichus),
and Charontidae (Charon, Stygophrynus). One species of Phrynus (Phrynidae) was
described from a cave in Flores (Indonesia). The most interesting record was the
description of a representative of the genus Phrynus from a cave on Flores (Indonesia)

by Harvey (2002). This genus (and the family Phrynidae) are exclusively American
and the finding of Phrynus in Flores is a zoogeographical riddle.
Opilioacarida From the small order of Opilioacarida (13 genera, 41 species) four
genera and seven species are known from caves (Beron 2007 and suppl.). Species of
the genera Caribeacarus and Neocarus have been found in caves of Cuba, Brazil,
and Panama (Juvara-Balş and Baltac 1977; Vasquez and Klompen 2009; Bernardi
et al. 2013). Most interesting are the two genera (Siamacarus and
Vanderhammenacarus) with three species, described by Leclerc (1989) from
Thailand caves that are endemic to these caves.
Acariformes
Trombidiformes – Prostigmata Fam. Eutrombiidae – an eyeless species that has

been described from a cave in Vietnam (Caecothrombium deharvengi Mąkol and
Gabryś 2005).
9.2 D
istribution of Cave Arachnida in Different
Geographical Areas
Europe Some orders (Schizomida, Amblypygi, Uropygi, Ricinulei, and

Holothyrida) do not live in Europe. Some others are represented on this continent,
but either do not live in caves (Opilioacarida) or are only accidental visitors of caves
or not typical cave dwellers (Scorpiones, Acariformes, Parasitiformes). From
Pseudoscorpiones to Opiliones and Araneae in the well explored European caves,
there are many species with different degrees of specialization. Troglobites are
represented only in Southern Europe. This distribution is connected with the glacial
cover of North and Central Europe during the Pleistocene glaciations. European
cave Arachnida have been analysed many times (see Encyclopaedia Biospeologica,
Beron 2015 for Bulgaria, Beron 2016 for Greece, Zaragoza 1986 for the Iberian
cave pseudoscorpions, and others).
Africa There are very few troglobitic Arachnida, especially in tropical Africa.
From the well explored caves of South Africa have been recorded such interesting
troglobites as Cangoderces lewisi Harington (Araneae, Telemidae), Purcellia [syn.
Speleosiro] argasiformis (Lawrence) (Opiliones, Pettalidae), Speleomontia caver-
nicola Lawrence (Opiliones, Triaenonychidae), and Chthoniella cavernicola
Lawrence (Pseudoscorpiones, Chthoniidae). From Botswana an endemic pseudo-
scorpion of the family Ideoroncidae has been described – the blind Botswanoncus
ellisi (Harvey et Du Preez). Spiders and other Arachnida have been described from
the caves of Kenya (Shimoni and others), one of the most remarkable being the
9.2 Distribution of Cave Arachnida in Different Geographical Areas 843
blind Opilion (Cyphophthalmi of unclear family attachement) Marwe coarctata

Shear (1985) from Cobra Cave. Troglobitic spiders and pseudoscorpions have been
described from the Kulumuzi Caves in Tanzania. The spider Paraleptoneta bellesi
Ribera et Lopez and the opilion Nemastoma tunetanum Roewer (troglobites) have
been described from Tunisia. Other troglobitic pseudoscorpions, close to the
European fauna (Neobisium, Roncus) live in the caves of Algeria.
North America north of Mexico (mostly USA) Many papers by Muchmore (see
lists in Harvey 1992 and Heurtault 1994) on cave pseudoscorpions (Chthoniidae,
Syarinidae, Garypidae, Bochicidae, Neobisiidae,). Opilions of genus Erebomaster
Cope (Cladonychiidae, syn. Erebomastridae) have been described from the caves of
the USA. Briggs (1969, 1971, 1974) published interesting data concerning relict
Laniatores from lava caves in North America.
Caribbean Islands Troglobitic pseudoscorpions from the families Bochicidae and
Chthoniidae have been described from Cuba (Dumitresco and Orghidan 1977 –
Antillobisium vachoni, A. mitchelli), Jamaica (Muchmore 1984 1991 – Troglobochica
jamaicensis, T. pecki, Tyrannochthonius hoffi). Chernetidae, living in guano, are
known from Cuba (Bitulochernes, Epactiochernes).
Rambla (1969) published three new Laniatorids from Jamaican caves, including
the true cavernicoles Stygnomma fiskei and Cynortina pecki.
Opilionids have been described from Cuban caves by Avram (1973).
Also from Cuban caves have been recorded such interesting Arachnids like
Opilioacarida (Caribeacarus vanderhammeni and Neocarus orghidani) (Juvara-
Balş and Baltac 1977) and Ricinulei (Dumitresco and Juvara-Bal).
Mexico, Guatemala, and Belize Extremely rich in caves and cave fauna, Mexico
has been explored as early as 1866 by D. Bilimek and then by Italian, American, and
other biospeleologists. Especially remarkable are the monumental studies of J. R.
Reddell and his team of the Association for Mexican Cave Studies in Austin (Texas)
(Reddell 1981). A review of the cavernicole fauna of Mexico, Guatemala and Belize
and the following volumes of this series). In his monograph Reddell (1981) enumer-
ated 122 troglobites and 420 other species from the caves of the three countries.
Since then many others have been added. Recorded are many troglobites and troglo-
philes in the following groups:
Palpigradi – fam. Eukoeneniidae.
Ricinulei – fam. Ricinoididae (Cryptocellus).
Amblypygi – fam. Phrynidae (Phrynus, Paraphrynus) – often found, but non-
troglobitic. Some have reduced or absent eyes.
Schizomida - fam. Protoschizomidae (Agastoschizomus, Protoschizomus), fam.
Hubbardiidae (Heteronochrus, Pacal, Stenochrus).
Scorpiones – the most remarkable cave scorpions in the world. Fam. Typhlochactidae
(Typhlochactas, Stygochactas, Sotanochactas, Troglocormus, Alacran), Buthidae
(Centruroides), Scorpionidae (incl. Diplocentridae) (Diplocentrus), Vaejovidae
(Vaejovis).
Pseudoscorpiones – fam. Chthoniidae (Aphrastochthonius), Bochicidae (Leucohya,

Mexobisium, Troglohya), Ideoroncidae (Albioryx, Typhloroncus), Vachoniidae
(Paravachonium, Vachonium - end. family for the caves of Mexico and Belize),
Syarinidae (Pachychitra), etc. According to Reddell (1981), 24 species of troglo-
bitic pseudoscorpions and 17 other species in the caves of Mexico has been
found at the time of writing. There are certainly more now.
Opiliones – Until 1981 there had been published 15 troglobitic opilions and 35 other
species (with troglobites are fam. «Sironidae » and Phalangodidae) from the
caves of Mexico. Remarkable is Shearogovea [Neogovea] mexasca Shear – the
only member of Cyphophthalmi in Mexico, with an uncertain position, endemic
genus, and species.
Araneae – Reddell (1981) enumerated 38 families of spiders from Mexican caves,
including 47 troglobites (Dipluridae, Theraphosidae, Agelenidae, Leptonetidae,
Nesticidae, Ochyroceratidae, Oonopidae, Pholcidae, Telemidae, and
Tetrablemmidae) and 252 other species. Most troglobites are contained in the
family Pholcidae (19). Since this time, there have certainly been many new pub-
lished discoveries.
Acari – the calculation of Reddell (1981) shows two troglobitic acari (Rhagidiidae:
Rhagidia) from Mexican caves and 81 other acari. Species of Argasidae
(Antricola and especially the endemic genus and species Nothoaspis reddelli) are
often found in bat guano.
South America
Continental South America Cave pseudoscorpions are known from Brazil

(Mahnert 2001 – 26 species, including the new genera Spelaeobochica and
Spelaeochernes).
Cave opilions have been observed from Ecuador (Metagovea philipi,
Neogoveidae; also from the collection of P. Beron), Venezuela (Stygnomma mona-
gasiensis, Stygnommatidae, Phalangozea bordoni, Agoristeridae), Brazil
(Pachyospeleus strinatii, Gonyleptidae), Argentina (Picunchenops speleus,
Triaenonychidae), Peru (Aukayacuella bordoni, Gonyleptidae), and others.
Ricinulei have been recorded from the caves of Brazil (Pinto-da-Rocha and
Andrade 2012 – Cryptocellus canga).
Schizomida are known from caves in Venezuela (Armas and Colmenares 2006,
Villareal and Teruel 2006 – Hansenochrus, Wuayuuzomus, Surazomus), and Ecuador
(Reddell and Cokendolpher 1984).
Amblypygi from caves have been recorded from French Guiana, Venezuela,
Brazil, and Colombia (Charinus, Trichodamon, Phrynus, Heterophrynus).
Scorpions are recorded from caves of Ecuador (Troglotayosicus vachoni
Lourenço) and Venezuela (troglobites).
9.2 Distribution of Cave Arachnida in Different Geographical Areas 845
Cave spiders (Araneae) are known from Ecuador, Peru, Brazil, Venezuela, and
Chile (the families Ctenizidae, Ochyroceratidae, Oonopidae, Theridiosomatidae,
Hahniidae, Loxoscelidae, Pholcidae, etc).
Galapagos Islands Juberthie (1970) described from lava caves the opilion
Galanomma microphthalma (Gonyleptidae, Prostygninae), with reduced eyes.
Troglobitic spiders (Lygromma anops Peck et Shear, L. gertschi Platnik et Shabad)
have also been described.
Hawaiian Islands Muchmore (1979, 1983) described the troglobitic pseudoscor-
pions Tyrannochthonius howarthi, T. stonei, and T. pupukeanus from Hawaiian lava
tubes
Temperate Asia This includes Asia Minor, Arabian Peninsula, the Levant, Central
Asia, Siberia, Mongolia, Korea, North China, Japan, and Afghanistan. The biospe-
leological research in Turkey, South Korea, Afghanistan, Japan, and the Asiatic part
of the former USSR revealed many cave spiders, some mites, pseudoscorpions, and
opilions. The thermophilic orders are generally missing from these areas and totally
missing from the caves. Two exceptions are Charinus ioanniticus (Amblypygi)
from caves in Greece, Turkey, and Israel and Ch. israelensis in Israel (Miranda et al.
2016).
Indomalayan Asia For a long time the main information on the cave dwellers of
tropical Asia was obtained from the thorough investigations in Siju Cave (India) and
Batu Cave near Kuala Lumpur. Together with the material collected in some African
caves, this circumstance led even prominent zoologists like Jeannel and Vandel to
think that tropical caves do not contain terrestrial troglobites. The expeditions in
Indonesia, Malaysia, South China, Vietnam, Laos, and some other countries changed
this view completely. Now we know many troglobites from these areas, including
Arachnids.
Australia, Tasmania, New Guinea The very intensive research, especially in the
caves of Western Australia, resulted in many new data concerning cave Arachnida
(Harvey, Humphreys, and others; many papers).
In Australian caves have been recorded, among others:
Schizomida – found often in Australian caves (Draculoides vinai, troglobite, D.
bramstokeri – caves of Cape Range; out of the 53 species of Schizomida in
Australia are known from caves).
Pseudoscorpiones – many species of different families – Chthoniidae, Cheliferidae,
Chernetidae, Cheiridiidae, Hyidae, and Syarinidae (Harvey 1991, Heurtault
1994).
Scorpiones - Volschenk and Prendini (2008) described the first troglobitic uro-
dacid – Aops orthodactylus (Urodacidae) from Australia.
Opiliones – the many descriptions of cave Laniatores (Triaenonychidae) in the
Australian continent and Tasmania (Hunt 1990, 1991, 1992, and other papers;
Hunt and Hickman 1993) have been analyzed by Rambla and Juberthie (1994).
In Tasmania Hickmanoxyomma and Lomanella are widespread, some are
troglobiomorphic.
Araneae – in many Australian caves there are troglobitic spiders (Desidae –
Forsterina, Dipluridae – Troglodiplura, Linyphiidae – Chthiononethes,
Dunedinia, Nesticidae – Nesticella, Theridiidae – Steatoda, Pholcomma,
Symphytognathidae – Anapistula), also Cycloctenidae, Filistatidae, Pholcidae,
and Stiphidiidae.
Acari – Uroobovella coprophila (Uropodidae) and other guanobionts live in many
bat caves.
During the British Speleological Expedition to Papua New Guinea in 1975 we col-
lected many remarkable cave animals in the high areas in the heart of the island,
but unfortunately most of them are still unidentified. Only some pseudoscorpions
have been identified by Beier (1982), including the new species Afrosternophorus
cavernae (Beier), Sternophoridae. A preliminary list of many new species of
spiders was published by Brignoli (1982), but his untimely death stopped the
final descriptions. Similar was the fate of the opilions, sent to Dr V. Šilhavý.
New Zealand Very interesting and very limited in Arachnid orders, the cave fauna
here remain mostly Araneae, Opiliones, and some mites.
Opiliones – some opilions (Hendea Roewer, Forsteropsalis Taylor) live in the caves
of Waitomo and other caves (Hunt 1972; Rambla and Juberthie 1994; Taylor
2011). Among the Pettalidae (Cyphophthalmi), in the caves are found members
of Rakaia Hirst.
Araneae – spiders of Dipluridae, Hexathelidae, and Holarchaeidae.
9.3 Conclusions
Exploring the amazing cave fauna of the Hawaiian lava tubes, Howarth (1980) came
to the conclusion that the troglobites are not limited to the ice-free regions of the
Northern Hemisphere, but, on the contrary, terrestrial troglobites can now be
expected to be found in any region that is old enough and in which there are exten-
sive caves with an adequate moisture supply and a continuous equitable environ-
ment and food supply for colonization. This bioclimatic model has been demonstrated
many times since then in different countries, including tropical countried.
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Chapter 10
High-Altitude Arachnida (Partly
from Beron 2008, Updated in Beron 2016)
Abstract The study of extreme habitats and the animals adapted to them (e.g., high
mountain fauna) is interesting in many ways. It gives us the opportunity to reveal many
ecophysiological mechanisms for adaptation to extreme environments. The recent
detailed studies from large and difficult to explore mountains have provided specialists
with the opportunity to describe hundreds of new taxa and to clarify the composition of
entire faunas, unknown until then. A good example is the campaign in the Himalayas
of the German Zoologist Prof. J. Martens, who in 3–4 years managed to significantly
contribute to the knowledge of some groups living in the huge mountain systems that
form the boundary between two zoogeographical kingdoms. Little by little, the altitu-
dinal limits for the separate taxa and whole groups are being discovered, thanks to
fieldwork by such intrepid travelers as H. Janetschek, H. Franz, and others.
10.1 Introduction
Enormous areas of Northern Eurasia are occupied by plains, with a larger surface
area than Europe, but there is no point higher than 2000 m. The Alpo-Himalayan
system provides countless possibilities for speciation and the creation of specific
communities with restricted areas. By studying the high mountain species, we can
detect the refugia from the Pleistocene and pre-Pleistocene and follow the
connections with the northern species, disrupted after the climatic changes in the
Holocene. The so-called Arctic-Alpine and Boreomontane species are witnesses to
changes which cannot be established by paleontological evidence. Many neoendemic
species (living often only on one mountain) have remained, until recently,
undescribed and this is reducing the overall knowledge of local species.
In discussing the species of the Old World, it is worth mentioning the Afrotropical
mountains, where ice-covered summits over 5000 m rise like islands amidst the
completely different environment of the savannah. The fauna of these mountains
provides a treasury of relicts from former connections with the mountains of
Eurasia. Generally speaking, the study of mountain fauna in light of the theory of
island biogeography would provide the key for understanding the affinities and
differences between them. This is yet to be achieved as the theory is relatively new

https://doi.org/10.1007/978-3-319-74418-6_10
854 10 High-Altitude Arachnida (Partly from Beron 2008, Updated in Beron 2016)
(MacArthur and Wilson 1969) and the degree of study was (and in many cases still
is) incomplete to allow adequate comparison.
While the Alps are relatively well explored and their fauna has been known since
the beginning of the 20th century, knowledge of the fauna from the mountians of
Middle Asia, the Himalayas, and the African mountains is being accumulated at a
much slower rate. Many mountains remain unexplored, even in Europe. A typical
example is Pirin in Bulgaria – up until 1915, this remarkable mountain remained
virtually unknown, being much less explored than New Guinea or Kilimanjaro.
In the last several decades, some important monographs (mostly on high-altitude
insects) have been published. We should mention the monumental books of
Prof. H. Franz “Die Nordostalpen im Spiegel ihrer Landtierwelt. Eine
Gebietsmonographie” (1954), “Ökologie der Hochgebirge” (1979), and of Mani
“Introduction to High Altitude Entomology” (1962), “High Altitude Insects” (1963),
“Ecology and Biogeography of High Altitude Insects”(1968), and “Ecology and
Biogeography in India” (1974, Ed.). These monographs, however, give little
attention to species other than insects. In “Ecology and Biogeography of High
Altitude Insects” Mani (1968), the discussion on the non-insect Arthropods (mainly
on Myriapoda and Arachnida) takes up four of 527 pages. Until now, the terrestrial
non-insect Arthropods have been rather neglected and no analysis has been made
concerning their presence in high mountains. One of the reasons for this is limited
knowledge on the taxonomy of terrestrial Isopods, Arachnids, and Myriapods
compared to most groups of insects. For these species, there was either no data or
the existing data were obsolete and needed revision. With extensive research in the
Himalayas and other mountains and modern taxonomical revisions in the last
decades, it has become possible to undertake a general review of the accumulated
data. This was done by Beron in 2008.
In terms of high-altitude Arachnida, we have to define the criteria for the con-
cepts of “high mountain” and “high mountain environment.”
In these definitions, altitude is important, although according to Mani (1968),
“…altitude as such is really of very little biological significance, except as merely
an indirect measure of certain complex ecological conditions and specializations in
organisms.” What we usually term a “high mountain environment” (i.e., lack of tall
forest, long-standing snow cover, and low temperatures), in Scandinavia starts at an
altitude of as low as 400 m whereas in the Himalayas high forests grow at 4200 m.
Is the forest flora and fauna at 4000 m a high mountain environment? Or are the
Arctic flora and fauna in the low Lapland? De Lattin (1967) includes the polar and
high mountain areas without tree vegetation in the bigger domain (biocycle)
oreotundral, with only the high mountain part called “oreal.”
Mani (1968, p. 8) presents a clear definition for high altitude: “Considered from
the point of view of ecology, biogeography and evolution, high altitude is the region
of mountains that is sufficiently elevated above the surrounding lowlands to be
characterized by significant climatic differences, different flora and fauna.
10.1 Introduction 855
Pronounced ecological differences and characteristic high-altitude specializations

appear among the mountain autochthonous insects in the so-called alpine zone that
commences at elevations above 2000–2500 m only.”
An altitude of 2000 m is most often the beginning of the high-altitude environ-
ment because we are more familiar with European high mountains. These moun-
tains are situated in Central and South Europe. North of the Pyrenees, in the Alps
and Tatra mountains, there are no mountains higher than 2000 m, except for the
Halhopingen (2469 m) in Norway and Kebnekayse (2123 m) in Sweden.
In the Alps, the Pyrenees, Caucasus, Tatras, Rila, and Pirin, the upper limit of the
closed high forest is usually 1900–2200 m, most often 2000 m. Higher we find areas
covered with Mountain Pine Pinus mugho (Alps, Rila) or Rhododendrons (Pyrenees,
Caucasus). Above these areas, the pastures or, as in Corsica, stony areas start. In
some places, the upper forest limit is either artificially lowered by human activities
(e.g., Stara Planina or Balkans, Bulgaria) or climbs in exclaves up to 2300–2400 m
(Pirin). In some southern mountains (Sierra Nevada, Taygetos, Crete), the landscape
above 2000 m is quite different and is more similar to that of the mountains of North
Africa or Middle Asia with their Cushion Plants of Tragacantha sp.
There is disagreement on what is forest and what is the upper forest limit. There
are different numbers regarding the height of the trees and the distance between
them. Specialists argue about whether the curved-stem Mountain Pine (Pinus
mugho) and the Rhododendrons (sometimes more than 3 m tall) could be considered
part of the treeless zone. This discussion concerns the European mountains, as in the
Himalayas Rhododendrons grow up to 10 m tall, even above 4000 m. This high-
altitude forest is unique to this extreme environment. Such forest is subject of the
actions of universal factors (ultraviolet radiation, reduced atmospheric pressure,
sharp variation of the nycthemeral temperature). These factors form the basis for
dealing with such heterogenous complexes as the treeless European orophyte belt,
the high Asiatic steppe, the arid “moonscapes” of Karakorum and Kunlun, and the
mountain forests and bushes of the Afrotropical mountains and the Himalayas.
As we must nevertheless state an arbitrary limit above which the fauna could be
listed and analyzed as high-altitude fauna, and in the present study an altitude of
2200 m was selected. In Europe, this limit eliminates almost everywhere where
there are inhabitants of the ecotone (forest dwellers, rarely appearing above the
upper forest limit or being found in the treeless zone). In other mountains, this limit
is usually in the forest zone and is not of any special importance, but it would not be
justified to compare the fauna living at 2000 m in Europe with that at 4000 m,
(where the treeless zone starts in other mountains). Therefore, we do not have the
opportunity to compare the influence of universal factors.
To emphasise the high-altitude dweller (the euhypsobionts), we have selected
another arbitrary boundary (3500 m) above which almost everywhere we find
orophyte environments, most often treeless. At this altitude, the action of the
universal factors is much stronger and the fauna consists of species that are, beyond
a doubt, well adapted to these factors.
Varga (1975) stated: “The concept of the oreal fauna covers two main eco-
geographical groups: the oreal fauna of the humide mountains with definite alpine
(glacial-fluviatile) geomorphological features, called Alpid, and the oreal fauna of
the arid-semiarid high mountains (with predominating physical corrosion and frost-
fluctuattion, resp.) called Xeromontan, where the high regions are bordered by no or
a very scattered timberline. The expansive taxa of the alpid fauna are connected –
caused by quaternary faunal fluctuations – with the tundral fauna, while the
expansive members of the xeromontan fauna penetrated towards the steppe and
semi-desert (eremic) zones” (Varga 1975).
With regard to the high mountains of New Zealand, we notice the categories
outlined by Fleming (1963), who concluded that the flora and fauna now endemic
to the Alpine Zone have only had a short existence as alpine species in New Zealand.
Some Terms and Notions:
Different sources and various languages have used different terms and notions,
sometimes in arbitrary, different, and unclear ways. We therefore consider it
necessary to clarify and put some order in the terminology.
Alpine (1) pertaining to the Alps; (2) belt or zone in the high mountains, usually
between the subalpine and the subnival belts.
Alticolous Living in a high-altitude environment.
Afroalpine Part of the Afromontane region, altitudinal belt, sometimes taken for a
region on its own, found only on the highest mountains of Africa (Kilimanjaro, Mt.
Kenya, Ruwenzori, Meru, Elgon, Semien), above 3400–4100 m, and in the
Drakensberg higher than 2860 m (Austro-Afro-alpine belt). Afroalpine can also
refer to the typical flora and fauna of this area.
Afromontane Region in the mountains of tropical Africa, from Sierra Leone to
Somalia and from Sudan to Cap. In phytogeography, some parts of these mountains
are called “Afroalpine regions,” within or separated from the Afromontane region.
Afromontane can also refer to the fauna and flora of this area.
Arctic Zone Term used instead of “nival zone” for the area above the permanent
snowline. Term should be avoided when speaking of areas outside the arctic region.
Arcto-Alpine Type of distribution of species, living in the Arctic and Subarctic
tundra and in the oreal of the mountains in the middle and southern parts of Europe,
Asia, and North America, above the upper forest limit. The origin of this disjunct
distribution is in the postpleistocene withdrawal of the glaciers from uninterrupted
areas. Some of these species have been mentioned in papers published in Europe by
Holdhaus (1912 1954) and other authors and in Bulgaria by Beron (1969) under the
name “Boreo-Alpine”.
Biochore A group of similar biotopes (such as temperate forests) (Merriam-

Webster Dictionary).
Boreo-Alpine Term from older publications, used for species living in the northern
parts of Eurasia and North America and in the mountains of the southern parts of
these continents. This term describes two types of distribution: the Arctic-Alpine
and the Boreomontane. It is preferable to use the latter terms and to avoid the unclear
term “Boreo-Alpine.”
Beron (1969) included a list and analysis of the Boreo-Alpine (s. lato) fauna of
Bulgaria. The problem with the Boreo-Alpine type of distribution has been discussed
by Holdhaus (1912, 1954).
Boreomontane Type of distribution including the northern coniferous forests of
Eurasia and North America and the mountains of South Europe, Caucasus, and
Central Asia.
Cryonival Living in the upper nival zone in very low temperatures.
Euhypsobiont Term introduced here to name the fauna species permanently living
at or higher than an altitude what is considered high worldwide (subject to the influ-
ence of universally acting factors like atmospheric pressure, radiation, low tempera-
tures (at least at night), etc.). In this study, the limit for this high altitude is 3500 m.
Using this threshold, we eliminated the temporary or not typical high-altitude spe-
cies, most of the forest forms, the inhabitants of the ecotone immediately above the
forest line, and part of the forest or lowland species blown by the wind in the nival
zone.
Hypsobiont Fauna species living permanently in areas of high altitude (in Europe
generally above 2000 m, in other areas, this can be much more).
Hypsoendemic Endemic to the given area, but living only or mostly in the oreo-
tundral area.
Montanmediterranean Autochthonous element of the mountain fauna of the
Mediterranean.
Nival Belt or zone above the permanent snowline in the high mountains (in Europe
above ca. 3000 m), the highest zone of the mountains, usually above the upper
alpine or above the subnival zone. Hypsobionts in Europe live above 2000 m, in the
alpine, subnival, and nival zones.
Nunatak Inuit word, meaning an isolated rocky outcrop in the glacier. The nuna-
taks are essential for the survival of some fauna species in the high mountain (“nun-
atak fauna”).
Oreal Definition from De Lattin (1967) and Sedlag and Weinert (1987):
1 . Biochor of the high mountain area above the upper forest limit.
2. The flora and the fauna of this domain of life.
Oreotropical belt A notion proposed for the belt of mountain forest above the
upper forest line in tropical and subtropical countries.
Oreotundral According to De Lattin (1967) and Sedlag and Weinert (1987):
1. Biochor of the high mountain area above the upper forest limit and the area north
of the polar forest limit.
2. The flora and the fauna of the described area.
Oreotundral is often used as a synonym for “Arcto-Alpine.” However, despite
some common features in the general habitats and some shared species (the Arctic-
Alpine elements), there are substantial differences between the Oreal and the
Tundral. The photoperiodism is different, as is the radiation, the atmospheric
pressure, and oxygen contents; sometimes there is no forest belt at all and the steppe
goes uninterrupted from the lowland to over 4000 m. The upper parts of the
mountains are richer in shelters, with rock debris, many stones etc., which is not the
case in the usually low and level tundra.
Orobiome Synonym of Oreal (the area or the high mountain above the upper forest
line).
Orophyte zone all belts above the upper forest limit. In this zone, there is typical
low vegetation containing many high-altitude endemics. Most species do not occur
in the lower belts.
Preglacial relict Taxon living in glaciated areas since before the Pleistocene
glaciations.
Subalpine Belt, zone or subzone between the high mountain and the alpine zones.
Subnival Belt, zone, or subzone between the nival (eunival) and the upper alpine
zone, or belt. A permanent but parcial snow cover is typical for this zone.
Taxogradient the change in the number of taxa with the increase of the altitude.
In the papers of La Greca (1955, 1958) have been analysed the climatical and
paleoclimatical factors for the forming of the high mountain entomofauna. Meyer
and Thaler (1995) outlined the animal diversity at high altitude in the Alps. The
present alpine ice cover, analysed be Ostrem (1974), is strongly reduced since.
8000
7000
6000
5000
4000
3000
2000
1000
0
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19
High-Altitude Arachnida – the different orders

1. Solifugae – up to 4000 m
2. Schizomida – up to 2600 m
3. Scorpiones – up to 4600 m
4. Pseudoscorpiones – up to 5000 m
5. Opiliones – up to 5600 m
6. Laniatores – up to 4600 m
7. “Palpatores” – up to 5600 m
8. Araneae – up to 6700 m
9. Mygalomorphae – up to 2600 m
10. Araneomorphae – up to 6700 m
11. Opilioacarida – up to 2500 m
12. Acariformes – up to 5800 m
13. Acaridida – up to 3500 m
14. Prostigmata – up to 5100 m
15. Oribatida – up to 5800 m
16. Parasitiformes – up to 5488 m
17. Holothyrida – up to 2650 m
18. Mesostigmata – up to 4550 m
19. Ixodida – up to 5488 m
10.2 M
ajor Groups of Arachnida in the Old World Living
At or Above 2200 m
Arachnida All eight orders of Arachnida known in the high mountains of the Old
World are also represented in the oreal of the Americas. The orders not represented
in the alticolous fauna of Eurasia and Africa (Palpigradi, Amblypygi, Uropygi,
Ricinulei, Opilioacarida) are also absent from the high mountains of the Western
Hemisphere, although these orders do occur there.
Scorpiones The Scorpiones order contains nine families (Polis 1990), eight fami-
lies (Nenilin and Fet 1992), 18 families (Fet et al. 2000), or 20 families in the pres-
ent book. Six families in the Old World are known to contain species living at
altitudes over 2200 m. Euscorpiidae are only represented in the high mountain
environment by genus Euscorpius Thorell (Balkan Peninsula 2569 m (Albania) and
Anatolia 2600 m). Scorpiopidae can reach 5000 m in the Himalayas (genus
Scorpiops Peters). Several genera of Buthidae are known to live up to 3500 m on
Meru and in Central Asia and the genus Scorpio L. (Scorpionidae) has been recorded
at 2300 m in Atlas. From Tibet (4600 m) the Tibetiomachus himalayensis Lourenco
et Qi, 2006 has been described. This is a strange and unique representative of the
family Liochelidae in the harsh Tibetan plateau. In Asia, scorpions have been found
up to 4600 m (Tibetiomachus himalayensis Lourenco et Qi, 2006, Tibet, Hormuridae)
and in Africa – up to 3500 m (Uroplectes fischeri Tullgren, 1910, Meru, Buthidae).
Polis (1990) writes: “Such high-elevation species are all small. They feed on a
diverse array of arthropods that are also found at these heights (Mani 1968), and
their small size may be due to the short period during which they are able to forage).
Surprisingly, high-altitude scorpions live under rocks, in scrapes, and in relatively
short burrows (…), rather than in deep burrows with terminal chambers below the
frost line.”
Solifugae From the 12 families in the Solifugae order, nine are known in the Old
World and three are known only in the Americas. At least five of the Old World
families contain species that live at altitudes higher than 2200 m: Gylippidae (up to
3500 m in Afghanistan), Karschiidae (up to 3500 m in Afghanistan), Galeodidae (up
to 4000 m in Tajikistan), Daesiidae (up to 2350 m in Anatolia), and Solifugaee. We
know only about ten species of these warm-loving Arachnids that live in the high-
altitude environment, all of them in Asia and Caucasus. The highest record for the
Old World is of Galeodes setulosus Birula from Tajikistan (4000 m).
Schizomida These small Arachnids are warm-loving and clearly avoid the high
mountain environment. They have been found up to 2200 m in Vietnam (“Schizomus”
peteloti Remy), 2600 m in Tanzania (Reddell and Cokendolpher 1995), and have
been recorded by Kraus (1957a, b) at 3100 m in Columbia (Surazomus cumbalensis).
This order does not live in Europe or in the Central Asian mountain systems.
10.2 Major Groups of Arachnida in the Old World Living At or Above 2200 m 861
Pseudoscorpiones The Pseudoscorpiones order in the Old World includes about

3000 species of approximately 435 living genera (429 in Harvey, 1990). According
to the latest revision of Harvey (1992), there are 24 families. Our analysis (Beron
2002b and more recent) of Pseudoscorpions in the Old World found that the altitude
of 2200 m is reached by 174 species belonging to at least 71 genera and 16 families.
Only a small number of Pseudoscorpions live in the European mountains higher
than 2000 m: two in the Pyrenees, one in Sierra Nevada, six in the Alps, one in the
Apennines, two in the mountains of the Balkan Peninsula, and 12 in Caucasus. Only
four species reach or live higher than 3000 m: Neobisium jugorum L. Koch (Alps,
3600 m), N. nivale Beier (Sierra Nevada, 3481 m), N. anatolicum Beier (Caucasus,
3000 m), and N. carcinoides Hermann (Alps, 3000 m and Pirin, 2914 m. The
members of Neobisium are adapted to the highest parts of European mountains.
Within the 2000- to 3000-m belt in Europe (including Caucasus), species belonging
to the genera Chthonius (Ch. tetrachelatus Preyssler, in Caucasus up to 2500 m, in
Iran up to 2900 m), Roncus (R. microphthalmus Daday up to 2200 m in Caucasus),
Chernes (Ch. montigenus Simon up to 2740 m), from the genus Neobisium also N.
bernardi Vachon (up to 2800 m), N. delphinaticum Beier (up to 2850 m), N. noricum
Beier (up to 2500 m), and N. dolomiticum Beier (up to 2400 m) have been found.
The studies of Tullgren (1910), Beier (1955 and many other papers), Mahnert
(1981 and the series about Kenya), and other researchers and our own investigations
from Kenya, Tanzania, and Uganda have shown that in the East and Central African
mountains there are at least 26 species of Pseudoscorpiones that live at or above
2200 m. At least 13 of these species reach 3000 m, four reach 3500 m, and only
Titanatemnus palmquisti is known to live above 4000 m.
The high-altitude Pseudoscorpions in South and North America belong to the
families Chthoniidae (Austrochthonius), Pseudogarypidae (Pseudogarypus),
Olpiidae (Olpiolum, Progarypus, Serianus, Stenolpiodes, Stenolpium), Cheliferidae
(Parachelifer, Haplochelifer, Dactylochelifer, Hysterochelifer), Chernetidae
(Lustrochernes, Parachernes), and Withiidae (Parawithius). The families
Chthoniidae, Olpiidae, Cheliferidae, Chernetidae, and Withiidae are common to the
Old World. Pseudogarypidae occurs in North and South America and in Australia.
None of the species and only two genera of Cheliferidae (Hysterochelifer and
Dactylochelifer) are shared with the oreal of the Old World.
Of the remaining families represented in the high-altitude fauna of the Old
World, Lechytiidae, Tridenchthoniidae, Geogarypidae, Ideoroncidae, Syarinidae,
Cheiridiidae, and Sternophoridae are represented in the Americas, but are not known
there above 2200 m. Hyidae does not live in the Western Hemisphere. Garypidae
reaches high altitudes in South America, but not in the Old World.
The highest altitude reached by Pseudoscorpions in South America is 4100 m
(Teratolpium andinum Beier).
Examples of Pseudoscorpions living in the Americas at or higher than 3500 m:
Fam. Olpiidae: Teratolpium andinum Beier – 4100 m (Peru).
Fam. Garypidae: Progarypus peruanus Beier – 3900 m (Peru).
Fam. Chernetidae: Parachernes loeffleri Beier – 1500–3800 m (Peru).
Opiliones The Opiliones order has more than 6500 species in four suborders. The
short-footed Cyphophthalmi do not live above 2000 m. The numerous species and
genera, belonging to Laniatores, are confined mostly to tropical countries, including
high in the mountains. In Europe, only Holoscotolemon oreophilum Martens is
known to reach 2000 m. All other European Opilionids found above 2000 m (33 spp.
in the Alps, 11 in Bulgaria) belong to suborders Eupnoi and Dyspnoi (both in the
former Palpatores) and they prevail in the Holarctic. In the mountains of tropical
Africa, Laniatores are predominant. In South America, we observed very high
(about 5000 m in Peru) representatives of the Neotropic families e.g., Gonyleptidae
(not identified yet).
In the Old World, at least 266 Harvestmen species (109 Laniatores and 157
Palpatores) reach or go higher than 2200 m. In Europe, 47 species are known above
this altitude. They belong to six families: Phalangiidae (25 spp. of the genera
Dicranopalpus, Gyas, Lacinius, Leiobunum, Megabunus, Metaplatybunus, Mitopus,
Odiellus, Opilio, Parodiellus, Phalangium, Platybunus, and Rafalskia),
Sclerosomatidae (four spp. of Astrobunus), Ischyropsalididae (five spp. of
Ischyropsalis), Sabaconidae (one Sabacon), Trogulidae (two Trogulus), and
Nemastomatidae (ten spp. of Mitostoma, Nemastoma, and Paranemastoma). Only 14
species reach or go above 2500 m and only two members of genus Mitopus cross into
the subnival and nival zones (over 3000 m and 3500 m, respectively). The alpine
endemic Mitopus glacialis has been found at altitudes up to 3675 m (around the
height reached by its analogue among the Pseudoscorpions, Neobisium jugorum).
The widespread mountain species Mitopus morio goes up to 3300 m (Beron 2002c).
Above altitudes of 2200 m in the Old World, there are 11 families, but not all of
these families are the true high mountain dwellers. Three families of Laniatores
(Triaenonychidae, Oncopodidae, and Podoctidae) live in tropical countries below
3000 m (in the zone of tropical forests) and are not considered to be true hypsobionts.
Only three families within the Old World Laniatores include true members of the
high mountain fauna: Phalangodidae, Biantidae, and Assamiidae. Only
Phalangodidae also live in Europe. The mostly tropical Laniatores could be
considered as members of the hypsobiont species, inhabiting areas above at least
3000 m. The two last mentioned families also live above 4000 m (Biantidae up to
4250 m in Nepal, Assamiidae up to 4600 m on Kilimanjaro). Ascending from 2200–
2999 m and from 3000–3999 m the number of both the genera and species decreased
almost twice as much with every 500 m. Only one species lives higher than 4500 m –
Hypoxestus accentuatus Sörensen (Assamiidae) on Kilimanjaro.
If we look closely at the “Palpatores”, which are predominant in the Palaearctic,
we find some peculiarities. From the five high mountain families, two
(Ischyropsalididae and Nemastomatidae) do not reach 3000 m. Some species of
these two families live in the European mountains. In the Alps, Pyrenees, and
mountains of the Balkan Peninsula at 2700–2800 m, the environment is purely
alpine and cannot be compared with the tropical rainforest at the same altitude on
the slopes of Kilimanjaro.
The Sabaconidae family has one member living up to 2300 m in the Pyrenees,
but also six in the Nepal Himalayas, including one candidate for the world record in
10.2 Major Groups of Arachnida in the Old World Living At or Above 2200 m 863
altitude. Sabacon dhaulagiri Martens has been recorded up to 4250 m, but Prof.
J. Martens (in litt.) has found one species of Sabaconidae even above 5000 m. The
Sclerosomatidae family has one species in Nepal reaching 3200 m.
Phalangiidae is the family containing the bulk of the high-altitude Opilions
(more than half of all genera and species known in the order occur above 2200 m).
The number of genera decreases with a certain regularity from 2200 m (51) to
2500 m (40), 3000 m (33), 3500 m (20), and 4000 m (10). The same regularity is
observed with number of species: 2200 m (126 spp.), 2500 m (94 spp.), 3000 m (60
spp.), and 3500 m (33 spp.). Very few species occur at altitudes higher than 3500 m.
Only the representatives of Mitopus live over 3000 m in Europe. These are
endemic to the Alps (M. glacialis), and are found as high as 3675 m; as are the
widely distributed M. morio.
If we analyze the high-altitude fauna of Opiliones in the Himalayas, we come to
the following conclusions:
1. Only two genera (Opilio and Sabacon) and none of the species are in common
with the European fauna. Out of the seven Himalayan families (in the high
Himalayas), three (Phalangodidae, Phalangiidae, and Sabaconidae) also live in
Europe, the last two in the high mountains of this continent.
2. The best represented (50 out of 87 species living above 2200 m, or more than 57
%) is the large Phalangiidae family. Its representatives are the highest living
Opilions in the world, e.g., Homolophus (= Euphalangium) nordenskioeldi
(L. Koch), 5600 m and Himalphalangium palpale (Roewer), 5540 m.
3. The ratio of the genera of Laniatores versus “Palpatores” is 9:21, of the species
is 29:58 (for both approx. 1:2). For comparison: in the mountains of Central and
East Tropical Africa, the ratio of the Laniatores and Palpatores genera is 19:6, of
the species is 61:22 (for both approx. 3:1). In the high mountains of Europe,
Laniatores do not live higher than 2000 m and the dominance of “Palpatores” is
undisputed.
4. Despite the fact that in the Himalayas the altitudinal span of 2200–3500 m is
entirely in the forest zone, with this 1300 m ascent, the number of Opilionid
species decreases more than six times (from 87 to 14 species). Only five species
reach the altitude of 4000 m, on which in the Himalayas grow tall forests. Higher
than 4250 m, there are only two representatives of “Palpatores”. At this altitude,
the highest living member of Laniatores (in Asia) was found – Biantes perne-
palicus Martens. Only one of the Laniatores of the Old World – the African
Hypoxestus accentuatus Sörensen on Kilimanjaro (up to 4600 m) – lives at alti-
tudes higher than that.
Among the many interesting Opilionids recorded from the high mountains of
Central and East Africa, two species have been found above 4500 m, 12 species
above 4000 m, 26 species above 3500 m, and 34 species above 3000 m (on the
mountains Kilimanjaro, Kenya, Meru, Elgon, Aberdare, Uluguru, Oldeani,
Ruwenzori, Hanang, Semien).
10.3 L
evel of Knowledge on Opiliones in the High Mountains
and Credibility of Conclusions
In Europe and in the Himalayas (partly also in the mountains of Central Asia),
Harvestmen are well known and we consider the data presented here as credible. In
the high mountains of China or New Guinea, many new taxa are likely to be found
and some older descriptions should be revised. In the European mountains, this has
been done by specialists (firstly by Prof. J. Martens) and the data are reliable. In the
mountains of Central and East Africa, the publications of Roewer (1961), Lawrence
(1962), Sörensen (1910), Loman (1902), Goodnight and Goodnight (1959), and
other older workers are based on material collected by non-specialists and are con-
sidered incomplete. They also need taxonomical revision. Considerable collections
from the higher parts of Ruwenzori, Kilimanjaro, Elgon, Karakorum, New Guinea,
Sumatra, Borneo, and the Himalayas by the authors remain unstudied. Nevertheless,
we believe that the basic ratio between Laniatores and “Palpatores” and between the
families will remain unchanged.
1. Triaenonychidae – up to 2500 m 6. Phalangiidae – up to 5600 m

2. Oncopodidae – up to 2600 m 7. Sclerosomatidae – up to 5200 m
3. Podoctidae – up to 2410 m 8. Ischyropsalididae – up to 2700 m
4. Biantidae – up to 4250 m 9. Sabaconidae –more than 5000 m
5. Assamiidae – up to 4600 m 10. Nemastomatidae – up to 3800 m
10.4 M
aximum Altitudes of Spiders Living Above 2200 m
in the Old World
Araneae The authors’ list of spiders living at or higher than 2200 m in the Old
World contains data concerning 1381 species of 315 genera and 42 families (out of
112 families of spiders in the world, according to the system of Platnick – Electronic
version). These data lead us to make some interesting conclusions:
1. High-altitude spiders belong almost exclusively to the suborder Araneomorphae.

The only four species (African and Asian) of infraorder Mygalomorphae are not
known from localities higher than 2500 m, at which altitude there is still rainforest
in tropical Africa. In the real high mountain environment (in the tropics of the
Old World, this is usually above 3500 m), only representatives of Araneomorphae
are found.
2. From 42 families of spiders, 18 do not reach 3000 m. According to the altitude
reached by them, the families are distributed as follows:
10.4 Maximum Altitudes of Spiders Living Above 2200 m in the Old World 865
42 fam. reach or live higher than 2200 m

33 fam. “ “ “ 2500 m
24 fam. “ “ “ 3000 m
22 fam. “ “ “ 3500 m
15 fam. “ “ “ 4000 m
11 fam. “ “ “ 4500 m
6 fam. “ “ “ 5000 m
4 fam. “ “ “ 5500 m
One family (Lycosidae) lives higher than 6000 m.
Besides Lycosidae, the altitude of 6000 m is almost reached by members of the
family Salticidae (known up to 5947 m), and the altitude of 5000 m – also by the
Gnaphosidae (4980 m), Dictynidae (4930 m), and Thomisidae (4880 m).
3. The 11 families reaching or higher than 4500 m represent the habitats of the
spider species in the highest regions of the Old World. This altitude is reached in
Europe only by the Alps and Caucasus, and in Africa only by the mountains
Kilimanjaro, Kenya, Ruwenzori, Meru, Semien, and Karisimbi. These 11
families contain 1099 from all 1381 species of spiders known in the Old World
above 2200 m (almost 84 %). Eight of these families contain more than 20
species each or 1081 species as a whole. The 11 families living at or above
4500 m are (with the altitude reached by them):
Total species >2200 m max alt. species >4500 m

Lycosidae – 112 species – 6100 m 9
Salticidae – 136 species – 5947 m 12
Linyphiidae – 572 species – 5545 m 19
Hahniidae – 24 species – 5500 m 3
Gnaphosidae – 112 species – 4980 m 6
Dictynidae – 11 species – 4930 m 1
Nesticidae – 1 species – 4900 m 1
Thomisidae – 65 species – 4880 m 1
Tetragnathidae – 6 species – 4600 m 1
Theridiidae – 32 species – 4600 m 1
Araneidae – 31 species – 4500 m 1
Total 1102 species 55
4. From these data, the bulk of the species living in all high mountains of the Old
World belong to the family Linyphiidae. This family contributesat least 600 spe-
cies to the 1381 species of spiders living at or above 2200 m in the Old World.
Interestingly, almost the same proportion (~50 %) exists by the genera i.e., the
152 genera of Linyphiidae recorded above 2200 m represents 46.34 % of the
total of 324 genera. The altitude of 3500 m is reached or surpassed by 203 spe-
cies of spiders (from 61 genera). The Linyphiidae family contributes 107 species
recorded above 3500 m to this total, They are almost half of the 203 species of
spiders reaching this altitude in the Old World; 4000 m is reached or surpassed
by 46 species of Linyphiids of 21 genera (almost half of the 85 species of spiders

living at this altitude in the Old World), 5000 m – by 4 Linyphiid species, belong-
ing to 4 genera (Erigone, Gongylidium, Hilaira, Mughiphantes – 31 % of the 13
species of spiders living at this altitude).
Clearly none of the genera could be considered as dominant in the high moun-
tains of the Old World (Mughiphantes has at least 29 high mountain species and has
been recorded at altitudes of up to 5545 m). Pardosa and Xysticus are also numerous
and widespread at higher altitudes.
Concerning the mountains of Eurasia, the high-altitude spiders are relatively
well-known (Pyrenees, Alps, Pirin, and other Bulgarian mountains, Caucasus, and
the Himalayas). As a model for comparison, we use the most numerous and varied
family in all high mountains – Linyphiidae (Erigoninae included). Over 2200 m in
separate mountain systems, the following genera and species of Linyphiidae are
represented:
Pyrenees – 39 genera, 65 species
Alps – 44 genera, 101 species
Rila – Pirin – Vitosha – Stara planina – 30 genera, 55 species
Caucasus – 51 genera, 84 species
Himalayas – 21 genera, 52 species
In the mountains of Eurasia live at or above 2200 m representatives of at least
104 genera of Schenkel, 3200 m), Gnaphosidae (Drassodes cupreus Blackwall,
3100 m; D. heeri Pavesi, 3000 m; Gnaphosa petrobia L. Koch, 3200 m; G. tigrina
Simon, G. muscorum L. Koch, G. badia L. Koch, G. leporina L. Koch, 3000 m;
Haplodrassus signifer C.L. Koch, 3082 m; Micaria alpina L. Koch, 3000 m; Zelotes
devotus Grimm, Z. similis Kul., 3000 m), Miturgidae (Zorinae)(Zora nigrimana
Schenkel, 3148 m), Thomisidae (Xysticus bonneti Denis, 3100 m; X. desidiosus
Simon, 3035 m; X. ibex Simon, 3000 m), Salticidae (Chalcoscirtus alpicola L. Koch,
3400 m; Talavera petrensis (C.L. Koch), 3180 m; Sitticus longipes Canestrini, 3040
m). To our knowledge, only five species of spiders have been found in the Alps
above 3500 m.
In Bulgaria, 207 species of spiders from 18 families and 94 genera have been
found above 1900 m and at least 165 species – at or above 2200 m. Higher than
2400 m (on Rila and Pirin) live at least 96 species belonging to 14 families and to
60 genera: Zodarion (Zodariidae), Aculepeira, Araniella, Larinioides (Araneidae),
Antrohyphantes, Araeoncus, Centromerus, Ceratinella, Diplocephalus,
Drepanotylus, Erigone, Evansia, Improphantes, Incetophantes, Lepthyphantes,
Linyphia,Mansuphantes, Maso, Meioneta, Mecynargus, Metopobactrus, Micrargus,
Mughiphantes, Oedothorax, Oreonetides, Ostearius, Pelecopsis, Porrhomma,
Prinerigone, Poeciloneta, Scothinotylus,Tenuiphantes, Tiso, Walckenaeria
(Linyphiidae), Achaearanea, Enoplognatha, Steatoda, Theridion, Rugothodes
(Theridiidae), Pardosa, Xerolycosa (Lycosidae), Coelotes (Amaurobiidae),
Tegenaria, Mastigusa (Agelenidae), Cicurina (Dictynidae), Cryphoeca (Hahniidae),
Clubiona (Clubionidae), Micaria, Gnaphosa, Haplodrassus (Gnaphosidae),
Table 10.1 Species of Family Species

Araneae living in the Alps at
Linyphiidae 105
or above 2200 m (202 spp.)
Lycosidae 23
Gnaphosidae 17
Salticidae 11
Thomisidae 11
Theridiidae 8
Araneae 7
Amaurobiidae 7
Phylodromidae 3
Clubionidae 2
Dictynidae 2
Agelenidae 1
Pisauridae 1
Hahniidae 1
Segestriidae 1
Miturgidae (Zorinae) 1
Tetragnathidae 1
Philodromus, Thanatus (Philodromidae), Ozyptila, Xysticus (Thomisidae),

Chalcoscirtus, Heliophanus, Pellenes, Sitticus, Ballus, Talavera (Salticidae).
The highest living spiders in the Alps (above 3000 m; those in bold above 3500
m) belong to the families Linyphiidae (Mughiphantes armatus Kulcz., 3769 m; M.
baebleri de Less., 3660 m; M. variabilis Kulcz., 3420 m; Improphantes complicatus
Emerton, 3281 m; Mughiphantes brunneri Thaler, 3200 m; M. handschini Schenkel,
3100 m; Incestophantes frigidus Simon, 3000 m; Diplocephalus rostratus Schenkel,
3540 m; D. helleri L. Koch, 3280 m; Erigone tirolensis L. Koch, 3500 m; E.
remota L. Koch, 3280 m; E. atra Blackwell, 3140 m; E. dentipalpis Wider, 3100 m;
Ceratinopsis austera Simon, 3400 m; Oreonetides glacialis L. Koch, 3400 m;
Araeoncus anguineus L. Koch, 3100 m; Hilaira montigena L. Koch, 3450 m; Tiso
aestivus L. Koch, 3400 m; Walckenaeria clavicornis Emerton, 3400 m; W. vigilax
Blackwall, 3000 m; Janetschekia monodon O.-P. Cambr., 3370 m; Linyphia
triangularis Clerck, 3000 m), Theridiidae (Enoplognatha serratosignata L. Koch,
3100 m; Theridion petraeum L. Koch, 3000 m), Lycosidae (Arctosa alpigena Dol.,
3000 m; Pardosa nigra C.L. Koch, 3700 m; P. giebeli Pavesi, 3400 m; P. palustris
L., 3140 m), Amaurobiidae (Coelotes pastor pickardi Kul., 3148 m), Hahniidae
(Cryphoeca nivalis Schenkel, 3200 m), Gnaphosidae (Drassodes cupreus Blackwall,
3100 m; D. heeri Pavesi, 3000 m; Gnaphosa petrobia L. Koch, 3200 m; G. tigrina
Simon, G. muscorum L. Koch, G. badia L. Koch, G. leporina L. Koch, 3000 m;
Haplodrassus signifer C.L. Koch, 3082 m; Micaria alpina L. Koch, 3000 m; Zelotes
devotus Grimm, Z. similis Kul., 3000 m), Miturgidae (Zorinae)(Zora nigrimana
Schenkel, 3148 m), Thomisidae (Xysticus bonneti Denis, 3100 m; X. desidiosus
Simon, 3035 m; X. ibex Simon, 3000 m), Salticidae (Chalcoscirtus alpicola L. Koch,
3400 m; Talavera petrensis (C.L. Koch), 3180 m; Sitticus longipes Canestrini, 3040
m). To our knowledge, only five species of spiders have been found in the Alps
above 3500 m.
Acariformes Both Sarcoptiformes and Trombidiformes are represented in the
high mountains. The maximum recorded altitude for both Acariformes is 6100 m
(Nepal), but is probably higher.
Prostigmata are represented in the high-altitude environment by 96 genera and 32
families. The families are varied in altitudinal distribution. Of the 32 families, 18
reach 3000 m, 12 – 3500 m, 7 – 4000 m., 4 – 4500 m., 1 – 5000 m. No identified
Prostigmata have been recorded above 6100 m. Champions are Adamystidae
(5100 m in Hindu Kush), Anystidae (4950 m in Nepal), Rhagidiidae (4800 m on
Kilimanjaro), Pygmephoridae (4500 m in New Guinea), Erythraeidae (4260 m in
New Guinea), Trombidiidae (4200 m on Ruwenzori), and Trombiculidae (4155 m
in Pakistan). With more intense research, the upper limit of Bdellidae and other
families might be found to live higher. The bulk of the material collected remains
unidentified. Presently, there are at least 363 species of 119 genera (a tiny fraction
of the Prostigmatic mites known in the world – more than 14,000 species of 1100
genera and 140 families) higher than 2200 m in the Old World. Higher than 3500 m,
there have been only 14 families, 31 genera, and 55 species recorded. From the
Prostigmatic mites found above 2200 m, the richest in species are the families
Trombiculidae (73), Trombidiidae (44), Erythraeidae (56), Bdellidae (27), and
Rhagidiidae (23). The most numerousgenus (20 spp.) is Balaustium (Erythraeidae).
The best studied are the Prostigmata of the Alps (studies by Irk, Willmann,
Schweizer, and other specialists) and of the mountains of tropical Africa, mainly
due to Marc André. According to Russian specialists, 36 species of tetranychoid
mites (including 30 Bryobriinae) live in Pamir between 2500 and 4500 m (species
not indicated).
Our extensive collections from various mountain systems are now under study.
Among the high-altitude Acariformes, most numerous are the soil-inhabiting
species of suborder Oribatida, from which 45 species have been found above 1900 m
(Beron, MS) in Bulgaria. Above 2200 m (mainly from Rila, Vitosha, and Pirin),
there have been 25 species of Oribatida recorded. The highest point of the Balkan
Peninsula (Mussala, 2925 m) is reached by the species Anachipteria deficiens
Grandjean (Achipteriidae), Trichoribates monticola (Trägårdh) (Ceratozetidae),
and Niphocepheus nivalis baloghi Travé (Niphocepheidae).
Acaridida are not well studied.
With (at least) 79 families found at or above 2200 m, the Oribatids are the most
diversified suborder of all high-altitude Arthropods, even more varied than the spi-
ders. The total worldwide species count is more than 7000 Oribatid mites, belong-
ing to 177 families. Their importance in the natural ecosystems is enormous (in
some places more than 70 % of all soil mites) and together with the theoretical
interest they represent, has led to intensive research of this group by many special-
ists. Thanks mainly to the publications of Maria Hammer (1977), Balogh (1970),
Mahunka (1991), Kunst (1957), Aoki (1965), Schatz (1978 and others), Piffl (1971)
and some Russian specialists, we now have knowledge of the high mountain parts
of South America, Hindu Kush, Himalayas, Pamir, the Alps, the mountains in
Bulgaria, tropical Africa and New Guinea. The 79 families known to occur above
2200 m contain 548 species. Most species are from the families Oppiidae (65),
Ceratozetidae (33), Brachychthoniidae (29), Phthiracaridae (27), Damaeidae (25),
and others. With the increase in altitude, the number of families and species
decreases as follows:
> 2200 m – 79 fam., 547 species > 4500 m – 21 fam., 26 species;
> 4000 m (incl. > 3950 m) – 27 fam., 42 species.
The Oribatids have been recorded in the Himalayas as high as 6100 m (Janetschek
1990). This author has also found in Nepal (up to 5800 m) unidentified representatives
of the families Brachychthoniidae, Hermanniidae, Damaeidae, Ceratoppiidae,
Tectocepheidae, Suctobelbidae, Oribatulidae, Ceratozetidae, Trichoribatidae,
Scheloribatidae, Oribatellidae, and (?) Niphocepheidae. There is a low level of
knowledge on the composition of Oribatida at altitudes above 5000 m.
Suborder Oribatida is represented, especially in the Andes, by a number of
high-altitude species, comparatively well-known, thanks to specialists L. Beck, P.
and J. Balogh, M. Hammer, W. Niedbala, etc. These high-altitude species belong to
families, known also in the European oreal (Phthiracaridae, Brachychthoniidae,
Nothridae, Camisiidae), but also to some other mountains in the Neotropical Region
(Anderemaeidae). It would be appropriate to make a comparison between the
“euhypsobionts” of the Andes and of Central Asia (including Himalayas, Karakorum,
Hindu Kush, Pamir), as in Europe we have not found Oribatids living higher than
3500 m, and in Africa this fauna is less studied.
Parasitiformes All three orders in this superorder are known from the high moun-
tains. Only one member of the small order Holothyrida has been recorded above
2200–2660 m in New Guinea (Hammenius ingii Leht.).
The Gamasid mites (order Mesostigmata) are the bulk of the representatives of
superorder Parasitiformes, but they are understudied, except in the Alps. Several
species of these mites have been recorded on Hindu Kush up to 4550 m and
Kilimanjaro up to 4285 m (mostly parasites and commensals on mammals).
Members of at least eight families live higher than 3000 m: Parasitidae,
Macrochelidae, Halolaelapidae, Zerconidae, Rhodacaridae, Hypoaspididae,
Dermanyssidae, and Laelapidae.
Two of the three families of order Ixodida are known from the high-altitude
environment (parasites on birds and mammals). Thanks to the intensive research of
the Himalayan fauna in the 1960s and 1970s (mainly by H. Hoogstraal and
colleagues) in Nepal the Argasid ticks (Argasidae) have been found up to 4575 m
and the Ixodid ticks (Ixodidae) have been found as high as 5488 m.
From Argasidae, four species are known in the high mountains of the Caucasus
to the Himalayas. From Ixodidae, at least 35 (out of about 800 species of this family)
members of the widespread genera Ixodes, Haemaphysalis, Dermacentor,
Rhipicephalus, and Hyalomma and of the genus Anomalohimalaya (endemic to the
Himalayas and the mountains of Central Asia) are known from the mountains of
Asia and Africa above 2200 m. Fourteen species have been recorded from altitudes
above 3500 m (Nepal, Hindu Kush, Tien Shan, Meru). The champion Ixodes berlesei
(5488 m, on the Snow Partridge Lerwa lerwa in Nepal) is the highest found
representative of Parasitiformes.
10.5 H
igh-Altitude Arachnida in the Oreal of the Himalayas,
Compared with the Alps
The dominating mountains in Europe and Asia belong to the same system
(Alpohimalayan). They are on a similar geographical latitude but are very different
in height (up to 4807 m in the Alps, up to 8848 m in the Himalayas). They have
different sources of fauna, different climates, and vegetation. The position of the
Himalayas on the border between two realms creates a mix of fauna from Palearctic
and Indo-Malayan elements. While the basic information on the high mountain
fauna of the Alps has been obtained as early as the beginning of the twentieth
century and took shape in the 1960s, knowledge on the rich Himalayan fauna has
been obtained in the second half of 20th century and is due largely to the meticulous
research campaigns of the German Zoologists (J. Martens and others).
Situated to the north and in the moderate climatic belt, the Alps offer the
Arachnida minimum conditions for existence up to about 3600–3700 m. However,
as part of Palearctical Region and Alpohimalayan System, they have many elements
in common with the higher Himalayas.
Scorpiones In Europe, only representatives of Euscorpiidae (Euscorpius) live in
the mountains of the southern part of the continent, up to 2569 m (Albania) and
2400 m (Mount Olympus). In the Alps, this genus and the Scorpions as a group do
not live above 2000 m. In the Himalayas, altitudes above 2200 m (even as high as
5000 m in Scorpiops) are reached by members of the families Scorpiopidae
(Scorpiops), Chaerilidae (Chaerilus), Hormuridae (Tibetiomachus), and Buthidae
(Himalayotityobuthus).
Pseudoscorpiones According to Schawaller (1987, 1991), from 37 known species
of Pseudoscorpions, 29 live above 2000 m, 17 above 3000 m, seven above 3500 m,
and two above 4000 m (Orochernes nepalensis – 4000 m and Stenohya (=
10.5 High-Altitude Arachnida in the Oreal of the Himalayas, Compared with the Alps 871
Levigatocreagris) martensi – 3300–4700 m) in Nepal. To these figures, we have to

add Dactylochelifer macrotuberculatus Krumpál‚ 1987 (3400–4000 m). Therefore‚
in Nepal, from 38 species of Pseudoscorpions, 30 live above 2000 m‚ 18 above 3000
m‚ 11 above 3500 m, and three reach 4000 m. The locality of S. martensi at
Meropapa La S Tudam (4700 m) in the Arun Valley, East Nepal is the highest record
of identified Pseudoscorpions. There is a higher record of unidentified young
Pseudoscorpions, found at 5000 m and published by Schawaller (1991) as
“Levigatocreagris – Bisetocreagris spec. juv.”
Following the widely accepted zonal subdivision of Nepal, the zone above
3000 m is called subalpine, above 4000 m, alpine. In the upper alpine subzone
(higher than 4500 m), there has been only one species of Pseudoscorpion recorded
(Stenohya martensi).
As previously mentioned, above 2000 m in the Alps there five species of the
genus Neobisium (Neobisiidae) and one species of Allochernes (Chernetidae) have
been found. Most species are alpine high-altitude endemics, not living in the
lowland. Neobisium jugorum (1700–3523 m) is particularly typical. The remaining
species live between 2400 and 3000 m. The oreals of the Alps and the Himalayas
have two families in common and none of the gerera and species (the genus
Allochernes has been recorded on Kashmir).
Some of the 11 families of Pseudoscorpions found in the high Himalayas do not
occur in the Alps (e.g., Lechytiidae, Tridenchthoniidae, and Hyidae). Some other
families live in the Alps, but under 2000 m (Chthoniidae, Geogarypidae, Olpiidae,
Cheiridiidae, Atemnidae, Cheliferidae, and Withiidae).
Opiliones In the Alps at or above 2000 m (the orophyte zone), there are at least 32
species of Opiliones, including one from suborder Laniatores (up to 2000 m) and
representatives of the Trogulidae family (up to 2000 m). Higher than 2200 m, there
have been at least 30 species of the families Nemastomatidae, Ischyropsalididae,
Phalangiidae s.l., and Sclerosomatidae recorded. Thanks to the recent studies of
Suzuki and especially of Martens, we now have information about at least 87 spe-
cies of Opiliones (29 Laniatores and 58 “Palpatores”) living in the high Himalayas
(above 2200 m). These species belong to the families Oncopodidae, Phalangodidae,
Assamiidae, Biantidae, Phalangiidae, Sclerosomatidae, and Sabaconidae. Only
Phalangiidae and Sclerosomatidae are represented in the oreal of the Alps.
Oncopodidae, Assamiidae, and Biantidae do not occur in Europe. Phalangodidae
does not occur on this continent at altitudes above 2000 m. High mountain species
of the Sabaconidae live in the Pyrenees, but not in the Alps. The families
Nemastomatidae and Ischyropsalididae do not occur in the Himalayas.
Of the 15 genera of Opiliones living above 2200 m in the Alps (all belonging to
“Palpatores”) and 21 genera of Himalayan “Palpatores”, there are no genera in com-
mon (only two of the Opilionid genera in the high Himalayas also live in Europe –
Opilio and Sabacon). The profusion of endemic genera containing many
species each in the high Himalayas is proof of the active speciation on this mountain
(Martens 1980, 1984, 1993).
In the Alps, six species of Opiliones have been recorded at or above 3000 m.
Only Mitopus glacialis is known to live above 3500 m (up to 3675 m). At least 42
species of the Himalayan Opiliones live above 3000 m (in the Himalayas, forests
grow as high as 4600 m). The altitude of 3500 m is reached by 15 species, 4000 m
by eight and only two reach 4500 m and go above 5000 m (Himalphalangium
palpale – 5540 m and Sabacon sp. – > 5000 m). In the Alps (above 3800 m), there
is snow everywhere and arthropod life stops at this altitude (mites reach 3774 m,
spiders reach 3769 m).
Araneae We have discussed elsewhere in this analysis the spiders of the high Alps
(196 species belonging to 86 genera and 15 families). The studies on the high-
altitude spiders of the highest mountain system on Earth – Himalayas (Karakorum
excluded) – resulted in the publishing of at least 192 species from 38 genera and 11
families (there is no doubt that their real numbers are much higher). From the
Himalayan spiders, as seen in the other mountains of the Old World, the most
numerous is the family Linyphiidae with 67 species and 32 genera. The Alps and the
Himalayas have in common eight families, at least 18 genera (Agyneta, Asthenargus,
Bathyphantes, Erigone, Hilaira, Linyphia, Mughiphantes, Oedothorax,
Palliduphantes, Piniphantes, Porrhomma, Tenuiphantes, Walckenaeria,
Acantholycosa, Pardosa, Xysticus, Chalcoscirtus, and Euophrys) and no species.
Absent from the oreal of the Alps are the Himalayan families Anapidae,
Tetrablemmidae, and Sicariidae. In the high Himalayas the genera Himalaphantes,
Martensinus, Oia, Paragongylidiellum, Hubertia, Saloca, Indicoblemma,
Brignoliella, Lysiteles, Plexippoides, Yaginumaella, and Synagelides have been
recorded and these genera are not known from the European oreal or in Europe as a
whole. Most of these genera are endemic and demonstrate the importance of the
Himalayas as a center of intensive speciation and creation of new taxa (Figs. 10.1
and 10.2).
m
6500
6000
5500
5000
4500
4000
3500
3000
2500
2000
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20
Fam.
Fig. 10.1 Families of spiders (Araneae) in the Old World at or above 2200 m
1. Dipluridae – up to 2600 m
2. Migidae – up to 2200 m
3. Pholcidae – up to 3500 m
4. Tetrablemmidae – up to 2730 m
5. Segestriidae – up to 2500 m
6. Dysderidae – up to 3270 m
7. Oonopidae – up to 3800 m
8. Orsolobidae – up to 2850 m
9. Archaeidae – up to 2650 m
10. Palpimanidae – up to 2300 m
11. Mimetidae – up to 2200 m
12. Nesticidae – up to 4900 m
13. Theridiidae – up to 4600 m
14. Anapidae – up to 2930 m
15. Linyphiidae – up to 5700 m
16. Tetragnathidae – up to 4600 m
17. Araneidae – up to 4500 m
18. Lycosidae – up to 6100 m
19. Pisauridae – up to 2400 m
20. Oxyopidae – up to 3700 m
m
6500
6000
5500
5000
4500
4000
3500
3000
2500
2000
21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40
Fam.
Fig. 10.2 Families of spiders (Araneae) in the Old World at or above 2200 m
21. Zoropsidae – up to 2165 m
22. Ctenidae – up to 2900 m
23. Agelenidae – up to 4000 m
24. Cybaeidae – up to 2200 m
25. Hahniidae – up to 5500 m
26. Dictynidae – up to 5700 m
27. Amaurobiidae – up to 4450 m
28. Titanoecidae – up to 4400 m
29. Liocranidae – up to 3460 m
30. Clubionidae – up to 4550 m
31. Zodariidae – up to 3500 m
32. Gallienellidae – up to 2400 m
33. Prodidomidae – up to 2745 m
34. Gnaphosidae – up to 4980 m
35. Miturgidae (Zorinae) – up to 3148 m
36. Selenopidae – up to 2400 m
37. Sparassidae – up to 3800 m
38. Philodromidae – up to 3850 m
39. Thomisidae – up to 5300 m
40. Salticidae – up to 5947 m
Acari The mites and ticks of the two mountain systems (Alps and Himalayas) have
not been studied in full, especially the ones of the Himalayas. From the Alps, thanks
to the researchers Willmann, Irk, Schweizer, Franz, Piffl and others, we already
know about many species of Prostigmata belonging to at least 28–30 families
(excluded are parasites like Myobiidae, Listrophoroidea, Sarcoptoidea, the endo-
parasitic mites and other groups not considered here). The maximum altitude is
reached by members of Bdellidae (3774 m). The Himalayan Prostigmata remains
almost unresearched, despite the considerable collections of Franz, Martens and the
author’s own. Janetschek (1990) has mentioned Prostigmata at altitudes of 6100 m
in Nepal. Himalayas and the high Alps share several families (Labidostomidae,
Rhagidiidae, Teneriffiidae, Anystidae, Trombiculidae, Trombidiidae), but there is
no doubt that most families known from the Alps will also be found in the oreal of
the Himalayas. The genera Shibaia (Rhagidiidae), Erythraeus, Leptus (Erythraeidae),
Leptotrombidium (Trombiculidae), perhaps also many others, live in both moun-
tains (Fig. 10.3).
5000
4500
4000
3500
3000
2500
2000
1500
1000
500
0
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18
Fig. 10.3 Families of Acari (Prostigmata) in the Old World at or above 2200 m
1. Pygmephoridae – up to 4500 m
2. Microdispidae - up to 3500 m
3. Scutacaridae - up to 3650 m
4. Tetranychidae – up to 3774 m
5. Tenuipalpidae – up to 2800 m
6. Raphignathidae – up to 2600 m
7. Stigmaeidae – up to 2925 m
8. Calyptostomatidae – up to 2800 m
9. Smarididae – up to 3100 m
10. Erythraeidae – up to 4260 m
11. Johnstonianidae – up to 3400 m
12. Trombellidae – up to 2500 m
13. Trombidiidae - up to 4200 m
14. Tanaupodidae – up to 2700 m
15. Eutrombidiidae – up to 2800 m
16. Microtrombidiidae – up to 3500 m
17. Trombiculidae - up to 4600 m
18. Leeuwenhoekiidae – up to 3600 m
Oribatida are still understudied in the high Himalayas, but at least ten families
have been recorded from altitudes above 5500 m (Janetschek 1990). From the Alps,
members of at least 37 families are known to occur above 2200 m. All families and
genera (except Nippobodidae, resp. Leobodes) known from the high Himalayas
have also been recorded in the Alps (Figs. 10.4 and 10.5).
m
6000
5500
5000
4500
4000
3500
3000
2500
2000
1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39
Fam.
Fig. 10.4 Families of Oribatida in the Old World above 2200 m

1. Aphelacaridae – up to 3200 m 21. Hermanniellidae – up to 2334 m
2. Brachychthoniidae – up to 5800 m 22. Gymnodamaeidae – up to 3400 m
3. Heterochthoniidae – up to 2600 m 23. Plateremaeidae – up to 3900 m
4. Hypochthoniidae – up to 3500 m 24. Licnodamaeidae – up to 2600 m
5. Eniochthoniidae – up to 3080 m 25. Damaeidae – up to 5800 m
6. Mesoplophoridae – up to 3000 m 26. Cepheidae – up to 3130 m
7. Parhypochthoniidae – up to 3209 m 27. Eutegaeidae – up to 4200 m
8. Lohmanniidae – up to 3400 m 28. Anderemaeidae – up to 3200 m
9. Eulohmanniidae – up to 3965 m 29. Microtegeidae – up to 4285 m
10. Oribotritiidae – up to 5000 m 30. Podopterotegaeidae – up to 5430 m
11. Euphthiracaridae – up to 3660 m 31. Niphocepheidae – up to 3300 m; 5800 m
12. Phthiracaridae – up to 3890 m 32. Microzetidae – up to 2850 m
13. Steganacaridae – up to 3890 m 33. Heterobelbidae – up to 2395 m
14. Trhypochthoniidae – up to 5000 m 34. Eremaeidae – up to 3400 m
15. Malaconothridae – up to 5000 m 35. Zetorchestidae – up to 2500 m
16. Nothridae – up to 3965 m 36. Eremobelbidae – up to 2500 m
17. Camisiidae – up to 4500 m 37. Damaeolidae – up to 3200 m
18. Nanhermanniidae – up to 3965 m 38. Ameridae – up to 3030 m
19. Liodidae – up to 2600 m 39. Caleremaeidae – up to 2550 m
20. Hermanniidae – up to 5800 m
m
6000
5500
5000
4500
4000
3500
3000
2500
2000
36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70
Fam.
Fig. 10.5 Families of Oribatida in the Old World above 2200 m - continuation
40. Astegistidae – up to 3900 m 60. Phenopelopidae – up to 3500 m
41. Ceratoppiidae – up to 5800 m 61. Unduloribatidae – up to 4500 m
42. Gustaviidae – up to 2850 m 62. Achipteriidae – up to 3400 m
43. Metrioppiidae – up to 2395 m 63. Tegoribatidae – up to 3100 m
44. Liacaridae – up to 3080 m 64. Oribatellidae – up to 5500 m
45. Xenillidae – up to 2775 m 65. Mycobatidae – up to 5430 m
46. Carabodidae – up to 3080 m 66. Ceratozetidae – up to 5800 m
47. Nippobodidae – up to 2800 m 67. Trichoribatidae – up to 5800 m
48. Tectocepheidae – up to 5800 m 68. Chamobatidae – up to 3360 m
49. Dampfiellidae – up to 2480 m 69. Galumnidae – up to 3000 m
50. Otocepheidae – up to 3900 m 70. Parakalummidae – up to 2920 m
51. Thyrisomidae – up to 3000 m 71. Zetomotrichidae – up to 3500 m
52. Quadroppiidae – up to 3820 m 72. Haplozetidae – up to 5000 m
53. Oppiidae – up to 5430 m 73. Fenicheliidae – up to 3900 m
54. Suctobelbidae – up to 5800 m 74. Oribatulidae – up to 5800 m
55. Cymberemaeidae – up to 3900 m 75. Scheloribatidae – up to 5800 m
56. Ameronothridae – up to 3900 m 76. Oripodidae – up to 3900 m
57. Licneremaeidae – up to 2800 m 77. Liebstadiidae – up to 3900 m
58. Scutoverticidae – up to 4438 m 78. Symbioribatidae – up to 2800 m
59. Passalozetidae – up to 3400 m
Parasitiformes The Gamasida of the Himalayas are also understudied so it would

be difficult to make comparisons with the families living in the high Alps (at least
24). Also almost all the families would be shared by both mountains. The Ixodida
order of Parasitiformes is more represented in the high Himalayas than in the high
Alps. In comparison, the genus Ixodes, barely entering the Alps above 2000 m, has
been discovered in the high Himalayas in least six high mountain species, including
the world record in Ixodida – Ixodes berlesei at 5488 m. The endemic genus and
species Anomalohimalaya lama has also been found. The widespread genus
Haemaphysalis in the Alps does not reach 2000 m but in the Himalayas has been
found as high as 4880 m (H. aponommoides) (Figs. 10.6 and 10.7).
m
6500
6000
5500
5000
4500
4000
3500
3000
2500
2000
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Fam.
Fig. 10.6 Families of Acari (Parasitiformes) in the Old World at or above 2200 m
1. Holothyridae – up to 2650 m
2. Ameroseiidae – up to 3897 m
3. Aceosejidae- up to 2850 m
4. Epicriidae – up to 2700 m
5. Ascidae – up to 2911 m
6. Parasitidae – up to 3209 m
7. Pergamasidae – up to 3450 m
8. Eviphiidae – up to 2500 m
9. Macrochelidae – up to 3500 m
10. Pachylaelapidae – up to 2850 m
11. Laelapidae – up to 3650 m
12. Haemogamasidae – up to 4550 m
13. Phytoseiidae – up to 2560 m
14. Podocinidae – up to 4300 m
15. Veigaiidae – up to 2900 m
16. Halolaelapidae – up to 3109 m
m
6500
6000
5500
5000
4500
4000
3500
3000
2500
2000
17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 Fam.
Fig. 10.7 Families of Acari (Parasitiformes) in the Old World at or above 2200 m
17. Parholaspidae – up to 3350 m
18. Zerconidae – up to 3987 m
19. Rhodacaridae – up to 3109 m
20. Hypoaspididae – up to 4285 m
21. Dermanyssidae – up to 4550 m
22. Spinturnicidae – up to 2770 m
23. Halolaelapidae – up to 3000 m
24. Uropodidae – up to 2300 m
25. Trachyuropodidae – up to 2500 m
26. Urodinychidae – up to 2700 m
27. Trematuridae – up to 2250 m
28. Trachytidae – up to 2600 m
29. Polyaspididae – up to 2680 m
30. Argasidae – up to 4575 m
31. Ixodidae – up to 5488 m
10.6 High-Altitude Arachnida in the Mountains

of Central Asia
The mountains of Central Asia are much higher than the Alps (8611 m in Karakorum,
7690 m in Hindu Kush, 7495 m in Pamir, 7439 m in Tien Shan, 4807 m in the Alps).
Even the Tibetan Highland is, in some places, higher than 4000 m, and the latitude
of the Central Asian mountains is similar to the latitude of the Alps. The vertical
zonation of these mountains is also very different. In Central Asia, often the moun-
tains lack forest belts and the cold steppe climbs to above 3000 m. On the other
hand, all these mountains are part of the same Alpo-Himalayan system, with com-
mon origin and development.
Solifugae Representatives of this order are well represented in Central Asia and at
least six to seven species live in the high mountains, up to 3500 m (Anoplogylippus,
Galeodila, Galeodellus) and even 4000 m (Galeodes) and 4300 m (Karschia). The
presence of Solpugids in the high mountains of Central Asia is one of their most
typical features. The champion of the Old World Solifugae is Karschia tibetana
Hirst (Tibet, 4570 m).
Pseudoscorpiones At least 12 species of Pseudoscorpiones, belonging to eight

families, have been recorded from Central Asia (above 2200 m), including 11 above
3000 m and four above 3500 m. The champions are Bisetocreagris kaznakovi
(Red.) – 4810 m (Neobisiidae, Tibet), Dactylochelifer brachialis Beier (4200 m,
Karakorum), Gobichelifer chelanops (Red. ) (3650 m, Karakorum), and “Chelifer”
baltistanus di Cap. (3950 m, Karakorum) (all four belonging to Cheliferidae). In the
Alps, the maximum altitude for Pseudoscorpions is also reached by members of the
family Neobisiidae – Neobisium jugorum L. Koch, 3523 m. Obviously, the Central
Asian high mountain Pseudoscorpionid fauna is much richer and varied compared
to the fauna of the Alps and many more families and genera are represented in the
Central Asian mountains. The family Cheliferidae, typical for the Central Asian
oreal is not represented in the high Alps.
Opiliones As in Europe (but unlike the Himalayas), all Opiliones known above
2200 m in Central Asia, belong to Palpatores (Eupnoi and Dysppnoi). Six species
occur above 3000 m and the five representatives of Phalangiidae live above 3500 m.
This family is also the best represented in the high Alps. The alpine families
Nemastomatidae and Ischyropsalididae are not represented at the higher altitude in
Central Asia. In the highest parts of Central Asia and the Alps, there are no genera
in common but some genera known from Europe, are present high in Central Asia
(e.g., Opilio, Egaenus). The highest living Opiliones in the Alps are the
representatives of Mitopus (up to 3675 m). In Central Asia, the champion is
Homolophus (= Euphalangium) nordenskioeldi L. Koch (5600 m in Karakorum).
This is also the highest living opilion in the world (identified to species level).
10.6 High-Altitude Arachnida in the Mountains of Central Asia 881
Araneae The research on high-altitude spiders in vast areas of Central Asia over
2200 m is inadequate. We managed to collate information on 180 species above this
altitude, using mostly the publications of di Caporiacco (1934-35) about Karakorum,
the papers of Denis (1958) and Roewer (1960, 1962) about Afghanistan and of the
Russian specialists about the mountains of the former Soviet Union. These 180
species belong to 18 families but only four families contain 149 of the speices:
Linyphiidae (88 spp.), Gnaphosidae (22), Salticidae (18), and Lycosidae (21). These
families are typical also for the European high mountains. The remaining 31 species
are distributed among 14 families, also represented in the European oreal. Even if
further research is to bring data about other species, the ratio between the families
in the current list is not expected to change.
The altitudinal distribution of these families, genera and species are as follows:
Above 2200 m – 18 Fam., 79 genera, 180 species
From 2200 to 4500 m, the number of spider species decreases almost nine times.
As seen throughout the mountains of the Old World, the Linyphiidae family is by far
the most numerous. With at least 88 species, it contains more than half of the high-
altitude spiders in Central Asia, and with 101 species in the high Alps – more than
half (51.8 %) of the spider species known in this mountain (196). In the mountains
of Central Asia, the best represented genus is complex Lepthyphantes (s.l., including
Mughiphantes, Tenuiphantes et al., altogether 30 spp.). Other numerous genera are
Parasyrisca (Gnaphosidae, 13 spp.) and Pardosa (Gnaphosidae, 13 spp.).
Acariformes The Acari in the mountains of Central Asia are not well-known,
except for some groups with medical importance (e.g., ticks, chiggers). At least 15
species of Trombiculidae have been recorded at altitudes of 2200–4155 m (mostly
Leptotrombidium, Helenicula, and Microtrombicula). Oribatids belonging to at least
35 families have been found to live in the oreal of Hindu Kush (M. Hammer 1977)
and Tajikistan (Khristov 1973). Almost all families and at least 20 genera
(Heminothrus, Nothrus, Trimalaconothrus, Nanhermannia, Hermannia,
Scheloribates, Sphaerozetes, Trichoribates, Chamobates, Eupelops, Oribatella,
Achipteria, Galumna, Acrogalumna, Neoribates, Eremaeus, Tectocepheus, Oribella,
Oppia, Phthiracarus) are shared by the two systems (Hindu Kush and Tajikistan).
There are even some species in common: Heminothrus targionii (Berlese), Nothrus
biciliatus (C.L. Koch), Nanhermannia nana (Nicolet), Hermannia gibba (C.L. Koch),
Scheloribates laevigatus (C.L. Koch), Oribatella meridionalis (Berlese). The highest
Trombiculidae in Central Asia has been recorded at 4155 m (Leptotrombidium puta
Womersley, Pakistan), the highest living Oribatida has been recorded at 5000 m (sev-
eral species in Pamir). It seems that the highest recorded member of Acariformes
from this area is Adamystis coinneaui Rafalski (Hindu Kush, 5100 m).
Parasitiformes Both in Mesostigmata and in Ixodida almost all genera in the high
mountains in Central Asia also live in the Alps. Examples from Mesostigmata
include Laelaps, Haemogamasus and Hirstionyssus, and examples from Ixodida
include Ixodes. The genera Haemaphysalis and Dermacentor also live in the Alps,
but below 2000 m, while in Central Asia they reach 4000 m (Haemaphysalis) and
3000 m (Dermacentor). In Central Asia, Alveonasus (Argasidae) reach 2900 m,
Ornithodoros – 2800 m. In the high Alps, Argasidae are missing completely.
10.7 Afrotropical High-Altitude Arachnida
Pseudoscorpiones Only 12 species of Pseudoscorpiones, belonging to three gen-

era (Chthonius, Neobisium, and Allochernes) are known from the European oreal
(excluding Caucasus). These genera are missing in the mountains of Equatorial
Africa. One exception is the European species Neobisium muscorum (= N.
carcinoides), recorded by Mahnert (1981) on the bases of tritonymphs from
Aberdare in Kenya (3203 m), the first representative of Neobisium in Africa south
of Sahara. In the extensive collection of Mahnert from the mountains of Africa, as
well as the collections of the older specialists (e.g., publications of M. Beier), we
could not find mention of Neobisium, so the authenticity and the authochthonous
character of this species is doubtful. This record has not been included in our zoo-
geographical analysis.
In the mountains of Equatorial Africa, all three European families are present. In
addition to this, there are ten other families, three of which do not live in Europe
(Lechytiidae, Tridenchthoniidae, and Ideoroncidae). Except for the doubtful
Neobisium record, none of the remaining Afrotropical genera live on the high
mountains of Europe. Only three of the genera (Lasiochernes, Withius, and
Apocheiridium) live on the European continent.
Opiliones All high mountain Opilions in Europe belong to five families of
“Palpatores” (28 species at altitudes higher than 2200 m). In the mountains of
Equatorial Africa (higher than 2200 m), there have been 83 species of Opiliones (61
Laniatores and 22 Palpatores) recorded and these species belong to three families.
In Europe, only two species live higher than 3000 m (both Mitopus), while in
Equatorial Africa 34 species do.
Araneae Much has been carried out in the study of Afrotropical spiders, but some
of the most important publications are older (E. Strand, A. Tullgren, R. de Lessert,
H. Fage, E. Simon, L. Berland) and need revision. Here we use data concerning 122
species belonging to 36 genera and seven families, recorded from altitudes over
2200 m. They include:
Above 3000 m – 18 genera

10.7 Afrotropical High-Altitude Arachnida 883

At altitudes over 3500 m, species belonging to the genera Callitrichia (4930 m),
Pelecopsis (4930 m), Heliophanus (4650 m), Araeoncus (4650 m), Microcyba
(4320 m.), Oreocyba (4300 m), Erigone (4200 m), Lepthyphantes (4000 m), Toschia
(3920 m), Walckenaeria (3820 m), Tybaertiella (3750 m), Asthenargus (3550 m),
and Mallinella (3500 m) exist. We can consider these species as true hypsobionts,
especially the species living above 3000 m. The Linyphiidae family is the most
numerous with 97 species and 30 of the 36 genera and is particularly well-studied
(Scharff, 1992). This family also reaches the highest altitude in Africa (4930 m,
Callitrichia and Pelecopsis). Next in number is the Salticidae family with the well-
represented genus Heliophanus (up to 4650 m). Besides Mallinella (up to 3500 m,
Zodariidae) and Heliophanus, the remaining 16 genera over 3000 m belong to
Linyphiidae.
Representatives of suborder Mygalomorphae (Migidae, Dipluridae) only live
over 2200 m in high altitudes in tropical Africa.
Out of 18 genera of spiders known in the Afrotropical oreal above 3000 m, more
than half are also found in the oreal of Europe (Lepthyphantes, Microlinyphia,
Erigone, Asthenargus, Araeoncus, Ceratinopsis, Pelecopsis, Meioneta,
Walckenaeria, Heliophanus). Some genera are endemic to Africa (Mallinella,
Aberdaria). From 12 genera known to occur above 4000 m, eight (Lepthyphantes,
Erigone, Araeoncus, Pelecopsis, Heliophanus, Meioneta, Ceratinopsis, Hahnia) are
also inhabitants of the higher parts of the European mountains. The other genera
(Bursellia, Microcyba, Callitrichia, Oreocyba) are typical for the Afrotropical oreal.
Acariformes The mites from suborder Acaridida are not well-known in both areas.
Genus Histiostoma (Histiostomatidae, or Anoetidae) is represented both in the
European and in the Afrotropical mountains. In Europe and especially in the Alps,
Prostigmata are better known (over 2200 m with at least 20 families, 35 genera and
more than 100 species, excluding the strict parasites, such as Myobiidae,
Listrophoroidea, etc.). This group is less known in the Afrotropical oreal. Shared
with the European oreal are families, genera, and even species (Anystis baccarum
L.). The data concerning the Afrotropical mountains concerns mainly the families
Anystidae, Bdellidae, Erythraeidae, Trombidiidae, and Microtrombidiidae. There is
no doubt that many other species will be found from the other families represented
in Europe, so no conclusions have been made. The highest record for Prostigmatid
mites in Europe is at 3774 m (Bdellidae in the Alps), in tropical Africa – at 4200 m
(Trombidiidae on Elgon).
From the mountains of tropical Africa, there have been many records of genera
either unknown in Europe or not found on the higher parts of European mountains.
Such genera are Compsothrombium, Dinothrombium, Dromeothrombium,
Eutrichothrombium (Trombidiidae), Camerotrombidium, Enemothrombium
(Microtrombidiidae), and others. The genera Prostigmata, living in the European
oreal but unknown in the Afrotropical, are Atomus, Podothrombium, Tanaupodus
(Trombidiidae), Eutrombidium (Eutrombidiidae), Microtrombidium,

Valgothrombium, Enemothrombium (Microtrombidiidae), etc.
We currently have considerable information on the representatives of Oribatida
in the oreals of Europe and tropical Africa. In the Afrotropical oreal, there have been
at least 19 families recorded, most of which also live in the European mountains.
Many genera are also shared (Liochthonius, Brachychthonius, Heminothrus,
Nothrus, Nanhermannia, Scheloribates, Ceratozetes, Galumna, Tectocepheus,
Suctobelba, Oppia, Quadropia, Rhysotritia). Scheloribates laevigatus C.L. Koch
lives in the Alps up to 2700 m and has been recorded on Kilimanjaro up to 4590 m.
This is also the maximum altitude at which Oribatids (and Acari in general) have
been found in Africa.
Parasitiformes The order Mesostigmata is not well known in these two regions,
especially in the Afrotropical region. The Ixodida order, is represented in both
regions with different species of genus Ixodes and in Africa with Rhipicephalus (up
to 3500 m on Meru).
10.8 Conclusions
In the mountains of the Old World (Eurasia, Africa, New Guinea, and the adjacent
islands), the Arachnida reach the following altitudes Solifugae – 4570 m,
Schizomida – 2600 m, Scorpiones – 4600 m, Pseudoscorpiones – 5000 m,
Opiliones – 5600 m, Araneae – 6700 m, Opilioacarida – 2500 m, Acariformes –
6100 m, Parasitiformes – 5488 m. Above 2200 m, there have been over 3070 species
recorded including: Solifugae – 15, Schizomida – two, Scorpiones – 26,
Pseudoscorpiones – 174, Opiliones – 266, Araneae – 1317, Opilioacarida – one,
Acariformes – 987, Parasitiformes – 282.
Altitudes above 3500 m have been reached by (at least) seven species of
Solifugae, eight species of Scorpiones, 25 species of Pseudoscorpiones, 60 species
of Opiliones, 203 species of Araneae, 155 species of Acariformes, 31 species of
Parasitiformes, altogether 489 species. These species could be regarded as true
hypsobionts, despite the fact that in some places (Himalayas) high forest is found at
4600 m. An altitude over 3500 m has specific parameters everywhere in the world,
mainly due to the atmospheric pressure and UV radiation.
10.9 C
hampions in Altitude (Above 4500 m) in the Old
World
Solifugae Karschia tibetana Hirst (4570 m, Tibet).

Scorpiones Scorpiops montanus Karsch (5000 m, Himalayas), S. hardwicki
Gervais (5000 m, Himalayas).
10.9 Champions in Altitude (Above 4500 m) in the Old World 885
Pseudoscorpiones Stenohya (=Levigatocreagris)/Bisetocreagris sp. (ca. 5000 m,

Nepal), Bisetocreagris kaznakovi (Redikorzev) (4810 m, Tibet), Stenohya martensi
(Schawaller) (4700 m, Nepal).
Opiliones Homolophus (= Euphalangium) nordenskioeldi (L. Koch) (5600 m,

Karakorum), Himalphalangium palpale Roewer (5540 m, Nepal), Homolophus
panpema Suzuki, H. luteum Suzuki, Octozaleptus harai Suzuki, Leiobunum mirum
Roewer (5200 m, Nepal), Sabaconidae gen. sp. (>5000 m, Nepal), Opilio sp. (4800
m, Karakorum), Hypoxestus accentuatus Sörensen, Rhampsinitus bettoni Pocock
(4600 m, Kilimanjaro), Phalangium sp. (4500 m, Karakorum).
Araneae Fam., gen., sp. indet. (6700 m, Nepal), Acantholycosa baltoroi di Cap.
(6100 m, Nepal), Euophrys omnisuperstes Wanless (5947 m, Himalayas), Sitticus
niveosignatus Simon (5570 m, Nepal), Mughiphantes yeti Tanasevitch (5545 m,
Nepal), Pardosa birmanica Simon (5300 m, Himalayas), Euophrys everestensis
Wanless (5185 m, Himalayas), Hahnia alini Tikader (5181 m, Nepal), Hilaira
dapaensis Wunderlich, Erigone atra Blackwall, Arctosa raptor (Kul.), Mughiphantes
alticola Tan. (5100 m, Nepal), Pardosa orealis Buchar (5000 m, Himalayas),
Gongylidium baltoroi di Cap. (5000 m, Karakorum), Gnaphosa stoliczkae
Cambridge (4980 m, Karakorum), Erigone dentipalpis Wider, Pardosa condolens
Cambridge (4950 m, Karakorum), Alioranus minutissimus di Cap., A. distinctus di
Cap., Dictyna consecuta Cambridge (4930 m, Karakorum), Callitrichia
ruwenzoriensis Holm (4930 m, Ruwenzori), Mughiphantes setifer Tan., M. sherpa
Tan., Agyneta yulungensis Wund. (4900 m, Nepal), Pardosa tridentis di Cap. (4900
m, Himalayas), Parasyrisca pshartica Ovtsharenko et al. (4900 m, Tajikistan),
Sitticus pubescens Fabr. (4900 m, Karakorum), Xysticus dolpoensis Ono (4880 m,
Nepal), Gnaphosa moerens O.P.-Cambr. (4850 m, Nepal), Pardosa tikaderi Buchar
(4850 m, Himalayas), P. thaleri Buchar (4800 m, Himalayas), Haplodrassus signifer
C.L. Koch (4800 m, Karakorum), Meioneta obscura Denis (4724 m, Ruwenzori),
Yllenus karnai Logunov et Marusik (4720 m, Ladakh), Y. baltistanus var.
shaksgamica di Cap. (4715 m, Karakorum), Araeoncus picturatus Holm (4650
m,.Kilimanjaro), Heliophanus crudeni de Lessert (4650 m, Kilimanjaro), H. dubius
C.L. Koch, Theridion glaciale di Cap. (4600 m, Karakorum), Euophrys yulungensis
Zabka, Mughiphantes falxus Tan. et Saaristo, M. restrictus Tan. et Saaristo (4600 m,
Nepal), Pardosa credula Cambridge (4590 m, Karakorum), Hahnia gigantes
Bosmans (4580 m, Ruwenzori), Callitrichia kenyae Fage (4530 m, M. Kenya),
Agyneta pseudofuscipalpis Wund., A. bieko Wund., Tenuiphantes plumipes Tan.
(4500 m, Nepal), Hahnia maxima di Cap., Zelotes baltoroi di Cap., Chalcoscirtus
glacialis di Cap., Araneus obscurissimus di Cap., Drassodes singularis di Cap.,
Phintella micans di Cap. (4500 m, Karakorum), Yllenus pamiricus Logunov et
Marusik (4500 m, Pamir)
arasitiformes: Ixodida (Ixodidae, Argasidae) Haemogamasus nidiformis
P
Bregetova (4550 m, Hindu Kush), Ixodes berlesei Birula (5488 m, Nepal),
Haemaphysalis aponommoides Warburton (4880 m, Nepal), Ixodes hyatti Clifford,
Hoogstraal et Kohls (4615 m, Nepal), I. ovatus Neumann (4615 m, Nepal), Argas

himalayensis Hoogstraal et Kaiser (4575 m, Nepal).
During the work on this comprehensive review of the high-altitude Isopoda,
Arachnida and Myriapoda of the Old World, a pattern has been detected. This
pattern was tested with the groups of Isopoda, Oniscidea, Pseudoscorpiones, and
Opiliones. Above 2500 m, the number of species decreased by 100 % with every
500 m increase in altitude (at a very similar rate with the three studied groups – one
of Crustacea, the others of Arachnida).
From 3000 m to 3500 m to 4000 m, the number of genera of Isopoda (22 – 14 –
7) and of Opiliones (44 – 27 – 13) decreased at almost the same ratio (roughly 3 –
2 – 1). Above 2200 m in all the studied mountains, there were 60 genera of Isopoda,
over 3500 m remain 14 and over 4000 m – seven. Over 2200 m, there were 61
genera of Pseudoscorpions, over 3500 m remain 16 and over 4000 m – six. Taking
into account the immense territory of the Old World with such a variety of mountains,
climates, belts, and natural zones, this phenomenon could hardly be considered a
coincidence. Higher than 4500 m in the Old World, there are an equal number of
genera and species of both Isopoda and of Opiliones – four. This Pattern is Known
as the “Taxogradient”.
10.10 S
ome Personal Notes on the High-Altitude
Arachnofauna of the Americas and Elsewhere
With my experience in the high mountains of the world, I can make first-hand
comparisons between the environment and the arachnid populations at similar alti-
tudes on different mountains. Around the hut “Whymper” under Chimborazo in
Ecuador (5100 – 5200 m), there are many animals to be found under stones, includ-
ing spiders. On the same altitude and latitude (almost on the equator) above Kibo
Hut on Kilimanjaro, it is hard to find any arachnid under the dry, frozen stones.
This situation is similar above 5000 m on Ruwenzori and Mount Kenya. In Peru,
under Huandoy, I was collecting in the highest tall forest in the world withPolylepis
trees more than 10 m tall and a rich litter layer. European mountains are not as high
(excluding Caucasus, less than 4708 m on the snow-covered Montblanc). There the
life in the hypolithon does not reach 4000 m. In the Himalayas, tall Rhododendron
forests are found even at 4500–4600 m, but the records in different groups of
Arachnida rarely reach 5000 m (Solifugae – 4570 m, Scorpiones – 5000 m,
Pseudoscorpiones – 5000 m, exceptions are Opiliones – 5600 m and Araneae –
6700 m).
I have also collected spiders on Mount Elbert (4401 m) in the Rockies and on
Djebel Toubcal (4167 m), the highest point of the Atlas Mountains. These summits
are lower but situated more to the north. At this altitude, there it usually snows every
night and this snow melts in the day. I collected rich fauna samples, including
spiders and opilions, above the hut Vicente Guerrero under Popocatepetl (Mexico),
at 4700–4800 m.
10.10 Some Personal Notes on the High-Altitude Arachnofauna of the Americas… 887
Solifugae In South America, other families of this order (Ammotrechidae) can

reach 5000 m (Dasycleobis crinitus Mello-Leitão in Argentina) (Maury 1982).
Scorpiones In South America, scorpions live much higher than in the Old World,
but this information has been corrected by Ochoa et al. (2011). They described six
new species of genus Orobothriurus Maury, 1976 and listed the scorpion species
recorded above 3000 m in the Andes. This list contains 39 species of the genera
Orobothriurus, Bothriurus, Pachkutej, and Brachistosternus (Bothriuridae, up to
4910 m in Peru), Tityus (Buthidae, up to 4200 m in Bolivia), Hadruroides (Iuridae,
up to 3379 m in Peru), and Teuthraustes (Chactidae, 3300 m in Ecuador). According
to these authors, the former maximum altitude, recorded for “Orobothriurus”
crassimanus – actually Pachakutej crassimanus (Maury 1976) by Lourenço (1997) –
5560 m at Nevado Huascaran (Peru) – is based on misidentification and errouneus
locality data. These authors accept the highest record of scorpions to be the newly
described Orobothriurus huascaran at 4910 m in Peru. In 1977, I visited the area
above Lake Paron and climbed on the slopes of Huandoy at altitudes over 5000 m.
The environment was free of snow and ice, with forests of Polylepis and it is quite
possible that scorpions could be found higher than 5000 m (Fig. 10.8).
Fig. 10.8 Zoogeographical subdivision of the palearctic fauna (According to Varga, 1975)
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Chapter 11
Zoogeographical Notes on Different
Orders of Arachnida
Abstract The distribution of some orders of Arachnida is determined by the pre-

ferred temperature of their environment (i.e., thermopreferendum). Thermophilic
orders (i.e., those that live at high temperatures) include: Schizomida, Amblypygi,
Thelyphonida (Uropygi), Opilioacarida, most of the scorpions, Solifugae, Opilions
Laniatores, and Cyphophthalmi. Thermophobic orders (i.e., resistant to cold tem-
peratures) include: Araneae, Opiliones Eupnoi, Dyspnoi, and some mites. These
thermophobic orders generally live in the Far North of Eurasia and North America,
including Greenland and high in the mountains. The distribution of some orders is
a result of paleogeographical events, whereas the distribution of other orders can be
difficult to explain (e.g., Holothyrida, spiders of suborder Mesothelae, with families
found in New Zealand and Chile; Ricinulei, the genera of Amblypygi e.g., Phrynus
in Indonesia, Trichodamon in Brazil; the presence of only one genus of Thelyphonida
in a restricted area in West Africa). Most of these scattered groups are clearly relics.
Similarly, in some groups (Scorpiones, Pseudoscorpiones) the presence of American
elements in the Mediterranean subregion is surprising and demands explanation.
11.1 Palpigradi
The most thorough analysis of this order was made by Condé (1996). The limits of
the order are 48° North and 40° South, with most of the northern species confined
to caves. In Europe, the troglobiomorphic species are close to intertropical lines and
could be considered relics from warm periods. Eukoenenia (60 spp., 27 in caves or
in the superficial underground is the most diversified genus with advanced cave
adaptation. The second most diversified genus is Koeneniodes, with eight species in
West Africa, the Mascarene Islands, Madagascar, New Guinea, and South China.
Allokoenenia has only one species and is found in Guinea, Leptokoenenia is found
in Brazil and Madagascar.
The family Prokoeneniidae has been found in Texas and California. There have
been unverified recordings in Chile. This Chilean record of the species Prokoenenia
chilensis (Hansen, 1901) led Jeannel (1942) to suppose that “il serait fort possible
que les Prokoenenia sud-américaines soient les restes d’une lignée paléantarctique,

https://doi.org/10.1007/978-3-319-74418-6_11
894 11 Zoogeographical Notes on Different Orders of Arachnida
dont la trace devra être recherchée en Australie ou en Nouvelle-Zélande.” However,

so far Palpigradi have not been detected in New Zealand and only two imported
Eukoenenia and one (so far considered native) species are known in Australia.
Palpigradi have not been found in Japan, despite intense research. The mapped dis-
tribution of Prokoenenia chilensis, according to Condé (1996), needs to be revised.
Triadokoenenia is found in Madagascar (the former “Prokoenenia” millotorum
Remy). However, true Prokoenenia live in Java (P. javanica, Condé, 1990), Thailand
(P. asiatica, Condé, 1994), and Sulawesi (Indonesia) (P. celebica, Condé, 1946).
Condé (1996) stated the following: “The Oriental region seems to have been an
important center of evolution in the history of the Palpigrades: all actual genera are
represented there”.
11.2 Schizomida
The distribution of the Schizomida order is mainly determined by climate. This

order lives at altitudes of up to 2600 m a.s.l. in Africa (Reddell and Cokendolpher
1995). The family Protoschizomidae (two genera, 12 spp.) is mostly endemic to
Mexico and includes many cave dwelling species. The much bigger family
Hubbardiidae (26 genera, more than 250 spp.) is distributed pantropically. The
subfamily Megaschizominae (two spp.) is endemic to Southern Africa. The
Hubbardiinae family includes all remaining species. Until 1992, only seven genera
had been recorded, often considered synonymous with each other (Artacarus Cook,
1898; Hubbardia Cook, 1899; Trithyreus Kraepelin, 1899; Stenochrus Chamberlin,
1922; Megaschizomus Lawrence, 1969; Agastoschizomus Rowland, 1971;
Protoschizomus Rowland, 1975). Harvey (1992) initiated a new approach to
Schizomid taxonomy, based on the spermatecae. Using this approach, Harvey
described five new genera (Australia). Thereafter followed the monograph by
Reddell and Cokendolpher (1995) and many new descriptions by Harvey,
Cokendolpher, de Armas, Tsurusaki and several South American authors
(Manzanilla, Giupponi, and others). Some endemisms seem plausible (Mexico,
Southern Africa, Seychelles, Cuba, Australia), while others may be due to inadequate
research. The “Schizomus” of Africa should be re-described. Species of this order
do not live in cooler areas such as the Palearctic and most of the Nearctic, as well as
New Zealand. Very rich fauna diversity, with three endemic and two non-endemic
genera, was recorded by Harvey (2001) in the Seychelles.
11.3 Amblypygi
Distribution of this order is a result of climate. This order lives mainly in warmer
environments. Many genera are widely dispersed (e.g., Charinus, Sarax, Charon,
Damon, Phrynichus, Paraphrynus, Phrynus), including across seas to many islands.
Some genera are more restricted – to Southern Africa (e.g., Xerophrynus,
11.4 Thelyphonida (Uropygi) 895
Phrynichodamon), North-west Africa (Musicodamon), and Myanmar (Catageus).

In general, Charinidae are widespread, Charontidae are found in South-east Asia
and Melanesia. Phrynichidae prefer tropical environments such as Southern Africa,
the Arabian Peninsula, Mauritius, Seychelles, Zanzibar, São Tomé and Príncipe,
Madagascar, Sri Lanka, and Brazil (one genus). Phrynidae live in tropical South and
Central America, the southern USA, the Caribbean, and Indonesia (one sp. of genus
Phynus). A relictual population of suborder Palaeoamblypygi is known from West
Africa.
Some Peculiar Distributions
Phrynichidae – Trichodamon Mello-Leitão (1935) has been described in Brazil
(two spp.). All other members of the family live in the Old World (Africa, Europe,
and Asia).
Phrynidae – One species of Phrynus has been described from a cave in Indonesia
(Flores Island).
Paracharontidae – Endemic suborder (Palaeoamblypygi), family, genus
(Paracharon Hansen, 1921) and species from Guinea Bissau.
11.4 Thelyphonida (Uropygi)
This order lives in warm environments and is almost unknown in the Palearctic and
very limited in the Nearctic. It is not found in Australia. These ground-dwelling
organisms are not able to travel easily from one island to another (unlike Amblypygi)
and their bigger size makes them even more difficult to import with earth or plant
material (unlike Palpigradi or Schizomida). The Uropygi populations on islands
(e.g., Hispaniola, Cuba, Hainan, Philippines, Sri Lanka, New Ireland, Java) probably
originated when these islands were connected with other continents.
Hypoctoninae are found mostly in South and South-east Asia, with one genus in
South America and one in the Dominican Republic.
Uroproctinae – there are three genera in North and South Americas, one in India
and one in the Philippines.
Hypoctoninae – Etienneus Heurtault (1984) is the only member of Uropygi in
Africa. Considered a relict in West Africa, the following questions regarding its
distribution have been raised: (1) With so many Uropygi in the other parts of the
tropics, why are there no other members of this genus in Africa and Australia; (2)
Why did these West African Uropygids not spread to the rest of Africa, or at least
more widely in West Africa? (3) If Etienneus is a relict, what restricts its distribution?
In both subfamilies, there are genera in North and South America and South-east
Asia, but not in Africa. Therefore, no Gondwanian explanation seems appropriate.
This same disjunction (present in South America and South-east Asia, but absent in
Africa) is observed in the Opilion family Zalmoxidae.
Uroproctinae – one species of the American genus Mastigoproctus (M. transoce-

anicus Lazell, 2000) was described from Hong Kong, but it was most probably
transported there by error.
11.5 Ricinulei
The distribution of this order in North and South America (two genera) and in West
Africa (one genus) indicates its relictual nature, as well as the old age of the order
(i.e., from before the formation of the Atlantic Ocean). The ability of the Ricinulei
to travel (actively or by human transport) seems very limited.
11.6 Solifugae
The Solifugae order prefers warm environments (i.e., climate-dependent distribu-

tion). There is a sharp distinction between the North and South American families
(Mummuciidae, Eremobathidae, Ammotrechidae) and the families of the Old World
(Galeodidae, Karschiidae, Solpugidae, Ceromidae, Melanoblossiidae, Gylippidae,
Hexisopodidae, Rhagodidae). The only family living on both sides of the Atlantic
Ocean is Daesiidae (Europe, Central Asia, the Middle East, Africa, and South
America). From the eight families of the Old World group, two (Ceromidae and
Hexisopodidae) are proper to Southern Africa.
The Solifugae order are desert-dwelling Arachnids. The reason for the absence of
this order from Australia is unclear. Most eastern living sun spiders are far from
Australia (Dinorhax rostrumpsittaci, Melanoblossiidae, from Vietnam and
Malucus). It is unclear why the order is absent from Madagascar, which is only a
few hundred miles across the Mozambique channel and not far from one of the
richest areas in Solifugae (Southern Africa) with200 species from six families. Sun
spiders are not good candidates for rafting or other means of water transport or
human transport, but they somehow managed to reach the Galapagos Islands, which
are much further from the mainland than Madagascar is from the African mainland.
Some distributions seem rather disjunct (e.g., Melanoblossiidae – South-east
Asia and Southern Africa; Gylippidae – Western Asia and Southern Africa). The
explanation of some present-day distributions resides in the past fauna – Ceromidae
is presently limited to Africa, but one fossil species has been found in Brazil.
11.8 Pseudoscorpiones 897
11.7 Scorpiones
The countries with the most species of scorpions are Mexico (272 spp.), Venezuela
(230 spp.), Brazil (165 spp.), USA (111 spp.), and India (117 spp.). Huge territories
are almost devoid of these warm-loving animals (Russia – five spp., Canada – one
sp.). In China, there are fewer scorpion species (50) when compared to the much
smaller Cuba (55 spp.).
The blind European genus and species Belisarius Simon is related to Neotropical
fauna, but the specialists cannot agree in which family to place it – Troglotayosicidae
(Lourenço 1998) or Chactidae (Soleglad and Fet 2003). In American scorpions, the
unique (and still only known from dead specimens) genus Akrav Levy was found
in a Palestinian cave (a monotypic family Akravidae). Another surprise was to find
another new and relic family Pseudochactidae in Central Asia (Gromov 1998).
Two new genera (Troglokhammouanus Lourenço and Vietbocap Lourenço et
Pham) and four new species have been discovered in the caves of Indochina (Laos
and Vietnam). Other peculiar disjunctions, like the distribution of Heteronebo
Pocock in the Caribbean and on the islands Abd-el-Kuri near Yemen, have been
discussed earlier.
11.8 Pseudoscorpiones
The Pseudoscorpiones is a large order that is widespread and has little endemism at
the family level. The richest (or best studied?) countries are Italy (246 spp.), Spain
(196 spp., including the Canary Islands), South Africa (135 spp.), and Australia
(181 spp., Tasmania excluded). These figures are from Harvey (2000), Lissner
(2014), Gardini (2000), Dippenaar-Schoenman and Harvey (2000), and other
sources.
This order is well-represented in caves.
As with some other groups (e.g., Isopoda, Scorpiones), the presence of American
elements in the Mediterranean subregion is surprising and demands explanation. An
example of this was to find a representative of Bochicidae (known from the
Antilleans, Mexico, Texas, and Guatemala) in a cave in Portugal. As with many
other cave animals, the genus Titanobochica Reboleira is considered to be a relict.
Other such relicts in the Mediterranean (all from caves) are Troglobisium Beier
(Spain), Troglochthonius Beier (Dalmatia, Croatia), and Neobalkanella Ćurčić
(Serbia). Another cave endemic (from Botswana) is Botswanoncus ellisi Harvey
and Du Preez, Ideoroncidae. The presence of the family Syarinidae in Europe is also
considered relictual. The American family Gymnobisiidae has one genus
(Gymnobisium Beier) in South Africa.
11.9 Opiliones
Cyphophthami
In Europe, there are two areas of intense speciation of Cyphophthalmi, but of differ-
ent types. In Spain and Portugal (i.e., the Iberian Peninsula), there are four genera,
three of which (Paramiopsalis Juberthie, Iberosiro de Bivort, Odontosiro Juberthie)
are monotypic and endemic. On the Balkans there are about 17 species of
Cyphophthalmus Joseph (including Tranteeva Kratochvil).
As pointed out by Giribet and Kury (2007), “each of the six recognized families
has a well-characterized biogeographic distribution.” Sironidae exhibit Laurasian
distribution, Pettalidae is distributed following a temperate Gondwanian model,
Troglosironidae is endemic to New Caledonia, etc. Cyphophthalmi are found on all
continents and major islands (including New Zealand, Madagascar, Sri Lanka, and
New Caledonia).
From the distribution map is clear that Cyphophthalmi are limited by climate and
ecological factors (cold, aridity, etc.). They are absent in most of the Palearctic
(except for Europe, Turkey, and Japan), missing in Palearctic Asia, North Europe,
North Africa, internal Australia, and most of Canada. In some areas (Balkans,
Iberian Peninsula, New Zealand, New Caledonia, and South-east Asia), an active
speciation is observed.
To quote Clouse and Giribet (2010), “Cyphophthalmi, which are known poor
dispersers and yet widespread and species rich, are here shown to be excellent
models for studying the ancient history of Southeast Asia” (author addition: also of
other regions), and also: “Molecular dating suggested that Stylocellidae separated
from other Cyphophthalmi 295 Ma and began diversifying 258 Ma, and the lineage
that inhabits mostly Borneo today began diversifying between 175 and 150 Ma.
Stylocellidae originated on Gondwana, arrived in Southeast Asia via the Cimmerian
palaeocontinent, and subsequently diversified north, then south. Their present
distribution in the Indo-Malay Archipelago is explained largely by a diversification
over the Sundaland Peninsula before western Sulawesi departed and the peninsula
was extensively inundated”. Based on Clouse (2012), Stylocellidae are the most
ancient animal group in South-east Asia.
Sri Lanka was detached from India only recently, but none of the members of
Pettalidae is known from the continent. The Cyphophthalmi, however, remained in
New Caledonia (Troglosironidae) and in New Zealand, where many orders of
Arachnida are missing. The connections of the New Zealand opilions and spiders
with the fauna of Patagonia (Chile, Argentina) are most intriguing.
Eupnoi
This rich suborder (ca. 1700 spp.) is predominant in the Northern Hemisphere
(Phalangiidae, Sclerosomatidae). More interesting is the family Protolophidae
(eight spp. endemic to the western USA) and Neopilionidae (including Monoscutidae)
found in South Africa, Chile, Argentina, Brazil, Australia, New Zealand, Snares,
Campbell Islands (20 genera, 64 spp.) – the “Southern end of the world.”
11.9 Opiliones 899
Among the genera with disjunct distributions is Caddo Banks (Caddidae), which
is found in the USA and Japan, with fossils found in Europe.
“The Boreotropic Concept is potentially relevant to sclerosomatid biogeography.
It is derived from evidence that global temperatures were unusually high during the
late Paleocene and early Eocene, that the northern subpolar regions had tropical to
subtropical climates during this time, and that there was substantial biotic
interchange between continents via Beringian and North Atlantic terrestrial
corridors. As the northern climate cooled during the Tertiary, the tropical elements
within each continent retreated southward, lost their connections to each other due
to latitudinal climatic and longitudinal oceanic barriers. An interchange of temperate
groups persisted to varying degrees until it was terminated by Pleistocene glaciation.
This scenario appears to explain certain similarities in the biotas of the New World
and Asian tropics as well as between temperate North American and East Asia. The
Boreotropic Concept has been developed and applied most thoroughly by plant
systematists…although there are zoological examples as well …. including non-
sclerosomatid harvestmen (Suzuki et al. 1977; Shultz and Regier 2009). The
existence of circumboreal tropics in the early Tertiary may have allowed movements
of ancestral gagrelline-like sclerosomatids between Eurasia and North America.
These taxa would then have followed the tropics southward, producing the modern
disjunction between the largely tropical New World and Asian “Gagrellinae” of
today” (Hedin et al. 2012).
Dyspnoi
After the important revision of Schönhofer (2013), we know that most members of
the suborder inhabit the Holarctic Kingdom (Ischyropsalididae, Sabaconidae,
Taracidae, Dicranolasmatidae, Trogulidae, Nemastomatidae, Niponopsalididae).
Some of the seven families of the Dyspnoi suborder are missing from the warmer
parts of the world, and the entire suborder is missing from peninsular India, tropical
and Southern Africa, tropical South America, Sri Lanka, Australia, New Guinea,
etc.
The Taracidae family includes genera found in North America and far east Russia.
Laniatores
The Laniatores is the largest suborder of Opiliones and has 23 families and more
than 4180 species (Kury, 2013). Most of these species live in the tropics. In the
Neotropics, there are 19 families, 13 of them are endemic (Kury, Catalogue). In the
Palearctic countries, very few genera (Lola, Paralola, Dinaria, Travunia, Abasola,
Haasus, Trojanela, Buemarinoa, Proscotolemon, Ptychosoma, Holoscotolemon) of
the families Phalangodidae, Travuniidae, and Cladonychiidae, live in caves. These
genera have been considered relicts in Europe, but some specialists (Martens,
1978a) disagree.
The Pyramidopidae family are found almost exclusively on the African continent
and near islands (Bioko, São Tomé and Príncipe), with one endemic genus
(Maiorerus Rambla) found in a cave in Fuerteventura (Canary Islands) and another

genus (Jarmilana Cruz-López, Proud et Pérez-Gonzalez) found in Belize. This
species is a troglobite, the first member of fam. Pyramidopidae in the New World
(North and South America).
Sharma et al. (2011) states the following: “Fuerteventura, which is 1800 km
north of the tropical belt, has a sub-Saharan climate and is the oldest island in the
archipelago, dating back 20.6 million years (Carracedo et al. 1998). The dynamics
of arthropod diversity on another group of Atlantic islands, the Azores, suggests that
the number of endemic cavernicolous species declines with the age of the island,
due to erosive processes (Borges and Hortal 2009). If this phenomenon has occurred
in the Canaries, Maiorerus randoi may represent the remnant of a formerly diverse
lineage of Pyramidopidae that inhabited the erstwhile extensive cave systems of
Fuerteventura” (Sharma et al. 2011).
Another more startling finding with regard to Pyramidopidae was published by
Cruz-Lopez et al. (2016). They discovered that the former troglobitic “Stygnomma”
pecki (Goodnight and Goodnight, 1977) was actually a member of the Pyramidopidae
family, the first one discovered in the New World. This lead Cruz-Lopez et al. (2016)
to describe the new genus Jarmilana. This finding raises the question of whether
this represents transoceanic dispersal or a relict of an ancient widespread tropical
Gondwanian distribution.
11.10 Araneae
Of the 112 families of the Araneae order, about 21 are very widespread, almost
cosmopolytic, and they are of little zoogeographic interest. Among them are
Scytodidae, Sicariidae, Pholcidae, Segestriidae, Oecobiidae, Uloboridae, Araneidae,
Tetragnathidae, Theridiidae, Lycosidae, Oxyopidae, Pisauridae, Agelenidae,
Dictynidae, Hahniidae, Gnaphosidae, Philodromidae, Thomisidae, Salticidae,
Corinnidae, and Liocranidae. Notes on the distribution of the remaining families are
below. It will be easier for the reader to analyze these distributions using a map.
Fam. Mecicobothriidae – distributed in the southern-most part of South America
and Mexico and California.
Fam. Microstigmatidae – South America from Panama (Micromegalinae) to
Argentina, one genus in South Africa (Microstigmata)
Subfam. Micromygalinae – Panama (one sp.)
Subfam. Microstigmatinae – Brazil, Argentina, Uruguay, Colombia, Venezuela,
Ecuador, South Africa (15 spp.)
Fam. Hexathelidae (Hexathelinae) – New Zealand, Australia, Chile, Argentina
(Mediothele)
Fam. Atypidae – missing in South America, Madagascar and Australia
Fam. Antrodiaetidae – USA (30 spp.), Japan (two spp.) A transpacific distribution.
Two species (Antrodiaetus roretzi and A. yesoensis) are endemic to Japan. They
are considered relict species (Miller and Coyle 1996).
11.10 Araneae 901
Fam. Idiopidae (Genysinae) – Central and South America, Madagascar, India, Sri
Lanka, Seychelles. Missing in Africa.
Fam. Migidae (Calathotarsinae) – Chile, Argentina, Australia
Fam. Actinopodidae – South America, Panama, Australia
Fam. Hypochilidae – China, USA
Fam. Austrochilidae – Chile, Argentina, Australia (Tasmania - nine spp.)
Subfamily Austrochilinae – Chile, Argentina (eight spp.)
Subfamily Hickmaniinae (as family Hickmaniidae) – Australian (Tasmanian
cave)
Fam. Drymusidae – South Africa, Brazil, Argentina, Caribbean
Fam. Leptonetidae – USA, Mexico, Panama, Mediterranean, Portugal, Algeria,
Japan, China (Mediterranean – Far East disjunction)
Fam. Telemidae – Sumatra, Malaysia, China, Vietnam, Japan, France, Spain,
Tanzania, Cameroon, South Africa, Seychelles, New Caledonia, New Guinea,
Guatemala, USA (Oregon, Alaska, ) – patchy distribution, relict. Missing in
South America, Australia, and New Zealand
Fam. Orsolobidae – Australia, New Zealand, Southern Africa,, Brazil, Chile,
Argentina, Falkland Islands (177 spp.)
Fam. Eresidae – Africa, Madagascar, Brazil, Eurasia. Widespread in Palearctic,
missing in Nearctic.
Fam. Mecysmaucheniidae – Chile, Argentina, Falkland Islands, Juan Fernandez
Islands., New Zealand
Fam. Micropholcommatidae – New Zealand, Australia, Auckland Islands, Campbell
Islands, New Guinea, Brazil, Chile
Fam. Palpimanidae – Africa, Socotra, South America, Cuba, South Asia, Uzbekistan,
Mediterranean, Seychelles, Sri Lanka. Missing in Nearctic.
New Guinea, Fiji, Borneo, Vietnam, India. Missing in Australia.
Fam. Malkaridae – Australia, Chile, Argentina
Fam. Mimetidae (Melaenosiinae) – São Tomé and Príncipe, India
Fam. Cyatholipidae – Africa, Madagascar, New Zealand, Australia, and one species
(Pokennips dentipes) in Jamaica
Fam. Synaphridae – Canary Islands., Southern Europe, Egypt, Turkmenistan, Spain,
Madagascar (three spp.)
Fam. Synotaxidae – Australia, New Zealand, South America
Fam. Stiphidiidae – Australia, New Zealand, Madagascar, Mauritius
Fam. Trechaleidae – South and Central America, Mexico, USA, Japan
(transpacific)
Fam. Udubidae – Sri Lanka, Madagascar, Africa, USA to Panama
Fam. Zoropsidae – Australia, New Zealand, Sri Lanka, China, Korea, Japan, Cyprus,
Mediterranean, North and South Africa. Missing in the Americas and in tropical
Africa
Fam. Amphinectidae – Chile, Australia, New Zealand
Fam. Zoropsidae – U.S.A., Mexico, South America, Costa Rica, Madagascar, New
Zealand
Fam. Titanoecidae – Missing in Africa, present in New Guinea and the Marquesas
Islands, but missing in Australia
Fam. Ammoxenidae – Australia, Southern Africa
Fam. Gallieniellidae (Meedoinae) – Australia (five genera), Argentina (one genus)
Interesting is the case of the relict family Liphystiidae, the sole member of the
suborder Mesothelae. This family is found in Japan, China and South-East Asia.
According to Xu et al. (2015), these spiders are “living fossils” and the suborder
Mesothelae is an ancient clade sister of all modern spiders. According to Xu et al.
(2015), Liphystiidae genera originate in Asia in the Paleogene (4–24 Ma).
This timing is relatively recent, taking into account the old age of the spider
divergence (297.6 Ma) between the Mesothelae and the Opisthothelae
(Mygalomorphae and Araneomorphae). The existence of Mesothelae in Japan
(Kyushu and Ryukyu Islands) is explained by Haupt (2003) as follows: “through
vicariant origins in the Tertiary when the Japanese island arc separated from
mainland Asia, or alternatively, as a consequence of dispersal events over land
bridges from east China during the Pleistocene.”
11.11 Opilioacarida
This order includes one family with 13 genera, 41 rec. species found in Yemen,
Cuba, Dominican Republic, Panama, India, USA, Mexico, Nicaragua, Venezuela,
Italy, Greece, Algeria, Angola, Ivory Coast, Kazakhstan, Gabon, South Africa,
Mozambique, Tanzania, Madagascar, India, and Thailand (i.e., the Afrotropical
Region (including Yemen), Madagascar, North Africa, South- east Asia, South
Europe, Kazakhstan, South and Central America, Caribbean Islands (Beron 2014).
The distribution of this family is dependent on climate.
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Chapter 12
Zoogeographical Subdivisions
of the World
Abstract But what is biogeographical region? There are different definitions, one
of which is written by Wallace (1894): “…those primary divisions of the earth’s
surface of approximately continental extent, which are characterized by distinct
assemblages of animal types.” Review and analysis of many zoogeographical
schemes for easier checking how the orders of Arachnida fit into them.
But what is biogeographical region? There are different definitions, one of which is
written by Wallace (1894): “…those primary divisions of the earth’s surface of
approximately continental extent, which are characterized by distinct assemblages
of animal types.”
Other sources: Dickerson et al.(1928), Illies (1974), Kolosov (1980), Lydekker
(1911), Müller (1973, 1974a, b), Paulian (1951), Schmidt (1954), Wallace (1876),
Weber (1902)
Review of some of the more recent zoogeographies and biogeographies. Included
are also several Russian authors, less used in the Western literature.
Subdivision of Sclater (1858) – six regions, Palearctic, Ethiopian, Indian,
Australian, Nearctic, and Neotropical, combined in two bigger “Creacio” –
C. Paleogeana (Old World) and C. Neogeana (New World)
Huxley (1868) united the four more northern regions (Africa, Eurasia, and North
America) in “Arctogea” and South America and Australia in “Notogea.”
Wallace (1876) renamed the Indian Region Oriental.
Heilprin (1887) united the Palearctic and the Nearctic Regions into Holarctic
Region.
Several authors try to alter the classical scheme of Sclater-Wallace, based on vertebrates. In one of
the recent articles, Rueda et al. (2011) used modern calculations. Again based on mammals, birds,
and amphibians, after many efforts, they concluded that “Using modern analytical techniques and
our current knowledge of distributions and generic concepts, we have shown that when Wallace’s
principles are followed, it is possible to obtain a map of zoological regions quite similar to the map
he generated more than a century ago.” Maybe we have to pay tribute to the people like Sclater and
Wallace. Our task is to compare the classical subdivision, based on vertebrates, with the distribu-
tion of Arachnida.

https://doi.org/10.1007/978-3-319-74418-6_12
906 12 Zoogeographical Subdivisions of the World
Blanford (1890) defined the three groups: Arctogea, South American, and
Australian. Lydekker (1896, 1911) called them “Kingdoms.”
Bobrinskiy et al. (1946) recognize eight regions, grouped into four “dry lands”
(Gea) (in Russia by 1946, it was not advisable to use the word “kingdom”):
Notogea
New Zealand region
Australian region
Polynesian region
Neogea
Neotropical region
Paleogea
Madagascan region
Ethiopian region
Indomalayan region
Arctogea
Holarctic region
Darlington (1957) renamed Arctogea as Megagea. He accepted the following
subdivision of land:
Kingdom Megagea (Arctogea)
Ethiopian region
Indomalayan region
Palearctic region
Nearctic region
Kingdom Neogea
Neotropical region
Kingdom Notogea
Australian region
De Lattin (1967): quite simplified subdivision, still recognizing the three
“Kingdoms” (Faunenreich):
Kingdom Megagea or Arctogea
Holarctic region
Subregions Palearctic and Nearctic
Oriental (Indian) region
Ethiopian region
Subregions African and Malgashian
Kingdom Neogea
Neotropical region
12 Zoogeographical Subdivisions of the World 907
Kingdom Notogea
Australian region
Subregions: Continental-Australian, New Zealandian, and Polynesian
Map 12.1 Biochors of the land biocycle: 1, arboreal; 2, eremial; 3, oreotundral (De Lattin 1967)
Müller (1974a, b) subdivided the Earth into five realms and nine regions:
1. Holarctic
(a) Nearctic
(b) Palearctic
2. Paleotropical
(a) Ethiopian
(b) Madagascan
(c) Oriental
3. Australian
(a) Australian
(b) Oceanic
(c) New Zealand
(d) Hawaiian
4. Neotropical
5. Archinotic (Antarctic, southwestern South America, and southwestern New
Zealand)
Map 12.2 Zoogeography of the land regions and subregions (Buchar 1983)
Kingdom Region Subregion

A. Notogea 1. Australian a. Australian Cont.
b. New Zealandian
c. Polynesian
d. Hawaiian
e. Wallacea
B. Neotropic 2. South American a. Brazilian
(Neogea) b. Chilean-Patagonian
c. Antillean
d. Middle American
C. Paleotropic 3. Ethiopian a. African
b. Madagascarian
4. Indomalayan a. Indian
b. Malayan
c. Indonesian
D. Holarctic 5. Nearctic a. Canadian
b. East Nearctic
c. West-Central
6. Palearctic d. Californian
a. Eurosiberian
b. Mediterranean
c. Middle European
d. East Palearctic
Map 12.3 (Müller 1974a, b, p. 42)
Map 12.4 Subdivision of Holarctic after Schilder (1956)
1. Hyperboreal region
2. Sonoran region
3. Palearctic region
Map 12.5 Subdivision of Holarctic after Krijanovskiy (1965)
1. Circumpolar region
2. Sonoran region
3. Region of the Old Mediterranean (pre-Mediterranean)
4. East Asian region
Lehtinen (1980) analyzes the arachnological zoogeography of the Indo-Pacific
region. His subdivision (centers of speciation):
1. Indo-Pacific region
(a) South India and Ceylon
(b) East-Himalaya – Indochina
(c) Malayan Archipelago
(d) New Guinea
(e) Hawaii
2. South-Gondwanan region
(a) South Africa
(b) Tasmania and Southwest Australia
(c) New Caledonia
(d) New Zealand
(e) South Chile
3. Madagascan region
Map 12.6 Centers of speciation in the new subdivision of Australasia (After Lehtinen 1980)
Map 12.7 Zoogeographical subregions of the Palearctic Region after Sergeev (1993)
I – the Eurosiberian subregion (two provinces)

II – the Scythian subregion (at least four provinces)
III – the Manchurian subregion (at least four provinces)
IV – the Orthrian subregion (at least four provinces)
V – the West Mediterranean subregion (at least three provinces)
VI – the East Mediterranean subregion (at least four provinces)
VII – the Saharan-Gobian subregion (at least 22 provinces)
Map 12.8 Zoogeographical provinces of North and Central Asia (After Sergeev 1993)
(1–2) The Eurosiberian subregion

1 – the Arctic province
2 – the Boreal province
(3–4) The Manchurian subregion
3 – the Amurian province
4 – the Sakhalin-Hokkaido province
(5–6) The Scythian subregion
5 – the Russian-Siberian province
6 – the Sarmathian province
(7–14) The Saharan-Gobian subregion
7 – the Kazakhstan province
8 – the Mongolian province
9 – the Turanian province
10 – the Gissar-Tien Shan province
11 – the Inner Tien Shan province
13 – the Pamirian province
14 – the Turkmeno-Iranian province
Lopatin (1980), specialist of Coleoptera, made the following (quite different)
subdivision of land fauna (Map 12.8):
Map 12.9 Zoogeography of the world, according to Lopatin (1980)
I Kingdom Paleogea
I1 Ethiopian region (with subregions East African, West African, and South
African or Cape Subregion)
I2 Indomalayan region (with subregions Indian, Malayan, Indochinese, and
Papuan)
I3 Madagascan region
I4 Polynesian region
II Kingdom Arctogea
Subkingdom Palearctic
II1 European-Siberian region (with subregions European-Obian and Angaran)
II2 Region of the Ancient Mediterranean (with subregions Mediterranean and
Saharo-Gobian)
II3 East-Asiatic region
Subkingdom Nearctic
II4 Canadian region (north of 50oN, with subregions, or provinces Alaskan,
Labradorean, and Greenland)
II5 Sonoran region (with subregions Californian, Rocky Mountains, and Atlantic)
III Kingdom Neogea
III1 Neotropical region (with subregions Guyano-Brazilian, Central American,
and Pampas)
III2 Caribbean region
IV Kingdom Notogea
IV1 Australian region
IV2 New Zealand region
IV3 Patagonian region
Morrone (2015) proposed a new biogeographic regionalization of the world:

Three kingdoms and nine regions:
Holarctic Kingdom (Nearctic and Palearctic regions)
Holotropical Kingdom (Neotropical, Ethiopian, and Oriental regions)
Austral Kingdom (Cape, Andean, Australian, and Antarctic regions)
Additionally, the following five transition zones are recognized: Mexican
(Nearctic-Neotropical transition), Saharo-Arabian (Palearctic-Ethiopian transition),
Chinese (Palearctic-Oriental transition), Indomalayan (Oriental-Australian transi-
tion), and South American (Neotropical-Andean) (Map 49).
Krizhanovskij (Крыжановский) (1980) analyzed the Paleotropical (Paleogean)
Dominion in an interesting article (in Russian, what makes it not very used in the
Western literature). According to him, the Dominion is subdivided in four
regions: Afrotropical, Madagascan, Indomalayan, and Papuan. If we look closer
at the map of the Dominion (p.62), some interesting conclusions are to be noted.
Together with the south of Arabian Peninsula, in the Afrotropical Region are
included southern Iran and western India. Included in the Papuan Region are
Sulawesi and the whole of Wallacea. Most of South China and Taiwan are
excluded from the Indomalayan Region, so is the chain of Riu-Kyu.
Map 12.10 The “Paleotropical Dominium” of Krzhanovskiy (1980) I, Afrotropical region; II,
Madagascan region; III, Indomalayan region; IV, Papuan region
Map 12.11 Zoogeographical subdivision of the Earth (After Krizhanovskij 2002)
1. Notogean Kingdom
1.1. Australian region
1.2. New Zealandian region
1.3. Chilean-Patagonian region
2. Neogean Kingdom
Neotropical region
3. Paleotropical Kingdom
3.1. Afrotropical region
3.2. Madagascan region
3.3. Indomalayan region
3.4. Papuan region
4. Holarctic Kingdom
4.1. Boreal region
4.2. Region of the Ancient Mediterraneum
4.3. East Asian region
4.4. Sonoran region
Rapoport (1968) proposed another revolutionary subdivision of the Earth in three
“cinturones biogeograficos” (biogeographical belts): Holartica (approx. equal to the
classical Holarctica), Holotropical (Neotropica plus Paleotropica) and Holantártica
(Fuegopatagonia, Cape area, Australia, New Zealand, New Guinea, and Antarctica,
roughly equal to Notogea).
Crosskey and White (1977) suggested the need to replace the term “Ethiopian”
with Afrotropical, which is now favored by most specialists.
Krizhanovskij (1980) analyzes the “Paleotropical Dominion” (= Kingdom), con-

sidering four components (regions): Afrotropical (Ethiopian), Madagascar, Papuan,
and Indomalayan (Oriental).
Map 12.12 Secondary faunistical centers within the primary arboreal Mediterranean center: 1.
Atlantic-Mediterranean, 2. Adriatic-Mediterranean, 3. Pontic-Mediterranean, 4. Tyrenian, 5.
Canarian, 6. Mauretanian, 7. Kyrenaian, 8. Cretan, 9. Cyprean (De Lattin 1967)
Ficetola et al. (2017) – defining the global determinants of zoogeographical bound-

aries (“shallow, deep, and intermediate”), these authors outline the following bio-
geographical regions: African, Amazonian, Arctico-Siberian, Australian, Chinese,
Eurasian, Guineo-Congolian, Indomalayan, Japanese, Madagascar, Mexican, North
American (= Nearctic), Novozelandic, Oriental, Panamanian, Papua-Melanesian,
South American, Saharo-Arabian, Tibetan, and Polynesian. The present author finds
this subdivision (calculated by clustering after deferral formulas) unacceptable.
Especially strange are the names like “Panamanian,” “South American” (different
from Amazonian), “Chinese,” etc. Moreover, the title of the article is “zoogeograph-
ical” and not “biogeographical” boundaries. The idea to weigh the importance of
the determinants of the boundaries (climatic, historical, tectonic, altitudinal, flora,
barriers, etc.) is interesting, but the output is unnecessary complex and
unacceptable.
Map 12.13 Main biochors of the land biocycle: 1, arboreal; 2, eremial; 3, oreotundral (De Lattin
1967)
La Greca (1962) outlined the following “types of distribution” of elements in

the Italian fauna:
Cosmopolitan (the species limited by the tropical climate are indicated as tropico-
politan or pantropical)
Holarctic
Holopalearctic
Euroasiatic
Eurosiberian
Eurocentroasiatic (including Euroturanic and Centroasiatico-pontian)
Holomediterranean (Circummediterranean), including Mediterranean-
Macaronesian, Mediterranean-Atlantic, Mediterranean-Iranic, Mediterranean-
Pontic, and Mediterranean-Turanic
Other distributions within the Mediterranean are Occidental-Mediterranean,
Nordmediterranean, Circumtyrrenian, Circumadriatic, and Transadriatic
European
Medioeuropean
Southeuropean
Centromeridional-European
Westeuropean
Easteuropean
Boreoamphiatlantic
Boreoalpine
Eurosouthmediterranean
Euroanatolian
Euromaghrebian
Alpine
Appeninian
Alpine-Appeninian
Ionian
Siculan (Sizilian)
Appeninian-Southmediterranean
Sardinian and Corsosardinian
Euro-African
Euro-Afroindian
The South African biogeographers Procheş and Ramdhani (2012) based their
analysis on the 821 ecoregions, defined by WWF. With good reason these authors
ascertain that “Genus-level data are preferable to species- or family- level data.”
They also incorporate the clustering method and say that “The clusters of ecore-
gions that were found to be consistently congruent and were well defined across the
four analyses…are recognized here as zoogeographical regions.” They also tried to
propose an updated global zoogeographical regionalization. “Eleven vertebrate-rich
(Nearctic, Caribbean, Neotropical, Andean, Palearctic, Afrotropical, Madagascan,
Indo-Malaysian, Wallacean, New Guinean, Australian) and three vertebrate-poor
(Arctic, Antarctic, Polynesian) zoogeographical regions were derived; the
Neotropical, Afrotropical, and Australian had the highest numbers of characteristic
tetrapod genera.”
Summary of the zoogeographical regions and subregions recognized here (in
Procheş and Ramdhani 2012), as compared with the zoogeographical scheme of
Darlington (1957)
Darlington (1957) Procheş and Ramdhani (2012)

Kingdom Megagea (Arctogea) Regions:
Ethiopian Region Nearctic
Indomalayan Region Caribbean
Palearctic Region Neotropical
Nearctic Region Andean
Kingdom Neogea Palearctic
Neotropical Region Afrotropical
Kingdom Notogea Madagascan
Australian Region Indo-Malaysian
Wallacean
New Guinean
Australian
Arctic
Antarctic
Polynesian
A chorotype classification of the Near East fauna has been proposed in 1999 by
Vigna Taglianti and ten other Italian specialists in Chilopoda, Coleoptera, Amphibia,
and Reptilia. This classification is widely used by people who make z oogeographical
interpretations of their results in different groups. It would be interesting to check
how the chorotypes of Italian colleagues would coincide with the distribution of
such zoogeographically important arthropods like the Arachnids.
Recently a group of authors (Holt et al. 2013) proposed a completely new subdi-
vision of land, analyzing the distribution of 21,037 amphibians, birds, and mam-
mals. The main difference from the system of Sclater-Wallace is that phylogenetic
relations are taken into account and that on the newly proposed scheme, the Earth’s
land is divided in 10 realms: Oceanian, Nearctic, Neotropical, Saharo-Arabian,
Afrotropical, Madagascan, Palearctic, Sino-Japanese, Oriental, and Australian.
Map 12.14 Subdivision of the World after Holt et al. (2013)
Kingdom Megagea (Arctogea)

Ethiopian region
Indomalayan region
Palearctic region
Nearctic region
Kingdom Neogea
Neotropical region
Holarctica
Heilprin (1887) – Holarctic region
Geptner (1936) – Holarctic region
Bobrinskij et al. (1946) – Holarctic (also written “Holoarctic”) region of the
Arctogea “Earth” (Gea)
Darlington (1957) – no mentioning
De Lattin (1967) – Holarctic region within Kingdom Megagea
Müller (1974a, b) – Holarctic realm

Lopatin (1980) – Kingdom Arctogea
Krizhanovskij (2002) – Holarctic Kingdom
Boreal region
Region of the Ancient Mediterranean
East Asian region
Sonoran region
Morrone (2015) – Holarctic Kingdom
Palearctic
Geptner (1936) – only conditional name within Holarctic region
Bobrinskij et al. (1946) – only conditional name within Holarctic region
Darlington (1957) – Palearctic region within Kingdom Megagea
De Lattin (1967) – Palearctic subregion within Holarctic region
Müller (1974a, b) – Palearctic region within Holarctic realm
Lopatin (1980) – Palearctic Subkingdom
Sergeev (1993) – Palearctic region
Morrone (2015) – Palearctic region
Subdivisions of Palearctic
Geptner (1936) – in the Palearctic section of Holarctic are recognized five subre-
gions European-Siberian, Mediterranean, Central Asiatic, Mandjuro-Chinese, and
Arktic (in common with Nearctic section)
Darlington (1957) – does not recognize clear regionalization
De Lattin (1967) – no subdivision
Müller (1974a, b) – no subdivision
Lopatin (1980, 1989)
Palearctic Subkingdom
Eurosiberian region
European-Obian subregion
Angarian subregion
Ancient Mediterranean region
Mediterranean subregion
Sahara-Gobi subregion
East-Asiatic region
Arctic
Geptner (1936) – Arctic subregion of Holarctic region (circumpolar)
Sergeev (1993) – Arctic province of the Eurosiberian region
European (Eurosiberian) subregion
Geptner (1936) – European-Siberian region
Bobrinskij et al. (1946) – no subregions
Darlington (1957) – no subregions
De Lattin (1967) – no subregions
Lopatin (1980) – European-Siberian region (with subregions European-Obian –

West of Enisey, Angaran)
Mediterranean subregion
Geptner (1936) – Mediterranean subregion
Darlington (1957) – no subdivision
Müller (1974a, b) – no subdivision
Lopatin (1980) – region of the Ancient Mediterranean (circummediterannean
lands and Central Asia). Two subregions: Mediterranean and Saharo-Gobian.
Saharo-Gobian subregion is divided into three superprovinces: Saharo-Sindian,
Irano-Turanian, and Central Asiatic.
Central Asiatic subregion
Geptner (1936) – Central Asiatic subregion
Lopatin (1980) – Central Asiatic superprovince of Saharo-Gobian subregion
Mandzhurian (Mandjuro-Chinese) subregion
Geptner (1936) – Mandjuro-Chinese subregion
Lopatin (1980) – East Asian (Himalayan-Chinese) region
Irano-Turanian subregion
Lopatin (1980) – superprovince of the Ancient Mediterranean
Nearctic
Geptner (1936) – section of Holarctic region, divided into Canadian and Sonoran
subregions
Darlington (1957) – Nearctic region of Kingdom Megagea (Arctogea)
De Lattin (1967) – Nearctic subregion within Holarctic region
Müller (1974a, b) – Nearctic region of Holarctic realm
Lopatin (1980, 1989) – Nearctic Subkingdom
Morrone (2015) – Nearctic region
Subdivisions of the Nearctic
Bobrinskij et al. (1946) – no subregions, only ecological complexes
Wallace (1876) – four subregions
Geptner (1936) – Canadian and Sonoran subregions of Holarctic region plus
Arctic (in common with Palearctic section)
Lopatin (1980, 1989)
Nearctic Subkingdom
Canadian region
Sonoran region – divided (as in Wallace 1876) into subregions (Californian,
Central, or Rocky Mts., and Eastern or Atlantic)
Neogea
Geptner (1936) – Neotropical region
Bobrinskij et al. (1946) – Neogean “Earth”

Darlington (1957) – Kingdom Neogea
De Lattin (1967) – Kingdom Neogea
Müller (1974a, b) – Neotropical realm
Lopatin (1980) – Kingdom Neogea
Krizhanovskij (2002) – Neotropical Kingdom
Subdivision of Neogea
Bobrinskij et al. (1946) – one region, Neotropical
Darlington (1957) – Neotropical region
De Lattin (1967) – one region, Neotropical
Lopatin (1980) – two regions, Neotropical and Caribbean; Patagonia excluded
Krizhanovskij (2002) – Neotropical region with three subregions
Morrone (2015) – Neotropical region, part of Holotropical Kingdom
Neotropica (Neotropical region)
Bobrinskij et al. (1946) – Neotropical region
Darlington (1957) – Neotropical region
De Lattin (1967) – Neotropical region
Lopatin (1980) – Neotropical region (does not include Caribbean and Patagonia)
Morrone (2015) – Neotropical region
Subdivisions of the Neotropica
Geptner (1936) – four subregions: Antillean (West-Indian), Central American,
Chilean, and Brazilian
Darlington (1957) – Kingdom Neogea, Neotropical region
Rapoport (1968):
La Region Neotropical
Centroamericana
Antillana
Guayano-Brasilena
Andino-Patagonica
Araucana
Lopatin (1980) – three subregions: Guiano-Brazilian, Central American, and
Chilean
Morrone (2001) – two regions with eight subregions (biogeographical) and
Neotropical region (4 subregions, 50 provinces). There is extensive synonymy of
the names given to each of the subregions and the provinces. What concerns
Arachnida, among the taxons, characterizing the Neotropical Region are listed only
one water mite, six genera of spiders and three genera of Opiliones.
Region Andina (4 subregions, 15 provinces)
Krizhanovskij (2002) – three subregions

Guiano-Brazilian subregion
Central American subregion (including Galapagos province)
Antillean subregion
Morrone (2005) outlined (based on insects) the biogeographic areas and the
transitional zones of Latin America and the Caribbean Islands. Actually, the term
“Latin America” has no relation to the biogeography. According to Morrone (2005),
the Nearctic part of Mexico comprises five provinces, California, Baja California,
Sonora, Mexican Plateau, and Tamaulipas, and follows the Mexican transitional
zone, again with five provinces: Sierra Madre Occidental, Sierra Madre Oriental,
Transmexican Volcanic Belt, Balsas Basin, and Sierra Madre del Sur. The
Neotropical region is divided into four subregions: Caribbean, Amazonian, Chacoan,
and Parana. The South American transition zone includes six provinces: North
Andean Paramo, Coastal Peruvian Desert, Puna, Atacama, Prepuna, and Monte.
The Andean region (with insects closely related with Australian fauna) includes
three subregions: Central Chilean, Subantarctic, and Patagonian. The Paramo-Puna
Subregion of Morrone (2001) is included into the transition zone.
From this subdivision (different from all so far existing) it is nevertheless clear
that the Mexican author divides South America into two biogeographical regions
(Neotropical and Andean) with a transitional zone in between.
Morrone (2015) – Andean and Mexican transitional zones
Position of the Caribbean (Antillean) archipelagoes
Geptner (1936) – Antillean (West-Indian) subregion in the Neotropical region
Lopatin (1980) – Caribbean (Antillean) region in the Kingdom Neogea
Krizhanovskij (2002) – Antillean subregion of the Neotropical region
Morrone (2015) – part of Neotropical region
Afrotropical (Ethiopian) region
Geptner (1936) – Ethiopian region
Bobrinskij et al. (1946) – Ethiopian region
Darlington (1957) – Ethiopian region
De Lattin (1967) – Ethiopian region
Müller (1974a, b) – Ethiopian region of Paleotropical realm
Lopatin (1980) – Ethiopian region
Krizhanovskij (2002) – Afrotropical region
Morrone (2015) – Ethiopian region (part of Holotropical Kingdom)
Subdivisions of the Afrotropical (Ethiopian) region
Geptner (1936) – four subregions: West African, East African, South African, and
Madagascan
Bobrinskij et al. (1946):
East African subregion
West African subregion
South African subregion

De Lattin (1967):
African subregion
Malgashian subregion
Lopatin (1980, 1989):
East African subregion
West African subregion
Cape subregion
Krizhanovskij (2002):
Guinea-Congo (West African) subregion
Sudan-Zambezian subregion
South African subregion
Saint Helena Island
Morrone (2015) – an extensive Saharo-Arabian territory in outlined as transition
zone
Madagascar
Some opinions concerning the rank of Madagascar:
Wallace (1876) – subregion within the Ethiopian region
Bobrinskij et al. (1946) – Madagascar region (including the island groups
Mascarene, Seychelles, Amirants, and Comoro)
Millot (1952) – strongly opposes the idea of merging the islands of the Indian
Ocean into one “Région malgache”
Darlington (1957) – analyzed separately from the “Ethiopian”
De Lattin (1967) – part of the Ethiopian region (Malgashian subregion)
Müller (1974a, b) – Madagascan region in the Palaeotropical realm (Madagascar
and its offshore islands)
Vtorov and Drozdov (1974) – Madagascar biophylotic Kingdom
Lopatin (1980) – Madagascar region in the Kingdom Paleogea
Gruev and Kuzmanov (1994) – Madagascar Kingdom (Madagascariensis),
including the island groups Mascarene, Seychelles, Amirants, and Comoro
Procheş and Ramdhani (2012) - Madagascan
Krizhanovskij (2002) – Madagascar region with two subregions:
Madagascan subregion
Mascarenian subregion
Indomalayan region
Sclater (1854) – Indian region
Wallace (1876) – Oriental region
Severtzov (1877) – Indomalayan region
Geptner (1936) – Eastern or Indomalayan region
Bobrinskij et al. (1946) – Indomalayan region
De Lattin (1967) – Oriental region
Lopatin (1980) – Indomalayan region
Krizhanovskij (2002) – Indomalayan region
Morrone (2015) – Oriental region
Subdivision of Indomalayan region

Geptner (1936) – two subregions: Indian and Malayan
Bobrinskij et al. (1946) – five subregions: Indian, Burmese-Chinese, Sundian,
Philippinian, and “Celebes” (Sulawesi)
Lopatin (1980) – four subregions (Indian, Malayan, Indochinese or Burman-
Chinese, and Papuan)
Krizhanovskij (2002): three subregions (Malayan, Indian, and Indochinese)
Morrone (2015) – including Transition Zone
Wallacea
Bobrinskij et al. (1946) – Celebes subregion
De Lattin (1967) – Oriental, Australian intermediate area
Notogea
According to Illies (1972): “Australia and the outlying continental islands of
Tasmania and New Guinea, together with New Zealand and the oceanic islands of
the Pacific, comprise the zoogeographical region of Notogaea” (p.75).
De Lattin (1967) – Australian region with subregions: Continental-Australian,
New Zealandian, and Polynesian
Lopatin (1980) subdivides the Kingdom Notogea into three regions – Australian,
New Zealand, and Patagonian
Polynesian region is part of the Kingdom Paleogea.
Papuan Subregion
Geptner (1936) – Papuan subregion of Australian region (? including Northern
Australia)
Darlington (1957) – part of the Australian region
Lopatin (1980) – Papuan subregion of the Indomalayan region
Krijanovskiy (1980) – Papuan region of the Paleotropical Dominion
Krijanovskiy (2002) – Papuan region of the Paleotropical Kingdom
New Zealand
Geptner (1936) – New Zealandian subregion of the Australian region
Darlington (1957) – undefined (not in Australian region)
De Lattin (1967) – New Zealandian subregion of the Australian region of
Notogea
Lopatin (1980) – New Zealandian region of Notogea
Krijanovskiy (2002) – New Zealandian region of Notogea
Patagonia
Geptner (1936) – Chilean subregion of the Neotropical region
De Lattin (1967) – the whole of South America forms the Neotropical region of
Kingdom Neogea
Lopatin (1980) – Patagonian region of Notogea
Müller (1974a, b) subdivided the Earth into five realms and nine regions:
1. Holarctic
(a) Nearctic
(b) Palearctic
2. Paleotropical
(a) Ethiopian
(b) Madagascan
(c) Oriental
3. Australian
(a) Australian
(b) Oceanic
(c) New Zealand
(d) Hawaiian
4. Neotropical
5. Archinotic (Antarctic, southwestern South America, and southwestern New
Zealand)
Morrone (2015) is recognizing the following three (biogeographical) kingdoms
and nine regions: Holarctic Kingdom (Nearctic and Palearctic regions), Holotropical
Kingdom (Neotropical, Ethiopian, and Oriental regions), and Austral Kingdom
(Cape, Andean, Australian, and Antarctic regions). Additionally, the following five
transition zones are recognized: Mexican (Nearctic-Neotropical transition), Saharo-
Arabian (Palearctic-Ethiopian transition), Chinese (Palearctic-Oriental transition),
Indomalayan (Oriental-Australian transition), and South American (Neotropical-
Andean transition). The present author thinks that the use of older names like
Ethiopian and Oriental is not a good idea. Zoologically it seems better to keep the
Neotropica separate from the Paleotropica. It is better to keep the Indomalayan
region and the name Wallacea as Indomalayan-Australian transition. The South
American transition should be Neotropical-Patagonian.
Biogeographical division of land areas of Udvardy (1975)
Biogeographical realm – the highest taxon, more or less corresponds to the region
of the faunists
Biogeographical province – more or less corresponds to the faunal province of
the faunists (the biotic province of Dice 1943)
The Biogeographical realms of Udvardy are eight:
1. Palearctic
2. Nearctic
3. Afrotropical
4. Indomalayan
5. Oceanian
6. Australian
7. Antarctic
8. Neotropical
The Forgotten Ideas of R. Jeannel
The prominent French coleopterologist Dr R. Jeannel (1879–1965) has been firmly
adept of the Wegenerian ideas, even in the period when the geologists were very
much against them. In his voluminous book “La genèse des faunes terrestres.
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Chapter 13
General Conclusions
Abstract Most of the “classic” zoogeographical regions are with very low-level
endemicity that concerns the Arachnida. Even in Kingdom Notogea, there are no
endemic orders or suborders. The distribution of several orders (Ricinulei,
Amblypygi, Thelyphonida (Uropygi), Schizomida, Solifugae, Scorpiones, Opiliones
of the suborders Cyphophthalmi and Laniatores) is defined by their thermopreferen-
dum (stenothermy), and this distribution, limited by isotherms, is important to char-
acterize the regions and subregions. The north of Holarctic is marked by the presence
of many spiders, mites, and Opiliones – Eupnoi and Dyspnoi – and some pseudo-
scorpions, as far north as 82°33′ (Grant Land spiders).
1. Most of the “classic” zoogeographical regions are with very low-level endemic-
ity that concerns the Arachnida. Even in Kingdom Notogea, there are no
endemic orders or suborders.
2. The distribution of several orders (Ricinulei, Amblypygi, Thelyphonida
(Uropygi), Schizomida, Solifugae, Scorpiones, Opiliones of the suborders
Cyphophthalmi and Laniatores) is defined by their thermopreferendum
(stenothermy), and this distribution, limited by isotherms, is important to
characterize the regions and subregions.
3. The north of Holarctic is marked by the presence of many spiders, mites, and
Opiliones – Eupnoi and Dyspnoi – and some pseudoscorpions, as far north as
82°33′ (Grant Land spiders).
4. The analysis of Beron (2008, 2015, 2016a) has shown that the highest found
Arachnida in the mountains of the Old World are as follows: Araneae (6700 m.),
Opiliones Eupnoi (5600 m.), Dyspnoi (>5000 m.), Laniatores (4600 m),
Solifugae (4570 m.), Scorpiones (?5000 m.), Acariformes (6100 m.),
Parasitiformes Mesostigmata (4550 m.), and Ixodida (5488 m.). The other
orders (Palpigradi, Ricinulei, Amblypygi, Thelyphonida (Uropygi), Schizomida,
Opiliones Cyphophthalmi, Holothyrida, Opilioacarida) do not climb over
3000 m.

https://doi.org/10.1007/978-3-319-74418-6_13
932 13 General Conclusions
5. In the amazing fauna of Madagascar are missing the Solifugae, abundant in

Southern Africa. The scorpions are 100% endemic but only up to genus level.
The Opiliones Eupnoi, Dyspnoi, and most families of Laniatores are also
missing. This is characterizing enough the separate status of Madagascar as a
region. This also is important for judging the limitations of the ability of many
groups to disperse across water barriers by rafting or air currents. From all
Arachnida on Madagascar, only one endemic family is known (the spiders
Halidae). Our analysis shows that there is considerable difference in the
Arachnids of Southern Africa and Madagascar.
6. The Arachnida of the Seychelles, the Mascarene, and the other islands of the
western part of Indian Ocean deserve special attention. In tropical South Africa
are found all orders and almost all suborders of Arachnida. They illustrate first
all the complex history of formation of the fauna of the granitic islands of the
Seychelles.
7. Another island group, characterized by the absence of even more orders of
Arachnida, is New Zealand. Absent from this ancient land are Scorpiones,
Amblypygi, Thelyphonida, Schizomida, Ricinulei, Palpigradi, Opilioacarida,
and Solifugae. Remarkable is the number of spiders (93 endemic genera) and
of Opiliones Cyphophthalmi. Nevertheless, among the Arachnida of New
Zealand there are no such high-level endemics like tuatara or the endemic
families of frogs and bats. The formation of the New Zealand fauna of Arachnida
remains quite a mystery.
8. One of the most intriguing patterns of distribution is the many genera of spiders
and Opiliones across the “Southern end of the world”: New Zealand, Australia,
and Patagonia. This distribution (Gondwanan, transantarctic, or even more
ancient) gives weight to the outlining of greater Notogea, including the
southernmost South America.
9. The cooler part of South America (South Chile and Argentina) is well charac-
terized by the presence of Notogean elements, by many endemic genera of
spiders, Opiliones and solpugids (even endemic solpugid family Mummuciidae),
and by the lack of Amblypygi, Thelyphonida, Schizomida, and Ricinulei.
10. The analysis of the value of lines of Wallace, Lydekker, and Weber that con-
cerns Arachnida has shown that the “lines” seem to “hold little relevance” not
only for Diplopoda (see Shelley and Golovatch 2011) but also for the Arachnida.
This confirms to some extent the conclusion of Lehtinen (1980) that “Various
lines limiting and dividing the Wallacea seem to have no equivalents [to
vertebrates] in the Arachnological zoogeography.” There is no order of
Arachnida living only of one side of the “lines.” Only the spider suborder
Mesothelae seems limited to the western side.
11. In tropical South America are found all orders and almost all suborders of
Arachnida: Palpigradi (no endemics above species), Solifugae (2 families, 1 of
them endemic for tropical South America), Ricinulei (17 spp. of 1 genus, only
end. species), and Scorpiones (3 end. genera;? 4 end. families in caves). Brazil
is one of the richest countries in scorpions – 165 spp. of 26 genera.
13 General Conclusions 933
Pseudoscorpiones (16 families, 18 endemic genera), Opiliones (Cyphophthalmi –

five endemic genera, Eupnoi – one endemic genus, Laniatores – huge variety,
with 15 fam., of them endemic), Amblypygi (many species, one endemic
genus, the only genus in the family Phrynichidae living in the Western
Hemisphere), Thelyphonida (two endemic genera), Schizomida (eight genera),
Araneae (spiders of 77 families are known south of Panama (72% of all spider
families). No family of spiders is endemic for the South American continent.
Opilioacarida (two genera), Holothyrida (two endemic genera), many mites.
Absent are the suborders Paleoamblypygi, Dyspnoi and Mesothelae. As a
whole, the level of endemism is too low to characterize a Kingdom, what
concerns the Arachnida.
12. The fauna of Arachnida in the Caribbeans is very rich (and well explored).
Represented are all orders, very different than the situations with the mammals
and reptiles: Scorpiones (130 spp., including 21 endemic, 17 genera, 4 endemic),
Pseudoscorpiones (in the 7 countries of Central America are recorded
Pseudoscorpions of 33 genera and 13 families; in the Caribbeans there are 39
genera of 15 families), Amblypygi (29 spp., 4 genera, 2 families; endemism
only on species level), Uropygi (1 endemic genus), Schizomida (very rich, or
well explored; 5 endemic genera), Opiliones Laniatores (117 spp. of 65 genera,
including 13 endemic, 11 families), Araneae (only endemic genera), Holothyrida
(1 endemic genus), Opilioacarida (2 genera with endemic species), and many
other mites. Absent are only the suborders Cyphophthalmi, Dyspnoi,
Palaeoamblypygi, and Mesothelae. These facts justify the outlining of the
islands as a separate Antillean (Caribbean) subregion. However, the comparison
between the Arachnida of Mesoamerica (between Mexico and Colombia) and
the Caribbean has shown that the difference is only on level of endemic genera.
13. Comparing the pseudoscorpions of Palearctic and Nearctic (20 families of
pseudoscorpions in Kingdom Holarctic, 17 in each region), 14 are in common
between Nearctic and Palearctic parts, and six families are not in common.
From the Holarctic are missing families Feaellidae, Gymnobisiidae, Hyidae,
Vachoniidae, Parahyidae, and Pseudochiridiidae. Sternophoridae is found in
Florida. In the Nearctic part are recorded 87 genera of pseudoscorpions and 79
genera in the Palearctic part, and only 18 genera are in common between the
two.
14. The forming of the arachnofauna (and other fauna) of New Caledonia is consid-
ered to be not older than 37 Ma and is derived from a few ancestors brought by
the three Ws (waves, wind, and wings). However, the presence of the endemic
family of cave Opiliones Cyphophthalmi- the Troglosironidae and of two spp.
of Holothyridae (both groups without relations to the respective Australian
families) speaks in favor of the idea that the island has not entirely sunk before
the Oligocene.
15. On the Australian continent live 13 orders (Palpigradi, Amblypygi, Schizomida,
Opiliones – Cyphophthalmi, Eupnoi, Laniatores, Pseudoscorpiones, Scorpiones,
Araneae, Opilioacarida, Holothyrida, Ixodida, Mesostigmata, Sarcoptiformes,
Trombidiformes), and missing are Solifugae, Thelyphonida, Ricinulei, and the

suborders Palaeoamblypygi, Mesothelae, and Dyspnoi.
Some peculiarities: Araneae, 73 families; Opiliones Cyphophthalmi (two
end. genera); Laniatores, Pseudoscorpiones (150 spp.); Oribatida, 340 named
species (256 endemic) and 237 unnamed species, or a total of 577 species.
There are no superfamilies of Oribatida indigenous to Australia.
Tasmania has also particularities in its arachnofauna. Many groups like
Schizomida, numerous in Australia, are absent (because of the climate?). There
are many endemic Laniatores.
As a whole, the level of endemicity of Arachnida in Australia is low, nothing
to compare with the vertebrates, especially mammals.
16. As with some other groups (Isopoda, Scorpiones), the presence of American
elements in the Mediterranean subregion is surprising and demands explanation.
Such surprise was to find representative of Bochicidae (known from the Antilles,
Mexico, Texas, and Guatemala) in a cave in Portugal. As many other cave
animals, the genus Titanobochica Reboleira is considered relict. Other such
relicts in the Mediterranean (all from caves) are Troglobisium Beier (Spain),
Troglochthonius Beier (Dalmatia), and Neobalkanella Ćurčić (Serbia). The
very presence of family Syarinidae in Europe is also considered relictual. The
South American family Gymnobisiidae has one genus (Gymnobisium Beier) in
South Africa.
17. Among the peculiar distributions of Amblypygi is the sensational presence of
Phrynus (purely American genus) in Indonesian caves (the only Phrynidae in
the Old World); from the family Phrynichidae, typical for the Old World, one
genus (Trichodamon Mello-Leitão) is living in South America. Especially
important is the relict Paracharon Hansen in Guinea-Bissau – the only
representative of the family Paracharontidae and of the entire archaic suborder
Palaeoamblypygi.
18. On Hawaiian Islands, far from any mainland and born without biota, are living
members of ten orders of Arachnida (Palpigradi, Scorpiones, Schizomida,
Araneae, Opiliones, Pseudoscorpiones, and various acari). Remarkable are the
blind arachnids of endemic genera of Araneae (132+ native spp. of 36 native
genera and 16 native families). Particularly interesting are the blind spiders,
found by Howarth in the relatively recent lava caves, Adelocosa anops Gertsch.
At least 128 species of spiders are endemic; remarkable are Pseudoscorpiones
(Vulcanochthonius Muchmore), described from lava tubes. How all these
arachnids reached the azoic islands and how long time was needed for their
amazing evolution remain a topic to discuss.
19. Another amazing island group is the Galapagos archipelago. Not so far from
the mainland South America (966 km), these islands have been volcanoes, and
the arachnofauna clearly came from the continent. Some of them formed local
species (the Amblypygi Charinus insularis Banks), the scorpions (Centruroides
exsul Meise and Hadruroides galapagoensis Maury), or genera (the solpugid
Neocleobis solitarius Roewer), but still the arachnofauna is no match to the
13 General Conclusions 935
remaining animals (reptiles, birds). On Galapagos are known 25 spp. of 9 gen-

era and 10 families of Pseudoscorpiones.
20. The cave arachnofauna is important for the assessment of many groups of
Arachnida. Examples: the European Laniatores, with many cave genera, the
numerous Schizomida known only from caves, the problems with the speciation
of the fauna of Hawaiian lava tubes and with the different models and rate of
troglobiomorphose. Important relicts are the only Syarinid on the Balkans
Hadoblothrus aegaeus Beron from Santorini, the unique families Troglosironidae
in New Caledonia, Typhlochactidae in Mexico, Akravidae in Israel,
Troglotayosicidae in Ecuador, Pseudochactidae in Vietnam and Laos,
Trogloraptoridae in North America, the relict members of Bochicidae in
Portugal, the Phrynus in Indonesian Flores, the only member of fam.
Pyramidopidae in the New World (Jarmilana Cruz-López et al.), the
Opilioacarida in Cuba and Thailand, and many other cave Arachnida.
21. The comparison between the arachnofauna of Indo-Malayan region and Eastern
Palearctic (belonging to different kingdoms!) has shown that the differences do
not support such high ranks (Beron 2016b). Here important is also the latitudinal
gradient in diversity.
22. Strange pattern is evident in the distribution of the ticks Holothyridae from the
Seychelles to New Zealand.
23. The comparison between the arachnofauna of Afrotropical Region and

Palearctic (belonging to different kingdoms!) has shown that there is
considerable difference, reaching the level or order (Ricinulei and Schizomida
in Africa) and suborders (Palaeoamblypygi in trop. Africa, Holothyrida on the
islands East of Africa, none in the Palearctic Mesothelae in the Afrotropical
Region). The families rarely are endemic (Halidae in South Africa), but the
suborders are most often different (Laniatores). Here important is also the
latitudinal gradient in diversity, due mostly to the thermopreferendum. The
paleogeographical history of both regions is totally different (glaciations,
Gondwana).
24. The Cape arachnofauna (south of Zambezi, Kunene) depending on the

Mediterranean-type climate is restricted to the extreme southwestern Cape. The
Cape flora is so distinct that it is recognized as floral kingdom of its own
(Capensis). In most groups of animals, the rank in Southern Africa is much
lower (subregion). The area contains 13 orders of Arachnida (missing are
Thelyphonida, Ricinulei, Holothyrida, and the suborders Palaeoamblypygi,
Mesothelae, Dyspnoi). There are one endemic family of spiders (Chummidae),
one endemic subfamily of Solifugae (Melanoblossiinae), one endemic subfam-
ily of Schizomida (Megaschizominae), one of Opiliones Eupnoi (Neopilioninae),
one of Opiliones Laniatores (Adaeinae), and one of Scorpiones (Lisposominae).
Some groups are numerous and with many endemic genera (163 spp. of
Solifugae, 64 families of spiders, 122 spp. of Pseudoscorpiones, 164 spp. of
Opiliones Laniatores, many endemic genera and species of scorpions and
mites). Almost endemic is the tick family Nuttalliellidae (Ixodida). With lack of
endemic orders or suborders, the arachnological results confirm the status of
the Cape area as a subregion. In South Africa there are elements connecting it
with Patagonia or Australia-New Zealand (Gondwanan remnants).
The purpose of this study was most of all to check how the arachnogeographi-
cal subdivision of the world would fit with the classical or modern regionaliza-
tion after other groups, mainly terrestrial vertebrates. The main conclusion is
that the level of endemicity in Arachnida is much lower than in the vertebrates.
It means that the differences between areas like Indo-Malayan and the Australian
regions are much less. They are defined not so much by the history of formation
of faunas but by the climatic (mostly thermic) conditions of the areas. The
major puzzles like the presence-absence in the arachnofaunas of islands
(Madagascar, Seychelles, New Zealand, New Caledonia) remain. Also remains
is the most intriguing history of the fauna of the Southern End of the World.
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Index
A Acaromorpha, 36
Abalakeus, 190 accentuatus, Hypoxestus, 862, 863, 885
Abaliella, 139, 416, 428, 434, 733, 749, Achaearanea, 840, 866
750, 781 Achaearyopa, 434
Abanatus, 468 Achipteria, 49, 881
abeli, Rowlandius, 665 Achipteriidae, 868, 877
Aberdaria, 883 Acihasta, 770, 780
Aberdereca, 465, 468 Acorigone, 366
Aboriscus, 399 Acrita, 489
Abracadabrella, 763 Acrobleps, 766
Abrolophus, 191 Acrobuninae, 156, 399, 413, 426, 737
Absonus, 494 Acrobunus, 426
abstrusum, Metaphalangium, 352 Acrographinotus, 492
Aburiplus, 468 Acromares, 635, 643
Aburista, 92, 468 Acromitostoma, 327, 347
Aburistella, 468 Acropsopilio, 80, 153, 488, 505, 506, 510,
abyssinicus, Caloctenus, 80 627, 635, 637, 770, 779, 780
Acaca, 468 Acropsopilionidae, 80, 153, 488, 505, 510,
Acanthepedanus, 426 635, 677, 757, 765, 770, 779
Acanthicochernes, 88, 748 Actinacantha, 430
Acanthocranaus, 489 Actinopodidae, 172, 646, 678, 738, 766, 901
Acanthocreagris, 336, 351, 355 Actinostygnoides, 494
Acanthogylippus, 84 acuaticus, Trogulus, 356
Acantholibitia, 488 Acuclavella, 92, 510
Acanthomegabunus, 372, 411 acuclavella, Ceratolasma, 92, 510
Acanthominua, 494 Aculepeira, 334, 866
Acanthopachylopsis, 492 Acuminochernes, 88, 509
Acanthophrynus, 98, 142, 513, 628, 637 Acumontia, 706
Acanthophrysella, 468 Acutipetala, 419
Acari, 36, 41, 45, 48–49, 66, 67, 69–71, 76, Acutisoma, 491
80, 97, 99, 184, 185, 333, 342, 343, Adaeinae, 695, 935
345, 351, 356, 361, 381, 405, 507, 515, Adaeulum, 695
522–524, 526, 711, 714, 726, 743, 744, Adamauna, 468
747, 754, 755, 764, 781, 844, 846, 874, Adamystidae, 193, 868
883, 884 Adamystis, 193, 881
Acaridida, 97, 193, 524, 744, 859, 868, 883 Adamystoidea, 193

https://doi.org/10.1007/978-3-319-74418-6
940 Index
Adelocosa, 521, 522, 840, 934 albimanus, Biantes, 716

Adelphochernes, 88, 432 Albionella, 649
Adenacarus, 93, 185–187, 381 Aldabrinus, 719
Adhynastes, 491 alejandroi, Antillostenochrus, 665
adictyna, 773, 780 alejandroi, Stenochrus, 666, 671
Adisomus, 85, 136, 498, 499 algerica, Minniza, 358
Admetus, 142 Algidia, 770, 779
adusta, Patelliella, 777 alini, Hahnia, 885
aegaeus, Hadoblothrus, 96, 935 Allereca, 468
aegatensis, Chthonius, 355 Allobunus, 765
aelleni, Acanthocreagris, 355 Allocaeculus, 192
Aenigmarachne, 647 Allochernes, 336, 351, 371, 871, 882
aenigmaticus, Lusoblothrus, 346 Allococalodes, 752, 763
aenigmus, (?) Opilioacarus, 49, 51, 67, 70 Allocranaus, 490
Aerogonyleptes, 491 Allodecta, 649, 668
Aesotrinoma, 491 Allogonyleptes, 492
aestivus, Tiso, 867 Alloixodes, 668
aetnensis, Roncus, 355 Allokoenenia, 83, 135, 455, 475, 681, 893
affinis, Feaella, 715 Allomicythus, 420
Afribactrus, 366, 699 Allonuncia, 765
africanus, Calathocratus, 356, 471 Allothyridae, 194, 198, 742–744,
Africepheia, 708 777, 779
Afrilobus, 697 Allothyrus, 194, 198, 764, 773, 779
Afroassamia, 92, 468 alpicola, Chalcoscirtus, 866, 867
Afroceto, 697 alpigena, Arctosa, 867
Afrogarypus, 715, 719 alpina, Micaria, 866, 867
Afroroncus, 86, 90, 463, 476 altenai, Bunochelis, 366
Afrowithius, 87, 89, 694 alticola, Anandia, 665
Afrozomus, 85, 136, 471, 472, 476 alticola, Antillostenochrus, 665
Agastoschizomus, 136, 628, 638, 841, alticola, Mughiphantes, 885
843, 894 alticola, Pseudocleobis, 676
Agelenella, 393 altimanus, Chelanops ? [Dinocheirus], 658
Agelenidae, 176, 328, 339, 341, 362, 402, Altobunus, 411, 425, 433
419, 422, 452, 647, 667, 678, 740, 772, amabilis, Reventula, 662
841, 844, 866, 874, 900 Amamanganops, 434
Aglaochitra, 509 Amatola, 695
Agoristenidae, 40, 156, 488, 501, 660, 662, Amaurobiidae, 9, 176, 328, 339, 341, 362,
674, 838 396, 404, 647, 648, 667, 678, 697, 740,
Agoristeninae, 662 766, 772, 841, 866, 867, 874
Aguaytiella, 489 Amaurochelifer, 87
Ahua, 772, 780 Amauromastigon, 85, 140, 497
Aisthetowithius, 87, 90, 463, 476 Amazonacarus, 93, 186, 188
Akamasia, 390 Ambatoiella, 489
Akdalima, 636, 662 Ambicodamus, 761, 767
Akrav, 79, 383, 834, 897 Amblyolpium, 336, 775, 836
Akravidae, 37, 65, 77, 144, 145, 380, 381, Amblyomma, 68, 71, 504, 710
834, 897, 935 Amblypygi, 36, 46, 50, 74, 76, 85, 95, 98,
Alabamocreagris, 87, 509 140–143, 333, 343, 354, 368, 371, 376,
Alacran, 627, 632, 834 381, 386, 387, 391, 392, 394, 397, 399,
Alaranea, 708 404, 414, 421, 427, 431, 433, 442, 443,
Alausius, 489 452, 454, 469, 476, 478, 495, 503, 507,
alayoni, Rowlandius, 671 513, 515, 516, 626, 628, 637, 640, 645,
Alayotityus, 654, 833 665, 669, 673, 676, 682, 690, 692, 695,
Albertops, 426 703, 707–710, 713, 714, 716, 721–727,
Index 941
743, 747, 749, 754, 757, 764, 769, 775, Anderemaeidae, 869, 876
778, 779, 841–842, 844, 894–895, andhraensis, Phrynichus, 399
931–934 Andocaeculus, 193
Ambohima, 708 Andoharano, 708
Americocreagris, 87, 509 Andrevella, 191
Americovibone, 80, 677 Andrias, 78
Ameridae, 876 Androctonus, 376, 377, 383, 448, 688
Ameronothridae, 97, 527, 877 Anduzeia, 489
Amhara, 468 Anelasmocephalus, 351, 352, 356, 361
Amilenus, 50, 338, 361 Anepsiozomus, 85, 136, 716, 721
Ammogarypus, 86, 89, 693 Angistrisoma, 489
Ammonius, 93 Angistrius, 489
Ammotrecha, 508, 631, 640, 669 Angka, 410, 419
Ammotrechella, 367, 508, 631, 640, 641, 653, Angolyppa, 468
654, 669, 673 Angopygoplus, 468
Ammotrechesta, 84, 631, 641, 669 anguineus, Araeoncus, 867
Ammotrechidae, 367, 481, 503, 508, 627, Angustaea, 420
631, 640, 653, 669, 673, 676, 896 angusta, Eukoenenia, 398, 400, 405
Ammotrechinus, 84, 653, 654, 669 Anhunga, 780
Ammotrechona, 84, 653, 669 Anisaspis, 648, 667
Ammotrechula, 508, 631, 669 Anisoditha, 86
Ammoxenidae, 176, 698, 731, 737, 741, 762 ankarafantsika, Grosphus, 704
Ammoxenus, 698 ankarana, Grosphus, 704
Ampheres, 490 Ankaratra, 91, 464, 701, 706
Amphibia, 78, 919 Ankaratrix, 706
Amphinectidae, 678, 679, 731, 737, 740, 759, Ankylonuncia, 765
761, 766, 772 annandalei, Meghalaya, 398, 410
amplum, Olpiolum, 657 annulatipes, Damon, 696
Ampycella, 490 annulatus, Boophilus, 393, 704
Ampycus, 490 annulatus, Grosphus, 704
amurensis, Typopeltis, 371 Anoetidae, 883
Anaceros, 706 Anomalobuthus, 382
Anacrobunus, 426 Anomalosa, 760
Aname, 760 Anophthalmoonops, 473
Anamota, 664 Anoplocheylus, 192
Ananeon, 763 Anoplogylippus, 880
ana, Neothyrus, 501 anops, Adelocosa, 934
Ananteris, 84, 482, 654, 655 Anopsolobus, 772, 780
Ananteroides, 450, 460 Antetriceras, 492
Anapidae, 177, 328, 341, 362, 396, 422, 474, Anthracomartida, 45
647, 678, 696, 739, 751, 760, 766, 767, Anthrenochernes, 360
771, 772, 776, 780, 840 antiguanus, Stygnoplus, 662
Anapogonia, 430 antillarum, Eucynortoides, 661
Anarchaea, 760 Antillattus, 649, 668
Anarthrotarsus, 327, 338, 339 Antillena, 647, 666
anasilviae, Rowlandius, 666, 671 Antillobisium, 86, 89, 657, 835, 843
Anatea, 776 Antillognatha, 647, 667
Anatemnus, 432, 720 Antilloides, 648, 667
anatis, Ixodes, 767, 773 Antillostenochrus, 46, 136, 646, 665, 666
Ancalochernes, 88, 632 Antillotrecha, 84, 653, 654, 669
Anchognatha, 761 Antongila, 706
Ancinosphenus, 647, 667 Antrodiaetidae, 172, 900
Ancistrochelifer, 88 Antrohyphantes, 340, 341, 866
andamanensis, Chaerilus, 404 Anystidae, 189, 192, 369, 868, 875
942 Index
Anystis, 192, 883 Araniella, 866

Anystoidea, 192 Arbanitinae, 173, 731, 737, 738
Aops, 755, 778, 845 arboreum, Planctolpium, 657
Aoraki, 91, 768, 770, 779 Arcanobisium, 86, 326, 346, 835
Aotearoa, 678, 772, 780 Archaeidae, 51, 177, 760, 873
Apatellata, 36 Archaeobatrachia, 78
Apatochernes, 770, 779 Archaeoroncus, 326, 336, 337
apejii, Ammotrechella, 653 Arctenus, 474
Aperittochelifer, 88, 90, 694 Arganotus, 636, 644, 662
Aphelacaridae, 876 argasiformis, Speleosiro, 842
apicoporus, Mahezomus, 716 argentatus, Dasylobus, 356
Apistobuthus, 383 argenteolunulatum, Histricostoma, 356
Apobrata, 434 argus, Hastocularis, 66, 70
Apocheiridium, 336, 368, 432, 882 Argyrodella, 717
Apohya, 86, 89, 632 Argyrodes, 521
Apolania, 718 Argyronetidae, 178, 328, 339
Apolpium, 388, 503 Ariaeus, 491
Apomerantzia, 189 Aridella, 717
aponommoides, Haemaphysalis, 878, 885 Arkyidae, 177, 760
Aporochelifer, 88, 90, 425 Arkys, 760
Aporoptychinae, 173, 731, 737, 738 armasi, Cubazomus, 665, 671
Apozomus, 136, 417, 418, 422, 515, 716, 734, armasi, Mexobisium, 657
759, 778 armatus, Leiothyrus, 752
Apygoplus, 749 armatus, Mughiphantes, 867
Apyretina, 709 Arminda, 493
Arabelia, 341 Arrallaba, 757, 779
Arabellata, 751 Arruda, 493
Arachnida, 1–3, 7, 11, 13–15, 33, 35–41, Artacarus, 85, 136, 471, 472, 894
45–51, 59–99, 135–199, 319, 321, 333, Arthrodes, 490
354, 358, 363–367, 394, 396, 398, 424, Aruana, 752, 763
442, 454, 503, 507, 514–519, 626, 652, arunachalicus, Schizomus, 399
677, 681, 688, 728, 744, 747, 775, 778, Arythaena, 49, 51, 67, 70
842, 854, 880–882, 898, 932, 933 Asadipus, 762
Arachnidea, 36 Asarcus, 490
Aracus, 420 Ascaphidae, 78
Araegeus, 700 Ascaphus, 78
Araeoncus, 866, 867, 883, 885 ascensionis, Withius, 368
Araguita, 494 Aschema, 709
Arandisa, 699 Ascuta, 772, 780
Araneae, 2, 25, 36, 41, 45, 75, 80, 82, 93, 96, asiatica, Prokoenenia, 405, 894
98–99, 344, 348, 352, 356, 362, 363, asiaticus, Iurus, 383
367–369, 373, 381, 392, 397, 402, 406, Asmea, 751
442, 452, 473, 478, 500–501, 503–505, Asproleria, 749, 781
515, 520, 524, 696–700, 726, 737, 747, Aspurochelifer, 88, 509
750, 755, 759–764, 771, 776, 777, 779, assamensis, Uroproctus, 399
846, 881–884, 900–902 Assamia, 399, 402
Araneidae, 41, 177, 339, 373, 396, 402, 404, Assamiidae, 40, 156, 396, 399, 401, 402, 412,
430, 443, 452, 504, 525, 647, 678, 707, 414, 433, 465, 695, 706, 712, 736, 748,
708, 759, 873, 900 757, 779, 781, 782, 862, 871
Araneina, 41 Assaphala, 396
araneoides, Galeodes, 98, 335, 371, 376 Astavakra, 434
Araneomorphae, 40, 41, 96, 176, 513, 628, Astegistidae, 877
638, 738, 743, 864, 902 Asterochernes, 505
Arangina, 773 Asteron, 762
Index 943
Asthenargus, 872, 883, Austromontia, 80, 695

Astia, 763 Austronuntia, 695
Astilodes, 763 Austrophaea, 700
Astrobunus, 327, 338, 344, 361, 862 Austropsopilio, 80, 91, 757, 765
Asygyna, 708 Austropurcellia, 91, 757, 778
Atelidea, 403 Austroteneriffia, 192
Atelolathys, 402 Austrotengella, 762
Atemnidae, 17, 38, 149, 326, 336, 343, 395, Autenriethia, 192
401, 406, 677, 715, 720, 747, 748, 775, Avelis, 700
777, 782, 871 aviculus, Bamazomus, 716
Atemnus, 336, 371, 432, 836 Avilaia, 494
Atherochernes, 88, 90, 487 Ayacucho, 491
Athleticatemnus, 88, 90 Azanialobus, 697
athos, Ausobskya, 96 Azulamus, 493
Atmetochilus, 419 Azulitaia, 494
Atomus, 883
Atopomelidae, 193
atra, Erigone, 867, 885 B
atramentaria, Gnaphosa, 348 Babycurus, 385, 460–462
Atrax, 760 baccarum, Anystis, 883
Atrophothele, 93, 393 Bacillemma, 419
Attaleachernes, 487 bacillifer, Phrynichus, 695
Attenuizomus, 85, 136, 734, 758, 759, 778 Bacillochilus, 473
Atylodes, 78 Baculigerus, 490
Atypidae, 172, 327, 339, 422, 737, 738, 900 Badessa, 749
Aucayacuella, 489 Badia, 474
Augusta, 708 badia, Gnaphosa, 866, 867
Augustsonella, 190 baebleri, Mughiphantes, 867
Auranus, 494 bagnalii, Zomus, 324, 417, 428, 716, 721
Aurecocrypta, 93 Baiami, 761, 841
aureum, Olpiolum, 657 bajacalifornicus chamelaensis, Neocarus, 638
Ausobskya, 92, 96, 339, 837 Bakala, 761
austera, Ceratinopsis, 867 Balabanus, 433
Austmusia, 761 Balaustium, 49, 67, 70, 191, 868
Austrachelas, 696 balearica, Acanthocreagris, 351
Australaena, 519 balearicum, Ptychosoma, 352
australasiae, Liocheles, 404, 436, 440, 442, balearicus, Anelasmocephalus, 351, 352
515, 688, 755, 775 balearicus, Chthonius (E.), 351
australianus, Charinus, 749, 775 balearicus, Dactylochelifer, 351
Australinocreagris, 509 balearicus, Euscorpius, 351
Australochelifer, 91, 756, 778 balearicus, Pselaphochernes, 351
Australoechemus, 367 balearicus, Trogulus, 351, 352
Australoonops, 698 balfouri, Gluviopsis, 392
Australothele, 760 Balkanoroncus, 336, 835
Australothyrus, 94, 194, 196, 198, 779 Ballarra, 757, 779
Austrammo, 762 Ballarrinae, 677
Austrarchaea, 760 Ballus, 836
austriaca austriaca, Eukoenenia, 334 Baloorthochirus, 688
Austrochilidae, 177, 678, 731, 737, 738, 766, Balsateres, 627, 632, 633
838, 901 baltistanus, “Chelifer”, 880
Austrochilinae, 177, 766, 901 baltistanus var. shaksgamica, Yllenus, 885
Austrochiloidea, 41 baltoroi, Acantholycosa, 396, 885
Austrodomus, 698 baltoroi, Gongylidium, 885
Austrohorus, 756, 778 baltoroi, Zelotes, 885
944 Index
Balzabamba, 489 Belicenozomus, 136, 670

balzanii, Ideobisium, 656 Belisariinae, 79
Bamazomus, 50, 136, 417, 418, 472, 513, 520, Belisarius, 79, 84, 321, 325, 345, 346, 483,
628, 706, 717, 721, 734, 758, 759, 778 834, 897
Bambereca, 465, 468 bellesi, Chthonius (E.), 351
Banaidja, 518 bellesi, Paraleptoneta, 843
Bancoella, 468 Beloniscellus, 426
Banconyx, 468 Beloniscops, 426
banksi, Kimula, 663 Belonisculus, 426
Banksula, 92, 512 Beloniscus, 426
Bannana, 419 bengalensis, Charinus, 399, 414
baracoae, Rowlandius, 666, 671 Bengalla, 760
Barahna, 761 benoiti, Anapistula, 182, 740
Baramella, 427 benoiti, Beierolpium, 715
Baramia, 427 benoiti, Chaussieria, 369
Barbaraella, 756 Benoitinus, 716
barbatus, Carabothyrus, 668, 674 Benoitodes, 369
Bardala, 717 Berlaia, 492
Barellea, 192 Berlandia, 840
Baria, 489 berlesei, Eukoenenia, 354, 653, 890, 891
Barinas, 488 berlesei, Ixodes, 396, 870, 878, 886
Barlovento, 488 berlesei, Thonius, 752
Barraina, 763 berlesei virginea, Eukoenenia, 653
Barrona, 644 berndti, Koeneniodes, 424
Barropelma, 647 beroni, Euscorpius, 335
Barrowammo, 762 besucheti, Dactylochelifer, 352
Barrus, 84, 448, 449 betschi, Pseudouroplectus, 705
Barrussus, 324, 334 bettoni, Rhampsinitus, 885
Barusia, 341, 839 Bezavonia, 706
Barychelidae, 172, 393, 402, 646, 647, 667, Biacumontia, 695
707, 737, 751, 776, 838 Biantella, 466
Barychelinae, 172, 738, 776 Biantessus, 695
Barycheloides, 93, 776 Biantidae, 40, 158, 392, 396, 412, 426,
Barychelus, 93, 776 433, 465, 660, 663, 664, 674, 684,
Basasteron, 777 695, 706, 712, 715, 716, 720, 723,
Baso, 427 736, 862, 864, 871
Basoides, 427 Biantomma, 466
Bassaniodes, 393 Biantoncopus, 92, 414, 433
Batesiella, 393 Bibundina, 468
Battalus, 762 Biconisoma, 492
Baustomus, 489 biconouros, Rowlandius, 666, 671
Baviola, 718 Bidoma, 663, 664
Bdellidae, 189, 526, 868, 875, 883 bieko, Agyneta, 885
Bdellophaga, 84, 692 bielawskii, Manahunca, 663
Bealeyia, 772, 780 Bigenditia, 762
Bebedoura, 493 bilineatus, Metacynortoides, 661
beccarii, Charon, 428 bimaculatus, Ibalonius, 716
Beckeresia, 492 Binamballeus, 490
beieri, Spelyngochthonius, 355 Binderia, 468
Beierobisium, 86, 677, 679 biocellatus, Oculozomus, 417, 428
Beierochelifer, 336 birmanica, Pardosa, 885
Beierolpium, 715, 720 Birrana, 761
Belemnodes, 493 birsteini, Taracus, 372
Belemulus, 493 birulai, Euscorpius, 335
Index 945
Birulatus, 383, 384 brachydactylus, Sarax, 414, 433

biseriatus, Acanthicochernes, 748 Brachylibitia, 489
Bisetocreagris, 371, 432, 871, 880, 885 brachyura, Hystrix, 199
Bishopella, 92, 512 Bradystichus, 776
bisignata, Cynortellana, 661 Branchia, 508, 631
Bissulla, 493 Branchioscorpionina, 37, 50, 66, 69
bistriatus, Grosphus, 704 Brandbergia, 693
Biton, 98, 324, 335, 354, 381, 449, 455–457 Brasilacarus, 93, 186, 188
Bitonota, 455, 458 Brasilogovea, 487
Bitonupa, 457, 458 braueri, Centrobunus, 716
Bitulochernes, 843 braueri, Lychas, 714
blanci, Neogrosphus, 704 Brazilatemnus, 88, 90, 487
blanda, Valifema, 664 breoni, Ibalonius, 713
Blanioonops, 473 Bresslauius, 493
Blantyrea, 468 Breviacantha, 757, 779
Blossia, 381, 449, 455–457 brevidigitatus, Oratemnus, 715
Blossiana, 455, 458 brevipatellatus, Antillostenochrus, 666
Bochica, 86 brevipatellatus, Rowlandius, 671
Bochicidae, 17, 38, 80, 96, 326, 345, 346, brevipes, Dicranopalpus, 355
483, 641, 657, 659, 671, 834, 835, 843, brevis, Chaussieria, 369, 414
844, 897, 934, 935 Briggsidae, 159
Bodunius, 488 brignolii, Apozomus, 417, 515
Bogdana, 490 Brignolizomus, 85, 136, 734, 758, 759, 778
Bokwina, 643 Bristoweia, 492
Bolama, 468 brolemanni, Eukoenenia, 345
Bolathana, 762 brosseti, Phalangiacarus, 475
Boliscodes, 420 Broteochactas, 484, 655, 656
Bonapruncinia, 369 brunneri, Mughiphantes, 867
bonariensis, Ammotrechella, 653 brunneus, Paranonychus, 511
Bonea, 433 Bryantina, 648, 666, 667
Boneta, 635, 643 Bryonuncia, 765
bonneti, Xysticus, 866, 867 Bucayana, 489
boreus, Paruroctonus, 98, 508, 627 Bucinozomus, 136, 324
Borneanapis, 429 Bueana, 468
Borneomma, 429 Buemarinoa, 91, 356, 838, 899
Borneoridion, 430 Buemba, 468
Borrala, 761, 772 Bugabitia, 644
Bothriuridae, 145, 395, 398, 407, 482, bulgarica, Dysdera, 334
485, 676, 681, 683, 693, 722, 755, Bullaepus, 492
764, 887 Bulolia, 752
Bothriuroidea, 37, 145 Bundukia, 468
Bothriurus, 485, 676, 887 Buniabia, 468
botosaneanui, Metakimula, 663 Bunicranaus, 489
Botswanoncus, 694, 842, 897 Bunochelis, 366
bouilloni, Eukoenenia, 345 Bunofagea, 706
boulengeri, Amblyomma, 504 Bunoplus, 492
Bourgnyia, 490 Bunostigma, 493
Bovidromus, 193 Buparus, 399
brachialis, Dactylochelifer, 695, 696, 880 buresi, Paralola, 83, 339, 388
brachialis, Damon, 695, 696 burmanicum, Palaeosiro, 48, 66
Brachionopus, 696 Burmezomus, 136, 417, 418
Brachistosternus, 485, 676, 887 Bursaustium, 191
Brachychthoniidae, 869, 876 Bursellia, 883
Brachychthonius, 884 Buruquelia, 494
946 Index
Buthacus, 383–385, 407, 448, 450, Calamoneta, 430

460–462, 688 Calathocratus, 327, 339, 356, 377, 378
Butheoloides, 450, 460–462 Calathotarsinae, 175, 901
Butheolus, 383, 385, 462 calcaratus, Piratrinus, 662
Buthidae, 95, 98, 145, 325, 335, 346, 367, Calcarsynotaxus, 760
371, 377, 380, 382–385, 395, 398, 401, Calchas, 385
405, 407, 408, 424, 432, 436, 442, 448, Calcitro, 46, 513
459–463, 482, 484, 485, 508, 631, 633, Calcitronidae, 24, 46
642, 652, 654, 656, 672, 676, 681, 683, Calcoschizomus, 46, 513
688, 691, 693, 703–705, 712, 719, 722, Calculus, 698
734, 742, 747, 755, 775, 781, 833, 843, Caldanatus, 492
860, 870, 887 Caldasius, 490
Buthiscus, 688 Caledanapis, 776
Buthoidea, 37, 145 Caledothele, 776
Buthoscorpio, 401, 407, 688 Caleremaeidae, 876
Buthotus, 383, 693 Calicina, 92, 512
Buthus, 325, 345, 346, 358, 383–385, 448, Calicynorta, 512
450, 460–462 Calidiochernes, 89, 90, 487
butleri, Caribchthonius, 655 californica, Prokoenenia, 508, 626
buxtoni, Apozomus, 417 Calima, 136, 498–500
Bwitonatus, 465, 468 Callcosma, 489
Bymainiella, 760 Callevophthalmus, 761
Bythocrotus, 649, 668 Callidosoma, 190
Callihamina, 757, 779
Callihamus, 757, 779
C Callitrichia, 883
cabreriensis, Chthonius, 351 Calliuncus, 765
Cacoxylus, 89 Calocaeculus, 193
Caddella, 91, 695 Calocheiridius, 336
Caddidae, 40, 48, 50, 79, 153, 411, 436, 510, Calocheirus, 365
677, 678, 684, 695, 723, 736, 780, 899 Caloctenus, 80
Caddo, 48, 51, 80 Calomyspoena, 504
Caddoidea, 38–40, 80 Calorema, 189
caeca, Catonetria, 369 Calymmachernes, 89
caeca, Microcreagrella, 365 Calyptostoma, 67, 70
Caecatemnus, 88, 90, 487 Calyptostomatidae, 191, 875
Caecogarypinus, 406 Calyptostomatoidea, 5, 189, 191
Caecoonops, 473 Camarana, 493
Caecothrombium, 420, 842 cambridgei, “Schizomus”, 417
caecula, Anapistula, 182, 740 Camelianus, 493
Caeculidae, 189, 192 Camerobunus, 465
Caeculisoma, 190 Camerotrombidium, 883
Caeculoidea, 192 Camisiidae, 869, 876
Caeculus, 193 campaneti, Chthonius, 351
caecus, Paracharon, 95, 469, 682 Camposicola, 492
Caelopygus, 490 Camposicoloides, 492
Caenoncopus, 92, 414, 427 Campostichomma, 403
Caenonetria, 430 Campostichommides, 751
Caesetius, 697 Campostrecha, 84, 481
Cajacayba, 491 Canala, 776
Cajamarca, 491 canariana, Bunochelis, 366
Calacadia, 679 Canarichelifer, 88, 365
calakmulensis, Neocarus, 638 Canariellanum, 366
Calamistrula, 707 Canariognapha, 366
Index 947
Canarionesticus, 366, 840 Cavasteron, 762

canestrinii, Dolichowithius, 658 cavernicola, Burmezomus, 417
canestrinii, Opilio, 358 cavernicola, Chthoniella, 842
Canga, 487 cavernicola, Sarax, 414, 427
Cangonia, 467 cavernicola, Speleomontia, 842
canhi, Vietbocap, 406, 834 cavicola, Apocheiridium, 368
Cantuarea, 780 Cayabeus, 489
Cantuaria, 771 Cazierius, 84, 655
Capichabesia, 492 Cea, 494
Caponiidae, 177, 390, 647, 678, Cecidopus, 191
685, 697, 724 Ceclusa, 420
capromydis, Ixodes, 668 Celimba, 468
Capromyidae, 668 Cenefia, 770, 779
Carabella, 649 Cenipa, 489
Carabodidae, 877 Centetostoma, 327, 347
Caracarana, 489 Centrobunus, 716
caraibicus, Cynortoides, 661 Centrocalia, 179, 739, 776
Carastrum, 191 Centrochelifer, 372
Carathea, 678, 766, 767 Centroina, 762
Carayaca, 495 Centroleptes, 491
Carbinea, 761 Centromerus, 839, 866
Cardiolpium, 336 Centrothele, 762
Cargaruaya, 491 Centrothelinae, 179, 739
Caribattus, 649, 668 Centruroides, 484, 485, 633, 642, 652,
Caribbiantes, 663, 664 654–656, 672, 704, 843
Caribeacarus, 186, 188, 649, 668, 674, 842 Centsymplia, 762
Caribena, 647, 666 Cepheidae, 876
Caribezomus, 137, 646 Ceratinella, 866
caribicus, Byrso.chernes, 658 Ceratinopsis, 883
Caribochernes, 89, 658, 660 Ceratochernes, 487
Caribothyrus, 94, 195, 197, 198 Ceratogyrus, 696
Caribula longimana, 664 Ceratolasma, 92, 510
carinatus, Ideoblothrus, 656 Ceratolasmatidae, 153
Carinostoma, 327, 361 Ceratolasmatinae, 510
Caripetella, 708 Ceratoleptes, 492
carlotanus, Galibrotus, 663 Ceratomontia, 80
Carlotta, 492 Ceratoppiidae, 869, 877
Carparachne, 699 Ceratozetes, 884
carpathicus, Euscorpius, 335, 354, 358, 360 Ceratozetidae, 868, 869, 877
carpathicus sicanus, Euscorpius, 358 Cercophonius, 395, 407, 755, 756, 778
Carrai, 760 cerdoso, Antillostenochrus, 666
Carsevennia, 489 Cereipes, 468
Carteroniella, 698 Cereodiscus, 467
carusoi, Roncus, 355 Cerionesta, 649, 668
casabito, Rowlandius, 666, 671 Ceromella, 692
Cassinia, 468 Ceromidae, 164, 455, 457, 458, 683, 692,
castaneus, Erginulus, 661 722, 896
castaneus, Pachylicus, 664 Ceropachylinus, 492
Catageus, 141, 414, 415, 732, 895 Ceropachylus, 492
caucasicus, Mesobuthus, 371, 376, Cersa, 664
377, 687 Ceryerda, 762
Caucnemastoma, 327, 377, 378 Cethegus, 760
Caudalia, 649, 666, 668 Cetratus, 752
Caurimare, 489 Chacoikeontus, 491
948 Index
Chactas, 482, 484, 486, 655, 656, 672 Cheliferidae, 38, 148, 326, 336, 343, 346, 351,
Chactidae, 79, 145, 325, 482, 508, 631, 360, 364, 365, 372, 376–378, 380–381,
633, 642, 654, 656, 672, 674, 834, 387, 388, 392, 395, 398, 401, 406, 422,
887, 897 425, 432, 436, 451, 463, 483, 509, 521,
Chactoidea, 37 641, 658, 660, 671, 672, 677, 685, 694,
Chaerea, 452 712, 720, 723, 735, 747, 748, 756, 769,
Chaerilidae, 145, 395, 398, 401, 405, 407, 777, 778, 836, 845, 861, 871, 880
408, 424, 432, 734, 834, 870 Chelypus, 458, 692
Chaeriloidea, 37, 145 Chenistonia, 760
chaibassicus, “Schizomus”, 399, 417 Chernes, 336, 371, 660, 861
Chajnus, 85, 139, 428, 733 Chernetidae, 38, 149, 326, 336, 343, 346, 351,
chalakudicus, “Schizomus”, 417 355, 360, 364, 365, 371–372, 376–378,
Chalcolemia, 752 380, 387, 388, 395, 398, 401, 406, 422,
Chalcoscirtus, 866, 872 425, 432, 442, 451, 462, 483, 487, 503,
chalumeaui, Dinochernes, 658 505, 509, 515, 521, 641, 658, 660, 671,
Chamaia, 494 672, 676, 677, 685, 694, 705, 712, 715,
Chamberlinarius, 88, 90, 463, 476 720, 723, 735, 747, 748, 756, 769, 770,
Chamobatidae, 877 775, 777, 778, 836, 843, 845, 861
Chanbria, 508, 631 Cherserigone, 452
Chaneke, 627, 632, 633 Chetronus, 489
Charinidae, 36, 37, 75, 140, 381, 387, 394, Chilenodes, 678–679, 767
399, 404, 414, 427, 433, 452, 469, 495, chilensis, Acropsopilio, 506
503, 640, 645, 665, 669, 682, 683, 707, Chileogovea, 91, 488
716, 722, 732, 742, 749, 757, 775, 778, Chileotrecha, 676
781, 841, 895 Chiliochthonius, 86
Charinides, 141, 733 Chilobunus, 765
Charinus, 74, 75, 98, 141, 399, 404, 414, 415, Chilumena, 762
469, 470, 495, 496, 503, 626, 645, 665, Chinacreagris, 87
669, 682, 733, 749, 757, 758, 778, 841, Chinchipea, 489
844, 894 Chiriboga, 490
Charletonia, 191 Chiridiochernes, 89, 90, 425
Charmus, 401, 407 Chirimena, 495
Charon, 141, 414, 415, 428, 434, 733, 749, Chirinosbius, 489
757, 758, 781 Chitrella, 835
Charontidae, 36, 37, 74, 141, 414, 428, 434, Chorizopella, 698
516, 732, 742, 749, 841, 895 Chorizopinae, 707
chartoni, Eukoenenia, 400, 703 Chresiona, 697
Chasmocephalon, 760, 840 Chrestobunus, 765
Chatzakia, 352 christiani, Eukoenenia, 358
Chaussieria, 192 Chrosioderma, 709
Chavesincola, 491 Chrysochernes, 89, 509
Chediminae, 181, 739 Chthoniidae, 37, 146, 323, 326, 336, 337,
Cheiridiidae, 38, 147, 326, 336, 343, 346, 343, 346, 351, 354–355, 360, 364, 371,
360, 364, 365, 368, 371, 372, 377, 378, 372, 376–378, 380, 387, 388, 392, 395,
380, 387, 395, 398, 406, 422, 432, 442, 398, 401, 406, 422, 425, 432, 442, 451,
463, 483, 503, 505, 515, 521, 657, 659, 463, 483, 503, 509, 515, 521, 641, 655,
677, 685, 723, 735, 747, 769, 836, 845, 659, 671, 672, 676–677, 684, 693, 705,
861, 871 712, 715, 719, 723, 735, 747, 765, 769,
Cheiridium, 336, 365, 371, 659 775, 777, 781, 782, 834, 835, 842–845,
Cheirochelifer, 336 861, 871
Chelanops, 505, 658, 660 Chthonius, 336, 365, 371, 719, 835, 861, 882
chelanops, Gobichelifer, 880 Chthonopes, 419
Chelicerata, 35, 36 Chumma, 696
Chelifer, 336, 372, 880 Chummidae, 177, 686, 696, 725, 935
Index 949
Chusgonobius, 489 Collonidium, 491

Ciba, 648, 667 Colombiazomus, 137, 499, 500
Ciboneya, 648, 668 comasi, Arcanobisium, 326, 346
Cicileiurus, 450, 451 Comboyus, 644
Cicileus, 450, 461 Comereca, 465, 467
Cicirra, 766 Commoris, 649, 668
Cicurina, 866 communis, Gomphochernes, 658
Cicynethus, 697 complicatus, Improphantes, 867
Cingulata, 199, 679 Compluriscutula, 49
Cirolanidae, 95 compressus, Parachernes
cirtanum, Metaphalangium, 358, 451 (Scapanochernes), 658
Cispinilus, 474 Compsaditha, 432, 715, 719
Cithaeronidae, 178, 687, 725, 741 Compsobuthus, 383–385, 407, 450,
Cladonychiidae, 48, 156, 159, 327, 360, 460, 462, 688
361, 372, 511, 684, 838, 843, 899 Compsothrombium, 883
Cladycnis, 366 Conakrya, 474
Clarinus, 491 condolens, Pardosa, 885
Clastes, 751 confundens, Olpiolum, 657
Claverythraeus, 190 Congochthonius, 86, 90, 463, 476
clavicornis, Walckenaeria, 867 Congolla, 467
Clavicranaus, 493 Conicochernes, 89, 91, 757, 778
claviger, Clavizomus, 417 Conoculus, 757, 779, 840
clavigerum, Pseudochiridium, 404 consecuta, Dictyna, 885
Clavismaris, 190 conspicillatum, Paecilaema, 636, 643
Clavizomus, 137, 418, 428, 733 copalensis, Palaeogrosphus, 705
clethrionomydis, Laelaps, 82 Coprochernes, 89, 641
Clinobiantes, 466 Coptophalangium, 465
Clinocippus, 489 coralensis, Typhloroncus, 657
Clipeosoma, 190 Corambis, 776
Clubiona, 866 Corasoides, 761
Clubionidae, 77, 178, 328, 339, 341, 362, Cordobulgida, 84
396, 402, 404, 430, 647, 678, 686, 698, Corinnidae, 178, 328, 339, 362, 402, 441,
725, 741, 751, 840, 866, 867, 874 452, 647, 678, 687, 700, 726, 741, 752,
Cluniella, 757, 765 763, 900
Cnemoleptes, 490 Cornifalx, 766
coarctata, Marwe, 836, 842–843 Cornigera, 494
Cobania, 491 Cornupalpatum, 49
Cobaniinae, 491 coronatus, Acanthophrynus, 513
coccinella, Holothyrus, 194, 713 Corosalia, 489
Coccorchestes, 752, 763 Corosoma, 89, 90, 487
cochinensis, Sarax, 399 corsa, Acanthocreagris, 355
Cochirapha, 495 corsicus, Chthonius (Ephippiochthonius), 354
Cocholla, 488 Corticettus, 649, 668
Cocinachernes, 89 Corynethrix, 763
Coelotes, 841, 866 Coryssiphus, 698
Cogonella, 467 Cosmetidae, 40, 156, 157, 488, 501, 512, 635,
coinneaui, Adamystis, 881 640, 643, 644, 660, 661, 674, 677
cokendolpheri, Antillostenochrus, 665 Cosmetus, 643
cokendolpheri, Kimula, 663 Cosmobunus, 327, 347, 510, 634
Cokendolpherius, 137, 646, 666, 670, 671 Costabrimma, 645
Colcarteria, 761 Costacosa, 760
Colima, 635 Couranga, 761
Collemboerythraeus, 190 Cousinea, 717
Collina, 766 cousinensis, Rowlandius, 666, 671
950 Index
Cranaidae, 40, 156, 488, 489, 501, 644, 674 cubanus, Agoristenus, 662
Cranainae, 489 cubanus, Caribbiantes, 663
Cranaostygnus, 490 cubanus cubanus, Cynortoides, 661
Cranellus balthazar, 663 cubanus, Santobius, 663
crassicauda, Androctonus, 376, 377 cubanus signatus, Cynortoides, 661
crassicaudatus, Schizomus, 324, 401, 417 cubanus, Tridenchthonius, 656
Crassicrus, 647 cubanus, Tyrannochelifer, 658
crassimanus, Pachakutej, 887 cubaorientalis, Decuella, 663
crassipalpis, Sabacon, 372 Cubazomus, 85, 137, 646, 666, 670, 671
crassipalpum, Mitraceras, 716 cubensis, Ethobunus, 664
credula, Pardosa, 885 cubensis, Reddellzomus, 665
crinis, Plesioballarra, 779 Cubiria, 494
crinitus, Dasycleobis, 887 Cucudeta, 752
Cristina, 465 Cucutacola, 493
Crosbycus, 92, 411, 511 cuevajibarae, Manahunca, 663
Crosbyella, 92, 512 Cultroribula, 49
Crotonia, 97 Cumbalia, 489
crucifer, Typopeltis, 415, 421, 440 Cunnawarra, 761
crudeni, Heliophanus, 885 cupeyalensis, Rowlandius, 666
cruzensis, Antillochernes, 658 cupreus, Drassodes, 866, 867
Cryphoeca, 866 Curculioididae, 508
Cryphoecina, 341 Curimaguanus, 494
Cryptadaeum, 695 curioi, Sarax, 433
Cryptobiantes, 695 Currala, 491
Cryptobranchidae, 78 Curteria, 190
Cryptobranchus, 78 Cutervolus, 490
Cryptocellus, 143, 482, 640, 654, 669, 673, Cyatholipidae, 51, 178, 647, 667, 686, 699,
841, 843 708, 725, 731, 737, 739, 760, 901
Cryptocreagris, 509 Cyathura, 95
Cryptoditha, 86, 90, 483 Cybaeidae, 178, 328, 339, 341, 362, 686, 725,
Cryptoerithus, 762 740, 874
Cryptogeobius, 493 Cyclochernes, 89, 748
Cryptognomae, 36 Cycloctenidae, 178, 429, 741, 773, 780,
Cryptomaster, 511 841, 846
Cryptopygoplus, 695 Cyclosa, 521
Cryptowithius, 87, 90, 463, 476 Cymbachina, 773
Ctenidae, 178, 183, 402, 474, 647, 648, 667, Cymberemaeidae, 877
686, 707, 718, 725, 740, 760, 874 Cynorta, 635, 643
Cteniogaster, 473 Cynortellana, 643
Ctenizidae, 9, 172, 327, 339, 341, 422, Cynortellina, 489
436, 439, 646, 685, 724, 738, Cynortellula, 489
838, 845 Cynortetta, 489
Ctenizinae, 172, 738 Cynortoides, 635, 661
Ctenolophus, 699 Cynortoperna, 643
Ctenophora, 46 Cynortosoma, 489
Cuangoblemma, 474 Cynortula, 635, 643
Cubacanthozomus, 137, 646 Cyphophthalmi, 2, 36, 38–40, 48, 50, 66, 74,
Cubachelifer, 88, 89, 658, 660 76, 149, 323, 326–327, 338, 343,
cubae, Ammotrechella, 653 345–347, 355, 358, 372, 376–378, 388,
cubanacan, Rowlandius, 666, 671 391, 395–398, 400, 401, 409, 422, 425,
cubana, Ibantila, 663 432, 436, 440, 442, 451, 454, 464, 465,
Cubanopillus, 649, 666, 668 476, 478, 487, 503, 509, 516, 625, 627,
cubanum, Maracaynatum, 662 628, 632, 637, 640, 653, 660, 683, 692,
cubanum, Mexobisium, 657 694, 706, 711, 712, 714, 715, 720, 722,
Index 951
726, 727, 742, 743, 748, 757, 764, 765, Deelemanella, 430
770, 775, 778, 779, 836, 842–844, 846, Defectrix, 649
862, 898, 931–934 defensa, Haversia, 524, 680
Cyphophthalmus, 327, 338, 339, 347, 388, deficiens, Anachipteria, 868
836, 898 deharvengi, Koeneniodes, 431
cyprianus, Mesatemnus, 388 Deinopidae, 178, 647, 678, 686, 724, 739, 776
Cyptobunus, 511 delamarei, Lechytia, 655
Cyriogonus, 709 deleta, Eukoenenia, 405
Cyrtaucheniidae, 172, 173, 419, 646, 678, Delicola, 426
685, 697, 724, 738 Deltaspidium, 491
Cyrtaucheniinae, 173 deltshevi, Euscorpius (E.), 335
Cyrtogrammomma, 93 Demendians, 760
Cyrtowithius, 87, 89, 694 Dendrochernes, 33, 372
Dendrolasma, 97, 410, 511
denticulata, Hemerotrecha, 98, 508
D Denticynorta, 512
Dacnopilio, 465 dentipalpis, Erigone, 867, 885
Dactylochelifer, 336, 372, 861 dentipes, Pokennips, 178, 901
Daesiella, 84, 693 Dentobunus, 411, 425, 433
Daesiidae, 162, 324, 335, 345, 354, 376, 380, Deriacrus, 490
381, 394, 398, 448, 449, 455–458, 676, desecheo, Rowlandius, 666, 671
683, 692, 722, 860, 896 Desidae, 178, 328, 339, 525, 678, 686, 725,
daitoensis, Apozomus, 417 740, 761, 766, 773, 776, 846
Dalmasula, 698 desidiosus, Xysticus, 866, 867
Dalquestia, 635 Desognanops, 762
Damaeidae, 869, 876 Desognaphosa, 762
Damaeolidae, 876 Despirus, 491
Dambullaeus, 191 Detigalus, 493
dammermani, Stygophrynus, 428 Devendra, 402
Damoetas, 763 Devonacaridae, 67, 71
Damon, 415, 469, 470, 696, 894 Devonacarus, 67, 71
Damoninae, 141, 695 devotus, Zelotes, 866, 867
Dampetrinae, 156, 412, 736, 781 Diaenobunus, 776
Dampetrus, 779 Diallomus, 402
Dampfiellidae, 877 Diaprograpta, 762
dapaensis, Hilaira, 885 Diaprograptinae, 181, 731, 737, 741
daphne, Orsa, 662 diaspora, Ammotrechella, 364, 367
darainensis, Grosphus, 704 Diatomyidae, 95
darainensis, Tityobuthus, 705 Diblemma, 717
Dardurus, 761 Dibuninae, 156, 399, 737
darlingtoni, Trinimontius, 662 Dibunus, 433, 516
darwini, Amblyomma, 504 Dicranolasma, 327, 338, 339, 344, 356, 361,
Darwinneon, 504 372, 377, 378, 389
dastychi, Tityobuthus, 705 Dicranolasmatidae, 40, 154, 327, 337–339,
Dasychernes, 89, 90, 487 344, 356, 358, 360, 361, 377, 378, 381,
Dasycleobis, 82 388, 389, 410, 684, 899
Dasylobus, 327, 337, 338, 361, 388 Dicranopalpinae, 488
Dasypodidae, 199, 679 Dicranopalpus, 48, 50, 327, 337, 338, 344,
Dasyponyssidae, 199, 679 347, 355, 361, 488, 862
Daviesa, 761 Dicrognathus, 94, 198
Decaphora, 649, 666, 668 Dictynidae, 178, 328, 339, 341, 362, 369,
Decarynella, 706 396, 402, 404, 419, 452, 474, 514, 518,
Decuella, 664 647, 678, 686, 697, 725, 740, 761, 773,
decui, Rowlandius, 666, 671 780, 865–867, 874, 900
952 Index
Didymocentrus, 84, 642, 655, 656, 673 Draculoides, 85, 137, 734, 758, 759, 778
Digalistes, 490 Drassodella, 696
digitiger, Rowlandius, 666, 671 drenskii, Euscorpius (E.), 333, 335
Diguetidae, 178, 678 Drepanochthonius, 86, 90
Diguetinus, 635 Drepanotylus, 866
dimentmani, Ayalonia, 388 Dromeothrombium, 883
Dimizonops, 393 Dromopoda, 47
Dinaria, 327, 339, 838, 899 Drymusidae, 178, 646, 678, 685, 724, 901
Dingupa, 757, 779 dubius, Heliophanus, 885
Dino, 427 ducoudrayi, Rowlandius, 666, 671
Dinocheirus, 336, 658 duffeyi, Stenowithius, 368
Dinothrombium, 883 dufoureius, Iurus, 336, 383
Diolemus, 752 Dugdalea, 780
Diplocentridae, 145, 382, 383, 392, 508, 631, Dumaguetes, 433
655, 683, 834, 843 Dumoga, 430
Diplocephalus, 866 Duripelta, 772, 780
Diplogena, 697 Durodamus, 761
Diplotemnus, 336, 371 Dysdera, 178, 738, 839
Diplothrixochernes, 89, 91, 676 Dysderidae, 178, 328, 339–341, 356, 362,
Diplothyrus, 94, 194, 195, 197, 198 373, 379, 391, 422, 678, 686, 724, 738,
Diplotychus, 709 839, 873
Dipluridae, 173, 373, 402, 419, 422, 428, Dysderinae, 178, 738
646, 678, 685, 724, 731, 737, 760, 776, Dysderocrates, 340
838, 844, 846, 873, 883 Dysderomorphae, 41
Diplurinae, 173 Dyspnoi, 2, 38–40, 50, 80, 149, 153, 323, 327,
Dippenaaria, 696 338, 344, 346, 347, 352, 356, 358, 361,
Dipristes, 757, 779 372, 377, 378, 388, 389, 391, 396–401,
Diptyonius, 489 410, 411, 422, 425, 431, 433, 436, 440,
Discocyrtoides, 491 442, 451, 454, 465, 478, 488, 503, 510,
Discocyrtulus, 492 516, 625, 635, 640, 641, 653, 660, 673,
Discocyrtus, 492 677, 682, 684, 692, 695, 703, 706, 711,
disjunctodens, Antillotrecha, 653, 654 712, 714, 715, 723, 726, 727, 736, 742,
dispar, Hansenochrus, 666, 671 743, 747, 748, 754, 757, 764, 765, 770,
dispar, Salfacarus, 700 775, 779–782, 837, 862, 899, 931–935
dissimilis, Chaussieria, 369
distinctus, Alioranus, 885
Dithidae, 146, 705, 735, 777, 835 E
Djemia, 467 Eatoniana, 190
docolora, Eremobates, 98, 508 Echinobunus, 412, 425
Dolocosa, 369 Echinussa, 709
dolpoensis, Xysticus, 885 Ectoceras, 88, 90, 398
dominicanus, Parachelifer, 658 Edeala, 467
dominicanus, Parachernes, 658 Egaenus, 327, 337, 338, 344, 361, 372, 388,
Dongila, 467 396, 411, 880
Donuea, 709 Egyptobuthus, 450
doratodactylus, Troglochthonius, 355 ehrenbergi, Biton, 98, 335, 354, 381
doriai, Scotolemon, 356 Elassoctenus, 761
dorotheae, Pseudocellus, 50, 508, 627 elegans, Benoitinus, 716
dorsalis, Gluvia, 98, 345 elegans, Galeodes, 98, 334, 335
Dorymetaecus, 778 Eleleis, 698
Doryonychus, 522 Elleguna, 761
Dracochelidae, 66 ellenae, Haplochernes, 748
draco draco, Eukoenenia, 351 Elleriana, 488
draco zariquieyi, Eukoenenia, 345, 351 ellisi, Botswanoncus, 694, 897
Index 953
elongata, Kimula, 663 Eremorhax, 508, 631

elongatus, Paratemnoides, 657 Eremothera, 508, 631
Emplesiogonus, 709 Eresidae, 178, 327, 339, 362, 402, 686, 698,
emzantsiensis, Gondwanascorpio, 47 724, 725, 739, 901
Enantiobuninae, 677 Eresinae, 178
Enantiosarax, 141 Ergastria, 492
Encheiridium, 491 Erginiperna, 635
Encoptarthria, 762 Erginoides, 643
Encyocratella, 473 Erginulus, 635, 643, 661
Encyocrates, 707 Erigone, 866, 872, 883
Encyocrypta, 93, 776 Erigonops, 699
Enemothrombium, 884 Erigophantes, 430
engombe, Rowlandius, 666, 671 Ernstmayria, 326, 336, 337
Enielkenie, 422 Ero, 180, 739
Enigmazomus, 113, 471, 472, 476 eruptoclausus, Enigmazomus, 387
Eniochthoniidae, 876 Erythracarus, 192
Enoplognatha, 866 Erythraeidae, 49, 51, 67, 70, 189–191, 342,
Eopachyloides, 492 400, 475, 501, 638, 700, 763, 764, 768,
Eopachylus, 492 773, 776, 781, 868, 875, 883
Eordea, 430 Erythraeoidea, 5, 51, 67, 71, 189, 191, 768
Epactiochernes, 843 Erythraeus, 49, 67, 70, 190, 875
Epedanidae, 40, 156, 396, 398, 399, 412–414, Erythraxus, 190
422, 426, 433, 440, 516, 684, 737, 748, Erythrellus, 190, 763, 768
781, 782 Erythrites, 190, 763, 768
Epedaninae, 413, 426 Erythrocheylus, 192
Epedanoidea, 39, 40 Erythroides, 190, 763, 768
Epedanulus, 426 Escadabiidae, 40, 91, 158, 488, 490, 501
Epedanus, 426 Escadabius, 490
Epicharitus, 762 espanoli, Ptychosoma, 347
Epichernes, 89, 632 Ethobunus, 636, 644, 664
Epigastrina, 766 Etienneus, 75, 79, 85, 96, 139, 470, 476, 497,
Epimerata, 36 682, 895
Epiocheirata, 37, 146, 684, 723 Euagrinae, 173, 685, 737, 760
Ereala, 467 Euamblypygi, 140, 682
Erebalda, 467 Euasteron, 762
Erebomaster, 511, 838, 843 Eubiantes, 466
Erebomastridae, 838, 843 Eubunus, 765
Ereca, 465 Eucratoscelus, 473
Erecabia, 467 Euctenizidae, 173
Erecanana, 465, 712 Eucynorta, 635, 643
Erecananinae, 156, 399, 427, 465, 737, 749, Eucynortella, 643
781, 782 Eucynortoides, 635, 643, 661
Erecella, 465 Eucynortula, 635, 643
Erecongoa, 467 Eudasylobus, 347, 352
Erecops, 467 Eugagrella, 412, 425, 433
Erecula, 465 Eugnidia, 643
Eregonda, 467 Eugyndes, 492
Eremaeidae, 876 Eugyndopsiella, 492
Eremobates, 98, 508, 631 Eukoenenia, 135, 323, 334, 343, 359, 424,
Eremobatidae, 166, 167, 508, 627, 631 681, 703, 712, 718, 732, 833, 893, 894
Eremobelbidae, 876 Eukoeneniidae, 135, 323, 334, 405, 424, 520,
Eremochelis, 508, 631 626, 631, 681, 683, 703, 712, 722, 732,
Eremocosta, 508, 631 743, 747, 781, 843
Eremogarypus, 87, 89, 693 Eulibitia, 489
954 Index
Eulohmanniidae, 876 F
Eumecochernes, 89, 521 Fageibiantes, 706
Eumesosoma, 510 Fageicera, 647, 666, 839
Euminua, 494 Fageiella, 340
Euminuoides, 494 Fakoa, 467
eupeus, Mesobuthus, 98, 371, 376, 377, falcifemus, Rowlandius, 666
687, 688 Falcileptoneta, 93, 437, 839
Euphalangium, 396, 398, 442, 863, Falklandia, 680
880, 883 Falklandoglenes, 680
Euphrynichus, 142, 415, 469, 470, 841 fallax, Prasiana, 662
Euphthiracaridae, 876 falxus, Mughiphantes, 885
Eupnoi, 38–40, 50, 80, 149, 153, 323, 327, Fangensinae, 409
338, 344, 346, 347, 352, 355, 358, 361, Fangensis, 91, 409
365, 372, 377, 378, 388, 391, 395, 398, Farqua, 717
401, 411, 422, 425, 432–433, 436, 440, Feaella, 715, 719
442, 451, 465, 478,, 488, 503, 505, Feaellidae, 38, 146, 401, 463, 483, 684, 705,
510, 625, 634, 640, 641, 660, 673, 677, 715, 719, 723, 933
684, 692, 695, 703, 706, 712, 714, 715, Felsina, 475
722, 726, 727, 736, 742, 743, 748, 757, Femoracoelotes, 422
765, 770, 779–782, 862, 880, 898–899, Fenicheliidae, 877
931–933, 935 Feretrius, 516
Eupodoctis, 399 Ferkeria, 489
Eupoecilaema, 643 fernandezi, Parascleropilio, 366
Eupograpta, 762 ferruginea, Metazalmoxis, 716
Eurybunus, 635 ferrugineus, Eudasylobus, 352
Euryeidon, 420 Fessonia, 67, 70, 189
Eurypoena, 367 feti, Grosphus, 704
Eurypterida, 36, 46 Fijicolana, 749
Eurytromma, 401 Fijocrypta, 93, 751
Euscorpiidae, 145, 325, 335, 343, 346, 354, Filistatidae, 41, 178, 327, 339, 404, 646,
360, 371, 376, 378, 380, 382, 383, 385, 648, 667, 678, 685, 708, 724, 731,
386, 405, 407, 408, 450, 482, 485, 633, 737, 738, 846
642, 673, 674, 683, 688, 860, 870 Filistatinae, 178
Euscorpiops, 394, 407, 408, 688 Filistatomorphae, 41
Euscorpius, 185, 325, 359, 385, 386, 448, 450, fiskei, Stygnomma, 662, 843
485, 833, 860, 870 Fissarena, 762
Eusidama, 465, 467 Fissilicreagris, 87, 509
Eusimonia, 98, 324, 364, 381, 449 Fissiphalium, 488
Eusitalces, 401 Fissiphalliidae, 40, 158, 490
Euso, 717 Fizibius, 467
Eutegaeidae, 876 Flavarchaea, 760
Eutichuridae, 178, 369, 402, 430 flavicaudis, Euscorpius, 343, 346, 354, 358
Eutimesius, 494 flavidus, Mizozatus, 375
Eutrichothrombium, 883 flavipes, Phalangodes, 663
Eutrombidiidae, 420, 875, 884 flavitarsis, Stygnoplus, 662
Eutrombidium, 884 flaviventris, Hinzuanius, 392
Euzaleptus, 412, 425 Flavonuncia, 706
Evansia, 866 flavopiceus, Grosphus, 704
everestensis, Euophrys, 885 flavopictus, Ibalonius, 716
Exechocentrus, 708 Flirtea, 643
Exlineia, 495 florenciae, Eukoenenia, 481, 508, 520, 626
Exopalistes, 751 florentiae, Rowlandius, 666
exsul, Phrynus, 75, 77, 427, 428, 496 Florichelifer, 88, 509
Index 955
floridanus, Tyrannochelifer, 658 Galaporella, 504

Folkia, 340, 839 Galeodellus, 880
Forania, 190 Galeodidae, 160, 161, 324, 334, 371, 376,
forbesii, Heteronebo, 79, 392 380, 381, 394, 398, 448, 449, 455–458,
Forfexia, 489 683, 722, 860, 896
formicoides,“Schizomus”, 417 Galeodila, 880
Forstarchaea, 780 Galeodopsis, 449
Forstera, 760 Galeodumus, 83, 382
Forsterella, 773 Galeosoma, 699
Forsterina, 761, 846 Galibrotus, 664
Forsteriola, 474 Galliena, 429
forsteri, S. (Neocharon), 749 Gallieniella, 707
Forsteropsalis, 770, 780, 846 Gallieniellidae, 178, 687, 696, 707, 731, 737,
Forstertyna, 773, 780 741
Forstrarchaea, 771, 780 Galumna, 881, 884
Fortalezius, 488 Galumnidae, 878
Fozustium, 191 Gamasomorphinae, 181, 717, 738
Francochthonius, 86, 91 garciai, Grosphus, 704
fraterna, Antillotrecha, 653, 654 Garcorops, 709
fraterna, Cynortula, 661 gardineri, Sitalcicus, 716
Frewena, 763 Gargenna, 427
Friburgoia, 491 garna, Cynortula, 661
frigidus, Incestophantes, 867 Garypidae, 38, 147, 326, 336, 343, 346, 351,
Friula, 430 367, 368, 372, 377, 378, 380, 388, 392,
Frizellia, 489 398, 401, 425, 451, 463, 483, 503, 657,
frondiger, Koeneniodes, 424, 712 660, 677, 685, 693, 705, 715, 719, 723,
Frontiphantes, 366 735, 765, 769, 843, 861
Fuchiba, 697 Garypinidae, 38, 148, 326, 336, 346, 380,
Fuchibotulus, 697 387, 388, 406, 425, 451, 483, 503, 521,
Fudeci, 494 641, 657, 659, 671, 676, 677, 685, 715,
fujuge fujuge, Hammenius, 752 719, 723, 735, 747, 765, 769, 775, 781,
fujuge iora, Hammenius, 752 782, 836
Fumontana, 511 Garypinus, 336
funafutensis, Haplochernes, 748 Garypus, 336
fundorai, Negreaella, 663 Gasparia, 525, 773, 780
Funkikoa, 422 gastrostictus, Hypoctonus, 428
furcichelis, Barrussus, 334 Gaucha, 84, 676
Furculattus, 752, 763 Gauchella, 84, 481
fuscimanus, Phrynus, 513 Geaya, 634, 641, 660
Gelachernes, 89, 748
Gelaninae, 180
G geniculata, Ammotrechella, 653
Gaditusa, 427 geniezi, Mauritanobuthus, 448, 451
gadorensis, Eukoenenia, 345 Genysa, 708
Gagrella, 396, 401, 412, 422, 425, 433, 440, Genysinae, 173, 708, 901
712, 748, 757, 781 Geogarypidae, 17, 38, 147, 326, 336, 343,
Gagrellina, 412, 425 346, 351, 360, 365, 371, 380, 387,
Gagrellinae, 411, 433, 634, 899 388, 392, 395, 398, 401, 406, 422,
Gagrellissa, 412, 425 425, 432, 451, 463, 483, 505, 515,
Gagrellula, 401, 411, 412, 425, 433, 440 521, 676, 677, 685, 715, 719, 723,
Gaibulus, 494 747, 748, 782, 861, 871
Galanomma, 495 Geogarypus, 336, 365, 432, 505, 715,
Galapa, 504 719, 748
galapagoensis, Ixodes, 504 Geospizinae, 502
956 Index
Geraecormobius, 491 Gomezyta, 467

Geralinura, 65 Gonazula, 491
Geralycosomorphae, 41 Goniosoma, 491
Gerdesius, 492 Goniosomoides, 491
gerlachi, Leptokoenenia, 381 Gonogotus, 493
gerlahi, Apozomus, 716 Gonyassamiinae, 491
germanus, Euscorpius, 98, 343, 360 Gonyleptellus, 491
Gertia, 493 Gonyleptes, 491
gertschi, Meriosfera, 662 Gonyleptidae, 40, 156, 488, 490, 501, 503,
gestroi, Dasylobus, 356 524, 644, 674, 677, 678, 680, 837, 844,
gibarensis, Antillostenochrus, 665 845, 862
gibbosus, Mesobuthus, 333, 335, 383 Gonyleptoidea, 38–40
giebeli, Pardosa, 867 Gonyleptoides, 493
gigantes, Hahnia, 885 Goodnightiella, 492
giganteus, Mastigoproctus, 513, 628, 637 goodnighti, Ethobunus, 664
Giljarovia, 327, 377, 378, 389 goodnightiorum, Kimula, 663
Ginosigma, 139, 416, 428, 733 Gorgasella, 649
Gint, 462 Goyenia, 773, 780
girgintiensis, Chthonius, 358 gracilis, Damon, 696
Giribetia, 409 gracilis, Rowlandius, 666
giustii, Chthonius (E.), 355 Gradungula, 770, 780
Glabropelmatinae, 176 Gradungulidae, 179, 738, 759, 771, 780
glaciale, Theridion, 885 Graecophalangium, 327, 337, 338
glacialis, Chalcoscirtus, 885 Graecotrombium, 191, 342
glacialis, Oreonetides, 867 graecus, Galeodes, 98, 333–335, 381
gladiger, Rowlandius, 666, 671 Graemontia, 695
gliroides, Dromiciops, 95 grafittii, Chthonius (E.), 355
Globibunus, 490 grafittii, Eukoenenia, 334
Globipes, 635 grafittii, Spelyngochthonius, 355
Globitarsus, 490 Grancanaridion, 367
Globocreagris, 87, 509 grandidieri, Grosphus, 704
Gluvia, 324 Grandjeanella, 191
Gluviema, 83, 455 grandjeani grandjeani, Hammenius, 752
Gluviopsylla, 381, 449, 455 Granobunus, 748
Glyptobunus, 765 granpiedrae, Orghidaniella, 662
Glyptogluteus, 85, 139, 416, 434, 733 granti, Heteronebo, 79, 392
Glysterus, 644 Granulaia, 494
Gnaphosa, 866 granulata, Cynortesta, 661
Gnaphosidae, 82, 179, 328, 339, 341, 348, granulatus, Vonones, 662
352, 362, 366, 367, 369, 373, 379, 396, Graphinotus, 492
399, 404, 420, 443, 504, 513, 514, 647, grassii, Trithyreus, 417
678, 687, 700, 717, 725, 741, 762, 773, Gravelyzomus, 137, 399
840, 865–867, 874, 881, 900 Graycassis, 762
Gnaricia, 489 Grayenulla, 763
Gnatopia, 489 grayi, Charon, 414, 421, 428, 434, 516, 749
Gnidiella, 489 greeni, Indothyrus, 198, 403
Gnoerichia, 475 greeni, “Schizomus”, 417
Gnomulus, 396, 399, 414, 433 Griswoldia, 697
Gnosippus, 381, 449, 455, 456 griswoldi, Tityobuthus, 705
Gobichernes, 89, 372 griveaudi, Neogrosphus, 704
gobiensis, Karschia, 371 Grosphus, 704, 705, 719
Godumops, 751 gruberi, Calchas, 383
Gohia, 525, 773, 780 gryllipes, Ammotrechinus, 653, 654
Goleta, 709 guadeloupensis, Tyrannochthonius, 655
Index 957
Guagonia, 494 haitensis, Vestitecola, 663

Guaiquinimia, 493 haitiensis, Pellobunus, 662
Guanazomus, 137, 646 Haitonia, 495
guantanamero, Rowlandius, 666 Hala, 93, 708
Guaraniticus, 492 Halidae, 93, 708, 721, 932, 935
Guasinia, 493 Halobisium, 509
Guasiniidae, 40, 158, 488, 493, 501 Hammenius, 94, 194, 196, 198, 718, 752, 781
guatemalae, Mayacentrum, 645, 670 handschini, Mughiphantes, 867
Guatubesia, 491 hanseni, Eukoenenia, 448, 508, 520, 626,
Guayania, 495 631, 712
Guayaquiliana, 489 Hansenochrus, 137, 498–500, 645, 646,
Gudalura, 92 670, 844
gudauricus, Hirstionyssus, 82 Haplochelifer, 88, 509, 861
Gueroma, 635 Haplochernes, 720, 748
Guerrobunus, 636, 637 Haploditha, 86, 90, 483
guillaumeti, Tityobuthus, 704 Haplodrassus, 866
Guiniella, 751 Haplothyrus, 94, 194, 196, 198, 742, 777, 782
gulosa, Meioneta, 82 Haplotmarus, 420
Gulufia, 467 Haplozetidae, 877
Gunasekara, 402 Hapona, 773, 780
Guruia, 465 Haptopoda, 45
Gustaviidae, 877 harai, Octozaleptus, 885
guzikae, Eukoenenia, 755 hardwicki, Scorpiops, 884
Gyas, 327, 338, 361, 372, 862 Harpachylus, 492
Gylippidae, 166, 376, 377, 381, 683, 722, Harpactira, 696
860, 896 Harpactirella, 696
Gymnobisiidae, 38, 147, 483, 676, 677, 679, Harpagella, 191
684, 694, 897, 933, 934 Harpagonellus, 426
Gymnobisium, 86, 89, 677, 694, 897, 934 hassleri, Cynorta, 661
Gymnodamaeidae, 876 Hauptmannia, 191
Gyndesoides, 492 Haurokoa, 773, 780
Gyndesops, 491 Havaika, 522
Gyndoides, 492 Haversia, 524, 680
Hedwiga, 770, 779
heeri, Drassodes, 866, 867
H Helebiona, 369
Haasus, 92, 389, 899 Helenactyna, 369
Habibiella, 382 Heliconilla, 420
Hadites, 341 Heliella, 493
Hadoblothrus, 96, 326, 336, 835 Heliophanus, 867, 883
Hadogenes, 693 Helladerythraeus, 190, 342
Hadogenidae, 145 hellenicus, Galeodes, 334
Hadrobunus, 510, 634 helleri, Diplocephalus, 867
Hadronyche, 760 Helsinia, 780
Hadrotarsinae, 751 Helvidia, 430
Hadruroides, 482, 486, 887 Hemerotrecha, 508, 631
hadzii, Euscorpius (E.), 335 Hemibuthus, 407, 688
Hadzinia, 327, 339 Hemicloeina, 762
Haedanula, 475 Heminothrus, 881, 884
Haemaphysalis, 710, 870, 878, 882 Hemiscorpiidae, 145, 380, 383, 385, 386,
Hahnia, 883 450, 459, 462, 652, 683, 688, 703
Hahniidae, 77, 179, 328, 339, 341, 362, 375, Hemiscorpius, 385, 386, 450, 462, 688
396, 402, 514, 647, 678, 686, 725, 741, Hemisolinus, 87, 89, 368
766, 773, 841, 845, 865–867, 874, 900 Hendea, 770, 779, 846
958 Index
Hendeola, 770, 779 Hexathelinae, 173, 731, 737, 759, 760,

Henriquea, 467 767, 900
Heradida, 697 Hexisopodidae, 169, 458, 683, 692, 722, 896
Heradion, 420 Hexisopus, 458, 692
Herbessus, 719 Hiboka, 708
Heriaesynaema, 475 Hickmanapis, 766
Hermannia, 881 Hickmania, 766, 838
Hermanniellidae, 876 Hickmaniinae, 177, 738, 766, 767, 901
Hermanniidae, 869, 876 Hickmanolobus, 766
Hernandaria, 644 Hickmanoxyomma, 765, 846
Hernandarioides, 644 himalayensis, Argas, 396, 886
Hersiliidae, 9, 179, 402–404, 647, 678, 686, Himalteneriffia, 188
699, 707, 724, 739 Hindreus, 465
Hesperochernes, 660 Hinewaia, 773
Hesperonemastoma, 511 Hinzuanius, 465, 712
Hesperopilio, 80, 91, 153, 757, 779 Hirriusa, 700
Hestimodema, 761 Hirstiosoma, 189, 190
Hesydrimorpha, 751 hirtus, Grosphus, 704
Hetaerica, 762 hispanica, Eukoenenia, 345
Heterobelbidae, 876 hispaniolana, Ammotrechella, 653, 654
Heterochernes, 89, 770, 779 hispaniolica, Ahotta, 662
Heterochthoniidae, 876 hispaniolicus, Chernes, 658
Heterocranainae, 490 Hispanognatha, 647, 667
Heterocranaus, 490 Histagonia, 698
Heterocubazomus, 137, 646, 666, 670 Histiostoma, 883
Heteroepedanus, 413, 426 Histiostomatidae, 883
Heteroibalonius, 413, 427, 749 Histricostoma, 50, 327, 356, 377, 378
Heterolacurbs, 466 Hoedillus, 648
Heteromeloleptes, 493 hoffi, Tyrannochthonius, 843
Heterometrus, 395, 408, 432, 688 Hoffmannichadrurus, 627, 632
Heteronebo, 79, 386, 392, 655, 656, Hoggicosa, 760
673, 897 Holactinochitinosi, 36
Heteronochrus, 137, 645, 646, 843 Holarchaeidae, 177, 739, 759, 841, 846
Heteronuncia, 757, 779 Holasteron, 762
Heteropachylinae, 491 Holcobunus, 634
Heteropachylus, 491 Holconia, 761
Heterophryninae, 142, 496 holguin, Antillostenochrus, 665
Heterophrynus, 142, 495, 496, 665, 669, 844 Holissus, 356
Heteropodoctis, 749, 781 Holonuncia, 757, 779
Heteropygoplus, 748 Holoscotolemon, 159, 327, 361, 372, 373, 684,
Heteroricinoides, 143, 841 862, 899
Heteroscorpionidae, 145, 722 Holothyrida, 2, 4, 35, 36, 66, 70, 76, 77,
Heterostygninae, 494, 662 94, 184, 185, 194–198, 319, 323, 333,
Heterovonones, 636, 661 343, 354, 364, 368, 371, 374, 376, 381,
heurtaultae, Phrynichus, 392 391, 397, 400, 403, 421, 424, 431, 436,
Hewittia, 475 439, 442, 454, 476, 478, 501, 625, 640,
Hexablemma, 474 650, 652, 668, 674, 687, 690, 692, 703,
Hexabunus, 490 711, 713, 714, 718, 721, 726, 727,
Hexachernes, 89 742–744, 747, 752, 755, 764, 773,
hexapodoides, Lichirtes, 662 775, 777, 779–782, 842, 859, 869,
Hexathele, 771, 780 931, 933, 935
Hexathelidae, 173, 422, 436, 439, 678, 685, Holothyridae, 194, 198, 714, 726, 742, 744,
724, 731, 737, 759, 760, 765, 767, 771, 747, 777, 781, 782, 878, 933, 935
780, 846, 900 Holothyrina, 194
Index 959
Holothyroidea, 194 Hyperwithius, 87

Holothyrus, 94, 194–198, 713, 721 Hypnoonops, 473
Holoversia, 491 Hypochilidae, 179, 685, 724, 901
Holovonones, 636, 643 Hypochiloidea, 41
Holozoster, 716 Hypochilomorphae, 41
holthuisi, Leiothyrus, 752 Hypochthoniidae, 876
Homalenotus, 327, 338, 344, 347, 352 Hypoctoninae, 37, 79, 139, 497, 733, 895
Homalonychidae, 179 Hypoctonus, 37, 75, 78, 139, 399, 415, 416,
Homocranaus, 490 428, 497, 733
Homoeothele, 762 Hypomegalopsalis, 757
Homolophus, 372, 388, 396, 411, 863, 885 Hypophyllonomus, 492
Homostela, 697 Hypoxestus, 465
Hoplobunus, 512, 636, 637 Hysterochelifer, 336, 861
Hoplodino, 433 Hystrichonyssidae, 199
Hoplolathys, 474
Hormuridae, 145, 395, 405, 417, 418, 424,
432, 436, 440, 442, 459–463, 482, 485, I
486, 515, 642, 654, 656, 672, 681, 683, Iandumoema, 492
705, 712, 714, 722, 734, 742, 747, 755, Ibaloniinae, 156, 399, 427, 465, 663, 737,
775, 781, 782, 860, 870 749, 781, 782
Horribates, 508 Ibalonius, 433, 712
hortuspalmarum, Bucinozomus, 324 Ibarra, 492
Hostus, 709 ibericus, Buthus, 346
Hottentotta, 383–385, 401, 407, 408, 450, Iberosiro, 91, 347, 898
460–462, 687, 688 ibex, Xysticus, 866, 867
Hovanoceros, 706 Icaleptes, 493
Hovanuncia, 706 Icaleptidae, 40, 158, 488, 493, 501
Hovops, 709 Ideoblothrus, 659, 672, 715, 719, 835
howarthi, Tyrannochthonius, 521, 845 Ideocaira, 698
Huadquina, 492 Ideoroncidae, 17, 38, 146, 387, 388, 392,
Huasampillia, 492 398, 401, 406, 463, 483, 641, 657, 659,
huascaran, Orobothriurus, 887 671, 677, 684, 694, 723, 734, 742, 743,
Hubbardia, 98, 137, 138, 513, 628, 631, 894 835, 842, 844, 861, 882, 897
Hubbardiidae, 46, 98, 136, 471, 499, 513, Ideoroncus, 86, 90, 483
515, 670, 683, 716, 722, 733, 734, 742, Idiochelifer, 88, 509
750, 775, 778, 782, 843, 894 Idiommata, 93
Hubbardiinae, 136, 471, 894 Idiopidae, 173, 402, 646, 678, 685, 699, 708,
Huitaka, 91 724, 731, 737, 738, 771, 901
Huka, 772, 780 Idiopinae, 174
Hulua, 773, 780 Idiothele, 696
Humbea, 467 Idjena, 427
Huntia, 761 Idomenta, 490
Huntiglennia, 763 Idzubius, 437
Huralvioides, 492 Iguarassua, 494
Huttonia, 771, 780 Iguassua, 492
Huttoniidae, 179, 739, 771, 778, 780, 782 Iguassuoides, 492
hyatti, Ixodes, 885 Iguatonia, 191, 501
Hydromantes, 78 Ikuma, 698
Hydrozetes, 67, 71 Ilipula, 419
Hygrochelifer, 88, 90, 398 Ilisoa, 699
Hyidae, 17, 38, 147, 398, 401, 406, 425, Illinichernes, 89, 509
432, 684, 705, 719, 723, 735, 845, imbulunga, Symphytognatha, 182, 740
861, 871, 933 imitatus, Tyrannochthonius, 655
Hyperpachylus, 492 Improphantes, 866
960 Index
Incasarcus, 491 ischyrum balearicum, Neobisium, 351

incertus, Sitalcicus, 716 Isela, 697
Incetophantes, 866 ishikawai, Anapistula, 182, 740
Indiacarus, 93, 186, 187, 741 Isocranaus, 490
indivisa, Bidoma, 663 Isolachus, 512
Indohya, 86, 719 Isometroides, 755, 756, 778
Indothyrus, 94, 194, 195, 198, 400, 742, Isometrus, 384, 401, 407, 408, 432, 450, 462,
777, 782 484, 486, 642, 655, 656, 688, 704, 747,
inexpectatus, Riftobuthus, 95 755, 756, 775, 781
Inferiores, 76, 97 israchanani, Akrav, 834
ingii, Hammenius, 752, 869 Israzorides, 390
Inhuma, 491 italicus, Euscorpius, 335, 343, 358, 371,
Innesa, 641, 669 376, 378
Innoxius, 494 italicus, Opilioacarus, 185, 186, 328,
Inola, 761 344, 357
inscriptus, Ibalonius, 716 Italustium, 191
insignis, Hansenochrus, 671 Itatiaincola, 492
insignis, Rowlandius, 666 Iuridae, 141, 325, 336, 376, 378, 380, 382,
insulaepinorum, Luisarmasius, 666, 671 383, 385, 482, 683, 887
insulae, Pseudochiridium, 657 Iurus, 325, 383, 385
insulana, Erecanana, 713 iviei, Antillotrecha, 653, 654
insulanus, Mirda, 664 Ivobiantes, 466
insulare, Cheiridium, 657 Ivocoryphus, 467
insularis, Charinus, 503, 934 Ivohibea, 706
insularis, Hammenius, 752 ivohibe, Tityobuthus, 704, 706
insularis, Heterovonones, 661 Ixamatus, 760
insularis, Lissochelifer, 748 Ixodes, 710, 870, 878, 882, 885
insularis, Orthochirus bicolor, 392 Ixodida, 3–5, 36, 49, 68, 69, 71, 72, 94, 97,
insularum, Ideoblothrus, 657 99, 184, 185, 198, 293, 362, 374, 396,
Insulocreagris, 87, 326, 336, 337 403, 421, 422, 442, 452, 501, 504, 514,
intertidalis, Grosphus, 704 515, 522, 524, 527, 638, 668, 687, 692,
Invexillata, 751 700, 710, 711, 726, 744, 764, 767–768,
Iocheirata, 37, 38, 146, 684, 723 773, 777, 779, 780, 859, 869, 878, 879,
Iomachus, 408, 462, 463, 705 882, 884–886, 931, 933–934
Iona, 519 Izecksohnopilio, 492
Iphoctesis, 709 Izunonychus, 437
Iporangia, 493
Iraponia, 390
Irileka, 761 J
Irnia, 467 Jacaena, 419
Irumua, 92, 467 jacksoni, Ixodes, 767, 773
isabel, Rowlandius, 666, 671 Jacksonoides, 763
Isaeinae, 156 Jajinia, 495
Isaeolus, 645 Jalkaraburra, 761
Isala, 763 jamaicana, Akdalima, 662
Ischnothelinae, 173 jamaicensis, Troglobochica, 657, 835, 843
Ischnotherus, 491 Jamaitidion, 648, 667
Ischnothyrella, 717 Jamara, 761
Ischyropsalididae, 40, 153, 327, 337, 338, Jamberoo, 761
344, 347, 360, 361, 372, 388, 410, 440, Janusia, 760
510, 684, 837, 862, 864, 871, 880, 899 Japetus, 427
Ischyropsalidoidea, 38–40 jaqueti, Holoscotolemon, 373
Ischyropsalis, 327, 338, 344, 347, 361, Jarmilana, 80, 644, 837, 900, 935
372, 862 jarmillae, Rowlandius, 666, 671
Index 961
Jaundea, 467 Katumbea, 474

Javazomus, 85, 137, 417, 418, 428 katyae, Calyptostoma, 67, 70
javensis, Sarax, 427 kaznakovi, Bisetocreagris, 881, 885
jerai, Anapistula, 182, 740 Keera, 761
jerseyi, Carios, 68, 71 Keilira, 761
Jim, 490 Kekenboschiella, 751
Jimeneziella, 664 kenyae, Callitrichia, 885
Jimeneziella negreai, 664 Kenyazomus, 85, 137, 471, 472, 476
jocasta, Chileogovea, 488 kerguelenensis, Ixodes, 527
Jocquella, 750 Kevonones, 636, 643
Johnstonianidae, 875 kharagpurensis, “Schizomus”, 399, 417
jonensis, Bituberochernes, 658 Kijabe, 473
Juanfernandezia, 506 Kilifina, 473
juberthiei cytheriaca, Eukoenenia, 334 Kilungius, 422, 440
juberthiei hellenica, Eukoenenia, 334 Kilyana, 761
Judalana, 763 Kimula, 48, 663, 664
judsoni, Tityobuthus, 704 Kimulidae, 40, 48, 158, 488, 494, 501, 660,
jugorum, Neobisium, 861, 862, 871, 880 663, 674
Julattenius, 85, 137, 734, 758, 759, 778 kinzelbachi, Iurus, 383
juncta, Cynortula, 661 Kituvia, 467
Junicus, 492 klapperichi, Parazaona, 658
Junquito, 495 Knoelle, 760
Juticus, 492 Koeneniodes, 135, 703, 712, 718, 732, 747,
jutisi, Ammotrechella, 653 781, 893
Juxtachelifer, 88, 509 Koinothrix, 367
Koloti, 627, 632, 633
Konetontli, 627, 632, 633
K Konfiniotis, 90, 338, 339
Kaapora, 494 koschewnikowi, Euscorpius (E.), 335
Kababina, 761 Koyanus, 426
kadleci, Iurus, 383 kozlovi, Galeodes, 371
Kainonychus, 437 Kraepelinia, 382, 384, 386
Kaitawa, 773, 780 kraepelini, Iurus, 383
Kakamasia, 190, 700 Kratochvilia, 473
Kakontwea, 467 kratochvili, Vampyrostenus, 663
Kamertonia, 190 krausi, Scotolemon, 352
Kangarosa, 760 Kraussiana, 189
Kapanga, 773 Krukt, 761
Karamea, 779 Krusa, 634
Karanga, 771, 779 Kuarapu, 627, 632, 633
Karangania, 467 Kuchingius, 426
Karaops, 762 Kulalania, 474
Karasbergia, 693 Kungwea, 467
karnai, Yllenus, 885 kunti, Buthus, 383
Karos, 636 kuntzi, Ixodes, 422
Karriella, 761 Kustarachnida, 45
Karripurcellia, 91, 757, 778 Kwangonia, 448
karschi, Ibalonius, 716 Kwonkan, 760
Karschiidae, 162, 324, 334, 364, 371, 376,
380–382, 394, 448, 449, 683, 860, 896
Karsenia, 78 L
Kasaina, 467 labarcae, Rowlandius, 666, 671
Kashimachelifer, 88, 91, 436 Labidognatha, 41, 638
Katableps, 709 Labidostomidae, 875
962 Index
Labochirus, 37, 139, 399, 416, 428, 497 Lathyarcha, 761

Labullinyphia, 402 Latifrons, 752
Lacinius, 50, 327, 337, 338, 344, 361, 377, Latimeria, 95
378, 388, 411, 862 latipes, Secozomus, 85, 716
Lacronia, 492 Laudetia, 648, 667
Ladantola, 490 laurae, Caribezomus, 666
Laelapidae, 49, 68, 71, 369, 869, 878 Lawrencega, 84, 693
laevigatus, Scheloribates, 881, 884 lawrencei, Maccabeesa, 712
laevis, Cynortesta, 661 lawrencei, Salfacarus, 700
Lagynochthonius, 365, 432, 835 Lawrencella, 695
Lakarobius, 750 Lawrencezomus, 137, 471, 472, 476
lalyae, Pseudouroplectus, 705 laysanensis, Ixodes, 522
lama, Anomalohimalaya, 878 Lechytia, 503, 505, 659
lamaika, Malaika, 699 Lechytiidae, 17, 37, 146, 380, 387, 395, 398,
Lamania, 429 406, 483, 503, 505, 521, 676, 684, 723,
Lamina, 773, 780 735, 835, 861, 871, 882
Lampertia, 709 leclerci, Koeneniodes, 405
Lamponata, 762 Leeuwenhoekiidae, 875
Lamponega, 762 legendrei, Holothyrus, 713
Lamponicta, 762 Legendrena, 707
Lamponidae, 179, 428, 739, 762, 776 Leiobunum, 48, 97, 327, 338, 347, 352, 361,
Lamponina, 762 372, 389, 411, 422, 440, 510, 634, 862
Lamponinae, 179, 739 Leiopelma, 78
Lamponoides, 762 Leiopelmatidae, 78
Lamponusa, 762 Leiosteninae, 488
Lamprochernes, 336, 720 Leiothyrus, 94, 194, 196, 198, 781
Landoppo, 434 Leiurus, 383, 384, 386, 450, 461
Lanesoares, 493 Lejokus, 427
Laniatores, 2, 38–40, 50, 96, 149, 154, 323, Leleupereca, 465
327, 339, 344, 346, 347, 352, 356, 358, Leleupiolus, 467
361, 365, 372, 376–378, 389, 392, 396, leporina, Gnaphosa, 866, 867
398, 399, 401, 412, 422, 426, 433, 437, Leptasteron, 762
443, 452, 465, 478, 487, 501, 503, 510, Leptathamas, 752, 763
511, 516, 625, 627, 628, 632, 635, 637, Lepthercus, 699
640, 643, 660, 674, 677, 680, 684, 692, Lepthyphantes, 839, 866, 881, 883
695, 706, 711, 712, 715, 723, 726, 736, Leptobunus, 97
742, 744, 748, 757, 764, 765, 770, 775, Leptocnema, 493
779, 781, 782, 836–843, 845, 859, Leptokoenenia, 135, 481, 833, 893
862–864, 871, 882, 899, 931–935 Leptonetidae, 179, 324, 339, 341, 437, 443,
lantiguai, Rowlandius, 666, 671 647, 685, 724, 839, 844, 901
Lanzaia, 476 Leptopsalis, 409, 410, 425, 781
Lanzatus, 85, 462, 463 Leptostygnus, 488
Laonastes, 95 Leptothele, 419
lao, Vietbocap, 406, 834 Leptotrombidium, 875, 881
Lara, 495 Leptus, 49, 67, 70, 190, 645
Larca, 336 Lessertina, 699
Larcidae, 38, 147, 326, 336, 346, 360, Leucohya, 86, 835, 844
685, 835 leucurus, Thelyphonus, 749
Larifuga, 695 Leuronychus, 97, 510, 634
Larifugella, 695 Levichelifer, 88
Larinioides, 866 Levigatocreagris, 871, 885
Lasiochernes, 336, 882 levii, Kimula, 663
Lasioerythraeus, 190 Leviola, 473
lateralis, Cynortoides, 661 Levymanus, 390
Index 963
lewisi, Cangoderces, 839, 842 littoralis, Rowlandius, 666

Liacaridae, 877 Lizamontia, 695
Libitiella, 489 Lobonychium, 426
Licneremaeidae, 877 Lohmanniidae, 876
Licnodamaeidae, 876 Lola, 92, 96, 327, 339, 899
Licornus, 489 Lomaita, 364, 648, 667
Liebstadiidae, 877 Lomanius, 422, 516
limbatus, Grosphus, 704 lombokensis, Ginosigma, 428
Lindosiella, 141 Lomeustium, 191, 475
Lindothyrus, 194, 196, 198, 742, 777, 782 longidigitatus, Geogarypus, 748
lineata, Meriosfera, 662 Longilyra, 647
lineatus, Proerginus, 662 longipalpus, Rowlandius, 666
Linga, 680 Longiperna, 491
Linnaeolpium, 756, 778 longipes, Sitticus, 866, 867
Linyphia, 866, 872 longipes, Thonius, 752
Linyphiidae, 2, 3, 82, 99, 179, 328, 339–341, Longrita, 762
348, 352, 362, 366, 367, 369, 373, 374, Lophodactylus, 88, 90, 483
379, 393, 396, 402, 404, 419, 430, 434, Lophopilio, 327, 337, 338, 361, 372
443, 452, 473, 506, 513, 514, 518, 521, Lordhowea, 777
525, 625, 647, 648, 667, 668, 678, 680, Lorisidae, 81
686, 699, 709, 724, 725, 739, 771, 776, lornani, Ibalonius, 719
839, 846, 865–867, 872, 873, 881, 883 Lossida, 467
Liobuthus, 382, 384, 386 Lossidacola, 467
Liochelidae, 145, 705, 860 Loxomphalia, 473
Liochthonius, 884 Loxoptygus, 473
Liocranidae, 179, 328, 339, 341, 362, 402, Luangnam, 419
419, 429, 473, 648, 667, 687, 698, 709, Lubinella, 751
726, 741, 763, 874, 900 Lubudia, 467
Liodidae, 876 Lucandamila, 467
Liogonyloptoides, 491 lucileae, Tityobuthus, 704
Lionneta, 717 Lucrinus, 699
Liopilio, 97 Luisarmasius, 85, 137, 646, 666, 670, 671
Liops, 493 lunatus, Ovozomus, 399, 417, 716, 721
Liparochrysis, 763 Lundulla, 427
Liphistiidae, 41, 96, 170, 171, 419, 441, 443, luquillense, Paecilaema, 661
685, 737 Lusoblothrus, 346
Liphistiomorphae, 40, 41 luteum, Homolophus, 885
Liphistius, 419, 838 lutzi, Neoscotolemon, 664
Lipophaga, 84 luzonensis, Orientzomus, 417
Lipoptena, 46 Lychas, 401, 407, 408, 432, 462, 688, 747,
Lisarea, 490 755, 756, 781
Lisna, 717 Lycosidae, 82, 179, 328, 339, 341, 348, 358,
Lisposidama, 467 369, 379, 396, 399, 402, 404, 420, 452,
Lisposoma, 693 521, 522, 525, 647, 678, 686, 700, 709,
Lisposomidae, 145, 683, 722 725, 740, 752, 759, 760, 766, 772, 840,
Lisposominae, 935 865–867, 873, 881, 900
Lissocreagris, 87, 509 Lygippulus, 465
Lissothus, 450, 451 Lygippus, 467
Listrophoroidea, 189, 193, 874, 883 lymphatica, Luxuria, 367
lithoclasica, Cynorta, 661 Lyogoniosoma, 491
Litochelifer, 88, 90, 694 lyrifer, Eukoenenia, 405
Litodamus, 761 Lyssanvira, 493
Litoralia, 489 Lystrocteisa, 776
964 Index
M Mandanapis, 776
Macaridion, 367 Mandaria, 467
Macarophaeus, 366 Mangaratiba, 491
Maccabeesa, 712 Mangareia, 773, 780
Machadoessa, 92, 467 Mangatangi, 770, 780
Macopaeus, 710 Mangua, 772, 780
Macrochelifer, 88 Maniapure, 489
Macrodampetrus, 749 manilanus, Minbosius, 428, 434, 750
macropalpus, Tyrannochelifer, 658 manilanus, Thelyphonus, 428
Macrothele, 173, 737 Manitherionyssidae, 199
Macrothelinae, 173, 685, 737 Manjala, 761
maculatus, Isometrus, 75, 368, 404, 436, 520, manonae, Tityobuthus, 704
688, 705, 714, 755 Mansuphantes, 866
maculatus, Trachelas, 334 Maoriata, 772, 780
Madacantha, 708 Maorichthonius, 769, 779
madagascariense, Xenolpium, 404 Maracandinus, 467
madagascariensis, Charinus, 707, 721 Marachernes, 89, 757, 778
madagascariensis, Erymnochelys, 78 Maratus, 763
madagascariensis, Grosphus, 704 marchingtoni, Trogloraptor, 514, 628
madagascariensis,“Podocnemis,” 78 mardua, Sarax, 414, 427
Maddisonia, 763 marginata, Assamiella, 92
madecassus, Koeneniodes, 400, 406, 424, 703, marginatus, Cynortoides, 661
714, 718 marginemaculatus, Phrynus, 513
madeira, Eukoenenia, 364 marianae, Rowlandius, 666
Madinahustium, 191 Mariblemma, 717
magna, Taphiassa, 777 maroccana, Eukoenenia, 448
magna, Titanobochica, 326, 346 Maromba, 492
Magnopholcomma, 760 Martensiellus, 414, 427
mahafaliensis, Grosphus, 704 martensii, Mesobuthus, 371, 442, 688
Mahafelytenus, 707 martensi, Stenohya, 871, 885
Mahezomus, 85, 137, 472, 716, 721 Martensolasma, 635
mahnerti major, Neobisium, 355 Marthana, 412, 425, 433
Mahura, 772, 780 Martibianta, 663, 664
Maijana, 429 martilli, Pararainbowia, 51, 67, 71
Mainosa, 760 Marwe, 91, 464–465, 476, 836, 842
Maiorerus, 92, 363, 837, 900 Masasteron, 762
Makolia, 190 Mascarenes, 711
Malaconothridae, 876 Mashimo, 474
Malarina, 761 Masirana, 93, 437, 839
Malayozodarion, 420 Maso, 866
Malcolmochthonius, 86 Masteriinae, 173, 731, 737
Malgaceros, 706 Mastidiores, 474
Malkara, 678, 760, 767 Mastigoproctinae, 139
Malkaridae, 179, 678, 731, 737, 739, 759, Mastigoproctus, 75, 79, 98, 140, 497, 498,
760, 766, 767, 901 628, 631, 637, 665, 670, 673, 896
Malkinola, 506 Mastigusa, 866
Mallinella, 883 Mastobunus, 324, 338, 347, 355
Maloides, 772, 780 Matachia, 773, 780
maltensis, Chthonius, 358 matiasis, Galibrotus, 663
Manahunca, 663, 664 Matilda, 760
Manampoka, 708 Matua, 773, 780
Manangotria, 91, 464, 706 Matundua, 699
Manaosbia, 493 Matyotia, 717
Manaosbiidae, 40, 156, 488, 493, 501, 644, mauriesi, Lustrochernes, 658
660, 663, 674 Mauritanobuthus, 448, 450, 451
Index 965
mauriticus, Hinzuanius, 712 Mesoanystis, 192

Maxanapis, 760 Mesobria, 648, 667
maxima, Hahnia, 885 Mesobuthus, 325, 382–384, 386, 395, 407,
Mayacentrum, 140, 498, 645 408, 688, 690
Mayazomus, 85, 137, 628, 631, 638, 670 Mesoceras, 749, 776
Mazarunius, 493 Mesoceratula, 413, 427, 776
Mecaphesa, 522 Mesoplophoridae, 876
Mecicobothriidae, 174, 678, 739, 900 Mesostigmata, 2, 36, 49, 67, 71, 97, 99, 184,
Mecritta, 490 185, 199, 293, 362, 374, 442, 501, 515,
Mecynargus, 866 522, 668, 687, 726, 744, 764, 767–768,
Mecysmaucheniidae, 180, 678, 739, 901 777, 779, 859, 869, 882, 884, 930, 933
Mecysmaucheniinae, 180, 739 Mesoteneriffia, 192, 362
Mediostoma, 327, 377, 378, 389, 410 Mesoteneriffiola, 192
Mediothele, 173, 900 Mesothelae, 40, 96, 170, 343, 354, 381, 391,
Meedo, 762 400, 405, 436, 439, 454, 478, 514, 630,
Meedoinae, 179, 731, 737, 741, 902 640, 653, 682, 685, 690, 692, 703, 711,
Megabulbus, 390 714, 723, 726, 727, 731, 737, 742, 744,
Megachernes, 371, 432 747, 754, 902, 932–935
Megacranaus, 490 Messatana, 489
Megadictyna, 773, 780 Mestonia, 765
Megadictynidae, 180, 771, 773 Mesudus, 773, 780
Megaloastia, 763 Metabiantes, 465
Megaloremmius, 709 Metacleobis, 84, 481
Megaoonops, 390 Metaconomma, 468, 636
Megaoryzomys, 502 Metacranaus, 490
Megapachylus, 491 Metacynorta, 636, 643
Megarhaucus, 489 Metadampetrus, 749
Megaschizominae, 85, 136, 696, 894, 935 Metadasylobus, 338
Megaschizomus, 85, 136, 472, 696, 894 Metadentobunus, 412, 422
Megateg, 761 Metadiscocyrtus, 492
Meghalaya, 91, 417 Metagagrella, 372, 398, 422, 433, 440, 442
Megistobunus, 465 Metagoniochernes, 720
Meiochernes, 515 Metagonyleptes, 491
Meiogarypus, 87, 89, 693 Metagovea, 487, 836, 844
Meioneta, 866, 883 Metagraphinotus, 492
Melaenosiinae, 180, 901 Metagyndoides, 492
Melanesotypus, 752 Metaltella, 679
Melanoblossia, 84, 693 Metalycomedes, 492
Melanoblossiidae, 74, 155, 406, 425, 683, Metalyogoniosoma, 491
693, 896 Metamermerus, 779
Melanoblossiinae, 84, 693, 935 Metamosoia, 749
Melanopella, 412, 425 Metampheres, 491
melici, Rowlandius, 666 Metanippononychus, 437
melitensis, Roncus, 358 Metanitobates, 491
Melloa, 491 Metanonychus, 511
mendi, Thonius, 752 Metanuncia, 770, 779
Mensamontia, 695 Metapachyloides, 492
Menthidae, 17, 38, 148, 380, 387, 388, 392, Metapachylus, 637
677, 685, 723, 735, 756 Metapellobunus unicolor, 664
Meridanatus, 490 Metaphalangium, 327, 337, 338, 347, 352,
Meringa, 772, 780 355, 358, 388, 451
Merostomata, 35, 36 Metaphareus, 494
Merredinia, 760 Metaplatybunus, 327, 337, 338, 372, 377,
Merucola, 467 378, 862
Mesatemnus, 88 Metapodoctis, 422
966 Index
Metarhagdopygus, 465 Micromerys, 761

Metarhaucus, 643 Micromesomma, 708
Metarthrodes, 491 Microminua, 494
Metasarcus, 491 Micromontia, 695
Metasclerosoma, 327, 344, 355 Micromygale, 647
Metasesostris, 467 Micromygalinae, 175, 900
Metasidama, 467 Micronelima, 327, 347
Metasiro, 91, 487, 510, 631 Micronimba, 468
Metasolpuga, 693 Micropachylus, 490
Metavonones, 636, 643 Micropholcomma, 177, 428, 678, 739, 760,
Metavononoides, 488 766, 767, 772, 777
Metazaleptus, 412, 425 Micropholcommatinae, 177, 760, 766,
Metazalmoxis, 716 767, 772
Metepedanulus, 426 Microsmarialla, 191, 400
Metepedanus, 426 Microsmaris, 191
Metereca, 465 Microstigmata, 696, 699, 898
Meterginus, 636, 643 Microstigmatidae, 175, 646, 647, 678, 685,
Meteusarcoides, 492 696, 699, 724, 898
Meteusarcus, 492 Microstigmatinae, 175, 900
Metibalonius, 516 Microsynotaxus, 760
Metinae, 176 Microtegeidae, 876
Metopilio, 635 Microtityus, 484, 655, 656, 672
Metopobactrus, 866 Microtrombidiidae, 875, 883–884
Metrioppiidae, 87 Microtrombidium, 884
mexicanus, Paraphrtnus, 513 Microzetidae, 876
Mexichelifer, 88, 89, 632, 836 Midgee, 761
Mexichthonius, 89, 632 Mierorchestris, 699
Mexotroglinus, 636 Migidae, 175, 678, 685, 708, 724, 738, 759,
Micalula, 649 873, 883, 901
micans, Phintella, 885 Miginae, 175
Micaria, 866 milleri, Minuides, 663
Michaelothyrus, 94, 195, 198, 718 Millomontia, 706
Michella, 636 Millotonyx, 706
Micradaeum, 695 Mimetidae, 180, 328, 339, 402, 473, 647,
Micrargus, 866 678, 686, 707, 724, 739, 776, 841,
Micrassamula, 92, 396 873, 901
Micro, 495 Mimetinae, 180
Microbiotheriidae, 95 Mimetus, 180, 739
Microbisium, 336, 659 Mimoscorpius, 85, 140, 434, 498, 645, 670
Microblossia, 84, 693 Minasteron, 762
Microbuthus, 386, 450, 462 Minax, 494
Microcaeculus, 193 Minbosius, 139, 416, 428, 434, 733, 750, 781
Microcharmidae, 145, 704, 705, 719, 722 mingrelicus caporiaccoi, Euscorpius, 335
Microchthonius, 86, 336 mingrelicus dinaricus, Euscorpius, 335
Microcina, 92, 512 Minicosa, 700
Microconomma, 466 Minicreagris, 87, 509
Microcranaus, 490 minimus, Biantes, 716
Microcreagrella, 326, 365 Minniza, 336
Microcreagrina, 365 minor, Parasiro, 355
Microcyba, 883 Minotauria, 341, 839
Microdispidae, 875 Minua, 494
Microdrassus, 717 Minuidae, 158, 488, 494, 501, 660, 663,
Microlinyphia, 883 664, 674
Micromatta, 648 minutissimus, Alioranus, 885
Index 967
minutus, Pseudomitraceras, 664 montana, Turquinia, 664

Miobunus, 765 montanus, Buthus, 346
Miopsalis, 409, 410, 425, 432 montanus, Cubazomus, 665
mirabilis, Eukoenenia, 334, 345, 354, 364, montanus, Scorpiops, 884
448, 755 montanus, Thonius, 752
Miratemnidae, 149, 735, 836 Montebello, 762
Mirochernes, 509 Montereca, 467
mirum, Leiobunum, 885 monticola, Rowlandius, 666
mirum, Myrmicotrombium, 342 monticola, Trichoribates, 868
mischi, Pseudochactas, 383 montigena, Hilaira, 867
Mischonyx, 491 mordax, Palaeokoenenia, 45, 50, 65, 69
Misumenops, 521 Morebilus, 762
mitchelli, Antillobisium, 657, 835, 843 Moreiranula, 493
Mitobates, 491 morio, Mitopus, 97, 507, 862, 863
Mitobatula, 491 Mormoopidae, 199, 501, 650, 668
Mitopernoides, 493 Moruga, 93
Mitopiella, 411, 425, 736 Moselabius, 489
Mitopus, 97, 327, 337, 338, 344, 347, 361, Mosfora, 422
372, 388, 411, 442, 862, 863, 880, 882 Mosoia, 749
Mitoscelis, 429 mossambicus, Megaschizomus, 696
Mitostoma, 50, 327, 361, 377, 378, 862 moultoni, Stenophrynus (Neocharon), 428
Mitraceras, 716 Mucrochernes, 509
Mitrager, 430 Mughiphantes, 866, 872, 881
Miturgidae, 178, 180, 328, 339, 341, 362, multidentatus, Chthonius (E.), 355
369, 390, 429, 647, 648, 678, 687, 699, Multumbo, 491
725, 731, 737, 741, 761, 762, 766, 773, Mummuciidae, 165, 481, 896, 932
776, 866, 867, 874 Mummucina, 82, 481
Miturginae, 180, 741 Mummuciona, 84, 481
Mitzoruga, 762 Mummucipes, 84, 481
mixtus, Rowlandius, 666 Mundochthonius, 336, 371, 659
modestus,“Schizomus,” 417, 734, 750 Muritaia, 772, 780
moerens, Gnaphosa, 886 Musaeus, 430
Moldoustium, 191 Muscicola, 770, 779
mollis, Nannobisium, 657 muscorum, Gnaphosa, 866, 867
Molycria, 762 Musicodamon, 85, 141, 469, 470, 626,
Momorangia, 191 682, 895
monae, Olpiolum, 657 Musola, 467
Monaga, 649 Mutadia, 92, 466
monasterii, Neobisium, 351 Mwenga, 466
monensis, Rowlandius, 666, 671 Myandra, 762
mongolica, Karschia, 371 Mycobatidae, 877
mongolicus, Galeodes, 371 Mygalarachne, 647
Monobiantes, 465, 466 Mygalomorphae, 40, 41, 96, 172, 422, 513,
monodenticulatus, Sarax, 749 646, 685, 724, 737, 742, 744, 859, 864,
monodi, Tityobuthus, 704 883, 902
monodon, Janetschekia, 867 mylonasi, Euscorpius, 335
Monomontia, 80, 695 Myobiidae, 189, 874, 883
Monoscutidae, 40, 153, 765, 770, 779, Myodalis, 142, 469
780, 898 Mypongia, 191, 763, 768
Monoscutinae, 770, 780 Myrmanu, 710
Monoscutum, 770, 780 Myrmele, 710
monstrosum, Nosomma, 403 Myrmicotrombium, 191, 342
Montadaeum, 695 Myrmochernes, 694
Montanapis, 776 Myrmochernetidae, 149, 735, 836
968 Index
Myrmopopaea, 429 Nemesiidae, 176, 327, 339, 341, 419, 478,

Myrmozercon, 49, 68, 71 646, 678, 685, 699, 724, 737, 760
Mysmenidae, 181, 328, 339, 341, 402, 403, Nemospiza, 698
473, 504, 697, 739, 751, 776, 840 Neoabrolophus, 191
Mystaria, 475 Neoancistrotus, 491
Neoaviola, 766
Neobaeorix, 466
N Neobalaustium, 191
Namandia, 766 Neobalkanella, 86, 326, 336, 337, 835,
Namanereis, 95 897, 934
Namaquarachne, 699 Neobisiidae, 17, 38, 96, 147, 326, 336, 337,
Namea, 760 343, 346, 351, 355, 360, 364, 371, 372,
Namibesia, 692 376–378, 380, 387, 388, 395, 406, 422,
Namirea, 760 432, 436, 442, 448, 463, 509, 641,
Namundra, 698 659, 671, 684, 723, 735, 834, 835,
Nanarchaea, 760 843, 871, 880
Nanepedanus, 426 Neobisium, 336, 351, 355, 371, 377, 378, 451,
Nanhermannia, 881, 884 835, 843, 861, 871, 882
Nanhermanniidae, 876 Neobuthus, 462
Nannocheliferoides, 88 Neocaeculus, 193
Nannoroncus, 86, 90, 463, 476 Neocalchas, 385
Nannowithius, 395 Neocarus, 185, 186, 188, 629, 631, 638, 649,
Nanodamon, 141 674, 842
Nanthela, 419 Neocheiridium, 505
Nanume, 717 Neocheylus, 192
Napometa, 369 Neochthonius, 86, 509
Napostygnus, 490 Neocleobis, 84
naranjo, Rowlandius, 666, 671 Neocoryphus, 466
Narcaeus, 430 Neocranaus, 490
nasutus, Tetrabalius, 428 Neocynorta, 489
Natgeogia, 93, 776 Neocynortina, 644
Nauhea, 773, 780 Neodiplothele, 93
naupliensis, Euscorpius, 335 Neoditha, 86, 90, 483
naxos, Eukoenenia, 334 Neogagrella, 412, 425
Neassamia, 92, 412 Neogarypus, 87
Neato, 762 Neogovea, 487, 632, 836, 844
Nebo, 383, 385, 386, 450 Neogoveidae, 2, 39, 74, 150, 152, 454, 465,
Necturus, 78 487, 510, 637, 684, 722, 836, 844
Negreaella, 663, 664 Neogrosphus, 704, 705
negreai, Rowlandius, 666, 671 Neomyro, 773, 780
Nelima, 97, 327, 355, 361, 372, 377, 378, 411, Neonothippus, 749
422, 440, 442, 634 Neonuncia, 524, 770, 779
Nelsoninus, 770, 779 Neoorsa, 664
Nemaspela, 327, 377, 378 Neopachyloides, 490
Nemastoma, 48, 97, 155, 327, 344, 360, 361, Neopachylus, 492
372, 377, 378, 389, 451, 843, 862 Neophanolophus, 189
Nemastomatidae, 40, 48, 50, 97, 154, 327, Neophrynus, 142
337, 339, 344, 347, 352, 356, 360, Neopilio, 695
361, 372, 377, 378, 381, 388, 389, Neopilionidae, 40, 80, 153, 488, 525, 677,
410, 436, 437, 451, 511, 635, 641, 678, 684, 695, 723, 736, 742, 757, 765,
673, 684, 736, 743, 837, 862, 864, 770, 779, 780, 898
871, 880, 899 Neopodoctis, 401
Nemastomella, 327, 451 Neoprolochus, 429
Nemastygnus, 491 Neopseudogarypus, 86, 765, 778
Index 969
Neopurcellia, 91, 770, 779 nivalis, Cryphoeca, 867

Neoramia, 772, 780 niveosignatus, Sitticus, 885
Neorhaucus, 489 Nkogoa, 466
Neoricinulei, 143 nohbecanus, Neocarus, 638
Neosadocus, 491 Nomana, 780
Neoscorpionina, 37, 66, 69 Nomindra, 762
Neoscotolemon, 664 Nomoclastes, 494
Neosmaris, 190, 768, 773, 781 Nomoclastinae, 494, 501
Neosparassus, 761 nondimorphicus, Sclerobunus, 511
Neostorena, 762 nordenskioeldi, Euphalangium, 863, 880, 885
Neoteneriffiola, 192 nordmanni, Calchas, 376, 378
Neothyridae, 194, 195, 198, 501 Nortanapis, 760
Neothyrus, 194, 195, 197, 198, 501 Nostera, 762
neotropicus, Roncus, 351 Notasteron, 762
Neowadotes, 647, 667 Nothridae, 869, 876
Neozomus, 85, 399, 417, 418 Nothrus, 881
Nepalobisium, 87, 395 Notiodrassus, 773, 780
Nepalsia, 92, 396 Notomatachia, 773, 780
Nepalsioides, 92, 396 Notonuncia, 765
Nephilinae, 177 Notozomus, 85, 137, 734, 758, 759, 778
Nephrochirus, 698 Notsodipus, 762
Nesiergus, 717 Novanapis, 771, 780
Nesopachylus, 644 novemtuberculatus, Sitalcicus, 713
Nesoryzomys, 502 Novobisium, 87, 509
Nesowithius, 720 Novodamus, 761
Nesticidae, 181, 328, 339, 341, 366, 402, 739, Nucina, 765
840, 844, 846, 865, 873 Nudochernes, 388
newbritainensis, Sarax, 749 Nuisiana, 773, 780
nicaraguensis, Neocarus, 649 Nukuhiva, 518
Niceforoiellus, 494 Nuliodon, 762
nicobarensis, Catatemnus, 404 Numipedia, 92, 466
nicobarensis, Garypus, 404 Nuncioides, 765
Nicodamidae, 180, 181, 428, 741, 759, Nuttalliella, 94, 198
761, 767 Nuttalliellidae, 94, 687, 700, 726, 935
Nicodamus, 761 Nyctimus, 430
nidificator, Paratemnoides, 657
nidiformis, Haemogamasus, 885
niger, Hammenius, 752 O
nigra, Pardosa, 867 Obatala, 697
nigrescens, Eusimania, 334 obscura, Meioneta, 885
nigrimana, Zora, 866, 867 obscurissimus, Araneus, 885
Nihoa, 93 obscurus dorsalis, Metacynortoides, 661
Nimbadus, 468 obscurus obscurus, Metacynortoides, 661
Niphocepheidae, 868, 869, 876 occidentalis, Vulsor, 80
Nippobodidae, 876, 877 occidentalis, Zalmoxis, 713
Nipponogarypus, 436, 440 Occitanobisium, 87, 360
Nippononychidae, 40, 156 occitanus, Buthus, 346, 358, 448
Nippononychus, 437 oceanusindicus, Anagarypus, 404
Nipponopsalididae, 40, 154, 410, 436, 437, ocellatus, Geogarypus, 715
440, 442, 684 ochropus, Chiromachus, 712, 714
Nipponopsalis, 410, 437, 442 Ochyroceratidae, 38, 181, 402, 647, 666,
Niquitaia, 495 685, 717, 724, 750, 839, 844, 845
nitidissimus, Leiothyrus, 752 Ocoita, 488
nivalis baloghi, Niphocepheus, 868 Octanapis, 760
970 Index
oculatus, Javazomus, 417, 428 Opilio, 327, 337, 338, 344, 355, 358, 360–361,
Ocyllus, 420 372, 375, 377, 378, 389, 396, 411, 436,
Odiellus, 338, 347, 352, 361, 372, 442, 862, 863, 871, 880, 885
377, 378, 862 Opilioacaridae, 186, 631, 687, 726, 744
Odomasta, 766 Opilioacariformes, 186
Odontobunus, 465 Opilioacarus, 49, 51, 67, 70, 185–187, 328,
Odontobuthus, 384, 386, 407, 688 342, 344, 357, 452, 682
Odontonuncia, 765 Opiliomorphae, 36
Odontosiro, 91, 347 Opiliones, 2–4, 9–12, 15, 18, 22, 25, 26, 28,
Odonturus, 462 29, 32, 35, 36, 38–40, 45, 47, 48, 50,
Oecobiidae, 181, 328, 339, 362, 473, 647, 66, 70, 74, 77, 79, 81–83, 91, 96–97,
678, 686, 697, 724, 739 149–170, 186, 319, 323,, 333, 337–339,
Oecobius, 739 343, 346, 347, 351, 352, 355, 358, 360,
oedipus, Chileogovea, 488 362, 364, 365, 372, 374–377, 381, 388,
Oedothorax, 866, 872 392, 395, 397, 398, 401, 405, 409–422,
oglasae, Euscorpius, 95, 354 425, 431, 432, 436, 439, 440, 442, 451,
Ogovea, 464–465, 476 454, 464, 476, 478, 487, 501, 503, 505,
Ogoveidae, 39, 91, 150, 465, 476, 509, 515, 516, 521, 524, 625–628, 630,
684, 722 632, 637, 640, 641, 660, 664, 673–674,
Ohvida, 648, 667 677, 678, 680, 683, 692, 694, 695, 703,
Oiclus, 84, 655, 656 706, 711–722, 726–728, 735, 742–744,
ojastii, Neocarus, 501 747, 748, 755, 757, 765, 769, 770, 775,
okadai, Sabacon, 372 778, 779, 781, 782, 836, 842, 844–846,
olgae, Grosphus, 704 859, 862–872, 880, 882, 884–886,
Olgania, 766 898–900, 931–935
Oligacanthophrynus, 141 Opisthacanthus, 460, 461, 463, 485, 486, 642,
Oligolophus, 97, 327, 337 655, 656, 672, 705
Oligovonones, 489 Opisthoplites, 490
Olpiidae, 38, 148, 326, 336, 343, 346, 351, Opisthopristis, 489
364, 367, 368, 371, 377, 378, 380, Opistoncana, 752
387, 388, 392, 395, 398, 401, 406, Opistophthalmus, 463
422, 425, 432, 436, 440, 451, 463, 483, Oppia, 881, 884
503, 515, 521, 641, 657, 659, 671, 672, Oppiidae, 526, 869, 877
677, 685, 705, 715, 720, 723, 747, 756, Oppodoctis, 433
769, 770, 777, 778, 781, 782, 834, 836, oppositum, Beierobisium, 677, 679
861, 871 Opserythraeus, 190, 475
Olpium, 336, 401, 432 Oramia, 772, 780
Ommatauxesis, 766 Oramiella, 772, 780
omnisuperstes, Euophrys, 885 Orchestrella, 699
Omucutia, 709 orealis, Pardosa, 885
Oncodamus, 761 Oreo, 762
Oncopodidae, 156, 396, 399, 412, 414, 426, Oreocyba, 883
427, 433, 737, 862, 864, 871 Oreonetides, 866, 867
Onychothelyphonus, 46, 513 Orepukia, 772, 780
Oonopidae, 181, 328, 339, 341, 362, 373, orghidani, Cubazomus, 665, 671
390, 393, 396, 402, 419, 429, 473, 647, orghidani, Eukoenenia, 653,
678, 686, 698, 717, 724, 738, 776, 839, orghidani, Neocarus, 668, 843
844, 845, 873 orghidani, Troglocubazomus, 666
Oonopinae, 181 Oribatella, 881
Oparba, 449, 457 Oribatellidae, 869, 877
Oparbella, 449, 456, 457 Oribatulidae, 869, 877
Opconomma, 468 Oribotritiidae, 876
Opconommula, 468 orientale, Conomma, 96
operculatus, Phrynus, 513 Orientzomus, 137, 417, 418, 434, 440, 515, 734
Index 971
Oripodidae, 877 Pachylicus, 637, 644, 664

Orituco, 495 Pachylinae, 492, 677, 678
Ornithoctonus, 419 Pachylomerinae, 172, 738
Orobothriurus, 887 Pachylospeleinae, 492
Orobunus, 427, 749 Pachylospeleus, 492
Orongia, 772, 780 Pachylusius, 492
Orsa, 662, 664 Pacifantistea, 375
Orsimonia, 466 Pacificana, 773, 780
Orsolobidae, 181, 680, 686, 697, 724, 731, Padangcola, 426
737, 739, 766, 772, 780, 873, 901 Paecilaema, 636, 643, 661
Orsonwelles, 521 Paecilaemana, 643
Orstom, 93, 776 Paedobisium, 87
Orthochiroides, 386, 462 Pagiopalus, 521, 522
Orthochirus, 382–384, 386, 392, 407, 408, Pahora, 772, 780
450, 462, 687, 688 Pahorinae, 182, 740, 767, 771, 780
Orthognatha, 41, 628, 638 Pahoroides, 772, 780
Ortholasma, 511, 635 Paisochelifer, 88, 509
Ortholasmatinae, 410, 511, 635, 643, Pakawops, 422
673, 736 Pakeha, 772, 780
Orthosterni, 37, 46, 65, 68, 69 Palaeogrosphus, 705
Orthothelae, 172, 685, 724 Palaeoncopus, 92, 414, 427
Ortonia, 494 Palaeoricinulei, 143
Orvilleus, 649 Palcabius, 488
Oryzomys, 502 Palcapachylus, 492
Ossinissa, 366 Palcares, 491
Ostanes, 475 palenquensis, Negreaella, 663
Ostearius, 866 Paleoamblypygi, 933
Otagoa, 773, 780 pallida, Ammotrechella, 653, 654
Othoes, 449, 456 Palmanella, 466
Otiothopinae, 181 palpale, Himalphalangium, 863, 872, 885
Otocepheidae, 877 Palpatores, 38, 39, 149, 327, 338, 346, 347,
Otuquisa, 493 352, 361, 372, 432, 509, 627, 632, 634,
ovala, Paraconomma, 664 637, 706, 720, 836, 859, 862–864, 871,
Ovalia, 495 880, 882
ovalis, Holozoster, 716 Palpigradi, 1, 3, 35, 36, 45, 50, 65, 69, 83, 98,
ovatus, Ixodes, 885 135, 319, 323, 334, 343, 345, 351, 354,
ovchinnikovi, Pseudochactas, 383 358, 359, 363, 364, 368, 376, 380, 381,
Ovozomus, 85, 137, 418, 472, 734, 758, 397, 398, 405, 406, 421, 424, 431, 436,
759, 778 442, 448, 454, 455, 475, 478, 481, 508,
Oxapampeus, 491 515, 520, 625, 626, 631, 640, 653, 669,
Oxyopidae, 9, 181, 328, 339, 362, 399, 402, 676, 681, 683, 692, 703, 711, 712, 714,
404, 429, 647, 678, 686, 709, 725, 740, 718, 722, 727, 732, 743, 747, 754, 755,
761, 873, 900 764, 778, 779, 781, 833, 843, 860,
Ozarchaea, 760 893–895, 931–934
Ozicrypta, 93 Palpimanidae, 9, 181, 328, 339, 341, 362,
Ozyptila, 867 390, 393, 402, 474, 678, 686, 698, 724,
739, 873, 901
Palpimaninae, 181
P palustris, Pardosa, 867
Pacal, 85, 137, 628, 631, 638, 843 Palystella, 699
pacei, Neobisium, 355 pamiricus, Yllenus, 885
Pachkutej, 887 Panalus, 490
Pachychelifer, 88, 509 panamensis, Caribeacarus, 649, 668
Pachylibunus, 492 Panaquire, 495
972 Index
Panaretella, 699 Paramatachia, 761

panayensis, Thelyphonoides, 434 Paramenthus, 87, 90, 388
Panchaetes, 185–187, 475, 476 Paramesoceras, 749, 781
Pandinus, 386, 460–463 Paramicrocranaus, 493
Panjange, 761 Paramicromerys, 709
Panoa, 773, 780 Paramigas, 708
Panopiliops, 644 Paramiginae, 175, 708
panpema, Homolophus, 885 Paraminuella, 495
Pantopsalis, 524, 770, 780 Paramiopsalis, 91, 347, 898
Papakula, 751 Paramitraceras, 636, 644
Papuaneon, 752 Paramo, 477, 495, 923
Papuchelifer, 88, 89, 748, 781 Paramontia, 695
Parabupares, 426 Parampherus, 491
Parabuthus, 386, 450, 462, 693 Paramyro, 772, 780
Paracarus, 49, 51, 67, 70, 185–187, 381 Paranaleptes, 492
Paracenobiopelma, 93 Paranapis, 771, 780
Paracharon, 85, 95, 143, 469, 470, 682, Paranelima, 510, 634
895, 934 Paranemastoma, 327, 339, 361, 372, 377, 378,
Paracharontidae, 32, 85, 95, 143, 469, 682, 389, 837, 862
683, 895 Paranonychinae, 437, 511, 690
Paracheiridium, 705, 720 Paranonychus, 511
Paracladycnis, 708 Parapalystes, 699
Paracrobunus, 413, 426, 433 Paraphalangodus, 492
Paracynorta, 643 Paraphanolophus, 190
Paradaeum, 695 Paraphareus, 494
Paradictyna, 773, 780 Paraphilaeus, 763
Paradiscocyrtus, 492 Paraphrynus, 37, 98, 142, 495, 496, 513, 628,
Paradonea, 698 631, 645, 665, 669, 843, 894
paradoxus, Haplochernes hebridicus, 748 Paraplatybunus, 327
paradoxus, Hebridochernes, 748 Paraplexippus, 649, 666, 668
Paradraculoides, 137, 734, 758, 759, 778 Parapostenus, 699
Paraembolides, 760 Paraprosontes, 492
Paragagrella, 412, 425, 433 Parapua, 780
Paragaleodes, 449, 455 Parapurcellia, 91, 464, 694
Paragaleodiscus, 83 Pararchaea, 771, 780
Paragarypus, 87, 705, 719 Pararchaeidae, 179, 678, 679, 739, 759, 760,
Parageaya, 634 767, 771, 776
Parageralinura, 65 Pararcheidae, 780
Paragoniochernes, 87, 89, 694 Pararezendesius, 493
Paragovia, 476 Pararoncus, 87, 91, 436
Paragryne, 488 Parasaitis, 649, 668
Paraharmochirus, 752 Parasamoa, 516
Paraheliophanus, 369 Parascleropilio, 366
Parahelpis, 763 Parasesostris, 466
Parahyidae, 38, 147, 406, 425, 735, Parasiro, 347, 355
775, 933 Parasitiformes, 36, 45, 67, 71, 94, 97, 99,
Parakalummidae, 877 184, 194, 293, 319, 342, 369, 393, 421,
Parakodaika, 466 422, 452, 501, 503, 504, 514, 522, 527,
Paralampona, 762 638, 640, 649, 668, 679, 700, 710–711,
Paraliochthonius, 336, 365 744, 764, 767–768, 773, 777, 842, 859,
Parallowithius, 87, 90, 694 869, 870, 878, 879, 882, 884, 885, 931
Paralobus, 772, 780 Parasmodix, 475
Paralola, 25, 92, 96, 327, 339, 837, 899 Parastalita, 341
Paraluederwaldtia, 492 Paratemnoides, 432, 657, 659, 672, 720, 748
Index 973
Parateneriffia, 192 Pelecopsis, 866, 883

Parathiodina, 649, 668 Pelinobius, 473
Paratricommatus, 493 Pelitnus, 433
Paratropididae, 176, 648, 667 Pellenes, 867
Paratropidinae, 176 Pellobunus, 636, 644, 662
Parattahia, 765 Peltonychia, 440, 838
Paratupua, 760 Penangolyna, 419
Paravachonium, 86, 844 Penestomidae, 698
Paravima, 488 Penestominae, 178
Paravoka, 780 Penestomus, 427, 698
Paravonones, 636, 643 Pentacranaus, 493
Parazaona, 505, 658, 660 penta, Liocheles, 747
Parazilia, 474 Pentanychidae, 159, 512
Pardosa, 866, 867, 872, 881, 885 Pentanychus, 512
Parepedanulus, 426 pentapeltis, Hubbardia, 138
Pareusarcus, 492 peradenyiensis, Ovozomus, 417
Parhypochthoniidae, 876 Periegopidae, 182, 739, 759
Parkocranaus, 490 Perilla, 419
Parmatergus, 708 Peripa, 490
Paro, 518 Perissopmeros, 760, 767
Parodiellus, 97, 862 Peritraeus, 403
Paroecobius, 697 Peromona, 716
Parogovia, 464–465, 487 perplexus,“Schizomus,” 401, 417
Paroligolophus, 327, 337, 361 Perthacantha, 757, 779
Paropilio, 352 pescotti, Charinus, 749
parriloi, Tityobuthus, 704 peteloti, “Schizomus,” 417, 860
parvulus, Biantes, 716 petraeum, Theridion, 867
parvus, Hummelinckiolus, 662 petrensis, Talavera, 348, 866, 867
parvus, Mesothelyphonus, 65 petrobia, Gnaphosa, 866, 867
parvus, Parachelifer, 658 Petrobunidae, 40, 156, 412, 413
parvus, Sclerobunus, 511 Pettalidae, 39, 74, 150, 152, 153, 400, 401,
Pasiasula, 475 410, 488, 684, 694, 706, 722, 736, 742,
Passalozetidae, 877 757, 770, 778, 779, 837, 842, 846, 898
Passosa, 492 Pettalus, 91, 400, 401, 410
pastor pickardi, Cryphoeca, 867 Petterchernes, 89
Patelliella, 777 Phalangiacarus, 93, 185–187, 475, 476, 682
Patelliellini, 777 Phalangiidae, 2, 40, 48, 50, 97, 153, 327, 337,
Patri, 717 338, 344, 347, 352, 355, 358, 360, 361,
patrizii, Eukoenenia, 354 365, 366, 372, 375, 377, 378, 381, 388,
Paulianyx, 706 389, 395, 398, 411, 422, 425, 436, 440,
Pebasia, 489 442, 451, 465, 488, 510, 684, 695, 723,
Pechota, 92, 412 736, 743, 838, 862–864, 871, 880, 898
pecki, Cynortina, 843 Phalangioidea, 39, 40, 80, 153, 757
pecki, Ethobunus, 664 Phalangiotarbida, 45
pecki, Troglobochica, 657, 835, 843 Phalangium, 97, 352, 862, 885
peckorum, Rowlandius, 666, 671 Phalangochilus, 491
Pectinibuthus, 382, 386 Phalangodes, 512, 663
Pedalocreagris, 87 Phalangodidae, 40, 48, 156, 327, 337, 339,
Pedinopistha, 522 344, 347, 352, 356, 358, 360, 381, 389,
Pedinopisthinae, 182, 522 396, 412, 413, 422, 436, 437, 440, 451,
Pedipalpides, 36 452, 512, 538, 636, 637, 660, 663, 674,
Peeto, 762 684, 715, 720, 757, 837, 844, 862, 863,
Peladoius, 490 871, 899
Pelechucia, 489 Phalangodinella, 495
974 Index
Phalangodoidea, 39, 40 Pichitus, 492

Phalangoduna, 644 Pickeliana, 494
Phalangomma, 512 picturatus, Araeoncus, 885
Phanerobunus, 765 Picuchenops, 677
Phanolophus, 191 pidgeoni, Pseudouroplectus, 705
Phanotea, 697 Piesocalus, 430
Phareicranaus, 644 Pijiguaia, 495
Phaulostyleus, 710 pijol, Mayacentrum, 645, 670
Phenasteron, 762 pileti, Bamazomus, 417
Phenopelopidae, 877 Pilochelifer, 88, 712, 720
Pherania, 492 pilorides, Capromys, 668
Philodromidae, 9, 182, 328, 339, 341, 362, Pilosa, 495
402, 514, 521, 522, 647, 678, 687, 700, Pilosoma, 188
725, 741, 751, 781, 867, 874, 900 Pimoidae, 182, 739, 840
Philodrominae, 182 Pionothele, 699
Philodromus, 867 Pirassunungoleptes, 495
phipsoni, Phrynichus, 399 Pirunipygus, 492
phipsoni, Rhagodes, 402 Pitonga, 761
Pholcidae, 182, 327, 339, 341, 362, 366, pityusensis, Allochernes, 351
402–404, 504, 647, 648, 667, 678, Plagiobothrus, 91, 402
685, 709, 724, 738, 761, 839, 844–846, planicauda, Antillostenochrus, 665
873, 900 Planiphalangodus, 492
Pholcipes, 717 Planophareus, 494
Pholeochthonius, 777 planus, Vonones, 662
Phonophilus, 452 Plastonomus, 709
Phormingochilus, 429 platensis, Neocarus, 501, 679
Phorochelifer, 88, 509 Plateremaeidae, 876
Phoxobunus, 765 Platnicknia, 648, 668
Phricotelus, 403 Platorish, 762
Phrurolithidae, 182, 402, 429, 778 Platybessobius, 338, 372, 377, 378
Phryganoporus, 761 Platybunus, 337, 372, 389, 862
Phrynichidae, 37, 79, 141, 381, 387, 399, Platygyndes, 492
452, 469, 478, 495, 682, 695, 713, 716, Platylampona, 762
722, 841, 895, 933, 934 platypedibus, Siro, 48
Phrynichinae, 142 Plectophanes, 773, 780
Phrynichodamon, 142, 470, 695, 895 Plectoptilus, 429
Phrynichosarax, 141, 733 Plectreuridae, 182
Phrynichus, 142, 387, 392, 399, 415, 469, 470, Plesioballarra, 757, 779
695, 841, 894 Plesiobuthus, 688
Phrynidae, 37, 75, 98, 142, 427, 428, 495, Plesiothele, 765
496, 513, 628, 637, 640, 645, 665, 669, Plesiothelinae, 765, 767, 778, 782
732, 742, 842, 843, 895, 934 Plesiowithius, 87, 451
Phryninae, 142, 637 Plethodontidae, 78
Phrynus, 75, 78, 79, 98, 142, 415, 428, 495, Pleurotus, 718
496, 513, 628, 631, 637, 665, 669, 732, Plicatiductus, 430
841–844, 894, 895, 934, 935 plumipes, Tenuiphantes, 885
Phthiracaridae, 869, 876 Plutonodomus, 474
Phyllarachne, 430 Plynnon, 429
Phylostomatidae, 199 Poachelas, 697
Physoglenidae, 182 poaensis, Cynorta, 661
Phyxelididae, 182, 429, 474, 687, 699, 708, Poaka, 772, 780
725, 741 Poala, 636
Pianoa, 771, 780 Poassa, 644
Piassagera, 491 Podacaridae, 97
Index 975
Podaucheniellus, 466 Prasmiola, 770, 779

Podocnemididae, 78 pratyushi, Indiacarus, 186, 400
Podoctidae, 40, 156, 396, 398, 399, 401, pretneri, Eukoenenia, 334
412–414, 422, 426, 427, 433, 436, 437, Prida, 717
440, 465, 516, 660, 663, 674, 684, 706, Prima, 707
712, 713, 715, 716, 720, 721, 723, 737, Primerigonina, 648
748, 749, 757, 781, 782, 862, 864 primibiconourus, Rowlandius, 666, 671
Podoctinus, 749, 781 Prinerigone, 866
Podoctis, 413, 427 Prionostemma, 634, 641, 673
Podoctomma, 427 Priperia, 521
Podoctops, 427 pristinus, Paracarus, 49, 51, 67, 70, 186, 391
Podopterotegaeidae, 876 Pristobunus, 770, 779
Podosmaridia, 190 Pristocnemis, 491
Podothrombium, 883 Prithinae, 731, 737, 738
Poecilaemula, 643 Proampheres, 491
Poecilarcys, 452 proboscideus, Labochirus, 401
Poecilipta, 763 Procaeculus, 193
Poecilocranaus, 493 Procambridgea, 761
Poeciloneta, 866 procerus,“Schizomus,” 417
Poecilosophus, 493 Proconomma, 465, 468
Poecilothomisus, 763 Proctobunoides, 491
Poessa, 710 Procydrela, 697
poeyi, Torreana, 663 Prodidomidae, 182, 328, 339, 474, 647,
Pogonowithius, 87, 90, 463, 476 649, 668, 678, 687, 698, 725, 739,
pohakuloae, Vulcanochthonius, 521 762, 840, 874
Pokennips, 178, 647, 667 Progonyleptoidellinae, 493
Poliocheridae, 508 Progonyleptoidellus, 493
Poliocheroidea, 143 Progonyleptoides, 491
Pollux, 191 Proholozoster, 749, 781
Polyacanthoprocta, 492 Prokoenenia, 135, 424, 718, 732, 893, 894
Polycoryphus, 695 Prokoeneniidae, 135, 405, 424, 676, 703,
Polylepis, 477, 886 718, 732, 743, 893
Pomerantzia, 193 Prolacurbs, 466
Pomerantziidae, 193 Promitobates, 491
Pomerantzioidea, 193 Promurricia, 403
Pongolania, 699 Proschizomus, 65
ponsi, Chthonius (E.), 351 Proscotolemon, 92, 437, 440, 899
Popassamia, 92, 412 Prosolpuga, 84, 693
Popcornella, 649, 668 Prosontes, 492
popovi, Euscorpius, 333, 335 Prostigmata, 2, 5, 97, 99, 189, 342, 362, 369,
Porioides, 773 400, 420, 475, 515, 522, 638, 692, 700,
Porius, 752, 763 744, 763, 768, 773, 776, 781, 859, 868,
Poroides, 780 874, 875, 883
Porotaka, 772, 780 Prostignidius, 490
Porrhomma, 839, 866, 872 Prostygnellus, 490
Porrhothele, 767, 771, 780 Prostygninae, 490, 845
portoricanus, Yunquenus, 663 Prostygnus, 490
portoricensis, Stenochrus, 138, 324, 345, 498, Proteidae, 78
503, 513, 645, 666, 670, 671 Protelyphonus, 65
poseidonis, Chthonius (E.), 355 Proteus, 78
Pothaeus, 420 Protimesius, 494
Pounamuella, 772, 780 Protochthonius, 67, 71
Praelibitia, 489 Protoiurus, 385
Prasma, 770, 779 Protolophidae, 153, 510, 635, 898
976 Index
Protolophus, 510, 635 Pseudoporrhopis, 709

Protoneobisium, 87, 326, 336, 337 Pseudopsellonus, 751, 781
Protoschizomidae, 136, 513, 628, 638, 841, Pseudopulcrolia, 491
843, 894 Pseudorhacochelifer, 88, 367
Protoschizomus, 136, 628, 631, 638, Pseudoscaphiella, 698
843, 894 Pseudoscorpiones, 2, 3, 16, 17, 27, 35–37, 45,
Protowithius, 87, 505 47, 66, 74, 80, 86, 95, 146–149, 319,
Protus, 489 326, 333, 336, 343, 346, 351, 353, 358,
proximus, Tyrannochthonius, 655 360, 362–364, 367, 371, 374, 376, 378,
Psalenoba, 770, 779 380, 381, 387, 388, 392, 395, 397, 398,
Psammobuthus, 384, 687 401, 404–406, 421, 422, 425, 431, 432,
Psammodnon, 697 436, 440, 442, 451, 454, 463, 476, 478,
Psammorygma, 697 493, 503, 504, 508, 515, 520, 524, 625,
Psathyropus, 372, 411, 412, 422 627, 632, 640, 641, 655, 659, 671, 676,
Psechridae, 182, 402, 404, 686, 725, 740, 679, 684, 692, 693, 703, 705, 711, 712,
761, 766, 772 714, 719, 720, 723, 726, 727, 734, 742,
Pselaphochernes, 336, 351, 355, 365, 371 743, 747, 748, 755, 756, 765, 769, 775,
Pselloninae, 182 777–779, 781, 782, 834, 842, 844, 845,
Pseudamyciaea, 430 859, 861, 870, 880, 882, 884–886, 897,
Pseudasteron, 762 933–935
Pseudatemnus, 87, 90, 694 pseudostrigilata, Pardosa, 334
Pseudauxinum, 697 Pseudosynagelides, 763
Pseudoacrographinotus, 492 Pseudotegenaria, 452
Pseudoblothrus, 96, 326, 365, 835 Pseudoteyl, 760
Pseudocellus, 143, 627, 631, 640, 654, 669 Pseudotrogulus, 491
Pseudochactas, 84, 95, 383, 406, 687, 834 Pseudotyrannochthoniidae, 37, 146, 372,
Pseudochactidae, 95, 145, 380, 382, 383, 387, 398, 401, 442, 684, 723, 765,
405, 406, 408, 683, 734, 834, 897, 935 834, 835
Pseudocheylus, 192 Pseudouroplectus, 705
Pseudochiridiidae, 17, 38, 148, 395, 425, pseustes, Ammotrechella, 641, 653, 654
432, 463, 657, 659, 685, 705, 715, 720, pshartica, Parasyrisca, 885
723, 933 Psoroptoidea, 193
Pseudochiridium, 404, 432, 657, 659, 720 Ptychosoma, 347, 352, 452, 899
Pseudochthonius, 655, 659, 672, 835 Pua, 772, 780
Pseudocleobis, 676 pubescens, Sitticus, 885
Pseudoepedanus, 413, 426 Puerilia, 489
pseudofuscipalpis, Agyneta, 885 puertoricense cavicola, Ideobisium, 656
Pseudogagrella, 411, 412, 422, 440 puertoricense, Ideobisium, 566
Pseudogarypidae, 509, 735, 742, 743, 765, puertoricensis, Americhernes, 658
835, 861 puertoricensis, Olpiolum, 657
Pseudogarypus, 86, 509, 835, 861 pugnator, Mimoscorpius, 645, 670
Pseudogyndesoides, 492 Pugnochelifer, 88, 509
Pseudohomalenotus, 411 Pukakia, 773
Pseudohostus, 76 Pulchandrus, 466
1, 776 pulchra, Mitraceras, 716
Pseudolampona, 762 Pumbaraius, 399
Pseudolamponinae, 179, 739 pumilus, Caribochernes, 658
Pseudolissothus, 450 Puna, 489
Pseudomelanopa, 412, 422 Punagraphinotus, 492
Pseudomenthus, 392 Punanus, 426
Pseudominua, 494 punctatus, Phalacrocorax, 768, 773
Pseudopachylus, 493 Pungalina, 763
Pseudophalangodes, 493 Punrunata, 492
Pseudopilanus, 505 pupukeanus, Tyrannochthonius, 521, 845
Index 977
Purcellia, 91, 464, 694, 837, 842 Raphignathidae, 875

Purcelliana, 698 raptator, Paraphrynus, 513
Puria, 92 raptor, Arctosa, 885
pusillus, Anelasmocephalus, 356 Rapua, 773, 780
Pussardia, 191, 763, 768 Rarahu, 519
Pycnaxis, 434 Rastellus, 698
Pycnoepisinus, 473 Ravilops, 79, 85, 139, 497, 498, 665, 670
Pyenganella, 765 Rebilus, 762
pygmaeus, Ideoblothrus, 657 Recifesius, 490
Pygmephoridae, 868, 875 recuerdo, Rowlandius, 666, 671
Pygocynorta, 489 reddelli, Nothoaspis, 638, 844
Pyramidopidae, 40, 80, 96, 156, 365, 465, Reddellzomus, 137, 646, 665, 666,
644, 674, 684, 723, 736, 837, 899, 670, 671
900, 935 Redonopistha, 521
pyrenaella, Eukoenenia, 345 Reichlingeria, 647
pyrenaica, Eukoenenia, 345 Reimoserius, 643
Pyza, 327, 389 Reinickella, 430
Rejanellus, 649, 668
remota, Americovibone, 80
Q remota, Erigone, 867
quadricristatus, Erginulus, 635, 643, 661 remyi, Chthonius (E.), 355
quadridentatus, Carios, 768 remyi, Eukoenenia, 334
quadrimaculata, Cynortellana, 661 Remyus, 706
quadrispina, Tavoybia, 92 renschi, Chajnus, 428
quadrispinosus, Cynortoides, 661 restrictus, Mughiphantes, 885
Quadropia, 884 Retropedis, 495
Quadroppiidae, 877 reunionis, Gagrella, 713
Queenslanapis, 760 Reventula, 662, 664
Queenvic, 762 reyesi, Rowlandius, 666
Quemusia, 761 Rezendezius, 493
Questo, 762 Rhacochelifer, 336, 365, 451
Questocrypta, 93, 776 Rhacocnemis, 718
quibijana, Cynorta, 661 Rhaebobates, 752
Quindina, 490 Rhagidiidae, 844, 868, 875
quinquesignata, Cynortula, 661 Rhagodalma, 456, 458
Rhagodes, 376, 377, 394, 402, 449,
455, 457
R Rhagodeya, 449, 456
Racata, 430 Rhagodidae, 170, 376, 377, 380, 381, 394,
racovitzai, Troglobisium, 326, 346 398, 402, 448, 449, 455–458, 683,
Rafalskia, 327, 337, 338, 389, 862 692, 896
rageaui, Charletonia, 768, 776 Rhagodippa, 455, 458
Rahavavy, 708 Rhagodira, 449
Rainbowia, 190, 763, 768 Rhagoditta, 449, 455
Rakaia, 91, 770, 779, 846 Rhampsinitus, 465
ralik, Orientzomus, 417, 515 Rhaucoides, 489
Ramblinus, 327, 337, 366 Rhaucus, 489
Ramonus, 488 Rhinoblemma, 518
ramosi, Cokendolpherius, 666 Rhinodromus, 193
ramosi, Rowlandius, 666 Rhion, 402
Ramsayella, 190, 768, 773, 781 Rhioxyna, 492
Randilea, 465, 466 Rhipicephalus, 710, 870, 884
randoi, Maiorerus, 96, 366, 837, 900 Rhodera, 340
Ransonia, 708 Rhodinae, 178
978 Index
Rhombonotus, 763 S
Rhondes, 776 Sabacon, 48, 97, 327, 347, 361, 372, 396, 410,
Rhopalochelifer, 88, 90, 694 437, 442, 511, 837, 862, 863, 871, 872
Rhopalocranellus, 493 Sabaconidae, 40, 97, 154, 327, 347, 360, 361,
Rhopalurus, 84, 484, 654–656, 672, 704 372, 396, 410, 436, 437, 442, 511, 684,
Rhynchobunus, 765 837, 862–864, 871, 885, 899
Rhysotritia, 884 Sabahya, 429
Rhyssoleptoneta, 93 Sabanilla, 488
Ricinoides, 143, 458, 459, 475, 681 Sabinebuthus, 462
Ricinoididae, 143, 482, 640, 683, 843 Saccarella, 327, 344
Ricinoidoidea, 143 Saccodomus, 763
Ricinulei, 2, 3, 35, 36, 45, 46, 50, 65, 69, 76, sachalinensis, Palaeoerythracarus, 67, 70
77, 143, 319, 320, 323, 333, 343, 354, Saetigerocreagris, 87, 509
364, 368, 371, 381, 397, 400, 421, 424, Saharobuthus, 85, 450, 451
431, 436, 439, 442, 454, 458, 459, 475, Saitissus, 752
476, 478, 482, 507, 508, 625–627, 631, Salfacarus, 185, 187, 475, 476, 682, 700,
640, 654, 669, 673, 676, 681–683, 692, 710, 721
703, 711, 714, 722, 726–728, 743, 747, salomonis, Paratemnoides, 748
754, 764, 769, 841–844, 860, 896, 931, Salticidae, 182, 339, 341, 348, 352, 362, 367,
932, 934, 935 373, 379, 392, 396, 399, 402, 404, 420,
riedeli, Galibrotus, 663 430, 443, 504, 513, 519, 521, 522, 647,
Riftobuthus, 95, 463, 476 649, 668, 678, 687, 700, 709, 718, 726,
Rilaena, 327, 337, 338, 372, 377, 378, 741, 750, 752, 759, 763, 773, 776,
389, 411 865–867, 874, 881, 883, 900
Rinawa, 773 Samoa, 75, 79, 138–141, 148, 158, 178, 181,
rioindiocubanicola, Negreaella, 663 183, 515, 516, 518, 519, 716, 733, 735,
Risdonius, 760 736, 738–740, 749, 750
Roche, 717 Samoidae, 40, 48, 158, 466, 488, 494, 636,
Roewereca, 466 644, 660, 662, 664, 674, 684, 706, 715,
Roeweria, 492 716, 720, 723, 736, 748, 749, 757, 779
Roeweriana, 695 Samooidea, 39, 40
roeweri, Cynortoides, 661 sanctaehelnae, Chaussieria, 192, 369
Roeweriscus, 84, 382 Sandokanidae, 40, 156, 396, 399, 412, 414,
romanus, Metacynortoides, 661 426, 427, 433, 737
Romerella, 634 sangkulirangensis, Sarax, 414, 427
Roncobisium, 87, 360, 835 Santhomea, 466
Roncus, 336, 351, 355, 358, 371, 451, 835, Santobius, 663, 749, 776
843, 861 Sanvincentia tarsalis, 664
Roquettea, 488 Saramacia, 493
rostratus, Diplocephalus, 867 Sarasinellinae, 156, 399, 737
Rostromontia, 695 Sarasinica, 426
rostrumpsittaci, Dinorhax, 74, 406, 425, Sarasiniciinae, 156, 399, 737
743, 896 Sarasinicinae, 413, 426
Rotundrela, 697 sarawakensis, Sarax, 427, 749
rowlandi, Cubazomus, 671 Sarax, 141, 387, 399, 404, 414, 415, 427, 433,
rowlandi, Mesoproctus, 65 733, 749, 781, 894
rowlandi, Troglocubazomus, 666 Sarcoptiformes, 36, 67, 71, 193–199, 362,
Rowlandius, 46, 137, 498–500, 645, 646, 665, 374, 442, 504, 687, 726, 744, 763, 768,
666, 670, 671 779, 868, 933
Rudaemannia, 191 Sarcoptoidea, 874
Rugothodes, 866 Sardostalita, 356
Runga, 772, 780 sardous, Spelyngochthonius, 355
Ruschia, 491 Sarotesius, 474
ruwenzoriensis, Callitrichia, 885 Sassandiothus, 688
Index 979
Sathrochthoniella, 89, 769, 779 Scutacaridae, 875

Satta, 752 Scutoverticidae, 877
sauteri, Apozomus, 409, 417, 422 Scytodidae, 182, 327, 339, 362, 396, 402,
sauteri, Suzukielus, 409, 417, 422 404, 646, 678, 685, 718, 724, 738, 776,
savignyi, Lamprochernes, 388, 451 839, 900
sawadai, Orientzomus, 417, 440 sechellana, Samoa, 716
Saxidromus, 193 Secozomus, 85, 137, 472, 716, 721
sayensis, Cynortula, 635, 643, 661 secunda, Platycynorta, 662
sayi, Vonones, 662 Sedna, 84
Sbordonia, 636 Segestriidae, 182, 328, 339, 341, 362, 402,
scaber, Phrynichus, 713, 716, 721 647, 678, 686, 724, 739, 776, 867,
Scabrosidama, 466 873, 900
scabrosus, Metacynortoides, 661 segmentatus, Opilioacarus, 185, 186, 328,
Scelidomachus, 393 342, 344, 452
Scharfenbergia, 192 segmentidentatus, Cacodemonius, 658
schatzi, Galapagacarus, 504 Segregara, 699
Scheloribates, 881, 884 Segundolus, 493
Scheloribatidae, 869, 877 Selachochernes, 89, 505
Schenkeliella, 403 Selachochthonius, 86, 89, 693
Schenkeliobunum, 510 Selencula, 466
Schizomida, 2, 18, 36, 45, 46, 65, 136–138, Selenopidae, 182, 420, 422, 434, 647, 678,
319, 625, 758, 841, 859, 894, 931 686, 709, 725, 741, 751, 762, 874
Schubartesia, 492 Selkirkiella, 506
schubarti, Diplothyrus, 501 sellnicki, Liochthonius, 884
Sclerobuninae, 511 Selmistomia, 95
Sclerobunus, 511 Semibulbus, 390
Sclerosomatidae, 40, 48, 153, 327, 337, 338, Semostrus, 493
344, 347, 352, 355, 360, 361, 372, 377, Senoculidae, 182, 647, 678
378, 381, 388, 389, 396, 398, 401, 411, Senoculifer, 751, 781
422, 425, 433, 440, 442, 510, 634, 641, Seothyra, 698
660, 673, 677, 684, 712, 713, 723, 736, Seqocrypta, 93
748, 757, 781, 838, 862–864, 871, 898 serbica, Trojanella, 339, 838
Sclerostygnellus, 490 sergeidedicatus, Sabacon, 372
Scopalio, 430 serrano, Rowlandius, 666
Scopticus, 430 serratosignata, Enoplognatha, 867
Scorpio, 383, 385, 386, 448, 450, 460, serratus, Lychas, 712
461, 860 Sesato, 717
scorpioides, Pselaphochernes, 355 Sesostrellus, 465
Scorpiones, 2, 12, 36, 45, 46, 65, 144–145, Sesostris, 465
320, 432, 625, 833, 859, 897, 931 seticauda, Tetrabalius, 428
Scorpionidae, 145, 380, 382, 383, 385, 386, setifer, Mughiphantes, 885
395, 405, 408, 424, 432, 448, 450, setulosus, Galeodes, 394, 860
459–463, 482, 485, 508, 631, 633, 642, Seuthesplus, 466
654–656, 672, 681, 683, 688, 693, 703, Seuthessus, 466
714, 722, 734, 834, 843, 860 sextuberculata, Cynorta, 661
Scorpionidea, 36, 37 Seycellesa, 717
Scorpionomorpha, 36, 46 seychellarum, Charinus, 716, 721
Scorteccia, 452 seychellensis, Afrogarypus, 715
Scothinotylus, 866 seychellensis, Compsaditha, 715
Scotolemon, 347, 352, 356, 361 seychellensis, Ideoblothrus, 715
Scotowithius, 87, 368 Shango, 697
Scufia, 468 sheari, Eophalangium, 66, 70
scullyi, Phrynichodamon, 695 Shearogovea, 91, 627, 631, 632, 637, 836, 844
scullyi, Phrynichus, 695 sherpa, Mughiphantes, 885
980 Index
Shibaia, 875 Snartia, 192

Shinobius, 183 sobrinus, Anepsiozomus, 716
Shravana, 86, 406 socotranus, Hemiscorpius, 392
shulovi, Paramenthus, 388 socotrensis, Hottentotta, 392
Siamacarus, 93, 186, 187, 741, 842 Socotroonops, 393
siamensis, Bamazomus, 50, 417, 420 Sodreana, 493
siamensis, Eukoenenia, 405 Sodreaninae, 493
Siamsporops, 420 Soelteria, 709
siankaanensis, Neocarus, 638 soerenseni, Dicranolosma, 356, 358
Sibambea, 157, 489 Soeuria, 718
Sibillus, 491 Sokodea, 718
Siblyia, 192 Soledadiella, 495
siboney, Rowlandius, 666 solegladi, Euscorpius, 333, 335
Sibundoxia, 490 Solifugae, 2, 3, 14, 15, 35, 36, 45–47, 50, 65,
sicanus, Euscorpius (E.), 335, 343, 354, 358, 69, 74, 76, 77, 83, 98, 159, 319, 320,
364, 448 323, 324, 333–335, 343, 345, 354, 359,
Sicariidae, 182, 327, 339, 362, 396, 646, 678, 364, 367, 371, 376, 381, 382, 392, 394,
685, 724, 738, 872, 900 397, 398, 402, 405, 425–431, 436, 448,
siciliensis, Chernes, 358 449, 454, 455, 476, 478, 481, 503, 507,
Sickesia, 494 508, 626, 627, 631, 640, 653, 654, 669,
siculus, Lasiochernes, 355 673, 676, 683, 692, 703, 711, 714, 718,
siculus, Roncus, 355 722, 726, 728, 743, 747, 754, 764, 778,
sierramaestrae, Mexobisium, 657 781, 859, 860, 880, 884, 886–887, 896,
signifer, Haplodrassus, 866, 867, 885 931–935
sijuensis, Trithyreus, 417 Solifugi, 159, 508, 692
silhavyi, Manahunca, 663 Solifugomorpha, 36
silvai, Peudocellus, 654 Solpuga, 449, 456, 457
Simaethula, 763 Solpugida, 74, 159
Simalurius, 433 Solpugidae, 163, 381, 448, 449, 456–458,
Simienatus, 466 683, 692, 722, 896
similis, Zelotes, 866, 867 Solpugista, 84
simoni, Grosphus, 704 Solpugistella, 457, 458
Simonobisium, 87, 360 Solpugisticella, 84
Simonus, 761 Solpuguna, 84, 693
Singalangia, 429 Soluta, 36
Singram, 492 Somalibuthus, 85, 462, 476
singularis, Drassodes, 885 Somalicharmus, 85, 462, 476
Sinistus, 426 Sondra, 763
Sinochelifer, 88 soqotranus, Charinus, 392
Sinoteneriffia, 192 Sorensenella, 779
Siponnus, 413, 426 Sororoditha, 86, 90, 483
Siro, 337, 347, 510 Sotanochactas, 627, 632, 633, 834, 843
Sironidae, 39, 50, 74, 81, 150–152, 326, 337, Sotanostenochrus, 86, 138, 631, 638
338, 347, 355, 360, 381, 388, 436, 465, Sotekia, 426
510, 628, 684, 836, 844, 898 southcotti, Proterythraeus, 49, 51, 67, 70
Sironoidea, 38, 150 spalacis, Nudochernes, 388
siscoensis, Chthonius (E.), 355 Sparassidae, 182, 328, 339, 362, 396,
Sitalcicus, 712, 713, 716, 721 399, 402, 474, 647, 649, 668, 678,
Sitalcina, 92, 512 686, 699, 709, 718, 725, 741, 751,
Sitticus, 866–868, 885 761, 841, 874
Sivianus, 495 spelaea hauseri, Eukoenenia, 334
Smarididae, 67, 70, 80, 189, 763, 768, 875 Spelaeobochica, 483, 844
Smaris, 189, 190 Spelaeochernes, 487, 844
Smionia, 700 Spelaeorhynchidae, 199, 501, 650, 668
Index 981
Speleoharpactea, 348, 839 Stenochrus, 98, 138, 498–500, 503, 513,

Speleomantes, 78 628, 631, 638, 646, 666, 670, 671,
Speleomaster, 511 843, 894
Speleomontia, 695 Stenohya, 371, 870, 871, 885
Speleonychia, 512, 838 Stenophareus, 494
Speleophrynus, 141 Stenoschizomus, 138, 498–500
Speleosiro, 91, 694, 837, 842 Stenostygnellus, 494
Spelungula, 771, 780 Stenostygninae, 660, 663, 664
Sphaerobothria, 647 Sternophoridae, 38, 148, 395, 398, 401,
Sphaerobunus, 491 406, 641, 657, 659, 671, 672, 685, 723,
Sphaeromides, 95 735, 747, 781, 782, 836, 846, 861, 933
Sphaerotarsus, 189, 190, 738, 763 Sternothyrus, 94, 194, 195, 198, 718, 721
Sphallowithius, 87, 368 Stewardpeckius, 86, 138, 646, 666, 670, 671
Sphenodon, 95, 767, 773 Sthelota, 648
sphenodonti, Aponomma, 767, 773 Stichius, 430
Sphoeroforma, 495 Stichoplastoris, 647
Spinanapis, 760 Stictocarbo, 767, 773
Spinasteron, 762 Stigmaeidae, 875
spinata, Torreana, 663 Stigmocheylidae, 192
Spinembolia, 717 Stigmocheylus, 192
Spinestis, 373 stimpsonii, Typopeltis, 437, 440
Spinicranaus, 489 Stipax, 718
Spinicrus, 765 Stiphidiidae, 740, 750, 751, 766, 841, 846,
Spiniella, 495 901
spinifera, Bunochelis, 366 Stiphropella, 700
spiniferum bolivari, Stygnomma, 662 Stoda, 717
spiniferum spiniferum, Stygnomma, 662 stoliczkae, Gnaphosa, 885
spiniger, Koeneniodes, 405 stonei, Tyrannochthonius, 521, 845
Spinimontia, 706 Storenosoma, 761
spinipalpis, Ramblinus, 366 Storosa, 762
Spinixestus, 465 Strandiellum, 751
spinooculorum, Paraconomma, 664 strandi, Syarinus, 95
Spinopilar, 493 stridulans, Maracaynatum, 662
Spinotrachelas, 697 strigilata, Pardosa, 334
Spiroctenus, 699 Stygnicranaus, 490
Spirunius, 489 Stygnicranella, 490
squama, Cercophonius, 755, 764 Stygnidae, 40, 156, 488, 494, 501, 660,
srilankensis, Lychas, 401 662, 674
Staianus, 709 Stygnidius, 494
Stalagtia, 341, 839 Stygninae, 494
Stalita, 341, 839 Stygnobates, 493
Stalitella, 341 Stygnoleptes, 644
Stankiella, 327, 337 Stygnoleptinae, 158, 737, 779, 781, 782
Starengovia, 389, 410 Stygnomma, 512, 636, 644, 662, 900
Stasinoides, 474 Stygnommatidae, 40, 158, 159, 488, 512,
stator, Cubachelifer, 658 636, 644, 660, 662, 674, 736, 844
Steatoda, 840, 846, 866 Stygnopsidae, 40, 156, 488, 512, 636,
Stefanasia, 491 644, 674
Steganacaridae, 876 Stygnopsis, 636
steineri, Rowlandius, 406 Stygnus, 494
steineri, Troglokhammouanus, 406, 834 Stygochactas, 627, 632, 633, 834, 843
Stenochelifer, 88 stygochtobius, Charinus, 392
Stenochilidae, 182, 402, 739, 750 Stygopholcus, 341
982 Index
Stygophrynus, 141, 414, 415, 428, 733, 749, 841 T

Stylocellidae, 39, 74, 150, 152, 398, 400, 405, Tabuina, 752
409, 410, 425, 736, 742, 748, 781, 898 Tachusina, 493
Stylocellus, 409, 410, 425, 748 Taguaza, 495
Styloleptes, 490 Tagulinus, 420
Stylonuncia, 765 Tainonia, 648, 668
Stylopisthos, 490 Taivala, 430
Subantarctia, 772, 780 Tajumulcia, 643
Subasteron, 762 Takaoia, 422
subcerdoso, Antillostenochrus, 666 Talavera, 348, 866, 867
subcerdoso, Stenochrus, 666, 671 Tallonia, 708
subfuscus, Novohorus, 657 Tamin, 429
subterraneus, Charon, 428, 749 Tampiconus, 636
subterraneus, Sarax, 749 Tanalaius, 706
succineus, Ixodes, 68, 71 Tanaupodidae, 875
Suctobelba, 884 Tanaupodus, 883
Suctobelbidae, 869, 877 Tandikudius, 399
Sudharmia, 429 Tanganoides, 766
Sulcia, 341, 839 Tangata, 772, 780
Summacanthium, 430 tantalus, Mayacentrum, 645, 670
Superiores, 76, 97 tanzaniensis, Salfacarus, 475
superstes, Savignia, 348 Tapacochana, 491
Superstitioniidae, 633 Tapetosa, 760
Surasmaris, 189, 190 Tapinothele, 474
Surazomus, 138, 498–500, 645, 646, Tapinothelella, 698
670, 844, 860 Tapponia, 429
Suzukielus, 81, 91, 409, 436 Tarabulida, 449
Syarinidae, 17, 38, 95, 147, 326, 336, 337, Taracidae, 40, 154, 372, 410, 411, 510, 899
343, 346, 351, 360, 364, 371, 372, 395, Taracus, 372, 511
398, 401, 406, 425, 432, 442, 463, 483, Taranakia, 190, 768, 773, 781
509, 641, 656, 659, 671, 672, 684, 715, Tararua, 780
719, 723, 735, 747, 765, 769, 775, 781, targionii, Phalangium, 358
782, 834, 835, 843–845, 861, 897, 934 Tarmapachylus, 492
Syarinus, 96, 509 Tarodus, 752
sylviae, Damon, 696 Tarsolarcus, 192
Symbioribatidae, 877 Tarsotomus, 192
Symphanodes, 762 Tartarocreagris, 87, 509, 835
Symphytognathidae, 182, 430, 686, 725, Tartarus, 841
740, 846 Tasmabrochus, 766
Synaphridae, 182, 362, 686, 708, 725, 901 Tasmanapis, 766
Synatemnus, 88, 90, 463, 476 Tasmanicosa, 766
Syncranaus, 493 Tasmanonuncia, 765
Syncynorta, 489 Tasmanonyx, 765
Synotaxidae, 182, 183, 678, 740, 750, 760, Tasmanopilio, 80, 91, 757, 765
766, 767, 772, 840, 901 Tasmarubrius, 766
Synotaxinae, 182, 740, 767 tassili, Compsobuthus, 448
Synothele, 93 Tauala, 763
Synsphyronidae, 147, 735 tauricus, Euscorpius, 371
Synthetonychia, 91, 770, 779 Taurisa, 371
Synthetonychiidae, 40, 91, 156, 736, 742, Tautukua, 780
744, 770, 778, 779, 782 taymirensis, Mesoanystis, 67, 70
Syroloma, 522 Tayos, 86, 138, 498–500, 834
Systariinae, 181, 741 Taypaliito, 434
Szczurehia, 489 Tchapinius, 372
Index 983
Tectocepheidae, 869, 877 Thaiphantes, 419

Tectocepheus, 881, 884 thais, Eukoenenia, 405
Tecution, 369 Thalassiopsis, 708
Tedia, 391 thaleri, Pardosa, 885
Teeatta, 766 Thaloe, 648, 667
Teemenaarus, 760 Thanatus, 867
Tegenaria, 176, 740, 841, 866 Thapsagus, 709
Tegipiolus, 494, 495 Thasyraea, 761
Tegoribatidae, 877 Thaumastochilus, 697
Tegyra, 493 Thaumatocranaus, 489
Tekellatus, 760 Thaumatoleptes, 491
Tekelloides, 772, 780 Thaumatopachylus, 490
Telechelifer, 88 Thaumatowithius, 87, 712, 720
Telemidae, 183, 647, 685, 724, 738, 750, 776, Thelyphonus, 139, 399, 416, 428, 434, 443,
781, 839, 842, 844, 901 742, 749, 750, 781
Teloleptoneta, 93, 839 Themacrys, 699
Teltus, 140, 415, 733 Theraphosidae, 9, 176, 399, 402, 419, 422,
temmincki, Manis, 199 429, 473, 646, 647, 666, 685, 696, 707,
Templar, 770, 780 717, 725, 737, 838, 844
Tencateia, 192 Theraphosina, 41
Teneriffia, 192 Theraphosomorphae, 41
Teneriffiidae, 189, 192, 362, 875 Thereza, 491
Tengellidae, 184 Theridiidae, 77, 183, 328, 339, 341, 348, 352,
Tenuipalpidae, 875 362, 367, 369, 373, 379, 399, 402, 404,
Tenuiphantes, 866, 872, 881 430, 434, 443, 473, 506, 514, 521, 525,
Tepotztlana, 190, 638 647, 648, 667, 678, 686, 698, 708, 717,
Teranodes, 760 725, 740, 751, 760, 776, 781, 840, 846,
tergestinus, Euscorpius, 335 865–867, 873, 900
termitarium, Apozomus, 417 Theridion, 504, 521, 840, 866, 867
Termitoonops, 473 Theridiosomatidae, 183, 328, 339, 362, 402,
terricola, Scotolemon, 356 419, 647, 686, 725, 740, 840, 845
tertiarius, Ixodes, 68 Therlinya, 761
teruel, Rowlandius, 666 Theromaster, 511
Tetebius, 466 Theuma, 699
Tetrabalius, 139, 416, 428, 733, 750, 781 Theumella, 474
Tetrablemmidae, 183, 396, 402, 419, 429, Thexmus, 756
474, 518, 647, 648, 717, 724, 738, 776, Thianella, 430
844, 872 thibaudi, Aphelolpium, 657
Tetrachelifer, 88, 406 thienduongensis, Vietbocap, 406, 834
Tetracranaus, 490 thienmuschanensis, Oligolophus, 375
Tetragnatha, 521 Thomecola, 466
Tetragnathidae, 183, 328, 339, 341, 362, 390, Thomisidae, 73, 183, 328, 339, 341, 348, 362,
399, 402–404, 429, 514, 521, 522, 647, 369, 373, 379, 393, 396, 399, 402–404,
667, 678, 686, 717, 725, 772, 776, 865, 420, 430, 434, 475, 514, 521, 522, 647,
867, 873, 900 649, 668, 678, 687, 700, 709, 725, 741,
Tetranychidae, 875 752, 759, 763, 865–867, 874, 900
Tetrastigmata, 194 Thonius, 94, 194–196, 198, 752
Tetrophthalmi, 48, 66, 70 Thrasychiroides, 488
Teuthraustes, 482, 485, 486, 887 Thrasychirus, 525, 677
texanus, Neocarus, 185, 514, 629 Thyreobaeus, 709
Texella, 512 Thyrisomidae, 877
Teyl, 760 Thyropoeus, 708
Teyloides, 760 Tiara, 495
Thainetes, 419 Tibangara, 493
984 Index
tibetana, Karschia, 394, 884 Travunia, 327, 339, 838, 899

Tigidia, 707 Travuniidae, 40, 81, 156, 327, 337, 339,
tigrina, Gnaphosa, 866, 867 356, 361, 436, 437, 440, 443, 512,
tikaderi, Neozomus, 399, 417 684, 838, 899
tikaderi, Pardosa, 885 Travunioidea, 38–40
Timesius, 494 Trechaleidae, 183, 647, 678, 686, 725,
Timotesa, 490 740, 901
Tingomaria, 492 Trencona, 427
Tinytrema, 762 Trephopada, 700
tirolensis, Erigone, 867 Trhypochthoniidae, 876
Tiso, 866, 867 Triacumontia, 706
titanius, Garypus titanius, 368 Triadokoenenia, 83, 135, 703, 718, 894
Titanobochica, 79, 85, 94, 323, 345, 346, 835, Triaenomontia, 706
895, 934 Triaenonychidae, 38, 156, 436, 442, 443,
Titanodamon, 141 511, 524, 677, 678, 684, 695, 706, 720,
Titanoeca, 741 723, 727, 736, 742, 744, 757, 764, 765,
Titanoecidae, 183, 328, 339, 362, 402, 428, 770, 775, 776, 779, 838, 842, 844, 845,
647, 678, 687, 725, 741, 874 862, 864
Tithaeidae, 40, 156, 412, 413 Triaenonychinae, 511
Titus, 700 Triaenonychoidea, 39, 40
Tityobuthus, 705 triangularis, Linyphia, 867
Tityopsis, 84, 655, 656, 672 Tribunosoma, 491
Tityus, 482, 484, 486, 633, 642, 655, 656, Trichocyclus, 761
672, 887 Trichodamon, 79, 142, 495, 496, 844, 895, 934
Tjurunga, 766 Trichognathiella, 696
Tmeticides, 709 Trichominua, 491
Tobotanus, 489 Trichopelmatinae, 172
Togwoteeus, 510 Trichoribates, 868, 881
Tokunosia, 422, 440 Trichoribatidae, 869, 877
toldo, Rowlandius, 666 Trichosmaris, 189, 190
Tolma, 93, 708 Trichothyse, 700
Toloella, 649 Trichotoma, 84, 692
Tolus, 92, 512 Trichotowithius, 87, 90, 463, 476
Tomobella, 710 Tricommatinae, 493
Tomocyrba, 710 Tricommatus, 493
Tonkouinatus, 468 Triconobunus, 776
Toreus, 84, 692 Tridenchthoniidae, 146, 395, 406, 425, 432,
Toschia, 883 463, 483, 509, 515, 641, 656, 659, 676,
Toxoniella, 473 684, 705, 715, 719, 723, 735, 747, 775,
Toxops, 766 781, 782, 835, 861, 871, 882
Toxopsiella, 773, 780, 841 tridentis, Pardosa, 885
Toxopsoides, 773, 780 Triglochinura, 492
Trachelidae, 183, 402, 697 Trigonobunus, 427
Trachycosmus, 762 Trigonotarbida, 45
Trachyrhinus, 634 Trilasma, 511, 643, 673
Trachyspina, 762 trinitatis, Lechytia, 656
Trachytrema, 762 Trionyxellinae, 402
Tragacantha, 855 Tripilatus ., 490
Tragulidae, 81 Triregia, 770, 779
Traiania, 495 Trisetobisium, 87, 509, 835
transoceanicus, Mastigoproctus, 497 Trithyreus, 86, 138, 417, 418, 434, 733, 894
transversalis, Metacynortoides, 661 Trittame, 93
Tranteeva, 25, 91, 327, 338, 339, Trochanteriidae, 183, 678, 687, 725,
836, 898 741, 762
Index 985
Troglobisium, 80, 96, 326, 345, 346, 835, Tuberocreagris, 87, 509
897, 934 Tugana, 648, 667
troglobius, Stewardpeckius, 666, 671 tunetanum, Nemastoma, 843
Troglobochica, 86, 657–659, 835 tunetanus, Buthus, 358
Troglochthonius, 86, 326, 336, 337, 355, Tungari, 93
835, 897 Tupacarana, 491
Troglocormus, 627, 632, 633, 843 Tupua, 766, 840
Troglocubazomus, 138, 666 turkestana, Eusimonia, 371
Troglodiplura, 760, 846 turquinensis, Kimula, 663
troglodytes, Hickmania, 766, 838 Turretia, 772, 780
Troglohya, 86, 835, 844 Tusipulla, 466
Troglokhammouanus, 85, 95, 406, 408, Tuxoctenus, 761
834, 897 Tybaertiella, 883
Trogloraptor, 514, 628 Typhlobunus, 92, 466
Trogloraptoridae, 183, 507, 514, 628, 935 Typhloburista, 92, 466
Troglosiro, 91, 151, 775, 782 Typhlochactas, 627, 632, 633, 834, 843
Troglosironidae, 38, 39, 74, 91, 96, 151, 153, Typopeltinae, 37, 140
736, 742, 743, 775, 778, 782, 837, 898,
933, 935
Troglostygnopsinae, 156 U
Troglostygnopsis, 636 Urodacus, 755, 756, 778
Troglotayosicidae, 79, 145, 325, 346, 482, Urodiabunus, 491
483, 485, 486, 683, 834, 897, 935 Uroecobius, 697
Troglothele, 93, 647, 666, 667 Urogelides, 763
Trogulidae, 40, 48, 154, 327, 337–339, 344, Uropachylus, 492
347, 352, 356, 360, 361, 372, 377, 378, Uroplectes, 460–463, 693
381, 388, 389, 410, 411, 451, 684, 838, Uroplectoides, 462, 463
862, 871, 899 Uropodina, 49, 67, 71
Trogulocratus, 338, 389 Uroproctinae, 139, 497, 628, 638, 895
Troguloidea, 38–40 Uroproctus, 85, 140, 416, 428
troguloides, Arythaena, 67, 70 Uropygi, 2, 3, 12, 36, 37, 45, 46, 50, 65, 69,
Trogulus, 48, 155, 327, 339, 344, 347, 351, 73, 75–79, 85, 96, 98, 138–140, 319,
352, 356, 361, 372, 377, 378, 389, 320, 323, 343, 361, 366, 371, 376, 381,
451, 862 391, 397, 399, 401, 415, 416, 421, 428,
Trojanella, 92, 96, 327, 339, 838 431, 434, 437, 440, 442, 443, 454, 470,
Trombellidae, 875 471, 476, 478, 497, 507, 513, 625, 626,
Trombiculidae, 189, 868, 875, 881 628, 631, 637, 638, 640, 645, 665, 670,
Trombidiidae, 189, 868, 875, 883 673, 676, 682, 683, 690, 703, 722, 726,
Tropasteron, 762 727, 733, 742, 743, 747, 749, 750, 754,
Tropidowithius, 87, 90, 487 759, 764, 769, 778, 779, 781, 782, 842,
Troya, 490 860, 895–896, 931
Trujillina, 648, 667 urquinensis, Calmotrinus, 662
Truncattus, 649, 668 usingeri, Amblyomma, 504
truncatum, Hyalomma, 393 Uspallata, 84
truncatus, Ideoblothrus, 657 Uviranus, 466
Tryferos, 450 Uzakia, 773, 780
Tschaidicancha, 491
Tuakana, 773, 780
Tuapoka, 772, 780 V
tuberculata, Kimula, 663 vachoni, Antillobisium, 657, 835, 843
tuberculata, Mystacina, 768, 773 Vachoniidae, 146, 835, 844, 933
tuberculatus, Stygnoplus, 662 Vachoniolus, 385, 386
Tuberculosa, 760 Vachonium, 86, 89, 632, 844
Tubereca, 466 Vachonobisium, 85, 91
986 Index
Vachonus, 407, 688 Vononella, 489

Vaejovidae, 145, 508, 627, 631, 633, 642, Vonones, 512, 636, 637, 646
673, 674, 843 Vononesta, 643
Vaejovoidea, 37, 145 Vononissus, 489
v-album, Cynortoides, 661 Vononula, 643
Valdesia, 676 Voraptipus, 698
Valeriophonus, 140, 497, 498, 645, 670 Vuattouxia, 474
Valgothrombium, 884 Vulcanochthonius, 521, 934
vallentini, Hoggellula, 524, 680 Vulsor, 80
valleorum, Siro, 343 Vytfutia, 429
Vandaravua, 399
Vanderhammenacarus, 741
vanderhammeni, Caribeacarus, 843 W
Vankeeria, 341 Wabarra, 761
variabilis, Mughiphantes, 867 Wabua, 761
variegates, Damon, 695, 696 wahrmani, Myrmecowithius, 388
Variratina, 752 waigiensis, Hormurus, 515
Vazaha, 708 Waiporia, 772, 780
velox, Biton, 98, 354 Wairua, 780
Venator, 760 Waitkerra, 780
Vendaphaea, 700 Walckenaeria, 866, 872, 883
Ventrifurca, 490 Walrencea, 698
Ventripila, 490 Walzia, 192
Ventrisudis, 490 Wanniyala, 403
veracruzensis, Neocarus, 638 Wartookia, 191, 763, 768
Verrucaditha, 86, 509 Wesmaldra, 762
Verrucastygnus, 494 Wesolowskana, 367
vespertilionis, Hesperochernes, 658 Wespus, 92, 512
Vestiferum, 327, 377, 378, 389 Westrarchaea, 760
Vestitecola, 664 Weyrauchiana, 495
Viacha, 495 wheeleri, Prokoenenia, 508, 626
Vibone, 695, 779 wiehlei, Dicranopalpus, 355, 356
Victanapis, 760 willeyi, Sarax, 749
Victoriaincola, 492 williamsi, Amblyomma, 504
vidali, Roncus, 351 Wiltonia, 772, 780
Vidole, 699 Withiidae, 17, 38, 148, 326, 336, 343, 346,
Vietbocap, 85, 95, 408, 897 351, 360, 364, 365, 367, 368, 372,
vigilax, Walckenaeria, 867 377, 378, 381, 387, 388, 392, 395,
Vilhena, 466 398, 401, 406, 422, 425, 432, 451,
Vima, 488, 838 463, 483, 487, 503, 505, 515, 521, 641,
Vimina, 488 658, 660, 671, 672, 677, 685, 705, 712,
vinai, Negreaella, 663 715, 720, 723, 735, 747, 769, 781, 782,
vinai, Rowlandius, 666 836, 861, 871
virginiae, Rowlandius, 666 Withius, 336, 365, 372, 432, 720, 882
virginsulana, Martibianta, 663 Wuayuuzomus, 138, 844
Viribestus, 752 Wugigarra, 761
Viridasiidae, 183 wunderlichi, Eusimonia, 364
Viridictyna, 773, 780 Wydundra, 762
viridis, Rowlandius, 666 Wyochernes, 371
Viroqua, 763
vitellinum, Ptychosoma, 344, 356, 358, 448,
452 X
vittatus, “Schizomus,” 417 xambeui, Belisarius, 79, 321, 346, 834
Vlachiolus, 665 Xamiatus, 760
Vononana, 636 Xenillidae, 877
Index 987
Xenochelifer, 88, 509 Zalmoxoidea, 39, 40

Xenolpium, 720 Zalmoxomma, 712, 713
Xerolycosa, 866 Zamora, 488
Xerophrynus, 142, 469, 470, 894 Zamorinae, 488
Xevioso, 699 Zantheres, 420
Xyphinus, 429 Zaraxes, 489
Xysticus, 73, 866, 867, 872 Zatavua, 709
Zealanapis, 771, 780
Zealoctenus, 773, 780
Y Zearchaea, 678, 772, 780
Yacambuia, 495 Zearchaeinae, 180, 739
yaksha, Electrokoenenia, 45, 50, 65, 69 zebroides, Ethobunus, 664
yamazakii, Apozomus, 422 Zelanda, 773, 780
Yania, 490 Zercidium, 369
Yapacana, 494 Zeria, 449, 456, 457
Yatala, 757, 779 Zetomotrichidae, 877
yayukae, Sarax, 414, 427 Zetorchestidae, 876
yeti, Mughiphantes, 885 zhensis, Apozomus, 417
yezoensis, Nipponopsalis, 375 zherikhini, Jurarchaea, 51
Yiinthi, 761 Zillimata, 762
Yilgarnia, 760 Zodariidae, 9, 184, 328, 339, 341, 362,
Yoroa, 751, 781 396, 402, 420, 474, 647, 678, 680,
Yulella, 695 686, 697, 709, 725, 741, 762, 773, 777,
yulungensis, Agyneta, 885 866, 874, 883
yulungensis, Euophrys, 885 Zodarion, 866
yumuriensis, Negreaella, 663 zoiai, Roncus, 355
yunquense, Ideobisium, 656 Zombis, 84, 382
yunquensis, Luisarmasius, 666, 671 Zomus, 138, 418, 515, 733
Yuria, 81, 91, 437, 690 Zophorame, 93
Zophoryctes, 93, 707
Zora, 181, 741
Z Zorinae, 181, 328, 339, 341, 362, 741, 761,
Zabkattus, 752 766, 776, 866, 867, 874
Zachaeus, 337, 338, 389 Zorocratidae, 184
Zachria, 761 Zorodictyna, 708
Zairebiantes, 466 Zoroides, 776
Zalanodicus, 493 Zoropsidae, 184, 328, 339, 390, 402, 647,
Zalmopsylla, 493 678, 686, 697, 707, 725, 731, 737, 740,
Zalmoxidae, 40, 158, 412–414, 488, 494, 741, 750, 761, 762, 773, 874, 901
516, 636, 644, 660, 664, 674, 677, Zortalia, 493
684, 706, 712, 713, 715, 716, 720, Zosis, 183, 739
723, 737, 748, 749, 757, 775, 776, zuluanus, Megaschizomus, 696
779, 781, 782, 895 Zulunigma, 700
Zalmoxis, 413, 516, 712, 749, 776, 781 Zuma, 511

Zoogeography of Arachnida

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Monographiae Biologicae 94

The Monographiae Biologicae provide a forum for top-level, rounded-off

More information about this series at http://www.springer.com/series/6304

ISSN 0077-0639 ISSN 2215-1729 (electronic)

Library of Congress Control Number: 2018932984

© Springer International Publishing AG, part of Springer Nature 2018

Printed on acid-free paper

5.5 Endemics and Relicts in Arachnida���������������������������������������������� 83

7.7 Balearic Islands ���������������������������������������������������������������������������� 350

7.18 Socotra������������������������������������������������������������������������������������������ 391

7.29 Korea (North and South) �������������������������������������������������������������� 441

8 Some Peculiarities of the Distribution of Arachnida���������������������������� 625

8.13 Arachnofauna of Madagascar and the Adjacent Islands���������������� 701

9.3 Conclusions ���������������������������������������������������������������������������������� 846

Abstract The classical zoogeographical subdivision of the land fauna, done by

© Springer International Publishing AG, part of Springer Nature 2018 1

Schizomida: Harvey (1992), Reddell and Cokendolpher (1995)

New Zealand: Forster (1967–1973, Araneae; 1978, Opiliones), Dumbleton, 1953,

Gecheva (2013, Ixodidae of Bulgaria), Gregson (1956, Ixodida of Canada),

United States – E. Benedict, J.C. Chamberlin, J.C. Cokendolpher, J. Cracraft, V. Fet,

© Springer International Publishing AG, part of Springer Nature 2018 7

Linnaeus, Carolus (Carl von Linné) (1707–1778) – professor of medicine and

C. Linnaeus (Portrait by A. Roslin, 1775)

Sørensen, William Emil (1848–1916) – Danish arachnologist. Described 157

William Emil Sørensen (1848–1916)

Hansen, H.J. (1855–1936) – Danish arachnologist. Some important papers:

Hans Jakob Hansen

Mello-Leitão, Cândido Firmino de (1886–1948) – Brazilian arachnologist, pro-

Cândido Firmino de Mello-Leitão

Pocock, Reginald Innes (1863–1947) – English arachnologist and curator of the

R.I. Pocock (From P. Bonnet, 1945)

Kraepelin, Karl Matthias Friedrich Magnus (1848–1915) – German arachnolo-

Roewer, Carl-Friedrich Roewer (1881–1963) – German arachnologist, profes-

C.-F. Roewer (From O. Kraus)

Chamberlin, Joseph Conrad (1898–1962) – one of the founders of the system of

Joseph Conrad Chamberlin

Brignoli, Paolo Marcelo (1942–1986) – Italian arachnologist, professor in Rome

Paolo Marcelo Brignoli

Simon, Eugène (1848–1924) – French zoologist, working mainly on spiders and

Koch Carl Ludwig (1778–1857) – German arachnologist. His principal works,

Banks, Nathan (1868–1953) – American arachnologist and entomologist. Author

Šilhavý, Vladimir (1913–1984) – Czech physician and prominent specialist in

Vladimir Šilhavý (Contributed by P. Bezdĕčka)

Hadži, Jovan (1884–1972) – Slovenian arachnologist and academician. He stud-

Jovan Hadži (Slovenia Wikipedia)

Petrunkevitch, Alexander (1875–1964) – American arachnologist of Russian ori-

A. Petrunkevitch (Courtesy, Yale University)

Kratochvil, Josef (1909–1992) – Czech Zoologist and Academician, worked on

Lawrence Reginald Frederick (1897–1987) – the most prominent researcher of

Reginald Frederick Lawrence

Vachon Max (1908–1991) – French specialist in Scorpiones and Pseudoscorpiones.

Gonzalez-Sponga Manuel-Angel (1929–2009) – Venezuelan arachnologist and

Manuel-Angel Gonzalez-Sponga (Courtesy of R. Pinto-da-Rocha)

Berlese Antonio (1863–1927) – Italian Acarologist who described thousands of

Oudemans Anthonie Cornelis (1858–1943) – Dutch acarologist and one of the

Anthonie Cornelis Oudemans

Staręga Wojciech (1939–2015) – Polish arachnologist and specialist in Opiliones

W. Starega (Courtesy of J. Cokendolpher)

Birabén M (1949) Necrológio – Cândido F. de Mello-Leitão. Revista de la Socedad Entomologica

“Traditions die hard: it is difficult to understand why fishes or

Heymons (1901), the author of the term Chelicerata, united Merostomata

© Springer International Publishing AG, part of Springer Nature 2018 35

Suborder Apulvillata – families Phrynidae and Phrynichidae

5.5 Endemics and Relicts in Arachnida�� 83

7.7 Balearic Islands �� 350

7.18 Socotra�� 391

7.29 Korea (North and South) �� 441

8 Some Peculiarities of the Distribution of Arachnida�� 625

8.13 Arachnofauna of Madagascar and the Adjacent Islands�� 701

9.3 Conclusions �� 846

5.1 Some Basic Notions

5.1.1 Zoogeography and Biogeography: The New Approach

5.1.2 Centers of Origin and Dispersal

5.1.3 Development of Atlantic Ocean

5.1.4 Development of the Indian Ocean

5.2 Paleogeography and the Past Distribution of Arachnida

5.2.1 Age of the Faunas

5.3.1 Dispersal (Natural and by Man Activities)

5.4 Ecological Factors

5.4.1 Climatic and Other Barriers and Bridges

5.4.3 Old World-New World Disjunctions

5.4.4 West African-Indomalayan Disjunction

5.4.7 Boreomontane and Arctic-Alpine Disjunctions

5.5 Endemics and Relicts in Arachnida

Some Endemic Genera and Families in Different Orders

5.5.2 Relicts and Relictness

Are There Relicts at All? The Issue of Relicts

5.6 The Northern Limits of Arachnida