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I A MANUAL OF ACAROLOGY
BY
I G.WKrantz
Oregon State University, Corvallis
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DEFENSE PESr J4GlI1 INFO ANALYSIS cm
AFPMB. FOREST GLEN SECtION. WRAMC
WASHrl'f(?'l'ON. tlC 20301

I Published by

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O.S.U. Book Stores, Inc.
Corvallis, Oregon

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Copyright © 1970 I
G. W. Krantz
Third printing (emended)
September, 1971
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I TABLE OF CONTENTS

I Page

I~.
I. Introduction 1
II. Systematic Position of the Acari 5
~ III. Morphology and Function 11
I~ IV. Reproduction and Embryogenesis 31
V. Oviposition and Life Stages 33
I VI.
VII.
Habits and Habitats
Collection, Preparation and Mounting of Mite Specimens
37
45
I VIII. Classification
Key to the Orders and Suborders of Acari . . . . . . . . . . . . . .
55
58
I ~ Order Opilioacariformes
Suborder Notostigmata
62
62

I~ Order Parasitiformes. . . . . . . .
Suborder Tetrastigmata
Suborder Mesostigmata
. . . . . . . . . . . . . . 64
64
67
~'" Paras ito idea . 70
II ~ Sejoidea
Trachytoidea
74
74

I ~
Uropodoidea :-- . 75
Protodinychoidea 77

II~
Diarthrophalloidea 77
Cercomegistoidea . . . . . . . . . . . . . . . . . . . . . 78
Antennophoroidea 78

II~
Megisthanoidea 79
Celaenopsoidea. . . . . . . . . . . . . . . . . . . . . . 80
Fedrizzioidea 82
Key to Families of Mesostigmata 83

II ~ Suborder Metastigmata. . . . . .
Ixodoidea
. . . . . . . . . . . . . . 135
136

I~~
Key to Families of Metastigmata 138
Order Acariformes 141
Suborder Prostigmata 141
Pachygnathoidea 144
II Tarsonemoidea
Eriophyoidea
145
148

II Eupodoidea
Tydeoidea. .
Bdelloidea
150
151
152
II Labidostommoidea
Caeculoidea .
154
154
Halacaroidea 155
II ~ lolinoidea
Tetranychoidea. . . . . . . .
156
. . . . . . . . . . . . . . 156

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Raphignathoidea . . . . . . . . . . . . . . . . . . .
Anystoidea
. . 159
161
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162
Cheyletoidea
Erythraeoidea
Trombidioidea
165
167
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Hydrovolzioidea
Hydryphantoidea . . . . . . . . . . . . . . . . . . .
170
. . 171
171
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Hydrachnoidea
Hygrobatoidea
Key to Families of Prostigmata
172
. 174 I
Suborder Astigmata 228
Anoetoidea
Acaroidea. . . . . . . . . . . . . . . . . . . . . .
231
. . 232
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Canestrinioidea 238
Listrophoroidea
Ewingoidea .
239
240
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241
Psoroptoidea
Analgoidea
Sarcoptoidea . . . . . . . . . . . .
243
245
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Cytoditoidea
Key to Families of Astigmata
247
249
255
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Key to Deutonymphs (Hypopi) of Some Families of Astigmata .
Suborder Cryptostigmata
Supercohort Palaeacari
288
292 I
Supercohort Oribatei Inferiores . . . . . . . . . . . . . . . 293

Index
Supercohort Oribatei Superiores 295
323
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I LIST OF PLATES

I Plate
General
Page

I 1.
2.
Opiliones and Araneae
Scorpiones and Pseudoscorpiones
7
8
I 3. Gnathosoma of Acari (MACROCHELIDAE, OPILIOACARIDAE,
GLYCYPHAGIDAE) 13

I 4. Gnathosoma and gnathosomal structures (VEIGAIIDAE,

HYSTRICHONYSSIDAE, EREYNETIDAE, HALACARIDAE,
CHEYLETIDAE, HYGROBATOIDEA) 14
I 5. Chelicerae and tarsi (MACROCHELIDAE, DIPLOGYNIIDAE, PYEMOTIDAE,
EREYNETIDAE, TETRANYCHIDAE, TUCKERELLIDAE,

I 6.
HETEROCHEYLIDAE)
Dorsal shield configurations
15
16
7. Male genital apparatus (TENUIPALPI DAE, TETRANYCH I DAE,
II HARPYRHYNCHIDAE, EUPALOPSELLIDAE) 22
8. Internal and external idiosomal structures (MACROCHELIDAE,

II NANORCHESTIDAE, GLYCYPHAGIDAE, EREMAEOIDEA,

PHTHIRACAROIDEA) 23
9. Internal morphology (TRACHYTIDAE) 26
II 10. Alimentary canal (diagrammatic) and ovipositors (PARASITIDAE,
UROPODIDAE, HALACARIDAE, ANYSTIDAE, ERIOPHYIDAE,

II 11.
NANORCHESTIDAE, ERYTHRAEIDAE, ACARIDAE, NOTHROIDEA)
Immature stages (EUTRACHYTIDAE, ACARIDAE)
27
34

II 12. Collecting apparatus

Notostigmata, Tetrastigmata
46

II 13.
14.
OPI L10ACAR IDAE, HOLOTHYR IDAE
HOLOTHYRIDAE
65
66

II Mesostigmata

15. Structures of mesostigmatid mites (ARCTACARI DAE, PODOCI NIDAE,

II MACROCHELIDAE, EVIPHIDIDAE, LAELAPIDAE, SPINTURNICIDAE,
PLANODISCIDAE, UROPODIDAE, DIPLOGYNIIDAE, EUZERCONIDAE,
CE RCOMEG ISTI DAE, KLiNCKOWSTROEM II DAE) 95
II 16. Structures of mesostigmatid mites (U ROPODE LLI DAE, MICROGYN II DAE,
ARCTACARIDAE, VEIGAIIDAE, SEJIDAE, ZERCONIDAE,

I 17.
DIPLOGYNIIDAE)
MICROGYNIIDAE, UROPODELLIDAE
96
97

I 18.
19.
SEJIDAE
EPICRIIDAE
98
99

1.
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20. PARASITIDAE 100 I
21.
22.
23.
24.
25,
VEIGAIIDAE
MACROCHELIDAE, PARHOLASPIDAE
PACHYLAELAPIDAE, OMENTOLAELAPIDAE, LAELAPONYSSIDAE
PODOCINIDAE, HETEROZERCONIDAE
RHODACARIDAE
101
102
103
104
105
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26.
27.
ARCTACARIDAE
AMEROSEIIDAE
106
107
I
28.
29.
DIGAMASELLIDAE, PHYTOSEIIDAE
PHYTOSEIIDAE,OTOPHEIDOMENIDAE
108
109
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30.
31.
OTOPHEIDOMENIDAE, ZERCONIDAE
ASCIDAE
110
111 I
32. SPE LAEO RHYNCH IDAE 112
33. HALARACHN IDAE 113 I
34. ENTONYSSIDAE, RHINONYSSIDAE 114
35. RHINONYSSIDAE, DASYPONYSSIDAE 115 I
36. SPINTURNICIDAE, MACRONYSSIDAE 116
37.
38.
MACRONYSSIDAE, DERMANYSSIDAE
HYSTRICHONYSSIDAE, IXODORHYNCHIDAE, EVIPHIDIDAE
117
118
II
39.
40.
EVIPHIDIDAE, LAELAPIDAE
LAELAPIDAE
119
120
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41.
42.
DIARTHROPHALLIDAE, Uropodoidea, Trachytoide<l
PROTODINYCHIDAE, POLYASPIDAE
121
122
II
43.
44.
DISCOURELLIDAE, TRACHYTIDAE
THINOZERCONIDAE, DINYCHIDAE
123
124
I
45.
46.
EUTRACHYTIDAE, CI RCOCYLLIBANIDAE, COXEOUESOMIDAE
COXEQUESOMIDAE, PLANODISCIDAE, METAGYNELLIDAE
125
126 II
127
47.
48.
UROPODIDAE
CERCOMEGISTIDAE, KLiNCKOWSTROEMIiDAE 128 II
49. KLiNCKOWSTROEMIiDAE, FEDRIZZIIDAE, PARAMEGISTIDAE,

50.
HOPLOMEG ISTI DAE
MEGISTHANIDAE, DIPLOGYNIIDAE, SCHIZOGYN IIDAE
129
130
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51.
52.
CELAENOPSIDAE, ANTENNOPHORIDAE
EUZERCONIDAE, PARANTENNULIDAE
131
132
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Metastigmata I
53. IXODIDAE 139
54. IXODIDAE, ARGASIDAE 140 I
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Plate Page

I Prostigmata

55. ERIOPHYIDAE 183

I 56. NEMATALYCIDAE, DEMODICIDAE, CLOACARIDAE 184
57. CLOACARJDAE, SCUTACARIDAE 185
I 58. TARSONEMIDAE 186
59. PYEMOTIDAE 187
I 60.
61.
CROTALOMORPHIDAE, CRYPTOGNATHIDAE
RHAGIDIIDAE, HALACARJDAE
188
189

I 62.
63.
PENTHALODJDAE, EUPODIDAE
EUPODIDAE, PENTHALEJDAE, TENUIPALPJDAE
190
191

I 64.
65.
TENUIPALPIDAE,IOLINIDAE
8DELLIDAE, CUNAXIDAE
192
193

I 66.
67.
LABIDOSTOMMJDAE
PACHYGNATHJDAE, SPHAEROLICHIDAE
194
195

I 68.
69.
NANORCHESTIDAE, ALiCORHAGIIDAE
TERPNACARIDAE, LORDALYCHIDAE, PEDICULOCHELIDAE
196
197
70. PEDICULOCHELIDAE, PARATYDEJDAE, TYDEIDAE 198
I 71. TYDEIDAE, EREYNETIDAE 199
72. EREYNETIDAE, CAECULIDAE 200
I 73. CAECULIDAE, ANYSTIDAE 201
74. PTERYGOSOMIDAE, PSEUDOCHEYLIDAE 202
I 75. TENERIFFIIDAE 203
76. CHEYLETIDAE 204
I I 77.
78.
HARPYRHYNCHIDAE,OPHIOPTIDAE
OPHIOPTIDAE, HETEROCHEYLIDAE
205
206

I 79.
80.
MYOBIJDAE,SYRINGOPHILIDAE
PSORERGATIDAE
207
208

I 81.
82.
TETRANYCHIDAE
TETRANYCHIDAE
209
210

I 83.
84.
TUCKERELLIDAE, ALLOCHAETOPHORIDAE, L1NOTETRANIDAE
ALLOCHAETOPHORIDAE, TARSOCHEYLIDAE, POMERANTZIIDAE
211
212

I 85.
86.
RAPHIGNATHIDAE
STIGMAEIDAE, CALIGONELLIDAE
213
214
87. STIGMAEIDAE, CALIGONELLIDAE
I 88. NEOPHYLLOBIJDAE, EUPALOPSELLIDAE
215
216
89. ERYTHRAEIDAE 217
I 90. SMARIDIIDAE. CALYPTOSTOMIDAE 218

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Plate Page
91. JOHNSTONIANIDAE, TROMBIDIIDAE 219
92. JOHNSTONIANIDAE, TROMBIDIIDAE, TROMBICULIDAE 220
93. TROMBIDIIDAE 221
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94. TROMBICULIDAE, HYDROVOLZIOIDEA 222
95. HYDRYPHANTOIDEA,HYDRACHNOIDEA 223
96. HYDRACHNOIDEA,HYGROBATOIDEA 224
97. HYGROBATOIDEA
Astigmata
225
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98.
99.
ANOETIDAE
HEMISARCOPTIDAE
256
257
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100.
101.
L1NOBIIDAE, CANESTRINIIDAE
ROSENSTEINIIDAE, CHORTOGLYPHIDAE
258
259
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102.
103.
ACARIDAE
ACARIDAE, FUSACARIDAE
260
261
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104.
105.
HYADESIIDAE, LABIDOPHORIDAE
LABIDOPHORIDAE, SAPROGLYPHIDAE
262
263 I
106. SAPROGLYPHIDAE, CARPOGLYPHIDAE 264
107. CTENOGLYPHIDAE 265 I
108. CHAETODACTYLI DAE 266
109. CHAETODACTYLIDAE, GLYCYPHAGIDAE 267 I
110. GLYCYPHAGIDAE, LEMURNYSSIDAE 268
111. AUDYCOPTIDAE, YUNKERACARIDAE 269 II
112. PYROGLYPHIDAE, PSOROPTIDAE 270
113.
114.
PSOROPTIDAE
FREYANIDAE, LOBALGIDAE
271
272
II
115.
116.
FREYANIDAE
ANALGIDAE
273
274
II
117.
118.
ANALGIDAE, PROCTOPHYLLODIDAE
PROCTOPHYLLODIDAE, TURBINOPTIDAE
275
276
I
119.
120.
EPIDERMOPTIDAE, DERMOGLYPHIDAE
DERMOGLYPHIDAE
277
278
II
121. GASTRONYSSIDAE, CYTODITIDAE, LAMINOSIOPTIDAE 279
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122. PNEUMOCOPTIDAE, EWINGIDAE 280
123. CHIRORHYNCHOBIIDAE, RHYNCOPTIDAE, MYOCOPTIDAE 281
124. L1STROPHORIDAE, MYOCOPTIDAE 282
125. L1STROPHORIDAE, EVANSACARIDAE, KNEMIDOCOPTIDAE 283
126. KNEMIDOCOPTIDAE, TEINOCOPTIDAE 284 I
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I 127. SARCOPTIDAE 285

Cryptostigmata
I 128. Primitive and advanced shield states in the Cryptostigmata 304
129. Posteroventral shield states in the Cryptostigmata (PALAEACAROIDEA,
I 130.
NOTHROIDEA, PERLOHMANNOIDEA)
PALAEACAROIDEA, MESOPLOPHOROIDEA
305
306

I 131.
132.
PHTHIRACAROIDEA, PARHYPOCHTHONOIDEA
HYPOCHTHONOIDEA, PERLOHMANNOIDEA
307
308

I 133.
134.
PERLOHMANNOIDEA, NOTHROIDEA
NOTHROIDEA, GALUMNOIDEA, PELOPOIDEA
309
310

I 135.
136.
PELOPOIDEA, MICROZETOIDEA
ORIBATELLOIDEA, AMERONOTHROIDEA
311
312
313
I 137.
138.
PASSALOZETOIDEA,ORIBATULOIDEA
CERATOZETOIDEA, HERMANNIELLOIDEA 314
139. L10DOIDEA, POLYPTEROZETOIDEA 315
I 140. PLATEREMAEOIDEA, DAMAEOIDEA, ZETORCHESTOIDEA 316
141. ZETORCHESTOIDEA, CARABODOIDEA, CEPHEOIDEA 317
I 142. L1ACAROIDEA,OPPIOIDEA 318
319
143. OPPIOIDEA, HYDROZETOIDEA
I 144. EREMAEOIDEA 320

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I PREFACE

I Acarology, the study of mites and ticks, has experienced a meteoric growth during the
past 25 years. Recognition of the medical and agricultural importance of the Acari during
and immediately after World War 11 stimulated research both in applied and basic aspects of
I acarology throughout the world, broadening our understanding of the field and opening new
vistas for further exploration. While expansion has been particularly rapid and dramatic

I in the field of acarine systematics, no up-to-date textbook or manual devoted to the

recognition and identification of mite families is currently available. Certainly the
need for such a treatment exists among beginning students in acarology, as well as among

I those engaged in acarological research and teaching. The Manual of Acarology has been
written to fulfill this need.

I The composition of the Manual is primarily systematic, with the text designed to
provide only basic information on biology, morphology, collection and preparation.
References to useful works on these subjects are listed at the conclusion of each major
I section in the text. Keys include all the currently recognized families of mites except in the
water mite supercohort Hydrachnellae and the suborder Cryptostigmata, where only the

I superfamilial categories are distinguished. Diagnoses of suborders and of superfamilies are

included, along with a bibliography for each of these taxa. The Manual contains nearly
700 illustrations, most of which are original. Illustrations adapted from other sources are

I cited where applicable.

1 wish to thank my colleagues who so kindly provided specimens, advice, encourage-

I ment and criticism during the preparation of the Manual. All were essential to its final
completion. Special thanks go to my wife, Vida, for her help in proofreading both the
manuscript and final galley copy.
I G. W. Krantz

I Corvallis, Oregon
August 1, 1969

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I. INTRODUCTION

Few animal groups illustrate the enormous diversity in form, habitat and behavior seen
in the arachnid subclass Acari-the mites and ticks. Unlike other assemblages in the Class
I Arachnida, many representative groups of the Acari have evolved far beyond primitive pre-
dation. Some are exclusively phytophagous, while others include external and internal
parasites of both vertebrate and invertebrate animals. Acarines may be found in virtually
I any environment throughout the world including severe desert and tundra situations, moun-
tain tops and ocean floors. Mite species have colonized almost every terrestrial, marine and

I fresh water habitat known to man.

The high degree of diversity in habitat illustrated by the mites and ticks is no more

I astounding than is their range of form, size, structure and behavior. On the basis of these
variations, approximately 30,000 species of Acari, and more than 1,700 genera (Radford
1950), have been described to date. It has been estimated that up to a half million more
I species may exist today. Such an estimate seems well within reason when one considers
that random examination of virtually any organic substrate commonly yields mite specimens
representing undescribed species. Thus our knowledge of acarine systematics is little more
I than fragmentary, but no less so than is our information on ecology, life history and behav-
ior of these forms.

I The genesis of acarology as a discrete discipline may be traced to 18th and 19th cen-
tury Europe, but awareness of the Acari existed well before that time. A reference to "tick

I fever" was found on an Egyptian papyrus scroll dated 1550 B.C. Homer mentioned the
occurrence of ticks on Ulysses' dog in 850 B.C. while some 500 years later, Aristotle dis-
cussed a prostigmatid mite parasite of locusts in De Animalibus Historia Libri. Other early

I references to the Acari appear in the writings of Hippocrates, Plutarch, Aristophanes and
Pliny. Until about 1660, mites were referred to as "lice," "beesties" or "little insects."
Use of the terms "Akari" and "mite" originated about 1650.
I In 1735, Linnaeus used the generic name Acarus in the first edition of the Systema
Naturae for which he named the type, A. siro, in 1758. The tenth edition of the Systema
I included fewer than 30 mite species, all of which were grouped in the genus Acarus. In the
100 years that followed, major contributions to acarine systematics, particularly of higher

I categories, were made by Latreille, Leach, Duges, DeGeer and C.L. Koch. Michael (1884)
summarizes these and related pioneer works.

I The emergence of acarology as a modern science came in late 19th and early 20th
century Europe with the historic researches of Kramer, Megnin, Canestrini, Michael,
Berlese, Reuter, Vitzthum and Oudemans (Evans et al. 1961). Their work provided the
I basis for virtually all subsequent acarological research until the end of World War II.

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Among those who participated in the post-war awakening of American acarology

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were Drs. Edward W. Baker and George W. Wharton. Realizing the need for an up-to'
date basic text in acarology, Baker and Wharton pubiished their Introduction to Aca-
rology in 1952. The Introduction served as the standard systematics text for acarology
I
until 1958, when Baker and colleagues published the G~ide to the Families of Mites.
The Guide included many families not treated in the Introduction to Acarology, and
contained a number of major changes in higher categories.
I
Some outstanding reference works in acarology have be·m published subsequent to the
Guide to the Families (those of T.E. Hughes (1959) and of G.O. Evans and colleagues
I
(1961) are of special significance) as have many important research papers dealing with
specific problems in mite biology and systematics. Many of these works will be referred to I
in the sections to follow.
I
Useful References

Baker, E.W. and G.W. Wharton (1952). An Introduction to Acarology. MacMillan Co., New York:
I
465 pp. + xiii.
Baker, E.W.. J.H. Camin, F. Cunliffe, TA. Woolley and C.E. Yunker (195B). Guide to the Families of
Mites. Institute of Acarology Contr. No. 3:242 pp. + ix.
I
Berlese, A. (1882-190.3). Acari, Myriapoda, et Scorpiones hucusque in Italia reperta; fasc. 1-101. Padova.
Berlese, A (1899). Gli Acari agrarii. Riv. Pat. veg. 7:312-344.
Brennan, J.M. and E.K. Jones (eds.) (1968). A Directory of Acarologists of the World, 9th edition.
I
Rocky Mo. Lab., Hamilton, Montana: 131 pp.
Canestrini, G. (1891). Abbozzo del sistema Acarologico. Atti Istit. ven. 38:699-725.
DeGeer, C. (1778). Memoires pour servir a I'histoire des Insectes. Stockholm.
I
Duges, A (1839). Recherches sur I'ordre des Acariens. Ann. Sci. Nat. Zool. 1(2) :18-63.
Evans, G.O., J.G. Sheals and D. MacFarlane (1961), The Terrestrial Acar of the British Isles. Vol. J. Intro-
I
duction and Biology. British Museum (Natural History), London: ?19 pp.
Hughes, T.E. (19591. Mites or the Acari. University of London, Athlone Press: 225 pp. + viii.
Koch, C.L. (1842). Uebersicht des Arachnidensystems. Nurnberg: Fasc. 3.
I
Kramer, P. (1877). Grundzuge lOr Systematik der Milben. Arch. f. Natul"g. 2:215-247.
Latreille, P.A. (1806-1809). Genera Crustaceorum et Insectorum, Paris: 4 vols.
Leach, W.E. (1815). A tabular view of the external characters of four <:lasses of animals which Linne ar-
I
ranged under Insecta. Trans. Linn. Soc. London 11(2):306-400.
Megnin, P. (1876). Memoire sur I'organisation et la distribution zoologique des Acariens de la famille des
Gamasides. Paris.
I
Michael, AD. (1884-1888). British Oribatidae, Vols.I·II. Ray Society, L,ndon: 657 pp. + xvii.
Oudemans, AC. (1906). Nieuwe classificatie der Acari. Ent. Ber. 2:43-4<1. I
Oudemans, AC. (1926). Kritisch Historisch Oversicht der Acarologie 1:500 pp. + vii (850 V.C.-1758l.
E.J. Brill, Leiden.
Oudemans, A.C. (1929). Kritisch Historisch Oversicht der Acarologie 2:1097 pp. + xvii (1759-1804).
I
E.J. Brill, Leiden.
Oudemans, A.C. (1936-1937). Kritisch Historisch Oversicht der Acarolo)ie 3:3379 pp. + ci (1805-1850),
E.J. Bril/, Leiden. I
Radford, C.D. (1950). Systematic check list of mite genera and type species. Internat. Union Bio. Sci.
Ser. C (Ent.) 1:232 pp.
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Reuter. E. (1909). Zur Morphologie und Dntogenie der Acariden. Acta Soc. Sci. Fenn. 36(4):1·288.
Vitzthum. H.G. (1929). Acari. Die Tierwelt Mitteleuropas 3(7):1'112.
Vitzthum. H.G. (1931). Acari. Kiikenthals Handbuch der Zoologie 3(2):1·160.
Vitzthum. H.G. (1940·42). Acarina. Bronn's Klassen und Drdnungen des Tierreichs 5(4). Book 5:1·1011.

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II. SYSTEMATIC POSITION OF THE ACARI

The Phylum Arthropoda includes a myriad of forms which share the characteristics of
jointed legs and a chitinous exoskeleton. Within the Arthropoda is a group of animals
I which, unlike the insects or myriapods, have neither antennae nor mandibles. These are
the Chelicerata, of which the Class Arachnida makes up the largest part.

I The Arachnida comprises those chelicerates which may possess simple eyes (ocelli,

•
Plate 1-2), and which are primarily terrestrial forms. It includes such diverse forms as
scorpions, spiders, vinegaroons and the long-legged harvestmen so common to most
temperate and tropical regions. Of the eleven extant divisions of Arachnida listed by
Savory (1964) all but two are completely predaceous in habit, with representatives often

I displaying a variety of morphological characteristics well suited to a predatory existence.

The two pairs of mouthparts-the chelicerae and pedipalps (Plate 1-1, Plate 2-2,3) are
adapted for grasping, piercing, sucking or tearing. Specialized pedipalpal or chelicera I
I venom glands are present in some of these orders, while others have terminal venomous
stings (Plate 2-1) or acid glands.

I Although generally predaceous, members of the subclass Opiliones (Plate 1-1) have
been observed to feed on dead organic matter. They, along with the Acari, comprise the
exceptions to the rule of total predation in the Arachnida.
I The mites and ticks differ from most arachnids in that somatic segmentation generally
is inconspicuous or absent. 1 Abdominal segmentation is a primary attribute in all of the
I other subclasses except the Araneida-the spiders. Thus the mites, ticks and spiders may
easily be separated from other arachnids on this basis.

I lLimited secondary development of somatic segmentation may be seen in the TARSONEMIDAE

and PYEMOTIDAE (Plates 58, 59). Evidence of primary segmentation occurs in the Order Opilio·

I cariformes (Plate 13-11.

I CLASS SUBCLASS

I With abdominal-------
E
Scorpiones
Palpigradi

•
I ARACHNIDA
conspicuous
extension; with

Without abdominal
extension; without
sting
{ Palps
chelate

palPs
simp le or
Uropygi

Ricinulei

~ Schizomida
Amblypygi
Opiliones

I raptorial

Abdominal segmentation - - - - - - - - - - - - - - - l[
Solifugae
Araneae
A .

I inconspicuous or absent can

Fig. 1. Key to arachnid subclasses.

I
I
6
I
I
•
Mites and ticks may be distinguished from spiders through the use of the following key.

1. Mouthparts inserted anteriorly on the cephalothorax (Plate 1·2), which is comprised

of fused head and thoracic segments; legs also borne on cephalothorax, which is con·
nected to the abdominal portion by a narrow pedicel Subclass ARANEAE

2. Mouthparts contained in a discrete anterior gnathGsoma (Fig. 2); portion of the

I
body on which the legs are inserted (the podosoma) broadly joined to the portion
of the body behind the legs (the opisthosoma) to form the idiosoma .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subclass ACARI
I
I
I
gnathosoma

\\lIIc--'-'~--epistome
I
I
_~circumcaPitular suture
"J"/""

<\.i~:;\ I
propodosoma
. '"'\""\ I
\, \ tl\
I
,TI, .:~ podosoma

'.1 ;':, \J I
I
hysterosoma
"i idiosoma

j··1 ,\ Jf - I
~.\ )/ I
A~ r' ~'Z~\ opisthosoma

f..,~ r f /'}1
'r,,'>--r • • 1/;' I
-----------~Ik4--·-----
I
Fig. 2. Dorsum of Macrocheles merdarius Berlese (Subclass Acari) sh:>wing major body divisions.
I
•
I
I PLATE 1
7

I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I 1-1; subclass Opiliones (Brazill, dorsum: 1-2; subclass Araneae (Mexico), dorsum (leg extremities removed)

I
 I
8
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
•
I
I
~:2'and 2-3; subclass
. es (Africa), dorsu~2', palp:
l' subclass scorPJ~~eudoscorPiones. 2.3; chelicera
•
I
I 9
I While lack of comprehensive fossil evidence does not permit a phylogenetic evaluation
of the Arachnida in general, or of the Acari in particular, the development of a wide range
I of non-predaceous habits in the Acari indicates that it is a highly evolved group. Specializa-
tion in habit also is correlated with a high degree of morphological diversity not seen in
other arachnids. Basically, the Acari most closely resemble the arachnid subclass Opili-
I ones, although Sharov (1966) feels that the mites evolved from pedipalpid ancestral stock
during the middle Devonian. Grandjean (1935). Zakhvatkin (1952) and others are of the

I opinion that the Acari are of diphyletic origin, with the order Acariformes having a different
derivation than the Parasitiformes and related forms (Fig. 12). Literature on phylogenetic
theory pertaining to the Acari has been reviewed by Woolley (1961).

I Useful References

I Bekker, E.G. (1959). Concerning the Acarina as a natural grouping. Trud. Inst. morf. zhiv. 27:151-162.
Bekker, E.G. (1960). Systematics and comparative anatomy in solving the probiem of the phylogeny of
I ticks and mites (Acarina). Rept. No.1. Critical discussion of the views of acarologists-taxonomists
on the polyphylogeny of the order Acarina. Vest. Mask. Univ. Ser. 6: BioI. 15(4):13·20.
Carpenter, F.M., J.W. Folsom, E.O. Essig, A.C. Kinsey, C.T. Brues, M.W. Boesel and H.E. Ewing (19371.

I Insects and arachnids from Canadian amber. Univ. Toronto Studies, Geol. Ser. 40:7-62.
DUbinin, V.B. (1959a). Chelicerate animals (subphylum Cheliceropha W. Dubinin nom. n.1 and their
systematic position. Zool. Zh. 38(8) :1163·11 B9.

I Dubinin, V.B. (1959b). Phylogenesis of chelicerate animals of the subphylum Chelicerapha W. Dub. and
affinity of Chelicerata with pycnogonides. Trud.lnst. marl. zhiv. 27:134·150.
Grandjean, F. (1933). Observations sur les Acariens (2e Serie). Bull. Mus. Hist. Nat. (Paris), ser. 2,

I 7:201·208.
Hammen, L. van der (1964). The relation between phylogeny and post'embryonic ontogeny in actina·
trichid mites. Acarolgia fasc. h.s.; 1st Congo Acar" Ft. Collins: 85·90.

I Hammen, L. van der (1968). Introduction generaIe a la classification, la terminologie morphologique,

I'ontogenese et I'evolution des Acariens. Acarologia 10(3):401·412.
Petrunkevitch, A. (1949). A study of palaeozoic Arachnida. Trans. Connecticut Acad. Art. Sci.

I 37:69·315.
Savory, T. (1964). Arachnida. Academic Press, London: 291 pp. + viii.
Sharov, A.G. (1966). Basic Arthropodan Stock with Special Reference to Insects. Pergamon Press,
I Oxford: 271 pp. + xii.
Snodgrass, R.E. (19381. Evolution of the Annelida, Onychophora and Arthropoda. Smithsonian Misc.
Call. 97(6):159 pp.

I Tiegs, O.W. and S.M. Manton (1958). The evolution of the Arthropoda. BioI. Rev. 33:255-337.
Woolley, T.A. (1961). A review of the phylogeny of mites. Ann. Rev. Ent. 6:263-284.

I Zakhvatkin, A.A. (1952). Division of the Acarina into orders and their position in the system of the
Chelicerata. Mag. Parasitol. Moscow 14:5-46.

I
I
I
I
I
I 11
I III. MORPHOLOGY AND FUNCTION

I A. EXTERNAL

I The exoskeletal envelope of the typical acarine begins its development as undifferen-

I tiated tissue covered by a thin layer of cuticulin and separated from the underlying epider-
mis by an extremely thin, poorly defined granular Schmidt layer (Fig. 3). As development
proceeds, surface portions of the undifferentiated layer often become sclerotized to varying

I degrees through orthoquinone tanning. These portions, the epicuticle and exocuticle, may
then be distinguished from the underlying laminated layer, the endocuticle, as discrete
shields or plates (Belozerov 1960). The cuticulin surface layer may contain a profusion of
I micropores which are connected to pore canals. These canals appear to arise from the
epidermal cells underlying the Schmidt layer, and pass through both the endo- and exocutic-
ular layers (Wharton et al. 1968). A possible function of the pore canals may be to
I
I cement layer }
tectostracum epicuticle
cuticulin

I
;====f:dI~2;J---pore canal
I ~//::-==::--~~\

I exocuticle

I
I \...,,,*
endocutic!e
Schmidt layer
__- - _ epidermis

I
Fig.3. Diagrammatic cross section of acarine cuticle.

I
I transport an epidermal secretion to the cuticulin surface layer, where the secretion forms a
protective waxy coating referred to as a tectostracum, as well as a thin overlying cement
layer. The tectostracal and cement layers offer protection against excessive water loss from
I the body surface. Other pores occur in smaller numbers both on the body and appendages
of Acari. These structures may also have a secretory function, since they too are connected
internally to canals. Round, elliptical or Iyriform pores often occur dorsally, while only
I Iyriform openings, or Iyrifissures, are found on the venter or on appendages.

•
I
12
I
The body of the typical acarine is composed of a discrete anterior gnathosoma and
posterior idiosoma (Fig. 2). These are separated by a circumcapitular suture, a separa-
I
tion similar to that between the head and thorax of insects.
I
Gnathosoma
I
The gnathosoma (Gorirossi 1956, van der Hammen 1968b) resembles the head of the
generalized insect only in that the mouthparts are appended to it. The brain lies in the
idiosoma behind the gnathosoma rather than within it, while the eyes (when present) are
I
found dorsally or dorsolaterally on the propodosoma (Plate 13-1). The gnathosoma, then,
is little more than a tube through which food is carried into the esophagus. The roof of the
tube is called the epistome, and the lateral walls are made up of the fused coxae of the pedi-
I
palps, or palps (Plate 3-1). The floor of the gnathosomal tube is the subcapitulum. The
anterior coxal endites, or lateral lips (Knulle 1959) of the sUbcapitulum, along with associ-
ated anteroventral structures, is called the hypostome (Plate 3-2). Lying above the buccal
I
cavity are the paired chelicerae (Plate 3-4) which generally are three-segmented. The
chelicerae, along with the pedipalps, comprise the organs of food acquisition. I
The palps may be simple sensory structures equipped with chemosensory or thigmo-
tropic hairs which aid the acarine in locating its food (Plate 4-4). They are, however, often
I
modified into grasping or piercing raptorial organs similar to the mandibles of many
predatory insects (Plate 4-1). The number of free palpal segments vary from one or two
(most Astigmata, some Prostigmata) to five (many Mesostigmata and Cryptostigmata).
I
Variations in pedipalpal chaetotaxy in the Mesostigmata may be useful in determining re-
lationships between higher taxa, and in separating immature stages of single species (Evans
1963b).
I
The chelicerae vary considerably in structure, but are never primarily sensory. I
Generally the terminal third segment of the chelicera is modified into a movable digit which
opposes the fixed distal portion of the second segment. These opposed digits, or chelae,
are edentate or toothed for grasping or grinding. The chelicera may be attenuate and elon-
I
gate in some parasitic groups, serving as piercing organs (Plate 4-6), Reduction of the fixed
digit is not uncommon in predaceous and phytophagous species, with a concomitant
development of the movable digit into a stylettiform piercing structure (Plate 4-5,7), In
I
some acarine groups, the movable chelicera I digit of the male is modified to effect sperm
transfer to the female (Plate 5-1). Other modifications of chelicera I structure are referred
to in the keys to families.
I
The buccal cavity opens internally into a strongly muscled pharynx which acts as a I
suction pump for the ingested food materials (Blauvelt 1945). The pharynx is served by
several sets of muscles which, along with those muscles which control the movement of the
chelicerae and paJps, virtually fill the gnathosomal cavity (Mitchell 1962). Salivary glands
may be present, opening through paired ducts into the buccal cavity or through styli
somewhat anterior to the oral opening (Plate 3-1). These glands supply the various
enzymes which allow pre-oral digestion of food materials (Moss 1962).
••
I
I
I
I PLATE 3

•
13

I
••
I
• --+----I-i--deutosternum

•
3-2

tritosternal

I
lacinia

I•
I _ .......-.....lk.J.eupathidium

I
'.
J
1
1
'".. 3-1; Holostaspel/a berlesei Krantz {Mesostigmata, MACROCHELIDAEl. dorsum of gnathosoma: 3-2; H. berlesei, venter of
gnathosoma: 3-3; Opilioacarus sp. (Notostigmata. OPI L10ACARIDAE), venter of gnathosoma: 3-4; Glycyphagus sp.

/1 (Astigmata, GL YCYPHAGIDAEI, venter of gnathosoma

 PLATE 4
14 I
I
I
4-3
I
I
I
I
I
I
•
I
4-6
I
I
I
•
I
4-5
I
I
I
4-1; Cheletomorpha lepidopterorum (Shaw) (Prostigmata, CHEYlETIDAE), dorsum of gnathosoma: 4-2: Pionopsis lutescens
(Hermann) (prostigmata, HYGROBATOIDEA), palp: 4-3; Agaue sp. (Prostigmata, HALACARIDAE), palp: 4-4; Veigaia
sp_ (Mesostigmata, VEIGAIIOAE), palp: 4-5: Opsereynetes sp. (Prostigmata, EREYNETIOAEI, venter of chelicerae:
4.-6; Hystrichonyssus turned Keegan et af. (Mesostigmata, HYSTRICHONYSSIDAE), chelicera: 4·7; Brevipalpus cardinalis
I·
(Banks) (Prostigmata, TENUIPALPIOAE), gnathosoma

I
I PLATE 5
15

I sperm transfer organ

I
I 5-2
I 5-1

I
I -::7'-:=I1tl..... 'empodium---l~
'W\u..~--c1ew .....,...

I
I
I
I
5·4
I
I
empodial claw

I
I true
claw

I remnant

I
I
I
5-1; Parholaspe/la spatulata Krantz (Mesostigmata, PARHOLASPIDAE, chelicera of male: 5-2; ?genus (Mesostlgmata,

I DIPlOGYNIlDAEl. chelicera of female: 5-3;Pygmephorus sp. IProstigmata, PYEMOTIDAEl. tarsus II: 5-4; Ricardoella

•
limacum (Schrank) (Prostigmata, EREYNETIDAE), tarsus II: 5-5; Tuckerella sp. (Prostigmata, TUCKERElllDAE),
tarsus II: 5-6; Oligonychus sp. (Prostigmata, TETRANYCH IDAEl. tarsus III: 5-7; Heterocheylus sp. (Prostigmata.
HETEROCHEYlIDAEl. tarsus II: 5-8; OPPIOIDEA ICryptostigmatal. tarsus II
16 PLATE 6 I
I
I
I
:§:g
I
I~ I
I
I
I
I
I
I
I
I
.---~ I
I
I
I
I
6.1 to 6-9; common dorsal shield configurations in the Acari. 6-1,2,3,4,6; Mesostigmata: 6-2,7,8; Prostigmata: 6·5; Astigmata: •

•
6-9; Cryptostigmata
I 17

I Idiosoma

I The idiosoma of the Acari assumes functions parallel to those of the abdomen, thorax
and portions of the head of insects. It may be covered with heavily sci erotized shields or it
may be soft and virtually without sclerotization. The great diversity in shape, size and orna·

I mentation of the idiosoma is evident in the illustrations which accompany the keys to
families.

I Although the idiosoma is considered to be undivided, various grooves and furrows may
occur in those groups where extensive idiosomal sclerotization has not occurred (Plate 58-1).
These grooves may indicate primitive idiosomal divisions which have been lost or obscured
I in the majority of acarine forms.

I The idiosoma includes an anterior propodosoma and a posterior hysterosoma which

mayor may not be separated from each other by a sejugal furrow (Plate 99-4). The anterior
two pairs of legs are inserted ventrally in the propodosomal region, while legs III-IV are

I located on the hysterosoma. The prodorsal region may be separated from both the hystero-
soma and the venter of the propodosoma in the Notostigmata by a disjugal furrow. The
portion of the hysterosoma behind the insertions of legs 111-1 V, or the opisthosoma, may be
I delimited from the anterior portion of the hysterosoma by a postpedal furrow (Plate 70-4).

Shields or platelets commonly cover portions of the idiosoma. An anterior shield may
I cover the prodorsum or the entire propodosoma (Plate 6). A posterior shield or series of
shields also may be present, while in some groups a single dorsal shield covers virtually the

I entire idiosoma. Ventrally, the idiosoma may be divided by sejugal and/or postpedal fur-
rows, and mayor may not be provided with shields. The genital and anal orifices usually
are set within a sclerotized shield or protected by paired valves (Plate 66-1,5). The genital

I and anal shields may, in many instances, be expanded so as'to cover all or nearly all of the
genital-anal region (Fig. 14, page 68). Anteriorly, a sclerotized sternal area may be found
which often is incorporated into an overall ventral shield complex.

I The primary external organs found on the idiosoma are locomotory, respiratory,
copulatory and sensory in function.
I
Locomotory
I With few exceptions, adult and nymphal Acari possess four pairs of jointed legs,

I while the larva has three pairs. The most posterior pair appears with the first nymphal
instar. The legs are divided into seven primary segments: the coxa, trochanter, femur,
genu, tibia, tarsus and apotele (Fig. 4). Secondary division of the trochanter may occur

I (Plate 13-1), and tarsal and femoral sutures appear in some groups, giving the impression
of even further division. The tarsus usually is terminated by a claw-like or sucker-like
ambulacral apotele although it is not uncommon to find tarsus I armed only with terminal

I sensory setae. The apotele often is a complex of paired claws and a median pad-like or
claw-like empodium (Plate 5-3,4). The empodium may persist in the absence of true
claws as a claw-like or sucker-like pretarsal extension (Plate 5-7,8). In some groups one of
I the true claws may be lost, with the empodium assuming its position, structure and

I
•
18
I
function (Plate 92-2). While leg I usually serves an ambulatory function, it is often elon-
gated or antenniform, and is used as a sensory structure (Plate 24-2). Legs I of members of
I
other families are more or less raptorial, and serve to grasp prey (Plate 73-1).
I
The legs may be armed with ridges or spurs and usually are ornamented with a series
of tactile and sensory hairs, or setae (Fig. 4). These setae generally follow fixed patterns of
insertion in a given species, not only in position but also in number. The setal distribution
I
on any given leg segment may be reduced to a formula which often is useful in establishing
systematic relationships (Fig. 4) (Evans 1963a, Coineau 1964). I

='N
,,~~=,-/p ant. post.
I
I
(0 I
.:
11/ • 92 a~2
I
~',0 --
femur av 2 2
1- -,--0
\ a1
1 0 I
OJ
genu
S?: pd l J I
tibia

tarsus
al
aV~2~V
~;. pl- 1- -'--1
2 1
,
aJ-illI, Pd_ pJ
av pV I
I
I
a I: <0 1 1
: pil
I
I
I

distal
I
I
Fig. 4. Leg IV of Macrochelesplumosus Evans and Hyatt, with leg chaetotactic formulae expressed for the
I
I
genu and tibia.

I
I
I 19
I Respiratory

I Exchange of oxygen and carbon dioxide in the Acari is accomplished in several ways
which are so diverse as to rule out any theory of single line evolution of respiratory systems.
The presence or absence of spiracular openings, and the relative position of these openings
I when present, provides a major diagnostic feature for the separation of orders within the
Acari. Thus the position of the spiracles, or stigmata, on the lateromedian or mesal aspect

I of the idiosoma characterizes the members of the suborder MESOstigmata (Fig. 5c).

I
I stigmata
dorsal

I
I
I
I
I
I
Fig. 5. External respiratory and associated structures in the Notostigmata (5a), Tetrastigmata (5b) and

I Mesostigmata (5c).

Stigmatal openings in the PROstigmata are found at or between the bases of the chelicerae
I or on the "shoulders" of the idiosoma, (Figs. 6b,c; 7a,b) while stigmata are hidden in the
coxal regions of the higher CRYPTOstigmata (Fig. 7c). Stigmata are absent in the
Astigmata, in the region of coxae IV in META stigmata (Fig. 6a), and on the dorsum of the
I idiosoma in NOTOstigmata (Fig. 5a). A pair of lateral pores resemble true stigmata in the
TETRA stigmata (Fig. 7b, Plate 14-3), creating the impression of four stigmatal openings.

I The stigmata in the Parasitiformes may be surrounded by a stigmal plate (Fig. 6a),
or they may each be associated with an elongate anteriorly directed groove of questionable

I function which is inappropriately called the peritreme (Fig. 5c). The peritremes may serve
as extensions of the stigmata, thus ensuring that blockage of the stigmata will not necessar-
ily interfere with respiration (Radovsky 1968). Where stigmata occur, they open internally

I into a tracheal system which ramifies throughout the body to the various organ systems.
In those mites which have no apparent stigmata or tracheal system (the Astigmata and a

I
I
20
I
stigmata between
chelicerae I
I
I
I
I
I
I
I
Fig. 6. External respiratory and associated structures in the Metastigmata (6a). Prostigmata·Tarsonemina
(6b) and Prostigmata·Parasitengona 160). I
I
I
I
I
I
I
I
b I
Fig. 7. External respiratory and associated structures in the Prostigmata-Eleutherogonina (7a,7b) and the
Cryptostigmata (70),
I
I
I
I 21

I few families in other suborders), exchange of oxygen and carbon dioxide is thought to occur
through the integument. Lack of stigmata does not always connote lack of a respiratory
I system. Certain families in the Cryptostigmata possess what are believed to be tracheae
associated with the paired dorsal pseudostigmata and with the acetabula of legs I and III.
Genital tracheae occur in some of the Astigmata and Prostigmata.
I
Copulatory
I Fertilization methods in the Acari are easily as diverse as are the respiratory systems

I discussed above. While copulation in the usual sense is not uncommon in the Acari, sperm
transfer may be accomplished in a variety of other ways.

I In those mite groups where a male intromittent organ, or aedeagus, is present

(Plate 7) transfer of sperm may be made directly to the female genital opening (various
families of Prostigmata and Astigmata) or to a special female copulatory structure called the
I bursa copulatrix (Plate 8-1). The bursa, which may be seen in a number of astigmatid mite
families, may be an extrusible terminal tube on the idiosoma which is connected internally
to the female reproductive system or a posterodorsal opening, as in some of the analgoid
I feather mites (Popp 1967).

A type of "bursa" also may exist in various families of the Mesostigmata, although the
I male has no aedeagal transfer organ (Plate 8-4). Unlike the single posteromedian bursa of
the Astigmata, a pair of copulatory orifices is present in the female, one located ventrally

I between coxae III and IV on each side of the body. Sperm is transferred from the male
genital aperture to the sperm transfer organ on the chelicera (Plate 5-1) which then is
appressed to one or both of the female bursae (Costa 1966). Little is known of how the

I passage of sperm to the female ovary is effected, but it is likely that the system is analogous
to that seen in the Astigmata. Direct transfer of a sperm packet from the male to the female
genital aperture in the absence of an intromittent organ or a cheliceral sperm transfer organ
I occurs in the mesostigmatid family UROPODIDAE. Here a complex pre-mating ritual is fol-
lowed by venter-to-venter contact, with the packet being affixed to the anterior edge of the
epigynial shield. Assimilation of the contents of the packet occurs soon afterwards
I (Radinovsky 1965). Transfer of a sperm packet to the female genital opening by means of
the chelicera or other gnathosomal structure occurs both in the Mesostigmata and the
Metastigmata.
I The method of sperm packet transfer in certain of the Prostigmata and Cryptostig-

I mata differs from those discussed above in a peculiar and interesting way. The sperm
packet is placed on a stalk which is secreted as a fluid thread by the male, and which
hardens on contact with the air (Plate 8-6) (Mitchell 1958). This spermatophore supports
I the sperm packet which is deposited by the male and which may be picked up by the
female upon contacting it with her genitalia (Lipovsky et al. 1957). Spermatophore transfer
of this type also occurs in other arachnids, including scorpions and pseudoscorpions.
I
I
I
I
 PLATE 7 I
22
I
•
I
I
I
I
I
I
_ _ _ _ aedeagus -I 7-3
7-2
I
7-1
I
I
I
aedeagus

I
I
I

7·5
••
aedeagal articulation
I
7-1 to 7-7; male genital apparatus_ 7-1; Brevipalpus sp. (Prostigmata, TENUIPALPIDAE): 7-2; Eotetranychus carpini I
•
(Oudemans) (Prostigmata, TETRANYCHIDAE): 7·3; Harpyrhynchus sp. (Prostigmata, HARPYRHYNCHIDAE):
7.4,5; Glycyphagus destructor (Schrank), lateral and ventral aspects: 7·6; ?genus (Prostigmata, EUPALOPSELLIDAE)
I PLATE 8
23

I -i--seminal
, receptacle

I
I
I
I
8-'
1
I
I
I
I
I
I
I
I
I
I
I
I
I 8-6

I 8-1; Glycyphagus domesticus (DeGeer) (Astigmata, GLYCYPHAGIDAEl. posteroventral aspect of female: 8-2; Eremaeus sp.
(Cryptostigmata, EREMAEOIDEA), propodosoma: 8-3; 'Phthiracarus sp_ ICryptostigmata, PHTHIRACAROIDEA),
lateral aspect of larva: 8-4; Macrocheles merdarius (Berlese) (Mesostigmata, MACROCHELIOAE), sacculus foemineus and

I related structures: 8-5; Nanorchestes sp. (Prostigmata, NANORCHESTI DAE), propodosoma: 8-6; stalked spermatophore
and sperm packet (diagrammatic) of a trombiculid mite (Prostigmata, TROMBICULIDAE)
24
I
Sensory I
The idiosoma is well equipped with various sensory receptors, almost all of which are
setal. Those setae which occur on the idiosoma proper ,are primarily tactile structures, the
movement of which activate nerve cells located at the base of each seta. Tactile setae may
I
be simple, plumose or leaf-like but all lack protoplasmic extensions into the body of the seta
itself. A second type of seta is found dorsally in certain groups, but only one or two pairs I
occur, These setae are termed trichobothria (Plate 8-5), and differ from the simple tactile
setae in having a protoplasmic nerve cell extension into the seta. Trichobothria'probably
are chemo-receptors, and assume a variety of shapes on the idiosoma. They are called
I
pseudostigmatic organs in the Cryptostigmata, and may be simple, feathered, clavate or
bulbous in this group (Plate 8-2), In addition to setal receptors, one or two pairs of simple
eyes may be found laterally on the propodosoma in members of all suborders except the
I
Mesostigmata. The larvae of various Prostigmata and Astigmata, and all of the observed
Cryptostigmata; possess paired Claparede organs or urstigmata (Grandjean 1946) ventrally
between coxae I and II (Fig. 8, Plate 8-3). These variously shaped structures are thought to
I
be humidity sensors. Nymphal and adult mites in these groups have paired genital discs
(Plate 7-4) which probably serve the same function (Knulle 1959). I
The legs of acarines are provided
with many tactile setae (Grandjean
I
1935a; 1964), and legs I and /I often
have trichobothria as well. Where they
occur on the legs, trichobothria are
I
simple hairlike structures rather than
ornate as are the pseudostigmatic or-
I
gans of the idiosoma in many cases.
Other setae related to trichobothria-
the eupathidia and solenidia {Plate
I
3-4)-may be found on legs I and /I,
and on the pedipalpal tarsus. The I
famolus (Plate 104-4) is a terminally
expanded or spinose seta with a proto-
plasmic core which is found only on
I
tarsus I of various mites. Sensory
setae of four different types are found I
in the Haller's organ of tarsus I of
the Metastigmata (Plate 53-3) (Arthur
1956). A similar structure is found
I
in the Tetrastigmata and in the Noto-
stigmata (van der Hammen 1968a), I
Fig. 8. Venter of the larva of Calyptostoma sp. showing
the position of the urstigmata.
I
I
I
I
I 25
I Trichobothria, eupathidia and famuli contain a layer of optically active chitin, or
actinochitin, around the protoplasmic core which exhibits birefringence in polarized
I light (Grandjean 1935a). Other setae with no protoplasmic extension may have a solid
actinochitinous core. Mite groups which possess optically active setae (the Prostigmata,
Astigmata and Cryptostigmata) are classified together by some authors as the Actino-
I chitinosi, the Actinotrichida or the Actinochaeta. The Mesostigmata, Notostigmata,
Tetrastigmata and Metastigmata are grouped by these authors in the category Anactino-

I chitinosi, Anactinotrichida or Anactinochaeta (Grandjean 1935b, van der Hammen 1961,

Evansetal.1961).

I B. INTERNAL

I Internally, the idiosoma is a surprisingly complex series of organ systems bathed in a

colorless plasma of indeterminant composition. The plasma moves freely throughout the
body cavity, or haemocoel, and is often circulated by a flat ostiate heart located antero-
I dorsally. Circulation to the legs and other extremities may be facilitated by the flexing of
strong dorsoventral muscles which contract the haemocoel and force the plasma to move.
Extension of the legs also is brought about by dorsoventral contraction and subsequent
I hydraulic pressure exerted on these appendages (Manton 1958).

I The major internal organ complexes of the idiosoma are the digestive, reproductive and
nervous systems.

I Digestive

I While the post-oral digestive system varies somewhat in the different suborders, certain
features are more or less constant (Plate 9). The buccal cavity opens internally to the
pharynx, as described under the section on the gnathosoma. The pharynx opens posteri-
I orly into an esophagus which passes through the brain. The esophagus leads into a ventricu-
lus which may have two or more gastric caecae (Blauvelt 1945, Hughes 1952). The caecae

I apparently provide additional surface for digestive processes to take place. A short intes-
tine joins the ventriculus to a hindgut, on which one or two pairs of Malpighian tubules
may be inserted. The hindgut generally leads to a rectal cavity which opens externally at

I the anus. In certain Prostigmata and Metastigmata, the connection between the ventriculus
and hindgut has been IQst. Where this occurs the hindgut, along with the anteriorly directed
excretory canal, serves as a separate system for collecting guanine which is passed through a

I ventral uropore. The fate of solid byproducts of digestion in the ventriculus where the
hindgut has been lost, is bizarre in adults of some of the higher Prostigmata (Parasitengona).
Here the indigestible residues accumulate in the gut cells, which break away and move into
I the posterodorsal gut lobes (Mitchell 1964). When a lobe has become filled with gut cells it
loses its connection with the gut proper and, through a process known as schizeckenosy
(Mitchell and Nadchatram 19691, passes through a horizontal split of the posterodorsal
I cuticle. The split soon heals, leaving a scar which indicates that schizeckenosy has occurred.

I
I
I
 PLATE 9 I
26
cheliceral retractor
muscle
I
I
I
I
I
I
I
I
I
I
I
I
~_ _...._;lC=;;;::=-+-_ventriculus

I
f---j'--++-- accessory gland
uterus'---+--+-- I
Malpighian tubule
intestine ---4-----'\..-'1--
r-+--+++- oviduct
I
hindgut ---\-----4-**
9-2
I
ovary _ _-..l...... -'-_ _"il
I
....
~ ~tJ.::::-------- anus
I
9-1; Caminella peraphora Krantz and AinscQugh (Mesostigmata, TRACHYTI DAE), midsagittal section (after Ainscough 1960):
9·2; C. peraphora, dorsal internal aspect (after AinscQugh 1960)
I
I
I PLATE 10
27

I
I
II ~'"'-"..
testes

I unpaired
10-1

I
I
I
I
paired ovaries

I
;£)
uterus

I
paired

I
bursa
copulatrix oviducts 10-5
oviduct
10-6
I genital disc

I
I
I
I
I
I
10-1 to 10-6; diagrammatic representations of reproductive systems (after Hughes 1959). 10-1; PARASITI DAE (Mesostigmata),
I male: 10-2; UROPODIDAE (Mesostigmata), male: 10-3; ERYTHRAEIOAE (Prostigmata), male: 10-4; generalized
Mesostigmata and Prostigmata, female: 10-5; ACARIDAE (Astigmata), female: 10-6; ERIOPHYIDAE, HALACARIDAE,
ANYSTIDAE (Prostigmatal, female

I 10-7 and 10·8; ovipositoni. 10-7; Nanorchestes sp. (Prostigmata, NANORCHESTI DAE): 10-8; Nanhermannia sp. (Cryptostigmata,
NOTHROIDEA)
28
I
The evolution of a blind gut in the Parasitengona may be the result of a highly efficient I
pre-oral digestive capacity which allows absorption of virtually all ingested material in the
midgut. This in turn might obviate the need for a hindgut as a voiding route for fecal
material, with the minute quantities of fecal residue being stored permanently or eventually I
eliminated through schizeckenosy. The role of organ system function and arrangement in
the evolution of the blind gut of certain parasitengonid mites is discussed in a forthcoming
paper by Mitchell (in press).
I
Reproductive
I
The reproductive system of both male and female acarines consists of a series of
paired, fragmented, or fused elements. In the male, the testes may be an unpaired organ as
I
in many of the lower Mesostigmata (Plate 10-1) and the Metastigmata (Roshdy 1966), a
paired structure in the mesostigmatid family UROPODIDAE (Plate 10-2) and the astigmatid I
ACARIDAE (Rohde and Oemick 1967), or a multiple organ in some of the higher Prostig-
mata (Plate 10-3). The testes manufacture sperm cells which are passed through the paired
or fused vas deferens to the ejaculatory duct. Various accessory glands occur between the
I
duct and the vas deferens. It is probable that one of the functions of these glands is that of
a seminal vesicle. I
The female reproductive system has a single or paired ovary (Plate 10-4,5,6) which
passes to the oviduct, or oviducts. An unpaired uterus is found both in the Mesostigmata
and Prostigmata (Plate 10-4,6). The uterus opens into a vagina which may be median or
I
posterior on the venter of the idiosoma. A separate uterus is not apparent in some of the
Astigmata (Plate 10-5). Where a bursa copulatrix occurs, the bursa opens into a seminal
receptacle which is connected to the ovary (Plate 10-5). A pair of small, club-shaped pro-
I
tuberances has been observed in an internal fold at the posterior edge of the seminal
receptacle in members of the family G LYCYPHAG IDAE (Plate 8-1). Their function is I
unknown.

In those mesostigmatid mites where coxal insemination appears to occur (see page
I
21), each of the paired external orifices is connected to a vestibule which opens internally
into a tube (Plate 8-4). The vestibule is termed the sacculus vestibulus (Krantz and
Mellott 1968) while the tube into which it empties is the tubulus annulatus. It is be-
I
lieved that sperm deposited in the sacculus vestibulus by the male passes through the
tubulus to a medial sacculus foemineus (Michael 1892). This structure terminates in
the appended cornu sacculus or spermatheca which, in Haemogamasus ambulans
I
(LAELAPIDAE), joins the ovary posteriorly (Young 1968).
I
Nervous
I
Lying around the esophagus in the anterior portion of the idiosoma is a well devel-
oped central nervous system consisting of a brain and a series of radiating nerves (Plate
I
9). The legs, digestive system, musculature and genitalia appear to be innervated by
nerves arising on the subesophageal portion of the brain (Winkler 1888, Ainscough 1960, I
I
I
I 29

I Moss 1962). The mouthparts receive impulses from the dorsal ganglia. The structure and
extent of the peripheral nervous system as a receiver of stimuli from surface receptors has
I not been studied.

I Useful References

I Ainscough. B.D. (1£1601. The internal morphology of Caminel/a peraphora Krantz and Ainscough. with
descriptions of the immature stages. Oregon State University Thesis LD 4330:53 pp.
Arthur, D.R. (195EiI. The morphology of the British Prostriata with particular reference to Ixodes
hexagonus Leach. III. Parasitol. 46:261-307.
I Balashov, I.S. (195B). The excretion processes and activity of Malpighian tubes of the ticks.
Sborn Zool.lr,st.. Akad. Nauk USSR. 18:120-128.
Parazit.

I
Belozerov, V.N. (1960). Structure of the integument ofgamasid mites (Parasitiformes, Gamasoidea).
Ent. oboz. 3914):850-859.
Blauvelt, W.E. (1945). The internal morphology of the common red spider mite (Tetranychus te/arius
Linn.). Corneil Univ. Agr. Exp. Sta. Memoir 270:35 pp. + 11 plates.
I Clifford. C.M. and G. Anastos. (1960).
Jour. Parasitol. 46(5):567-578.
The use of chaetotaxy in the identification of larval ticks.

a I'etude
I
Co;neau, Y. (1964). Contribution des Caeculidae. Premiere serie: Developement postlavaire
de A/locaecu'us catalanus Franz 1954.·0euxieme partie: La chetotaxie des pattes. Acarologia
6(1 ):47-72.
Costa, M. (1966). Notes on macrochelids associated with manure and coprid beetles in Israel l. Macro-

I che/es robustulus (Berlese 19041. development and biology. Acarologia 8(4):532-548.

Dosse, G. (19581. Die Spermathecae, ein zusatzliches Bestimmungsmerkmal bie Raubmiiben (Acar.,
Phytoseiidae). Pflanz. Ber. 20(1·2):1-11.

I Douglas, J.R. (1943), The internal anatomy of Dermacentor andersoni Stiles. Univ. Calif. Publ. Ent.
7:207-272.
Evans, G.O. (196:~a). Observations on the chaetotaxy of the legs in the free·living Gamasina (Acari:

I Mesostigmatul. Buil. Brit. Mus. (Nat. Hist.1 Zooi. 10(5):277-303.

Evans. G.O. (1963b). Some observations on the chaetotaxy of the pedipalps in the Mesostigmata
(Acari). Ann. Mag. Nat. Hist. 6(13):513-527.

I Evans. G.O., J.G. Sheals and D. MacFarlane (19611. The Terrestrial Acari of the British Isles. Vol. i.
lntroductior and Biology. British Museum (Natural History), London: 219 pp.
Fain, A. (1963). La spermatheque et ses canaux adducteurs chez les acariens mesostigmatiques. Aca-

I rologia 5(4):463-479.
Gorirossi. F.E. (19561. The gnathosoma of Mega/olaelaps ornata (Acarina Mesostigmata Gamasidesl.
Amer. Midland Nat. 55(21 :357-362.

I Grandjean, F. (1934).
France 103:109-146.
Les organes respiratoires secondaires des Oribates (Acariensl. Ann. Soc. Ent.

Grandjean, F. (1935a). les poiles et les organes sensltifs portees par les pattes et Ie palpe chez les

I Oribates. Bull. soc. 2001. France 60(1) :6-39.

Grandjean. F. (Hi35bl. Observations sur les Acariens (1 ere ser.l. Bull. Mus. Hist. nat. Paris (2) 7:119-126.
Grandjean, F. (1946), Au sujet de I'organe de Claparede des eupathidies multiples et des taeindies

I mandibulaires chez les Acariens actinochitineux. Arch. Sci. phys. nat. 28:63-87.
Grandjean, F. (1 £164). La solenidiotaxie des Oribates. Acarologia 6(31 :529-556.

I
Hammen, L. van der (1961). Description of Holothvrus grandjeani nov. spec., and notes on the classifica-
tion of the mites. Nova Guinea n.s. 10(9): 173-194.
Hammen, L. van der (1963). The addition of segments during the postembryonic ontogenesis of the
Actinotrichida (Acarida) and its importance for the recognition of the primary subdivision of the

I body and the original segmentation. Acarologia 5(3) :443-454.

I
I
30
I
Hammen, L. van der (1964). The morphology of Glyptholaspis confusa (Faa, 1900) (Acarida, Gamasina).
Zool. Verhandl. 71:56 pp.
I
Hammen, L. van der (1968a). Stray notes on Acarida (Arachnida) I. Zool. Meded. 42(35):261-280.
Hammen, L. van der (1968b). The gnathosoma of Hermannia convexa (C.L. Koch) (Acarida: Oribatina)
I
and comparative remarks on its morphology in other mites. Zool. Verhandl. 94:45 pp.
Hughes, T.E. (1952). The morphology of the gut of Bdel/onyssus bacoti (Hirst 1913, Fonseca 1941). Ann.
Trap. Med. Parasit. 46:54-60. I
Knulie, W. (1959). Morphologische und Entwicklungsgeshichtliche untersuchungen tum phylogenetischen
System der Acari: AcariformesZachv.11. Acaridiae: Acaridae. Mitt. ZooI. Mus. Berlin 35(2):347-417.
Krantz, G.W. and J.L. Meliott (1968), Two new species of Macrocheles (Acarina: Macrochelidae) with
notes on their host-specific relationships with geotrupine beetles (Scarabaeldae: Geotrupinae).
I
Jour. Kansas Ent. Soc. 41 (1) :48-56.
Lipovsky, L.J., G.W. Byers and E.H. Kardos (1957), Spermatophores-the mode of insemination of
chiggers (Acarina: Trombiculidae). Jour. Parasitol. 43:256-262.
I
Manton, S.M. (1958). Hydrostatic pressure and leg extension in arthropods, with special reference to
arachnids. Ann. Mag. Nat. Hist. ser. 13, 1:161-182.
Michael, A.D. (1892). On the variations in the internal anatomy of the Gamasinae, especialiy in that of the
I
genital organs, and their mode of coition. Trans. Linn. Soc. London, ser. 2, 5:281-324.
Michael, A.D. (1896). The internal anatomy of Bdel/a. Linn. Soc. London (Zoo I.) 6(7):477-528.
Mitcheli, R.D. (1958). Sperm transfer in the water-mite Hydryphantes ruber Geer. Amer. Midi. Nat.
I
60(1):156-158.
Mitcheli, R.D. (1962). The structure and evolution of water mite mouthparts. Jour. Morphol. 110(1):
41-59.
I
Mltcheli, R.D. (1964). The anatomy of an adult chigger mite Blankaartia acuscutel/aris (Walch). Jour.
Morph. 114(3) :373-391.
Mitcheli, R.D. (71971). The evolution of a blind gut in trombiculid mites. Jour. Nat. Hist. (in press).
I
Mitcheli, R.D. and M. Nadchatram (1969). Schizeckenosy: the substitute for defecation in chigger mites.
Jour. Nat. Hist. 3:121-124. I
Moss, W.W. (1962), Studies on the morphology of the trombidiid mite AI/othrombium lerouxi Moss
(Acari). Acarologia 4(3) :313-345.
Popp, E. (1967). Die Begattung bei den Vogelmilben Pterodectes Robin (Analgesoidea, Acari). Z. Morph.
Okol-Tiere 59 :1·32.
I
Radinovsky, S. (1965). The biology and ecology of granary mites of the Pacific Northwest. IV. Various
aspects of the reproductive behavior of Leiodinychus kramer; (Acarina: Uropodidae). Ann. Ent.
Soc. Amer. 58(3) :267-272.
I
Radovsky, F.J. (1968). Evolution and adaptive radiation of Gamasina parasitic on vertebrates. Parasit.
(Leningrad) 2(2): 124-136.
Rohde, C.J. and D.A. Demick (1967). Anatomy of the digestive and reproductive systems in an acarid
I
mite (Sarcoptiformes). Acarologia 9(3):608-616.
Roshdy, M.A. (1966). Comparative internal morphology of subgenera of Argas ticks (lxodoidae,
Argasidae). 4. Subgenus Ogadenus: Argas brumpti Newmann, (1907). Jour. Parasitol. 52(4):776-
I
782.
Wharton, G.W., W. Parrish and D.E. Johnston (1968). Observations on the fine structure of the cuticle of
the spiny rat mite, Laelaps echidnina (Acari-Mesostigmata). Acarologia 10(2) :206-214.
I
Winkler, W. (1888). Anatomie der Gamasiden. Arbeit. ZooI. Inst. Univ. Wi en 7(3): 1-38.
Wright, K.A. and I.M. Neweli (1964). Some observations on the fine structure of the midgut of the mite
Anystis sp. Ann. Ent. Soc. Amer. 57(6) :684-693.
I
Young, J.H. (1968). The morphology of Haemogamasus ambulans II. Reproductive system (Acarina:
Haemogamasidae). Jour. Kansas Ent. Soc. 41(4):532-543. I
I
I
I
I 31
I IV. REPRODUCTION AND EMBRYOGENESIS

I Although reproduction in the Acari generally follows the classic pattern of fertilization
and subsequent production of male and female progeny, facultative parthenogenesis is
I known to occur throughout the subclass. Arrhenotoky, the production of males from un·
fertilized eggs, has, been observed in both the Mesostigmata and the Prostigmata (Filipponi

I 1957, Boudreaux 1956). The production of females from unfertilized eggs (thelytoky) is
common among certain Prostigmata. It has also been observed in ixodid ticks (Metastig-
matal and in some of the lower Cryptostigmata (Grandjean 1947, Woodring and Cook

I 1962). Both arrhenotoky and thelytoky have been noted in the Anoetoidea, suborder
Astigmata (Hughes and Jackson 1958).

I Embryonic development in the Acari is, at best, poorly known. From the scanty
information presently available, it appears that total cleavage of the primordial cytoplasmic
mass does not occur, although an exception has been observed in the family PYEMOTIDAE
I (Reuter 1909). The nucleus divides within the cytoplasm and migrates to the surface, or it
migrates to the surface prior to cleavage. The nuclei continue to divide until they form an
envelope called the blastoderm, within which is the yolk. A few blastoderm nuclei enter
I the yolk area where, as vitellophage cells, they liquefy the yolk and make it readily
available to the developing blastoderm. An anterior polar cap appears, providing the site

I for subsequent development of the central nervous system. A germinal band then differ-
entiates ventrally and a ventricular area develops dorsal to the band. The germinal area
eventually gives rise to both the gnathosomal and idiosomal appendages, the latter usually

I consisting of only three pairs of limb buds. Four pairs have been observed in early
developmental s':ages in some species (Sicher 1891, Supino 1895) but the fourth pair is
resorbed at the t.me of pedipalpal differentiation.
I The mature embryo normally hatches by rupturing the eggshell. In many cases,
however, a second membrane is formed inside the shell. The deutovarial membrane has
I been seen in Metastigmata, Cryptostigmata, and in some of the higher Prostigmata. Where
the membrane occurs it expands after rupture of the eggshell, resulting in a pre-larval or

I deutovum "stage." The developing larva thus has additional room for growth and expan-
sion prior to actual hatching (Michael 1884).

I Useful References

I Boudreaux, H.B. (1956). Revision of the two-spotted spider mite (Acarina, Tetranychidae) complex.
Tetranvchus telarius (Linnaeus). Ann. Ent. Soc. Amer. 49(1):43-48.
Fi1ipponi, A. (19!57). Arrenotochia in Macrocheles subbadius (Acarina, MesostigmataL Est. Rend. 1st.

I Sup. Sanit. 20:1037-1044.

Grandiean, F. (1847), Observationssur lesOribates (18eSerie). Bull. Mus. Hist. Nat. Paris 19(2):165-172.
Hansell, R.I.C., M. Mollison and W.L. Putman (1964). A cytological demonstration of arrhenotoky in

I three mites of the family Phytoseiidae. Chromosoma 15:562-567.

Hughes, R.D. and C.G. Jackson (1958). A review of the Anoetidae (Acari). Virginia Jour. Sci. 9. n.s.
(1):5-198.

I
I
I
32
I
Hughes, T.E. (1950). The embryonic development of the mite Tyrog/yphus farinae Linnaeus 1758.
Proc. Zool. Soc. London 119(4):873·886.
I
Michael, A.D. (1884). 8ritish Oribatidae. Ray Society, London 1:336 pp. + xi + plates.
Reuter, E. (1909). Zur Morphologie und Ontogenie der Acariden mit besonderer Berucksichtigung von
Pedicu/opsis graminum. Acta Soc. Sci. Fenn. 36(4): 1·288.
I
Sicher, E. (1891). Contributione alia embriologica degli acari. Atti Soc. Veneto-Trent., Padua 1, 12.
Supino, F. (1895). Embryologia degli Acari. Atti Soc. Veneto-Trent., Padua 2, 2(1):242·261.
I
Woodring, J.P. and E.F. Cook (1962). The biology of Ceratozetes eisa/pinus Berlese, Sche/oribates
/aevigatus Koch, and Oppia neer/andica Oudemans (Oribatej), with a description of all stages.
Acarologia 4(1):101·137. I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I 33

I V. OVIPOSITION AND LIFE STAGES

I While embryDnic or postembryonic oVDviviparity has been observed in various Meso-

stigmata and Prostigmata (Strandtmann and WhartDn 1958, Filipponi and Francaviglia 1963,
I Cross 1965) typical oviposition Dccurs in the majority of Acari which have been studied.
Oviposition is accomplished in several ways. The oval or ovoid eggs are passed through the

I genital valves and dropped either singly or in clusters. They may be smoDth or sculptured
and usually are Dpaque white, although eggs of many Prostigmata may be brightly colored
in shades of red, orange Dr green. A waterproof waxy coating may be applied tD each egg

I prior tD depDsitiDn (Lees and Beament 1948). Phytophagous and graminivorous mites tend
to oviposit haphazardly on their food substrate while predaceous soil-inhabiting mites gen-
erally secrete their eggs where IDss from predators is at a minimum. Parasitic mites often

I are larviparous but thDse which ovipDsit generally chDose a particular host tissue as an
oviposition site. Others place their eggs in prDtected situations where access of hatching
larvae tD the host will be virtually assured (Camin 1953). Dicrocheles phalaenodeetes
I (Treat). a mesD,;tigmatid parasite of noctuid moths, prepares the oviposition site by
kneading the tissue under which the egg is to be deposited (Treat 1958).

I Ticks may deposit several thDusand eggs at a time, while some of the Prostigmata make
a small cluster of 4-8 eggs. Such is the case with members Df the family CHEYLETIDAE.

I The female of Cheyletus eruditus Schrank, a grain-inhabiting predatDry mite, has been
observed tD brood her cluster of eggs, driving other cheyletids or larger arthropods from the
"nest" area. Hatching larvae, however, are eaten by the mother if they tarry tOD long after

I freeing themselves of their embryonic eggshell. Development of large numbers of eggs

simultaneously is made possible in certain prostigmatid groups through the phenomenon of
physogastry. Here the idiosoma may swell to many times its normal size SD as to accomo-
I date developin£ embryos. Members of the family PYEMOTI DAE commonly exhibit
extreme physogastry (Plate 59-2), with development Df the progeny proceeding within the
chDrion until tile adult stage is reached (Herfs 1926). Species Df the pyemotid genus
I Trichometridium, on the other hand, oviposit their eggs but hatching does nDt occur until
the adult stage is reached (Cross 1965). Free nymphal forms in the PYEMOTIDAE are
knDwn only in the genus Siteroptes.
I Females of the Cryptostigmata, and Df some of the higher Prostigmata, possess an

I extrusible ovipositor (Plate 10-7,81 through which the egg passes. By means of terminal
"fingers" on the ovipositor, the female holdS the egg while prDbing the substrate for a
likely SPDt tD place it. Aquatic mites of certain types utilize an ovipDsitor tD insert eggs

I in aquatic plants.

The Acari generally pass through three primary stages after hatChing Df the egg.
I These are the larva, nymph and adult.

I Larva

I Typically, the acarine larva is a 6-legged form with little Dr no sclerotization and
withDut indication Df external genitalia (Plate 11-2). Lack of development of the majority

I
•
 PLATE 11 I
34
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
•
I
'I.
35

of key structures makes identification of larvae difficult, except perhaps to suborder, A

notable exception is seen in the prostigmatid family TROMBICULIDAE, where species
determinations commonly are made on the basis of larval characteristics (Brennan and
Jones 1959).

I Nymph

I With the exception of the Metastigmata, a single nymphal stage is uncommon in the

•
I
AcarI. There are usually two or three individual nymphal stages, with as many as eight
occurring in certain ticks. Nearly all have eight legs and differentiation of shields. External
genitalia are undeveloped in the nymphal stage (Plate 11,3). but male and female nymphs
often may be determined prior to the adult molt.

The three major nymphal forms are known as the protonymph, deutonymph and
tritonymph. All three occur in the Cryptostigmata, and all may occur in certain Prostig,
I mata and Astigmat". Only two nymphal stages occur in the Mesostigmata (Plate 11,1,3).
while in the Prostigmata, members of the Tarsonemina may have no free nymphal form at
all. A single active nymphal stage is found in the prostigmatid supercohort Parasitengona,
•• Several nymphal molts may occur in the tick family ARGASI DAE (suborder Metastigmata)

•
but the majority of ticks pass through only one nymphal stage (Cooley 1932).

The second nymphal stage, or deutonymph, occurs only sporadically in the Astigmata,
and mayor may not appear within a given species, The astigmatid deutonymph is com,

I pletely unlike the preceding or succeeding nymphal forms both in morphology and behavior,
and is known as the hypopus (Plate 11.4). The hypopus is resistant to environmental stress
and usually has ventral suckers or claspers by which it may secure attachment to passing
I insects or other animals (Michael 1901), Inert hypopi lack suckers or claspers (Fig, 30,
p, 233) and often rely on air currents for transportation. The hypopi of the family
HYPODERIDAE are inactive, seeking shelter in the tissues of a variety of animal hosts (Fain
I 1965, 1967). Generally the hypopus is a non,feeding migratory form, as is the deuto-
nymphal stage in some of the Mesostigmata. When deutonymphs occur in parasitic

I Mesostigmata, the stage often is an inactive non-feeding phase (Evans and Till 1965).

I Useful References

Brennan, J.M. and E.K. Jones (19591. Keys to the chiggers of North America with synonymic notes and

I descriptions 01' two new genera (Acarina: Trombiculidae). Ann. Ent. Soc. Amer. 52(1):7·16.

•
Camin, J.H. (1953). Observations on the life history and sensory behavior of the snake mite, Ophionyssus
natricis (Gervais) (Acarina: Macronyssidae). Chicago Acad. Sci. Spec. Publ. 10:75 pp. + 3 plates.
Cooley, R.A. (19321 The Rocky Mountain wood tick. Montana State Call. Bull. 268:58 pp.
Costa, M. (1966). The biology and development of Hypaaspis (Pneumalaelaps) hyatti (Acari: Mesostig-
matal. Jour. Zool. 148:191-200.

I Cross, E.A. (1965), The generic relationships of the family pyemotidae (Acarina: Trombidiformesl.
Unlv. Kans. ~.ci. Bull. 45(21 :29-275.

,
Dasse, G. and I. Schneider (1957). Biologie und Lebensweise von Czenspinskia lord; Nesbitt (Acar.,

I Sarcoptifarmesl. Zeit. ang. Ent. 44:403-418 .

I
36
I
I
•
Evans, G.O. and W.M. Till (19651. Studies on the British Dermanyssidae (Acari: Mesostigmata). Pt. J.
External morphology. Bull. Brit. Mus. (Nat. Hist.) Zool. 13(8):249-294.
Fain. A. (1965). Un nouveau type d'hypope, parasite cuticole de rongeurs africa ins (Acari: Sarcopti-
formes). Zeit. f. Parasit. 26:82-90.
Fain, A. (19671. Les hypopes parasites des tissus cellula ires des oiseaux (Hypodectidae: Sarcoptiformes).
Bull. Inst. roy. Sci. nat. Belg. 43(4):1·139.
Fillpponi, A. and G. Francaviglla (1963), Oviparita e larviparita in Macrocheles peniculatus Berl. (Acari:
Mesostigmata) regolate da fattori ecologici. Parassitol. 24(2):81-104.
I
Griffiths, a.A. (1966). Nutrition as a factor Influencing hypopus formation in the Acarus siro species
complex (Acarina, Acaridae). Jour. Stored Prod. Res. 1:325-340. I
Herfs, A. (1926). Okologische Untersuchungen an Pediculoides ventricosus (Newp.) Berlese. Zoologia.
Gesamtgeb. Zool. 74:3-68.
Lees, A.D. and J.W.L. Beament (1948). An egg waxing organ in ticks. Quart. Jour. Micr. Sci. London
98:291-332.
I
Michael, A.D. (1901). British Tyroglyphidae. Ray Society, London 1:291 pp. + vii,
Strandtmann, R.W. and G.W. Wharton (1958). A Manual of Mesostigmatid Mites Parasitic on Vertebrates.
I
Inst. Acarology Contrib. 4:330 pp. + xi + plates.
Treat, A.E. (1958>' Social organization in the moth ear mite IMyrmonyssus phalaenodectes). Proc.l0th
Intntl. Congo Ent. 2:475-480.
I
I
I
I
I
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I
I
I

I•
••
I 37

I VI. HABITS AND HABITATS

I:
I
The tremendous diversity in acarine morphology is more than equalled by the variety
of behavioral characteristics seen in the subclass. Specialization in habitat is paralleled in
many cases by !.pecialization in structure. It is therefore essential that both habits and
habitats of the J\cari be known if an understanding of their classification is to be achieved.

The Acari may be grouped under two major habital headings: the free-living forms and
the parasitic forms. Species which may be beneficial or injurious to man occur in both
groups.

I Free-Living Forms

I The free-living acarines comprise a vast complex of species representing all of the mite
suborders except the Metastigmata. Here one may find predaceous mites of infinite variety,
I mites feeding on plants and fungi, and others utilizing dead organic matter as food. It is
possible to categorize the free-living mites on the basis of their habits and, at least in part,
of their morphology. Some common examples are listed below under appropriate habital
I headings.

I A. Predaceous Mites

1. Ground species. These mites are common in soil, moss, humus and animal waste

I products where they feed on small arthropods or their eggs, on nematodes, and occa-
sionally on each other. Predaceous ground forms are commonly long-legged, fast mites
equipped with chelate-dentate or stylet-like chelicerae. Where the chelicerae are styletti-
I form, the palps may be raptorial. Many have well developed shields, and eyes often are
present on the propodosoma. Typical representatives of this group may be found in the
families MACROCHELIDAE (Plate 22), RHAGIDIIDAE (Plate 61), PARASITIDAE
I (Plate 20), UIBIDOSTOMMIDAE (Plate 66), ASCIDAE (Plate 31) and CHEYLETIDAE
(Plate 76). IVlany predaceous ground species are considered beneficial to man, since they
may feed on housefly eggs (Axtell 1963) or on harmful arthropods {Hurlbutt 1958\.
I At least one moss-inhabiting species of Cryptostigmata is known to supplement its
normally saprophytic diet with live nematodes (Rockett and Woodring 1966).

I 2. Aerial species. Like the ground-inhabiting species, the predaceous aerial mites
usually are long-legged and fast, and prey on plant-feeding mites or their eggs (McGregor

I 1956\. While large dorsal and ventral shields occur commonly, they are weakly sci erotized
and often difficult to see. In many cases, mites of this group are brightly colored in shades
of red, yellow or green. Aerial species are found in several families among which are the
I PHYTOSEIIDAE (Plates 28, 29), ANYSTIDAE (Plate 73), RAPHIGNATHIDAE (Plate 85)
and TYDEIDAE (Plates 70, 71\. Various members of the PHYTOSEIiDAEare con-
sidered to be highly beneficial in controlling phytophagous mite species of the family
I TETRANYCHIDAE (Chant 1961).

I
I
I
38
I
3. Aquatic species. The supercohort Hydrachnellae contains the majority of pre- I
daceous mite species found in aquatic situations. A common but not constant feature in
this highly diverse group is the presence of long "swimming hairs" on the legs (Plate 96-3).
Many aquatic species are brightly colored in red, orange, green or blue tones (Soar and
I
Williamson 1925). Eyes usually are present and the palpi often are modified for grasping.
Sci erotized shields mayor may not be distinct. Predaceous hydrachnellid adults and
nymphs feed on other mites, and on small crustaceans, isopods and insects. Larval forms
I
commonly are parasitic on insects, molluscs or fishes (Sparing 1959). The prostigmatid
family HALACARIDAE (Plate 61) is primarily marine in habitat (Newell 1947), comprising
over 300 species of which many are predaceous.
I
B. Phytophagous Mites
I
1. Aerial species. Aerial phytophagous species are slow-moving or sedentary mites I
which are weakly sclerotized. The majority of species are red, yellow, or green in color
while others may appear white or translucent. They feed by inserting stylet-like chelicerae
into the cells of the plant host and sucking up the contents. Included in this group
I
are some of our most important arthropod pests of plants. The spider mites, family
TETRANYCHIDAE (Plates 81, 82), and the ERIOPHYIDAE (Plate 55) are of particular
concern to agriculture. Aside from their feeding injury, at least five eriophyid species have
I
been found to transmit plant viruses (Slykhuis 1963). Other important aerial phytophagous
species are found in the families TARSONEMIDAE (Plate 58) and TENUIPALPIDAE
(Plates 63,64).
I
2. Storage species. Stored grains and other stored products often are infested by I
various kinds of mites, some of which feed on the product itself (Hughes 1961). Storage
species are white or brownish-white in color, and commonly slow-moving and sac-like.
The chelicerae are blunt and toothed, and are useful for scraping and gouging the food
I
material. Graminivorous mites feed on the germ tissue of the grain and may move into the
surrounding endosperm as well. Dried fruit, linseed oil, stored tubers and bulbs also
are subject to injury through feeding and through their contamination of foodstuffs.
I
Common storage species are found in the families ACARIDAE (Plates 102, 103) and
GL YCYPHAGIDAE (Plates 109, 110). The grain mite, Acarus siro, an important acarid
contaminant of processed grain products throughout the world, was chosen by Linnaeus
I
in 1758 as the type for the first formally named mite genus.

3. Soil species_ This is a rather small group of mites which feed on growing root
I
tissue, corms or tubers in the soil. Like the storage species, most of the soil forms are
opaque white and sac-like, and the majority belong to the family ACARIDAE. At least one I
cryptostigmatid mite (family PER LOHMANN II DAE) has been implicated as a phytophagous
soil species (Evansetal. 1961).
I
C. Fungivorous Mites I
Aside from a tendency to be slow-moving or sedentary, the fungivorous mites do not
lend themselves to categorizing. Species of all suborders except the Metastigmata have been I
I
I
I 39

I found to feecf-en fungi in habitats ranging from tree buds to stored grain. Species of the
mesostigmatid family UROPODIDAE (Plate 47) feed on fungi in soil and storage situations

I as do some of the weakly sclerotized ACAR IDAE. Many representatives of the Cryptostig-
mata, including species of Hermannia and Scheloribates, are mycetophagous in woody plant
tissue (Woolley 1960). These mites often are found in association with wood-boring insects.
I Members of the pl'Ostigmatid families EUPODIDAE (Plates 62,63) and TARSONEMIDAE
(Plate 58) feed on soil fungi, along with various Cryptostigmata. Members of the genus
Linopodes (family EUPODIDAE) are important pests in commercial mushroom houses, as
I are species of Tyrophagus (family ACARIDAEI (Davis 19441.

I D. Coprophagous and Saprophagous Mites

I Dung offers an attractive habitat for many kinds of arthropods, among which are
mites which generally feed on other animals which in turn feed on dung. Foremost in this
group are the predaceous ground forms mentioned earlier. However, certain mites actually

I feed on dung, including members of the Mesostigmata and Cryptostigmata. Nymphs of

cryptostigmatid species of Pseudotritia and Hoplophora associated with bark beetles, may
require beetle feces as fo()d for continued development (Woolley 1961).
I Saprophagy occurs in most mite suborders, either as a way of life or as a facultative or
sporadic phenomenon. For example, the astigmatid mite Caloglyphus berlesei is found
I commonly as a ,;aprophage on dead soil insects but is equally at home in proteinaceous
stored products. Astigmatid mites also are found feeding on dead or dying plant tissue in a

I variety of habitats and at least one saprophagous species (Rhizoglyphus echinopusJ may
compound initial insect or fungal damage to narcissus and lily bulbs.

I E. Phoretic Mite:s

I Deutonymphs and adults of non-aquatic free-living mites may utilize insects or other
arthropods as a means of dispersal. This non-parasitic association is referred to asphoresy
(Rapp 19591. The hypo pal stage of astigmatid mites, discussed in the previous section,
I usually is a phoretic form. Phoretic deutonymphs of the mesostigmatid cohort Uropodina
may attach to their arthropod associates by means of an anal pedicel, a strand secreted

I from the anus of the mite which hardens upon contact with the air (Fig. 9). Other
uropodines (POLYASPIDAE, for example) grasp their host with claws and chelicerae.
Phoretic adults of the genus Dinogamasus (family LAELAPIDAE) may be found in a

I special abdominal pouch, or acarinarium, of their carpenter bee hosts (Le Veque 1930).
Nymphs and/or adults of the gamasine families PACHYLAELAPIDAE (Plate 23), PARA-
SITIDAE (Plate 20), RHODACARIDAE (Plate 25), LAELAPIDAE (Plates 39, 40),

I ASCI DAE (Plate 311 and MAC ROCHE L1 DAE (Plate 22) have been observed as phoretic
associates of a variety of arthropods. Phoretic adults of the tarsonemoid genus Iponemus
(suborder Prostigmata) have been collected from bark beetles (Lindquist 1969). The
I associations of the many species of Trigynaspida (suborder Mesostigmata) with insects
appear to be more intimate than simple phoresy, with the adult mites deriving food from
the insect host in the form of bodily secretions. A similar likelihood exists for some of
I the mongynaspid insect paraphages of ants, including members of the Uropodina and

I
I
 40
I
Gamasina. A phoretic association with hummingbirds has been observed in certain genera I
of the mesostigmatid family ASCI DAE, with the birds providing transportation for the
mites from flower to flower.
I
t

l'

t .;
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I 41

I and of military operations. Aside from transmission of pathogens, acarines may be

vectors for internal parasites such as tapeworms and filarial worms. Members of both the
I Cryptostigmatil and Astigmata may serve as intermediate hosts for a variety of cestodes
including the !;heep tapeworm (Allred 1954). The simple process of feeding by mites may
result in damage to their animal hosts through exsanguination, irritation, or by providing
I sites for invasion by secondary disease organisms.

I The parasitic acarines may be divided into two groups on the basis of their feeding
sites. A few examples of each group are cited below.

I A. Ectoparasitic Mites

I 1. VertEibrate ectoparasites. Almost every animal group has its complex of external
acarine parasites, many of which show varying degrees of host specificity (Nutting 1968).
Ectoparasitic mites and ticks are recovered from such animals as bats, armadillos, birds,
I ungulates, fishes and reptiles. They include those acarines which feed on blood, lymph,
sebaceous secretions or digested tissues of their hosts, either by puncturing the skin or by
invading surface pores of the host.
I The common chicken mite, Dermanyssus gallinae, mange and scab mites of the

I astigmatid families SARCOPTIDAE (Plate 127) and PSOROPTI DAE (Plates 112, 113). and
the many species of chiggers (which are larval forms of the prostigmatid family TROMBI-
CULIDAE (Plates 92, 94)) are only a few of the hundreds of ectoparasitic mites found on

I vertebrates (Baker et al. 1956), Demadex fallieularum (Simon), the prostigmatid follicle
mite of man (family DEMODICIDAE, Plate 56) (Spickett 1961). feeds on sebaceous
materials in hair follicles of the eyebrows and forehead of a majority of the world's
I population. Species of the family TROMBICULIDAE secrete a digestive enzyme at the
point of attack and ingest the dissolved host tissue (Fig. 23, p. 168) (Wharton and Fuller
1952). Sareoptid mange mites (family SARCOPTIDAE) burrow into the skin of their host
II while psoroptid scab mites (family PSO ROPTI DAE) attack the surface skin, causing a weep-
ing lesion which finally hardens into a protective scab. Other astigmatid mites are found on
the feat hen; of birds (e.g., the ANALGIDAE (Plates 116, 1171, PROCTOPHYLLODIDAE
1/ (Plates 117. 118) and DERMOGLYPHIDAE (Plates 119, 120)). while prostigmatid
mites of the family SYR INGOPH ILI DAE (Plate 79) are found within the feather qUills.

II Mesostigmatid mites of the families DERMANYSSIDAE (Plate 37), LAELAPIDAE

(Plates 39, 40). MACRONYSSIDAE (Plates 36, 371 and related groups feed primarily
on the blood or tissue secretions of their hosts. The ticks (Metastigmata). however,

II are essentially hematophagous.

Viruses, rickettsias, bacteria, spirochaetes, protozoans and helminths have been

II isolated from a wide variety of ectoparasitic acarines. Many of these organisms cause
virulent or debilitating diseases in man and in animals throughout the world (Audy 1968,
Philip 1963, Smith et al. 1944, Arthur 1962).
II 2. Invertebrate ectoparasites. Comparatively few mites have established a truly

II parasitic i1ssociation with invertebrates. Species of the prostigmatid families TROMBI-

DIIDAE (Plates 91,92,93). ERYTHRAEIDAE (Plate 89). SMARIDIIDAE (Plate 90) and

I
I
42
I
I
•
JOHNSTONIANIDAE (Plate 91) are parasitic on insects, but only in the larval stage
(Severin 1944, Southcott 1961). Nymphal and adult stages of these mites are predatory.
A similar situation exists in the Hydrachnellae, where the larval form may be found
parasitizing aquatic insects (Mitchell 1961) or in some cases molluscs (Mitchell 1955).
Nymphs and adults of certain hygrobatoid Hydrachnellae are parasites of molluscs and
sponges. Species of the prostigmatid families PYEMOTIDAE (Plate 59), TARSONEMIDAE
(Plate 58), PODAPOLIPODIDAE (Fig. 22, page 147) and PTERYGOSOMIDAE (Plate 74)
I
may be parasitic on invertebrates throughout their lives (Hirst 1921, Newell and Ryckman
1966). Ricardoella limacum (family EREYNETIDAE, Plates 71,72) feeds on the exudates
of slugs and snails, and occasionally causes the death of the host under laboratory conditions
I
(Turk and Phillips 1946). Members of the mesostigmatid genera Hemipteroseius and Treatia
(family OTOPHEIDOMENIDAE, Plates 29, 30) parasitize hemipteran insects (Krantz and I
Khat 1962), feeding in the conjunctival regions beneath the hemielytra. The astigmatid
mite Ewingia coenobita Pearse (family EWINGIDAE, Plate 122) is a parasite on the gills of
pagurid crabs in the Antilles and in Florida (Pearse 1929).
I
I
•
B. Endoparasitic Mites

1. Vertebrate endoparasites. The majority of vertebrate endoparasites show a,

reduction in sclerotization when compared to related external forms. Many have reduced
mouthparts and legs, and none has functional eyes. Most of the vertebrate endoparasites
are associated with the respiratory system of their host. Members of the mesostigmatid'
family HALARACHNIDAE (Plate 33) are found in the nasal passages of seals and walruses,
I
while others may be embedded in the sinuses, bronchiae, tracheae or lungs of their
mammal hosts (Chandler and Ruhe 1940). Species of the family ENTONYSSIDAE (Plate I
34) are recovered from the lungs and air sacs of reptiles, while astigmatid mites of the
family TURBINOPTIDAE (plate 118) may be found in the lungs of sea gulls (Turbinoptes).
Pneumocoptes (family PNEUMOCOPTIDAE, Plate 122) is an astigmatid genus found in the
I
lungs of rodents.

Endoparasitic mites may occasionally be found infesting portions of their vertebrate

I
hosts other than the respiratory system. A species of the astigmatid genus Laminosioptes
(family LAMINOSIOPTI DAE, Plate 121) is confined to subcutaneous infestations in
domestic fowl where dead mites provide sites for the formation of subcutaneous cysts.
I
These cysts may be felt through the skin of the bird host, particularly in the pectoral area.
The astigmatid mite Cytodites nudus (Vizioli) (family CYTODITIDAE, Plate 121) is found
primarily in the air sacs of chickens. However it also occurs in the body cavity or alimen-
I
tary canal of its host, where its presence may produce peritonitis or enteritis.
I
Accidental ingestion of live mites by vertebrates may result in a condition known as
acariasis, in which the mites may survive and reproduce in the alimentary tract. Acariasis is
not infrequent in cattle which are fed mite-infested grain (families ACARIDAE and
I
G LYCYPHAG IDAE). Symptoms include vomiting and diarrhea (Hinman and Kampmeier
1934). I
2. Invertebrate endo parasites. Relatively few mites are found as endoparasites of
invertebrates, and all of them are confined to the orders Prostigmata and Mesostigmata. I
I
I
I
43
I The prostigmatid genus Acarapis (family TARSONEMIDAE, Plate 58) comprises at least
three species which parasitize honeybees. One of these, Acarapis wood! (Renniel invades
I the tracheal svstem of the bee host, causing degeneration of injured host tissue and
eventual suffocation. This condition is called Isle of Wight disease, and is a serious
problem in commercial honeybee hives throughout much of Europe and Asia (Hirst 19211.
I The tracheae of other hymenopterans, of orthopterans and of coleopterans may be invaded
by genera of the prostigmatid family PODAPOLIPIDAE (Podapolipus, Locustacarus and

I others (Fig. 22:)). The HALACARIDAE (Plate 61) (Order Prostigmata) includes a variety of
marine mite genera of which the genus Enterohalacarus (Viets 1938) represents an endo-
parasite of delJp sea urchins. The mesostigmatid genus Otopheidomenis is parasitic in the

I tympanic and countertympanic cavities of phalaenid moths (family OTOPHEIDOMENIDAE,

Plates 29, 30) (Treat 1967).

I Useful ReferenCI}S

I Allred. D.M. (1!J541. Mites as intermediate hosts of tapeworms. Proc. Utah Acad. Sci., Arts and Letters
31:44-51.
Arthur, D.R. (1 !)62). Ticks and Disease. Pergamon Press, Oxford: 445 pp.
I Audy, J.R. (19681. Red mites and typhus. Univ_ London, Athione Press: 191 pp. + x.
Axtell, R.C. (19631. Effect of Macrochelidae (Acarina: Mesostigmatal on house fly production from

I dairy caHie manure. Jour. Econ. Ent. 56(3):317-321.

Baker, E.W., T.M. Evans, D.J. Gould, W.B. Hull and H.L. Keegan (1956). A Manual of Parasitic Mites of
Medical or Economic Importance. Natl. Pest Control Assoc. Tech. Publ.: 170 pp.

I Chandler, W.L. and D.S. Ruhe (1940). Pneumonyssus caninum n. sp., a mite from the frontal sinus of the
dog. Jour. Parasitol. 26:59-68.
Chant, D.A. (19611. An experiment in biological control of Tetranychus telarius (L) (Acarina: Tetrany.

I chidae) in a greenhouse using the predaceous mite Phytoseiufus persimilis Athias·Henriot (Phyto-
seiidae). Can. Ent. 93(6):437-443.
Chant, D.A. and C.A. Fleschner (1960). Some observations on the ecology of phytoseiid mites (Acarina:
Phytoseiidae) in California. Entomophaga 5: 131·139.
II Davis, A.C. (19441. The mushroom mite (Tyrophagus lintneri (Osborn)) as a pest of cultivated mush-
rooms. IJ.S.D.A. Tech. Bull. 879:26 pp.

II Evans, G.O., .I.G. Sheals and D. Macfarlane (19611. The Terrestrial Acari of the British Isles. Vol. I.
Introdu<:tion and Biology. British Museum (Natural History) London: 219 pp.
Hartenstein, Ii. (1962). Soil Oribatei. I. Feeding specificity among forest soil Oribatei (Acarina).
Ann. Enl. Soc. Amer. 55(2):202-206.
II Hinman, E.H. and R.H. Kampmeier (19341.
Amer. Jour. Trap. Med. 14:355·362.
intestinal acariasis due to Tyrophagus longior Gervais.

Hirst, S. (1921). On the mite (Acarapis woodi, Rennie) associated with Isle of Wight Bee Disease.
II Ann. M,g. Nat. Hist. ser. 9, 7:509-519.
Hirst, S. (19221. Mites injurious to domestic animals. Brit. Mus. (Nat. Hist.1 Econ. Ser. 13:107 pp.

II Hughes, A.M. (1961). The Mites of Stored Food. Ministry Agr .. Fish. and Food Tech. Bull, 9:287 pp.
+ vi.
Hurlbutt, H.W. (1958). A study of soil-inhabiting mites from Connecticut apple orchards. Jour. Econ.
Ent.5116):767-772.
II Krantz, G.W. and N.S. Khat (19621. A review of the family Otopheidomenidae Treat 1955 (Acarina:
Mesostigmatal. Acarologia 4(4) :532-542.

I Krombein, ~~. (1962), Biological notes on acarid mites associated with solitary wood-nesting wasps and
bees (Acarina: Acaridae). Proc. Ent. Soc. Wash. 64(1):11-19.

I
I
 I
44
Le Veque, N. (1930). Mites of the genus Dinogamasus (Dolaea) found in the abdominal pouch of
African bees known as Mesotrichia or Koptorthosoma (Xylocopidae). Amer. Mus. Nov. 434:1·19.
I
McGregor, E.A. (1956). The mites of citrus trees in southern California. Mem. Calif. Acad. Sci. 3(3):42 pp.
Newell, I.M, (1947). A systematic and ecological study of the Halacaridae of eastern North America.
I
Bull. Bingham Ocean. Call. 10(3): 1-232.
Newell, I.M. and R.E. Ryckman (1966). Species of Pimeliaphilus (Acari: Pterygosomidae) attaCking insects,
with particular reference to the species parasitizing Triatominae (Hemiptera: Reduviidae). Hilgardia
37(12) :403·436.
I
Nutting, W.B. (1968). Host specificity in parasitic acarines. Acarologia 10(2):165·180.
Pearse, A.S. (1929). Two new mites from the gills of land crabs. Pap. Tortugas Lab. 26:225-230.
I
Mitchell, R.D. (1955). Anatomy, life history and evolution of the mites parasitizing fresh-water
mussels. Univ. Mich. Misc. Publ. Zoo I. 89:28 pp. + plates.
Mitchell, R.D. (1961). Behavior of the larvae of Arrenurus fissicornis Marshall, a water mite parasitic on
I
dragonflies. Animal Behav. 9(3-4):220-224.
Philip, C.B. (1963). Ticks as purveyors of animal ailments: a review of pertinent data and recent
contributions. Advances in Acarology 1 :285-325.
I
Pulpan, J. and P.H. Verner (1965). Control of tyroglyphoid mites in stored grain by the predatory mite
Cheyletus eruditus (Schrank). Can. Jour. Zool. 43:417-432.
Rapp, A. (1959). Zur Biologie und Ethologie der Kafermilbe Parasitus coleoptratorum L. (Ein Beitrag
I
zum Phoresie - Problem). Zool. Jahr. Syst. 86(4-5):303-366.
Rockett. C.L. and J.P. Woodring (1966). Oribatid mites as predators of soil nematodes. Ann. Ent. Soc.
Amer. 59(4) :669-671.
I
Rodriguez, J.G., C.F. Wade and C.N. Wells (1962). Nematodes as a natural food for Macrocheles
muscaedomesticae, a predator of the housefly egg. Ann. Ent. Soc. Amer. 59(4) :669·671.
Severin, H.C. (1944). The grasshopper mite Eutrombidium trigonum (Hermann) an important enemy of
I
grasshoppers. So. Dakota Agr. Exp. Sta. Tech. Bull. 3:36 pp.
Slykhuis, J.L. (1963). Mite transmission of plant viruses. Advances in Acarology 1:326·340.
Smith, M.G" R.J. Blattner and F.M. Heys (1944). The isolation of the St. Louis encephalitis virus from
I
chicken mites (Dermanyssus gal/inae) in nature. Science 100:362-363.
Sparing, I. (1959), Die Larven der Hydrachnellae, ihre parasitische Entwicklung und ihre Systematik.
Parasit. Schrift. 10:1-168
I
Soar, C.D. and W. Williamson (1925). The British Hydracarina, Vol. I. Ray Society, London: 214 pp. + x.
Southcott, R.V. (1961). Notes on the genus Caeculisoma (Acarina: Erythraeidae) with comments on the
biology of the Erythraeoidea. Trans. Roy. Soc. So. Australia 84: 163-178.
I
Spickett, S.G. (1961). Studies on Demodex fol/iculorum Simon (1842). I. Life history. Parasitol. 51 :181-
192.
Treat, A.E. (1967). Mites from noctuid moths. Jour. Lepidop. Soc. 21(3):169-179.
I
Turk. F.A. and S.M. Phillips 11946). A monograph of the slug mite-Ricardoel/a limacum (Schrank).
Proc. Zool. Soc. London 115:448-472.
Viets, K. (1938), Eine merkwurdige, neue, in tiefsee-echiniden schmarotzende Halacariden·gattung
I
und-Art (Acari). Z. Parasitenk. 10(2):210-216.
Volkonsky, M. (1940), Podapo/ipus diander n. sp. acaden heterostygmate parasite du criquet migrateur
(Locusta migratoria L.) Arch. Inst. Pasteur Alger. 18:321-340.
I
Weis-Fogh, T. (1948). Ecological investigations on mites and collemboles in the soil. Nat. Jutland
1:135-270.
Wharton, G.W. and H.S. Fuller (19521. A manual of chiggers. Mem. Ent. Soc. Wash. 4:185 pp.
I
WOOdring, J.P. (1963). The nutrition and biology of saprophytic Sarcoptiformes. Advances in Acarology
1:89-111.
Woolley, TA. (1961). Some interesting aspects of oribatid ecology (Acarina), Ann. Ent. Soc. Amer.
I
53(2) :251-253.
Zumpt, F. (ed.) (1961). The arthropod parasites of vertebrates in Africa south of the Sahara (Ethiopian
region). S. Afr. Inst. Med. Res. 11(50):457 pp.
I
I
I
I 45

I
•
VII. COLLECTION, PREPARATION AND MOUNTING
OF MITE SPECIMENS

Collection
I Successful collection of mites and ticks depends to a great extent on proper selection
of collection sites. Yields of predaceous ground forms, for example, will generally be
I sparse in exposed dry humus or in sandy soil. Except for relatively few species, most
ground and aE'rial forms prefer a substrate which is not exposed to direct sunlight or wind,

I and one which provides enough moisture for the mite to maintain a satisfactory water
balance. Phytophagous species may be less dependent on a protected substrate, since they
have a constant source of moisture from the juices of their plant hosts. Ticks and other

I vertebrate ectoparasites often may be collected in nesting areas or on runway or watering

routes of their hosts, but only rarely through indiscriminant search.

I The smEdl size of most acarines often makes collection of individual specimens from
a given substrate rather impractical. Thus it has become customary, under most conditions,
to collect samples of the habitat for subsequent separation of the acarines hidden within.
I Predaceous, fungivorous and saprophagous ground and soil forms may be collected in
substantial n~mbers through the use of a modified Tuttgren apparatus (Plate 12-1) (Haarlov
I 1947). based on the Berlese funnel. Habitat samples are placed on a screen within the
funnel, and an incandescent light is appended over it. The heat from the light bulb

I desiccates the sample, forcing the arthropods in the sample to burrow deeper into the
substrate. E.ventually they reach the screen, fall through, and are collected in a screw-top
jar or vial suspended below. The strength of the light bulb used depends in part on the

I size and wetness of the sample. High wattage lights (over 100 watts) often kill large
percentages of lightly sclerotized slow-moving mites. Wattages of less than 40, however,
may demand an inordinately long period of time for a sample run. Generally speaking,
II samples should be no more than 3 gallons in volume, and should be separated using a light
bulb rated at no more than 100 watts. Collection of mites for preservation may be made
in 70% ethyl alcohol. However, prolonged preservation in alcohol (over one year) tends
II to harden internal tissues. Where prolonged preservation is anticipated, a mixture of
glycerine (~,O parts). water (40 parts) and glacial acetic acid is recommended. Collection of

II live mite specimens with the Tullgren funnel is accomplished by placing moistened wadded
paper towels in the collection bottle rather than a preservative. The paper towels provide
both moisture and hiding places, reducing mortality caused by desiccation and by predation.

II Phytophagous and predaceous aerial species may be collected with the Tullgren funnel
as described above, although other methods are available. Samples of mite-infested foliage
II may be beaten on a 1/2" mesh screen covering a large tray, and the dislodged mites picked up
with a finll brush and placed into preservative. This method is particularly useful in the
field, where large bulky samples of foliage are an inconvenience. At least one power-
I driven brushing apparatus has been manufactured for the collection and assessment of plant
mite populations on infested foliage (Chant and Muir 1955). A suction device utilizing a
portable electric blower has been devised by Johnson et al. (1957) for collecting plant
I mites from rough grassland and from low-growing plants. Handpicking with the aid of a

I
I
 PLATE 12 I
46
I
I
I
incurrent
tube I
I
removable
inner chamber -intake I
J) I
I
I
I
I
I
I
I
I
sink vacuum

I
I
12-3
I
12-1; a modified Tullgren apparatus: 12-2: a Singer aspirator (after Singer 1964): 12-3; a Buchner funnel apparatus for vacuum
I
sample filtration
I
I 47

I ,I
hand lens and fine brush is practiced on a variety of habitats, including piants. The use of
a bulb aspirator of the Singer type (Plate 12-2) is highly recommended for collecting from

I plant hosts sinGe it collects directly from the habitat into the preservative, eliminating the
necessity of handling the specimens (Singer 1964).

I Aquatic mites are collected in a variety of ways, depending on whether the mites are
free-swimming, benthic or parasitic. Birge net or bottom dredge samples should be
examined in a white porcelain tray in which the aquatic mites may easily ,be seen and from
I which they may be collected with an eye dropper. A small silk stocking net, or a tea
strainer on a IDng stick, is invaluable in collecting individual free-swimming Hydrachnellae

I from ponds or sluggish streams. The larvae of water mites often are parasitic on semiaquatic
or aquatic insects, or molluscs. Examination of hemipterans, beetles, dragon flies, mayflies
or damsel flies often reveals sac-like opaque

I white or bright orange nymphochrysafides,

quiescent nymphs in larval skins (Fig. 10)
attached to the wings or body of their

I host. The !Jill and the excurrent siphon

tissue of molluscs may contain nympho-
chrysalides and tefeiochrysafides (quiescent
I adults in nymphal skins) (Mitchell 1955).

Unlike many of the aquatic mites, the

I marine mites are non-swimming forms and
cannot be collected by conventional means.

I Most marine species adhere tightly to their

plant or animal hosts and are difficult to
see. Collection from mite-infested marine

II algae or coral, therefore, cannot be im-

plemented through the use of the porcelain
pan method outlined for fresh water mite
II species. Marine samples should be placed in a
bucket, imrr,ersed in sea water, and treated
with chloroform or one of the halogenated
II ether compounds (ca. 15 cc/gallon). This
anesthetizes the mites, causing them to re-

II linquish their hold on the substrate. The

procedure is hastened by pouring the treated
sample from one bucket to another so as to

II mix the anesthetic with the sea water. After

30 minutes have elapsed, the substrate mate-
rial should be shaken vigorously and re-

II moved from the treated sea water. The

water may then be filtered through a muslin
bag, with more sea water being added as
I necessary for retrieving all of the precipitate.
The precipitate may then be labeled and tied

I shut and placed in 95% alcohol for subse-

quent study (Newell, personal communica-
Fig. 1: O. Water mite nymphochrysalis (Oregon),
with a portion of the developing nymph
tion). showing through a break in the larval

I skin.

I
 I
48

Collection of mite parasites of vertebrates is accomplished both through collection of

I
the host habitat (nests, runway litter, etc.), and through examination of the host itself.
Nest material lends itself well to Berlese funnel separation techniques. Unattached ticks
may be collected in runway areas by dragging a flannel cloth over suspected infestation sites.
I
Ticks will adhere to the cloth as it is dragged along. Individual mites often are discovered at
nest entrances or in recently abandoned nest material. The use of a bulb-operated Singer
aspirator in place of the oral vacuum aspirator eliminates the possibility of accidentally
I
inhaling potentially dangerous disease vectors or acariasis-producing species while collecting.

Collection of host animals is advisable or necessary for many external and for all
I
internal parasites. Live collecting may be accomplished with baited live traps (rodents) or
with mist nets (birds, bats). Since many acarine ectoparasites are not obligate, they will I
detach from the host animal after feeding. If the host is maintained for 2-4 days in a
screen cage suspended over a pan of water, many of the ectoparasites may be collected
from the water surface. Reptiles captured alive may be isolated in a similar fashion for
I
ectoparasite collection.

Museum Special rodent traps are ideal for snap-trap collections in that the head
I
capsule of the host usually is not crushed when the host is caught. This allows subsequent
examination of the sinuses and nasal areas for respiratory parasites. The use of bird shot is
advisable for bird collection, since heavier shot disfigures the host and tends to make
I
examination for parasites rather difficult. Removal of mites from dead rodents and birds
may be accomplished by immersing the animal in water-detergent mixture, which then is
shaken vigorously (Lipovsky 1951). This detaches most of the ectoparasites, which may be
I
decanted into another container for collection with brush, eye dropper or forceps. The
liquid may also be poured into a Buchner funnel (Plate 12-3) in which filter paper has beem I
inserted, and drawn through the filter paper with a sink vacuum. The filter paper may then
be examined for mites (Watson and Amerson 1967). Mite parasites of furred animals may
also be recovered through vigorous fine-combing and brushing over a suitable pan or dish.
I
Removal of mites from within feather quills (family SYRINGOPHILIDAE) or from lung
tissue (families ENTONYSSIDAE, RHINONYSSIDAE and others) must be done through
examination and dissection of the infested structure or organ. Nasal mites are recovered by
I
flushing the nasal cavities with a stream of water under high pressure. Splitting the bill
between the nares often simplifies recovery. I
Invertebrate parasites are found on their hosts in a number of locations (Fig. 9).
Antennal bases, wing axillaries, coxal cavities, spiracular plates and abdominal conjunctiva I
of insects and other arthropods shelter a variety of mite associates including members of
over a dozen families. These may be removed with a fine brush, a pin, or a pair of forceps.
Recovery of mite parasites and phoretic associates of insects from museum collections is I
often difficult in that the mites are dry and brittle, and often are secreted in deep recesses
or cavities of the insect host. Breakage may be kept to a minimum by wetting the mites
with a small droplet of glycerin, carefully removing them with a small brUSh or pin and
I
placing them in 70% alcohol for temporary storage prior to restoration (Krantz 1967).
Internal parasites of invertebrates (families TARSONEMIDAE, HALACARIDAE and others)
may be recovered only through careful dissection of the host.
I
Additional collecting methods for free-living and parasitic Acari are discussed by I
I
I
I 49

I Evans et al. (1961) in the chapter entitled TECHNIQUES. An excellent review of culturing
techniques for mites may be found in the same section.
I
Preparation
I With thE- advent of phase contrast and interference microscopy, it has become more

I and more desirable to prepare mite specimens in such a way that a high degree of trans-
parency is achieved. Since the degree of clearing which occurs in most mounting media is
minimal, maceration or removal of opaque muscle and organ tissue from all but small

I weakly sclerotized mites should be attempted before the mite specimen is placed on the
slide. This is accomplished either by: 1) the use of a clearing or digesting agent, or
2) dissection.

I
1. Clearing t~gents

I Various_ chemicals are useful in macerating internal tissues of preserved mites with
little or no damage to the exoskeleton. One of the most popular for general use is lacto-
Ii phenol, which is prepared with the following ingredients added in sequence:

II Lactic acid
Phenol crystals
, 50 parts
25 parts
Distilled water. . . . . . . . . . . . . . . .. 25 parts

II Specimens placed in lactophenol at room temperature may be left for a week or more
without hal'm to exoskeletal structure. Lactophenol is an acid corrosive and does not tend
II to soften integument as do the basic corrosives (potassium hydroxide, for example).
Acceleration of maceration occurs at higher temperatures. Larger specimens are punctured
to allow easy entrance of the lactophenol into the body cavity. Blood-engorged mites or
II mites containing large amounts of pigment are punctured and gently squeezed so as to
remove much of these substances. Additional material may be squeezed out after immer-

f. sion in lactophenol for 24-48 hours. Immersion of dried or brittle mite specimens in
lactophenol for 48 hours at room temperature restores most treated specimens to a
condition ,'esembling freshly collected material. It is therefore extremely useful in preparing

II collections from dried insects, preserved bird or mammal skins, or from pressed plant
materials in herbarium collections.

I Other acid corrosive materials which may be recommended for general use are pure
lactic acic, and Andre's fluid (113 glacial acetic acid, 1/3 chloral hydrate, 1/3 water, all by
weight). Nesbitt's fluid is a powerful clearing agent which is recommended only for old,
I alcohol-stored specimens which do not yield to clearing by other methods. Nesbitt's fluid
is prepared as follows:

I Chloral hydrate. . . . . . . . . . . . . . 40 grams

Distilled water . _. . . . . . . . . . . . .. 25 cc
Concentrated hydrochloric acid.... , 2.5 cc
I
I
I
50
I
It should be mentioned that, although prolonged treatment with lactophenol and other I
comparatively slow-acting acid corrosives will not harm well-sci erotized specimens,
submersion of lightly sci erotized specimens for more than 48 hours tends to weaken leg
conjunctiva and shields. This breakdown makes dissection difficult, since the techniques
I
involved (see next section) require considerable manipulation and handling. Dorsal
shields are particularly prone to breakage during dissection of over-treated specimens. I
Enzymatic digestion of internal tissues of HALACARIDAE and other strongly
pigmented mites may be accomplished through the use of trypsin in a toluene atmosphere,
or with pepsin. As is the case with lactophenol, pigmented mites treated with trypsin or
I
pepsin solutions must first be punctured so as to allow easy entrance of the digesting agent
into the body cavity (Newell 1947). I
2. Dissection I
Heavily sci erotized mites, particularly those of the suborders Mesostigmata and
Cryptostigmata, often are difficult to study despite the use of clearing procedures outlined
I
above. The thickness of the dorsal and ventral shields prevents easy observation of surface
structures through the microscope and makes the use of phase contrast or interference
techniques impossible. Also, the growing use of microphotography in acarology demands
I
clarity of structure often beyond that obtainable through standard clearing procedures. A
satisfactory solution to this problem is to carefully separate the dorsum and venter with
appropriate dissection tools and, in cases where the shape or structure of the epistome or
I
chelicerae is of importance, to remove the chelicerae as well. The following tools are
recommended; I
Watchmaker's or microdissection forceps-straight shaft
Minuten pin, finely sharpened on carborundum stone, inserted in a wood I
matchstick base and fixed in place with paraffin wax
Microscalpel, made of razor blade fragment embedded in glass or wood
handle
I
Specimens to be dissected are placed in lactophenol for 24-48 hours in order to soften the
integument. They are then washed in several changes of water and transferred to 20-40%
I
ethyl alcohol in a spot plate or small watch glass for dissection under a stereoscopic
microscope. I
The specimen is held gently but firmly with the fine tips of the forceps, while the
minuten pin is inserted into the soft integument laterad of the dorsal shield or shields. If I
the integument has been sufficiently softened, it may be torn by the pin without harm to
the shields. The pin should be pulled through the integument from the vertex to a point
behind coxae IV. The forceps may then be released for the second part of the operation,
I
Since most of the delicate setae are found on the dorsum, the dissection should be
completed with the forceps holding only the venter of the mite. The minuten pin is
inserted into the integument on the intact side, where the tearing operation is repeated.
I
The gnathosoma thus remains with the venter.
I
I
...
I
I 51

I Once the dorsum and venter have been separated, the pin may be used to hold the
dorsal portion in place while the internal tissues are carefully picked out with the forceps.

I The vlmter is cleared in a similar manner, although the muscle tissues associated with the
sternal and coxal areas may be more difficult to extract. The chelicerae should be removed
from the gnathosoma by inserting the minuten pin into the open end of the cheliceral base
I and pushing the chelicera through the gnathosomal tube. The forceps are then used to pull
the chelicerae out the gnathosomal tube. Where the proximal cheliceral digit is elongate and
intact, the chelicera may be removed by drawing the proximal portion back through the
I gnathosomal tube with the forceps and removing the entire structure from the body
cavity.

I Separation of dorsum and venter often is difficult in weakly sclerotized acarines with
heavy integument; i.e., Metastigmata. Here it is advisable to use a microscalpel for the

I initial incision and for subsequent separation of dorsum and venter.

I Mounting

The small size of most acarines requires that they be observed under a compound
I microscope at substantial magnifications. Specimens to be studied must therefore be
placed temporarily or permanently on microscope slides in an appropriate fluid. The

I techniques and materials utilized in this operation vary considerably with each researcher
and with the type of acarine being observed. For example, it is extremely difficult to study
or photograph whole mounts of certain heavily sclerotized Cryptostigmata, primarily be-

I cause of body shape and cuticular tanning. Here a temporary mount utilizing a cavity
slide with lactic acid as the mounting medium has been found to be highly satisfactory.
The mite is placed in a drop of lactic acid near the side wall of the slide cavity and

I manipulated into any position by movement of the cover slip partially covering the
cavity. Additional movement is accomplished with the use of a pin inserted into the
exposed side of the cavity (Grandjean 1949). Because of the thickness of the mite's
I cuticle, a vertical illuminator operated through the microscope objective, or some
alternate method of direct lighting, is advisable for study of cavity slide temporary mounts.

I In recent years, most of the permanent or semi-permanent mounting media used by

acarologists have been aqueous; i.e., they contain, and are soluble in, water. Aqueous

I mounting media have replaced oil soluble resins such as balsam or dam mar primarily
because specimens may be mounted in aqueous media without preliminary fixing operations
as required for resin mounts. A serious disadvantage in using aqueous media is that they are

I hygroscopic, taking up water from the atmosphere. An aqueous mount thus is subject to
displacement of medium by atmospheric moisture, and to eventual breakdown. To
remedy this situation, it has become customary to ring the completed slide preparation with

I a non-soluble protectant, making the preparation more or less permanent.

The most satisfactory aqueous mountant now in use is Hoyer's medium, which is
I prepared as follows:
Distilled water . 50 grams

I Gum arabic (crystals)

Chloral hydrate
Glycerine .
.
. 30 grams
200 grams
20 grams

I
I
52
I
These ingredients should be added in the order shown. Solid ingredients should be
completely dissolved before addition of succeeding reagents. The resulting material
I
should be filtered through clean cheesecloth in order to remove bits of wood or other
impurities from the gum arabic used. Powdered gum arabic should not be substituted for I
crystalline gum arabic under any circumstances, since the minute particles are difficult or
impossible to wet. The resulting medium, therefore, contains many dry gum arabic
particles which make most preparations unsatisfactory.
I
The following mounting procedures are recommended: I
1. Remove lactophenol-treated specimens from lactophenol and clean in 3-4
changes of water in a porcelain spot plate. Continue to wash until cloudy
interface of lactophenol and water disappears.
I
2. Place a drop of Hoyer's medium in the center of a clean 1x3 microscope slide. I
3. Lift specimen from spot plate with a fine wire loop affixed to a matchstick base,
or by gently grasping leg II or III with watchmaker's forceps (taxonomically I
important setae on legs I or IV could be lost accidentally if held by forceps).

4. If forceps are used, focus stereoscopic microscope on forcep tip and slowly
I
separate tines. This will cause the mite to adhere to one or the other tine, and
dissipate the water trapped between the tines. I
5. Touch wire loop or forcep tine on which mite has adhered, to the droplet of
Hoyer's medium on the slide. I
6. Carefully press the mite to the bottom of the droplet and arrange it on a vertical
axis with a minuten pin probe. (If the specimen is on the surface of the droplet I
when the coverslip is applied, it will roll to the edge of the coverslip).

7. Using a clean pair of forceps, pick up a coverslip at its edge, apply the opposite I
edge to the edge of the Hoyer's droplet, and allow coverslip to fall into place.
Last second orientation of the specimen may be accomplished under the
stereoscopic microscope with gentle pressure of a probe on the coverslip surface.
I
A 12 mm a-thickness coverslip is recommended except for large specimens
(3-6 mm). Larger, somewhat thicker coverslips should be used here. I
8. Turn slide so that the gnathosoma of the mite is directed posteriorly. Mark the
right side of the preparation with identifying number or letter (a diamond pen is
recommended, since wax markings often become obliterated in succeeding steps).
I
9. Place slide in an oven at no more than 45°C for 4 days-one week. Temperatures I
above 55°C cause the medium to bubble at the edges of the coverslip, while
. those between 46-54°C may bring about excessive contraction and breakage of
coversl ips.
I
10. Heat-treated slides should be held for one week at room temperature to allow the I
I
•
 I
53
I extra-thin coverslip to return to its normal flat state (heating causes drying and
contraction of the medium at the edges, bringing about "bowing" or "bubbling"
I of the coverslip. Decrease of heat often results in pulling away of the medium

III
from the straightening coverslip, or in cracking). After one week, edges should be
filled if necessary, and cleaned with water.

11. A ring of "Zut" paint derivative, euparal or other essentially non-soluble

material should be applied so as to seal the edge of the coverslip to the slide
surface. A second coating applied after the first has dried is recommended to
assure an impervious seal.

I 12. A locality label should be placed where the slide originally was marked. The
locality label should include date, host or substrate and collector as well as the
I locality. An identification label should be affixed on the left side of the slide.
The mounting medium used in making the slide should be noted at the bottom of
the identification label (F ig. 11).
I
I Fanl. Ma.crochelidae
OREGON' Benfona.
LoC.5 mi. W. Larvo//is
I Gerl. Nacrocheles Date 20 Oclooe", rl6g

•
I
sp•.'iIJbbtldtils(getL) ~
Det. Nov. 19 G8
By GW j{ronfl-
oYERS
Coli. J. L, !fe/IoN
Habitat ex Ju/ar arw
Ace.
of 5-1ol77tJx!j2 ~
f( /0 6'il- ;J3

I Fig. 11. Completed microslide preparation.

I Dissected specimens should be cleaned as in step 1. The dorsum, venter and chelicerae
should be placed in the Hoyer's medium as in steps 3 and 4. The dorsum and venter should
be arranged side by side, with the chelicerae occupying a position anterior to the venter.

I "roperly mounted and ringed slides may be expected to last indefinitely. When, for
reaSOlS of inadequate cleaning or poor ringing, deterioration of the preparation occurs, the

I specimen is easily retrieved by soaking off the old coverslip in water. The freed specimen
may be remounted directly to a new slide.

I More assured permanency of slide preparations may be possible through the use of oil
soluble resins, or by employing a double coverslip technique using glycerine jelly as the
mountant between the coverslips and balsam as the permanent "bed" for the preparation
I (Mitchell and Cook 1952).

I Serial sections often are found desirable for embryological or internal morphological
studies. Useful sectioning and staining techniques are outlined by Rohde (1964).

I
I
 I
54

Specimens to be held for future study should be cleared in lactophenol or lactic acid,
I
placed in a small cotton stoppered shell vial of 70% ethyl alcohol and stored upside down in
a larger vial of 70% alcohol. Collecting data for each collection should be typed or written
in India ink on high quality paper and placed in the larger vial, which should then be
I
stoppered with cotton and placed in a one-pint screw-top museum jar for storage.
I
Useful References

Auerbach, S.1. and D.A. Crossley, Jr. (1960). A sampling device for soil microarthropods. Acarologia
I
2(3):279·285.
Balogh, J. (1959). On the preparation and observation of Oribatids. Acta Zool. Hung. 5:241·253. I
Chant, D.A. and R.C. Muir (1955). A comparison of the imprint and brushing machine methods for
estimating the numbers of fruit tree red spider mite, Metatetranvchus ulm; (Koch), on apple
leaves. Rept, E. Mailing Res, Sta. for 1954: 141-145.
Ciifford, C.M. and D.T. Lewers (1960). A rapid method for clearing and mounting the genitalia of
I
female ixodid ticks. Jour. Parasitol. 46(6):802.
Evans, G.O., J.G. Sheals and D. Macfarlane (19611. The Terrestiral Acari of the British Isles.
Introduction and Biology. British Museum (Natural History), London: 219 pp.
Vol. I. I
Grandjean, F. (1949). Observation et conservation des tres petits arthropodes.
Paris ser. 2, 21:363-370.
Bull. Mus. Hist. Nat.

Haarlov, N. (1947). A new modification of the Tullgren apparatus. Jour. Animai Ecol. 16(21:115-121.
I
Hirschmann, W. and O. Woelke (1960). Das preparieren von Milben. Mikrokosmos 49(41: 122·124.
I
•
Johnson, C.G .. T.R.E. Southwood and H.M. Entwistle (1957). A new method of extracting arthropods and
moiluscs from grassland and herbage with a suction apparatus. Bull. Ent. Res. 48:211-218.
Krantz, G.W. (1967). A review of the genus Holocelaeno Berlese 1910 (Acarina: Macrochelidae).
Acarologia 9, fasc. suppl.:1-89.
Lipovsky, L.J. (1951). A washing method of ectoparasite recovery with particular reference to chiggers
(Acarina-Trombiculidae). Jour. Kansas Ent. Soc. 24: 151·156.
Macfadyen, A. (1955). A comparison of methods for extracting soil arthropods.
Academic Press, New York:315-332.
In Soil Zoology; I
Mitchell, R.D. (1955). Anatomy, life history, and evolution of the mites parasitizing fresh-water mussels.
Univ. Mich. Misc. Publ. Zool. 89:28 pp. + plates. I
Mitchell, R.D. and D.R. Cook (1952). The preservation and mounting of water mites. Turtox News
30(9):4 pp.
Newell, I.M. (1947). A systematic and ecological study of the Halacaridae of eastern North America.
Bull. Bingham Ocean. Call. 10(3):1-232.
I
Rohde, C.J. (1964). Some techniques in the preparation of stained whole mounts and serial sections of
mite embryos and adults, Acarologia 6 fasc. h.s.:208·214. I
Sengbusch, H.G. (19561. A modified Tullgren funnel for the collection of small invertebrates (Mesobiota)
in soil. Turtox News 34(111 :226-228.
Williams, R.W. (1946). A contribution to OUf knowledge of the common North American Chigger
Eutrombicula alfreddugesi (Oudemans) with a description of a rapid collecting method. Amer.
I
Jour. Trap. Med. 26:243·250.
Singer, G. (1964). A simple aspirator for collecting small arthropods directly into alcohol. Ann. Ent. Soc.
Amer. 57(61 :796-798.
I
Watson, G.E. and A.B. Amerson Jr. (1967). Instructions for collecting bird parasites. Smithsonian Inst.
Mus. Nat. Hist. Information Leaflet 477:11 pp.
Woodring, J.P. (1968), An automatic collecting device for tyroglyphid (Acaridae) mites. Ann. En!. Soc.
I
Amer. 61 (4): 1030-1 031.
I
I
I
 I
55
I VIII. CLASSIFICATION

I The fragmentary state of knowledge presently existing in the field of acarological

I, systematics makes anv attempt at a familial classification of the subclass Acari a difficult
task. Discoveries of new mite species and genera occur by the hundreds every year, and
many of these make various family diagnoses obsolete or tenuous. Acarology is, in fact, in

II a state of systematic turmoil similar to that experienced in the field of entomology nearly a
century ago. A so-called "natural" classification for the Acari is impossible with our present
level of understanding, nor maya phylogenetic system ever be realized in this fossil-poor

II group. A number of arbitrary family classifications for the Acari have thus come into being,
most of which have proven useful for at least a period of time. The classification and keys
to families presented below attempts to bring our knowledge of the subclass up to date.
II It should be recognized, however, that the scheme presented provides only one of a
number of possible treatments.

II Cohort names in the Order Acariformes have been given "-ina" endings so as to
conform with those in the Parasitiformes. No attempt has been made to standardize

II supercohort names.

References to earlier literature on higher classification are included in the Introduction

II (page 2).

I CLASSIFICATION OF THE HIGHER CATEGORIES OF

THE SUBCLASS ACARI

I I. ORDER OPILIOACARIFORMES
A. Suborder Notostigmata
1. Superfamily Opilioacaroidea
I II. ORDER PARASITIFORMES

I A. Suborder Tetrastigmata
1. Superfamily Holothyroidea
B. Suborder Mesostigmata

I SuperGohort Monogynaspida
a. Cohort Sejina
1. Superfamily Sejoidea

I b. Cohort Uropodina
1. Superfamily Protodinychoidea
2. Superfamily Trachytoidea
I 3. Superfamily Uropodoidea
c. Cohort Diarthrophallina
1. Superfamily Diarthrophalloidea
I d. Cohort Gamasina
1. Superfamily Parasitoidea

III
I
 I
56

Supercohort Trigynaspida
I
a. Cohort Cercomegisti na
1. Superfamily Cercomegistoidea
b. Cohort Antennophorina
I
1. Superfamily Celaenopsoidea
2. Superfamily Megisthanoidea
3. Superfamily Antennophoroidea
I
4. Superfamily Fedrizzioidea
C. Suborder Metastigmata I
III. ORDER ACARIFORMES
A. Suborder Prostigmata
I
Supercohort Heterostigmata
a. Cohort Tarsonemina
1. Superfamily Tarsonemoidea
I
Supercohort Endeostigmata
a. Cohort Pachygnathina
1. Superfamily Pachygnathoidea
I
Supercohort Promata
a. Cohort Eupodostigmatina
1. Superfamily Eupodoidea
I
2. Superfamily Tydeoidea
3. Superfamily Bdelloidea I
4. Superfamily Halacaroidea
b. Cohort Stomatostigmatina
1. Superfamily Labidostommoidea
I
c. Cohort Eleutherogonina
1. Superfamily Caeculoidea
2. Superfamily Raphignathoidea
I
3. Superfamily lolinoidea
4. Superfamily Tetranychoidea
5. Superfamily Anystoidea
I
6. Superfamily Cheyletoidea
d. Cohort Tetrapodilina
1. Superfamily Eriophyoidea
I
Supercohort Parasitengona
1. Superfamily Erythraeoidea I
2. Superfamily Trombidioidea
Supercohort Hydrachnellae
1. Superfamily Hydrovolzioidea
I
2. Superfamily Hydryphantoidea
3. Superfamily Hydrachnoidea
4. Superfamily Hygrobatoidea
I
B. Suborder Astigmata
Supercohort Acaridia
1. Superfamily Acaroidea
I
2. Superfamily Canestrinioidea
3. Superfamily Anoetoidea I
I
I
II
57
II Supercohort Psoroptidia
1. Superfamily Ewingoidea
II 2. Superfamily Psoroptoidea
3. Superfam ily Analgoidea
4. Superfamily Sarcoptoidea
I 5. Superfamily Listrophoroidea
6. Superfamily Cytoditoidea

I C. Suborder Cryptostigmata
Supercohort Palaeacari
1. Superfamily Palaeacaroidea

I Supercohort Oribatei Inferiores

1. Superfamily Parhypochthonoidea
2. Superfamily Hypochthonoidea
I 3. Superfamily Mesoplophoroidea
4. Superfamily Phthiracaroidea
5. Superfamily Perlohmannoidea
I 6. Superfamily Nothroidea
Supercohort Oribatei Superiores
a. Cohort Pycnonoticina
I 1. Superfamily Hermannielloidea
2. Superfamily Neoliodoidea

I 3. Superfamily Plateremaeoidea
4. Superfamily Damaeoidea
5. Superfamily Cepheoidea

I 6. Superfamily Zetorchestoidea
7. Superfamily Eremaeoidea
8. Superfamily Liacaroidea
I 9. Superfamily Carabodoidea
10. Superfamily Polypterozetoidea
11. Superfamily Oppioidea
I 12. Superfamily Hydrozetoidea
b. Cohort Poronoticina

I 1. Superfamily Ameronothroidea
2. Superfamily Passalozetoidea
3. Superfamily Pelopoidea

I 4. Superfamily Oribatelloidea
5. Superfamily Microzetoidea
6. Superfamily Ceratozetoidea

I 7. Superfamily Galumnoidea
8. Superfamily Oribatuloidea

I
I
I
I
I
58 •
I
•
KEY TO THE ORDERS AND SUBORDERS OF ACARI

1. With one to four pairs of stigmata located dorsolaterally or lateroventrally on the

hysterosoma (Fig. 5, p. 19; Fig. 6, p. 20); specialized propodosomal sensory organs
absent ORDERS PARASITIFORMES AND OPILIOACARIFORMES
Without hysterosomal stigmata; propodosomal sensory organs, when present, often
2 I
in the form of trichobothria or more specialized structures in distinctive insertions
(Plate 8-2) , , , , ORDER ACARIFORMES 5 I
2. With a terminal, subterminal or basal simple or tined apotele on the pal pal tarsus
(Plates 4-4, 13-2, 14-1,2); hypostome serving only as part of the floor of the I
gnathosoma (Plate 3-2); tarsus I only rarely with dorsal sensory pit 3
Pedipalpal tarsus without apotele; hypostome modified into a piercing organ with
retrorse teeth (Fig. 18, p. 135);dorsum of tarsus I with a distinct sensory pit
I
(Haller's organ, Plate 53-3); stigmata behind coxae IV or laterad between coxae
II-IV, each surrounded by a stigmal plate (Fig. 6a, p. 20), elongate peritremes
absent SUBORDER METASTIGMATA 1
I
3. With one or two pairs of hysterosomal stigmata, palpal tarsal apotele never terminal; I
with or without peritremes ORDER PARASITIFORMES 4
Hysterosoma with four pairs of dorsolateral stigmata (Fig. 5a, p. 19), tarsus of palp I
•
with one or two terminal claws (Plate 13-2); peritremes absent .
.............. ORDER OPILlOACARIFORMES, SUBORDER NOTOSTIGMATA 2

4. Hypostome with a maximum of three pairs of setae (Plate 3-2); with a two-, three-,
or four-tined apotele near the inner basal angle of the palpal tarsus (rarely absent in
endoparasitic forms); tritosternum (Plate 3-2) usually present and commonly with I
one or two laciniae; anal valves nude or at most with a pair of setae; peritremes
generally present (Fig. 5c, p. 19; Fig. 14, p. 68); epistome (Plate 3-1) present,
roofing the gnathosoma . . . . . . . . . . . . . . . . . . . . . .. SUBO RDER MESOSTIGMA TA
I
Hypostome with more than three pairs of accessory setae (Plate 14-4); apotele
sometimes divided, inserted basally or medially on the pal pal tarsus (Plate 14-1,2);
anal valves strongly setate (Plate 13-4); peritremes present (Fig. 5b, p. 19; Plate
I
SUBORDER TETRASTIGMATA 3
14-3), epistome absent
I
5. Aquatic forms, found in fresh, brackish or salt water (occasionally in wet moss, etc.
or under surface film of water on land); with widely divergent form and structure,
commonly showing morphological adaptations for living in water; larvae either
I
parasitic on insects (Fig. 10, p. 47), molluscs or fishes, predaceous, or herbivorous;
propodosomal sensory organs absent
Primarily non-aquatic species, without adaptations as mentioned above; propo-
6 I
dosomal sensory organs (Plate 8-2,5) present or absent

1 Suborder IXODI DES of Baker el al. 1958.

7
I
2 Suborder ONYCHOPALPI DA, Superfamily Opilioacaroidea of Baker el al. 1958.
3 Suborder ONYCHOPALPIDA. Superfamily Holothyroidea of Baker el al. 1958; Suborder HOLO·
THYRINA of Johnston 1968.
I
I
I
I 59

I 6. Primarily marine species, generally herbivorous or predaceous in the larval stage;

with 3-4 palp,il segments; with 4 dorsal and 4 ventral shields; swimming hairs on
I legs never pres1mt; empodial elements present or absent (Plate 61)
· SUBORDER PROSTIGMATA. Superfamily Halacaroidea
.

I Primarily fresh and brackish water forms, commonly parasitic in the larval stage;
with 5 pal pal segments; dorsal and ventral shield condition variable; swimming hairs

•
present (Plate 96-3,4) or absent; empodial elements absent " .
· SUBORDER PROSTIGMATA, Supercohort HYDRACHNELLAE

7. Empodial elements of legs II-III commonly pad-like, rayed or broadly membranous

I (Plate 5-3,4,5). rarely claw- or sucker-like (Plate 5-6,7) and sometimes absent;
chelicerae typically stylettiform or hooklike (Plate 4-5,7), occasionally chelate
(Plate 66-2); palpi usually distinct (when tiny and indistinct, then the idiosoma is
I vermiform (Plate 56-2) or reduced, or the empodia of tarsi II-III are broad and
membranous [Plate 5-3)); palpi often modified into a thumb-claw process (Fig. 20,

I p. 142); stigmata, when present, opening at or between the bases of the chelicerae,
the base of the gnathosoma, or on the anterior propodosomal shoulders (Figs. 6b,c,
p. 20; Figs. 7a,b, p. 20); a strongly heterogenous group .

I · SUBORDER PROSTIGMATA 1

•
Empodial elements commonly claw- or sucker-like, never rayed, and rarely absent;
chelicerae typically chelate-dentate, palpi simple; tracheal openings absent or
indistinct 8

8. Generally w€,akly sclerotized species, without distinctive propodosomal sensory

I structures; tracheal system absent; true claws absent, the empodium being either
c1aw- or sucker-like and usually inserted on a distinct pretarsus or fleshy caruncle

I (Fig. 26, p. 229; Fig. 27, p. 229); usually with two pairs of genital "discs" flanking
the genital aperture (Plate 102-2). which may be shaped like an inverted "U," "Y"
or "V," or which may occasionally be transverse (Fig. 28, p. 230; Plate 112-1) .....

I ............................................. SUBORDERASTIGMATA2
Generally strongly sclerotized species, almost always with a pair of setate, clavate
or club-shaped sensory organs (pseudostigmatic organs) inserted in pseudostigmata
I on the propodosoma (Plate 8-2); hypostome with a pair of prominent terminal

•
rutella (Fig. 33, p. 289); tracheal system, when present, opening in the acetabular
cavities of legs I and III, on the legs themselves, or through the pseudostigmata (Fig.
7c, p. 20); true claws generally present (when absent. the empodium is claw-like).
pretarsi or caruncles absent; usually with three pairs of genital "discs" flanking the

I genital aper1:ure, which generally is covered by a pair of longitudinally opening

plates (Fig. 34, p. 289); anal aperture similarly covered .
........................................ SUBORDER CRYPTOSTIGMATA 3

I
I - - 1 Suborder TROMBIDIFORMES. Supercohorts Prostigmata, Heterostigmata and Parasitenini of
Baker et al. 195B.
2 Suborder SARCOPTIFORMES, Supercohort Acaridiae of Baker etal. 1958.

I 3 Suborder SARCOPTI FORMES. Supercohort Oribatei of Baker et al. 1958.

I
I
 I
60
I
•
I
•
I
I
Tetrastig ata I
I
I
I
I
I
- - - -- - diphyletic ??
I
•
I
Fig. 12. Dendrogram illustrating possible relationships within the subclass Acari.
I
I
I
I
I
---------------------
Order Suborder Supercohort Cohort Superfamily

Opilioacariformes--- N o t o s t i g m a t a - - - - - - - - - - - - - - - - - - - - - - - - Opi!ioacaroidea

Tetrastigmata HoJothyroidea

- S e j i n a - - - - - - - Sejoidea

Gamasina - - - - - - Parasitoidea

Monogynaspida
Uropodina ----r-= Trachytoidea

Uropodoidea

Mesostigmata
l{ Protodinychoidea

Parasitiformes Diarthrophallina - - - Diarthrophalloidea

..
Cercomeglstlna --- Cercomegistoidea

Celaenopsoidea

~
Trigynaspida - { Megisthanoidea

~
A'''""ooh",O' A'''""OOho'O'"oo
Fedrizzioldea

M e t a s t i g m a t a - - - - - - - - - - - - - - - - - - - - - - - - Ixodoidea

Fig. 13. Dendrogram illustrating possible relationships within the orders OPIUOACARIFORMES and PARASITIFORMES of the subclass ACARI. ~
62
I
I
•
ORDER OPILIOACARIFORMES

Suborder Notostigmata

The Notostigmata are large (1000 M+) elongate unsclerotized leathery mites which
resemble members of the arachnid group Opiliones, at least superficially. The notostigma-
I
tids are considered to be the most primitive acarine group since they possess a number of
characteristics which are peculiar to unspecialized arachnid forms. Some of the more
important identifying features of the Notostigmata are:
I
1. Two pairs of lateral simple eyes, or ocelli, on the propodosoma. I
2. A series of weak transverse sutures which are demarcated by round or elliptical
muscle attachment sites, and which divide the hysterosoma into 12 "segments." I
3. Four pairs of small dorsolateral stigmatal openings arranged in a crescent on the
hysterosoma behind coxae IV (Plate 13-1}. Peritremes are absent.
I
4. More than 200 Iyriform pores, or Iyrifissures, on the dorsum (Plate 13-1). I
5. A terminal pal pal apotele which allows the palp to be used as a grasping structure
(Plate 13-2) (the more advanced parasitiform mites have a subterminal apotele on
the palptarsus which is primarily sensory in function).
I
6. A pair of large hypertrophied setae laterad to the setate hypostome on the
venter of the gnathosoma (With's organs). A pair of strong sci erotized rutella
I
flank these organs (Plate 3-3). The rutella may aid the mite in cleaning its
chelicerae or in holding its food. I
7. A strongly developed raduliform labrum (Plate 3-3).
projecting epistome.
There is no anteriorly
I
8. A paired tritosternal base (Plate 3-3) rather than the fused base found in more I
•
advanced parasitiform mites.

9. Divided trochanters 111-1 V (Plate 13-1}, plus false articulations on femora,

tibiae and tarsi.

10. Transverse genital apertures in male and female uncovered, between coxae III-IV. I
Notostigmatids prefer dark protected semi-arid habitats, and are found under rocks
or stones where they apparently prey on other small arthropods. They may be omnivorous I
to some extent, since pollen grains have been recovered in the ventricular contents of
dissected specimens. The distribution of the Notostigmata includes the southwestern
United States, South America, Central Asia and the Mediterranean region.
I
The Notostigmata are considered to be unique in the Acari, but show definite I
I
I
I 63

I affinities with the Parasitiformes. They are grouped in a separate order, the Opilioacari-

•
I
formes (Fig. 13), and are relegated to a single family, the OPILIOACARIDAE .

Useful References

Chamberlin, R.V. and S. Mulaik (1942). On a new family in the Notostigmata. Proe. BioI. Soc. Wash.
55: 125-132.

I Hammen, L. van der (1966). Studies on Opilioacarida (Arachnida) I. Description of Opi/ioacarus texanus
(Chamberlin and Mulaik) and revised classification of the genera. Zool. Verh. 86:1·80.
Hammen, L. van der (1968). Studies on Opilioacarida (Arachnida) II. Redescription of Paraearus
I hexophthalmus (Redikorzev). Zool. Meded. 43(5):57-76.
With, C.J. (1904). The Notostigmata a new suborder of Acari. Vidensk. Medd. Natur. Foren. Kobenh.:
137-192.

I
I
I
••
I
I
I
I
•
I
I
I
I
I
64
I
ORDER PARASITIFORMES I
Suborder Tetrastigmata

Unlike the Notostigmata, members of the suborder Tetrastigmata are heavily sclero-
••
•
tized non-segmented species without propodosomal ocelli. They are extremely large
(2000-7000 p), and somewhat rounded in shape. Useful identifying features include:

1. A pair of lateroventral stigmatal openings opposite coxae III which are con-
nected to peritremes (Plate 14-3). A second pair of openings behind coxae IV
(air sac pores) may be homologous to the expulsory vesicles in certain free-
living Mesostigmata.
I
2. Only four pairs of Iyrifissures on the dorsum, the median pair corresponding to
the propodosomal fissures of the Mesostigmata (Plate 14-3). With at least eight
I
additional pairs laterally.
I
3. A subterminal pal pal apotele which may be useful in holding or grasping food
(Plate 14-1,2). I
4. Lack of With's organs, rutella and epistome. The labrum is strongly developed
and raduliform. Corniculi are present.

5. Six pairs of setae on the hypostome (Plate 14-4). ••

6. Lack of a discrete tritosternum.

7. A transverse genital aperture covered by four setate shields (Plate 13-3). The
posteroventral anal aperture is covered by a pair of setate valves, each of which
may have Iyrifissures (Plate 13-4).

The Tetrastigmata apparently are predators, and have been found in Australia, New
•
I
Zealand, New Guinea, Ceylon, Seychelles and Mauritius. They comprise a single family,
the HOLOTHYRIDAE.
I
I
•
I mportant References

Hammen, L. van der (1961). Description of Ho!othyrus grandjeani nov. spec., and notes on the classifica-
tion of the mites. Nova Guinea n.s. 10(9): 173-194.
Hammen, L. van der (1965). Further notes on the Holothyrina (Acarida) l. Supplementary description
of Holothyruscoccinella Gervais. Zool. Meded. 40(28):253·276. I
I
I
I
I
I PLATE 13
65
I
•
I
I
I
I
I
I
•
13-1

I
•
I
stigma --+~'-J4-0 median genital shield

weak transverse
sutures
13-3

I
•
I
l'
I
I
I
I 13-1 and 13-2; family OPI L10ACARIDAE, Opilioacarus sp. (Mexico). 13-1; dorsum: 13-2; palpal tibia and tarsus, showing
terminal apotele

I
13,3 and 13-4; family HOLOTHYRI DAE, Holothyrus sp. (New Guinea). 13-3; genital valves of female: 13·4; anal valves of
female
 PLATE 14 I
66
I
C¥'M'~~-
palpal apotele subterminal

•
I
palpal tibia ----'''''''''''''''''.....'
I
14·1
I
I
14-2 I
I
••
peritreme --------4P.:

stigma--- -..l.~

._!-1ft:.'·:·
I
•
I

~·",,·~~~Jl •
I
I
I
•
14-3
14-4

14-1 to 14-4; family HOLOTHYRIDAE, Holothyrus sp. 14-1; pa/pal tarsus (after Camin etal. 1958): 14·2; palpal tibia and
I
tarsus: 14-3; anterodorsal aspect: 14-4; hypostome

I
I 67

I
•
ORDER PARASITIFORMES

Suborder Mesostigmata

I The Mesostigmata is a large and successful group of acarines which occupies a variety of

•
habitats. While the majority of mesostigmatid species are predators, many are external or
internal parasites of mammals, birds, reptiles or invertebrates (Strandtmann and Wharton
1958, Treat 1967). Mesostigmatids range in size from 200 to over 2000 Jl and usually
possess a number of sci erotized shields or plates on the dorsum and venter. A discrete

I sternal shield may be present in any stage. Other important identifying features are:

1. A pair of lateroventral or laterodorsal stigmatal openings between coxae II-IV,

I usually associated with elongate peritremes (Fig. 5c, p. 19). Peritremes may be

••
reduced or absent in certain families.

2. A pair of propodosomal Iyrifissures generally discernible (Fig. 2, p. 6). Round

and elliptical pores also are found dorsally in many families.

3. An inner basal palpal apotele with two, three or four tines (Fig. 14, p. 68). The

••
terminus of the palpal tarsus usually is equipped with several sensory setae.

4. A pair of horn-like corniculi at the terminus of the hypostome (Fig. 14, p. 68).
The hypostome has three pairs of setae arranged in a triangle, or in a virtually
straight line.

5. A tritosternum ventrally behind the gnathosoma, with 1-3 laciniae (Plate 15-10,
I 11,12). The laciniae, or the entire structure, may be absent in certain parasitic
families.

I 6. An anteriorly projecting epistome which may be simple or highly ornamented

••
(Plate 15). The labrum may be well developed, but is not clearly raduliform as
in the Tetrastigmata.

7. A transverse genital aperture in the intercoxal region covered with one, three, or
four shields in the female (Plate 15-7,8,9) and by one or two shields in the male.
The posteroventral anal valves may have a pair of setae.

I 8. Lack of an aedeagus in males. The male genital aperture is located either at the
anterior edge of, or within, the sternitigenital region.

I Members of the Mesostigmata may be found throughout the world in association with

••
soil, litter, nests, plants, food storages and animals. The suborder is divided into two
supercohorts primarily on the basis of genital shield configurations. Approximately 60
families are included in the suborder, and these are grouped into six cohorts and eleven
superfamilies.

I
I
68
I
I
-t--f-:=f---corniculus
palpal genu

" ' - ' r - - - - pal pal femur

••
•
Ir-......! - - - - pal pal trochanter
r:p::;,j===';;;;- hypostomal setae
nr-hff---t---- deutosternum
\\"I11,r--t---- deutosternal seta
I
__~\ V,!=,..-''-----'''--tritosternum

I
••
parapodal sh leld
sternal shield

peritreme
coxal opening III
peritremal shield
~·~rr~~t1(jpt:= metasternal shield
••
\ stigma
="-4-l--endopodal shield I
coxal opening IV

epigynial shield
I
"'\_.. -+__ ventral shield I
F,\-----,f--- metapodal shield

I
•
# - - - - , f - - - - - anal shield
I
\\J;:tr-'+.f----r------- anal valve

••
Fig. 14. Venter of a generalized mesostigmatid mite.

•
I
---------------------

'6
R)

its
Fig. 15. Three commonly used dorsal chaetotaxies for adult Mesostigmata (lines connote assumed vertical or horizontal relationships between
setal bases so connected) 15a after Evans and Browning 1956; 15b after Hirschmann 1957; 15c after Lindquist and Evans 1965.
 I
70

Useful References

Camin, J.H. and F. Gorirossi (1955). A revision of the suborder Mesostigmata (Acarina), based on new
interpretations of comparative morphological data. Chicago Acad. Sci. Spec. Pub!. 11 :70 pp.
Evans, G.O. (1957). An Introduction to the British Mesostigmata (Acarina) with keys to the families and
••
genera. Linn. Soc. Jour. Zoo!. 43:203-259.
I
••
Filipponi, A. (1962). Metodi sperimentali nella sistematica degli acari macrochelidi (Acarine, Mesostig-
mata, Macrochelidae). Parassit. 4(2-3): 113·146.
Hirschmann, W, (1957), Gangsystematik der Parasitiformes, Teil I. Rumpfbehaarung und Ruckenflachen.
Schriftenreihe fur vergleichende Milbenkunde: 20 pp. + plates.
Johnston, D.E, (1968), An Atlas of Acari I. The Families of Parasitiformes and Opilioacariformes, Acar.
Lab" Ohio State Univ. 172:110 pp. + x.
Karg, W. (1965). Larvalsystematische und phylogenetische Untersuchung sowie Revision des Systems der
Gamaslna Leach, 1915 (Acarina, Parasltiformes), Mitt. Zool. Mus. Beriin 41(2):193-340.
Karg, W. (1965), Neue Erkenntnisse zum System der Gamasina (Acarina, Parasitiformes) durch Larval-
I
•
systematische Untersuchungen. Zesz. Problem. Post. Nauk Roln. 65:89-114,
Karg, W. (19651. Die Anwendung Systematisch - Phylogenetlscher Arbeitsmethoden bei einer Bearbeitung
der Gamasina (Acarina, Parasitiformes). Zesz, Problem. Poxt. Nauk Roln. 65:115-138.
Karg, W. (1965). Entwicklungsgeschichtliche Betrachtung zur Okologie der Gamasina (Acarina, Paraslti-
formes). Zesz. Problem. Post. Nauk Roln. 65:139-155.
Schweizer, K. (1961). Die Landmilben der Schweiz (Mittelland, Jura und Alpen). Parasitiformes Reuter.
I
••
Mem. Soc, Helvet. Sci. Nat. 84:1·207 pp. + vii.
Strandtmarin, R.W. and G.W. Wharton (1958). Manual of mesostigmatid mites parasitic on vertebrates.
Inst. Acarology, Unlv. Maryland, Contr. 4:330 pp. + vii + 69 plates.
Tra"gardh, I. (1946). Outlines of a new classification of the Mesostigmata (Acarina) based on comparative
data. Kung!. Fysiogr. Sa"lIska. Hand!. 57:1·37.
Treat, A.E. (1967). Mites from noctuid moths. Jour. Lepid. Soc. 21(3):169-179.

Superfamily Parasitoidea

DIAGNOSIS: With a single epigynial shield, typically with one pair of epigynial setae
(none in some cases); sternal shield entire, may be fused with metasternals;
usually with one or two dorsal shields; tarsi I with or without apoteles.
I
•
Male genital aperture at anterior margin of sternal shield or within it,
male chelicerae often with spermatodactyl for the transfer of sperm to
female.
I
The Parasitoidea is a large and somewhat heterogenous assemblage of families which I
assume a wide variety of free-living or parasitic modes of life. Many are predaceous ground
or aerial forms (Sellnick 1958, Bregetova 1961, van der Merwe 1968), or vertebrate ecto- or
endoparasites (Tipton 1960, Radovsky 1967, Fain 1967). Others may be found as associates
I
of arthropods IEvans 1962, Hurlbutt 1967) or as detritus feeders.
I
••
Members of the families VEIGAIIDAE, PARHOLASPIDAE, EPICRIIDAE, ZERCON-
DAE and PARASITIDAE (among others) may be considered free-living predaceous
species. Except for the PHYTOSE II DAE and ASCI DAE, essentially all members of these
families occur in soil and humus, in rotting logs and other habitats where environmental
conditions favor large populations of microarthropods. Species of the PHYTOSEIIDAE and
ASCIDAE are aerial predators in orchards, where they prey on phytophagous mites.

I
I
I
71
I
•
Certain ascid mites act as predators on mites in stored grain. Blattisocius tarsalis (Berlese) is
particularly common in mite-infested grain storages.

A number of parasitoid families contain species which, although predaceous, are

associated phoretically with other animals during one or more stages of the mite's develop-
I ment. Phoresy or paraphagy on insects or my ria pods is common among predaceous

•
members of the families MACROCHELIDAE (Evans and Hyatt 1962), EVIPHIDIDAE
(Karg 1963), RHODACARIDAE (Ryke 1961), LAELAPIDAE (Evans 1955), AMERO-
SEIIDAE (Evans 1963), ASCIDAE (Lindquist and Evans 1965), DIGAMASELLIDAE
(Hurlbutt 1967) and PACHYLAELAPIDAE (Vitzthum 1931). Members of the genera

I Holocelaeno, Macrocheles and Neopodocinum (MACROCHELIDAE) exhibit varying

degrees of phoretic specificity to their scarabaeid beetle (Krantz and Mellott 1968) or
muscid fly carriers. Specificity also exists in other phoretic families, especially in the
I EVIPHIDIDAE and PACHYLAELAPIDAE. While most phoretic parasitoid associates of
insects attach to· the gular or coxal regions of their hosts, members of the genus Dino-
gamasus (LAELAPIDAE) display a somewhat more intimate physical relationship with their
I hosts, carpenter bees of the genus Mesotrichia. Here the mites are carried in the abdominal
acarinarium of the bee host, from which they emerge to feed on detritus or pollen in the

I bee gallery. Various myrmecophilous species of the families LAELAPIDAE and HETERO-
ZERCONIDAE may derive food directly from their hosts, feeding on salivary secretions of

•
the ants upon which they ride.

Species of the genus Rhinoseius (ASCIDAE) are phoretic on hummingbirds, utilizing

them for transportation from flower to flower where the mites may feed on other arthro-

I pods or on pollen (Baker and Yunker 1964, Lindquist and Evans 1965). A similar relation-
ship may exist between bees and mites of the genus Neocypholaelaps (AMEROSEIIDAE),
which have been collected from both bees and flowers (Evans 1963). Adults of the genus
I Aspidilaelaps (LAELAPIDAE) are associated with hermit crabs in the southwestern Pacific
region.

I Parasitoid parasites are found throughout the world on mammals, birds, reptiles and
insects, feeding on tissues or secretions of their various hosts. Most parasitic families are

I more or less specific to particular animal groups; for example, the families SPINTURNI-
CIDAE (Rudnick 1960) and SPELAEORHYNCHIDAE (Fain et al. 1967) are restricted

•
to bats, the HALARACHNIDAE to mammals, the RHINONYSSIDAE to birds, and the
ENTONYSSIDAE, OMENTOLAELAPIDAE and IXODORHYNCHIDAE to reptiles (Fain
1961, Strandtmann and Wharton 1958). The families MACRONYSSIDAE. DERMANY-
SSIDAE and HETEROZERCONIDAE comprise species which may be found on mammals,

I birds or reptiles and, in the case of the heterozerconids, on arthropods also. Members of the

•
family OTOPHEIDOMENIDAE are parasites of insects, feeding on exudates at sites of
chelicerallaceration of host tissues.

Primary injury to host animals by parasitoid parasites may be occasioned by the

piercing of the skin of the host, as with members of the genus Dermanyssus (DERMANYS-
I SIDAE), or through secondary injury of broken skin as with Laelaps (LAELAPIDAE).

•
I
Exsanguination of bird hosts by species of Ornithonyssus, and of snakes by Ophionyssus
(MACRONYSSIDAE) may cause anemia or death of the host (Bakeretal. 1956). Respira-
tory parasites ofthe families RHINONYSSI DAE, ENTONYSSIDAE and HALARACHNIDAE

•
72
I
I
•
often bring about lung congestion, sinusitis or rhinitis in host species. Species of the
families DERMANYSSIDAE, LAELAPIDAE and MACRONYSSIDAE are proven trans-
mittors of disease in birds, reptiles and mammals-including man.

Useful References I
Athias Henriot, C. (1961). Mesostigmates (Urop. excl.) edaphiques Mediterraneens (Acaromorpha,
Anactidotrichida). Premo Ser. Acarologia 3(4) :381·509.
Athias Henriot, C. (1968), Observations sur lesPergamasus IV. Un assai de coordination de la taxonomie
I
at de la choroJogie du sQus-genre Pergamasus 5.S. (Acariens Anactinotriches, Parasitidae). Acarologia
10(2):181-190.
Baker, E.W., T.M. Evans, D.J. Gould, W.B. Hull and H.L. Keegan (1956). A Manual of Parasitic Mites of
I
Medical or Economic Importance. Natl. Pest Control Assoc. Tech. Publ.:170 pp.
Baker, E.W. and C.E. Yunker (1964), New blattisociid mites (Acarina: Mesostigmata) recovered from
neotropical flowers and hummingbird's nares. Ann. Ent. Soc. Amer. 57(1):103-126. [ASCIDAEj
I
Beriese, A. (1910). Brevi diagnosi di generi e specie nouvi di Acari. Redia 6:346-388. [HETEROZER-
CONIDAE]
Bregetova, N.G. (1961). The veigaiaid mites (Gamasoidea, Veigaiaidae) in the USSR. Parazit. sborn. Zoo I.
Inst. Akad. Nauk SSSR 20:10-107.
I
Bregetova, N.G. (1967). Ontogenesis in the gamasid mites as a criterion for the erection of their natural
system. Akad. Nauk SSSR, Parasit. 1(6):465-479. I
Chant, D.A. (1965). Generic concepts in the family Phytoseiidae (Acarina: Mesostigmata). Can. Ent.
97(4) :351·374.
I
•
Damrow, R. (1965). New laelapid nasal mites from Australian birds. Acarologia 7(3):430-460.
[RHINONYSSIDAE]
Dove, W.E. and B. Shelmire (1932), Some observations on tropical rat mites and endemic typhus. Jour.
Parasitol. 18: 159·168. [MACRONYSSI DAEj
Evans, G.O. (1955a). A review of the laelaptid paraphages of the Myriapoda with descriptions of three new
(Acarina: Laelaptidae). Parasitol. 45:352-368.
Evans, G.O. (1955b). A revision of the family Epicrlidae (Acarina: Mesostigmata). Bull. Brit. Mus. (Nat.
Hist.) Zool. 3(4):171-200.
I
Evans, G.O. (1958). A revision of the British Aceosejinae (Acarina: Mesostigmata).
London 131(1):177-229. [ASCIDAE]
Proc. Zool. Soc.

Evans, G.O. (1962). Observations on the classification of the family Otopheidomenidae (Acari: Mesostig-
I
mata) with descriptions of two new species. Ann. Mag. Nat. Hist. 5(13) :609-620.
Evans, G.O. (1963). The genus Neocvpholaelaps Vitzthum (Acari: Mesostigmata). Ann. Mag. Nat. Hist.
6(13):209-230. [AMEROSEIiDAEj
I
Evans, G.O. and K.H. Hyatt (1958). The genera Podocinum Berl. and Podocinella gen. nov. (Acarina:
Mesostigmata). Ann. Mag. Nat. Hist. Ser. 12, X:913-932.
~vans,G.O. and K.H. Hyatt (1960). A revision of the Platyseiinae (Mesostigmata: Aceosejidae).
I
Bull. Brit. Mus. (Nat. Hist.) Zool. 6(2):25-101. [ASCIDAEj
I
•
Evans, G.O. and K.H. Hyatt (1962). Mites of the genus Macrocheles Latr. (Mesostigmata) associated with
coprid beetles in the collections of the British Museum (Natural History). Bull. Brit. Mus. (Nat. Hist.)
Zoo I. 9(9):327-401. [MACROCHELIDAE]
Evans, G.O. and W.M. Till (1965). Studies on the British Dermanyssidae (Acari: Mesostigmata). Part I.
External morphology. Bull. Brit. Mus. (Nat. Hist.) ZooI13(8):249-294.
Evans, G.O. and W.M. Till (1966). Studies on the British Dermanyssidae (Acari: Mesostigmata). Part II.
Classification. Bull. Brit. Mus. (Nat. Hist.) Zoo!. 14(5):109·370. I
•
Fain. A. (1961). Une nouvelle famille d'Acariens. Rev. Zool. Bot. Afr. 66(3-4):283-296. [OMENTO-
lAELAPIDAEj

I
•
II 73

II Fain, A. {1967l. Les acariens paras~tes nasicoles des oiseaux de Trinidad Indes Occidentales. I. Rhinonvs-
sidae: Mesostigm,tes. Bull. Inst. roy. Sci. nat. Belgique 43:1-44.

1. Fain. A .• G. Anastos. J. Camin and D. Johnston (1967). Notes on the genus Spelaeorhvnchus. Description
of S. praecursor Neumann and of two new species. Acarologia 9(31:535·556. [SPELAEORHYN·
CHIDAEJ

II Farrier, M.H. (1957).

124:103 pp.
A revision of the Veigaiidae (Acarinal. N. Carolina Agr. Exp. Sta. Tech. Bull.

Hirschmann, W. (1960). Gangsystematik der Parasitiformes, Teil 3. Die Gattung Dendrolaelaps Halbert
Schriftenreihe fUr vergleichende Milbenkunde: 27 pp. + plates. [DIGAMASELLIDAE]
II 1915.
Hurlbutt, H. (1967). Digamasellid mites associated with bark beetles in North America.
9(3) :497·534. [DIGAMASELLIDAE]
Acarologia

II Karg, W. (1963). Systematische Untersuchung der Eviphididae Berlese 1913 (Acarina, Parasitiformesl
mit einer neuen Art aus Ackerboden. Zool. Anz. 17017·8) :269·281.
Keegan, H.L., C.E. Yunker and E.W. Baker (19601. Malaysian Parasites. XLVI. Hvstrichonvssus

II turneri n. sp., n.g., representing a new subfamily of Dermanyssidae (Acarina) from a Malayan
porcupine. Inst. Med. Res. Fed. Malaya 29:205·208. [HYSTRICHONYSSIDAEJ
Krantz, G.W. (1960). A re·evaluation of the Parholaspinae Evans 1956 (Mesostigmata: Macrochelidael.
Acarologia 2141 :393-433. [PARHOLASPIDAE]
II Krantz, G.W. (1962), A review of the genera of the family Macrochelidae Vitzthum 1930 (Acarina:
Macrochelidae). Acarologia 4(2): 143·173.

II Krantz, G.W. and J.L. Mellott (19681. Two new species of Macrocheles (Acarina: Macrochelidae) from
Florida, with notes on their host-specific relationships with geotrupine beetles (Scarabaeidae:
Geotrupinael. Jour. Kansas Ent. Soc. 41(1):48.56.

il Lindquist, E.E. and G.O. Evans (1965). Taxonomic concepts in the Ascidae, with a modified setal
nomenclature for the idiosoma of the Gamasina (Acarina: Mesostigmatal. Mem. Ent. Soc. Canada
47:64 pp.
Merwe, G. van der (1968). A taxonomic study of the family Phytoseiidae (Acari) in South Africa with

I contributions to the biology of two species. S. Afr. Dept. Agr. Tech. Service Ent. Mem. 18:198 pp.
Radovsky, F .J. (1967). The Macronyssidae and Laelapidae (Acarina - Mesostigmatal parasitic on bats.
Univ. Calif. Publ. Ent. 46:288 pp.

I Rapp, A. (19591. Zur Biologie und Ethologie der Kafermilbe Parasitus coleoptratorum L. 1758 IEin
Beitrag zum Phoresie·Probleml. Zoo I. Jahr. Syst. 86(4·5) :303·366. [PARASITIDAEj
Rudnick, A. (1960), A revision of the mites of the family Spinturnicidae IAcarina). Univ. Calif. Publ.

I Ent. 17(21: 157284.

Ryke, P.A.J. (1961). A review of the genus Saintdidieria Oudemans (Acarina: Rhodacaridae) with re-
marks on the genus Lobocephalus Kramer. Acarologia 3(3) :250·255.

I Ryke, P.A.J. (1962l. The subfamily Rhodacarinae with notes on a new SUbfamily Ologamasinae

•
IAcarina: Rhocacaridae). Ent. Ber.8(22,1):155·162.
Sellnick, M. 119581. Die Familie Zerconidae Berlese. Acta. Zool. 3(3-4) :313·368.
Stammer, H.J. (edJ (1963). Beitr<3ge zur Systematik und Okologie Mutteleuropaischer Acarina.
Zoo I. Inst. Freidrich - Alexander - Univ, Bd. II. Mesostigmata 1:804 pp. + vii.
Strandtmann, R.W. and G.W. Wharton (1958). Manual of mesostigmatid mites parasitic on vertebrates.

I Inst. Acarology, Univ, Maryland, Contr. 4:330 pp. + vii + 69 plates.

•
Till, W,M. (1963). Ethiopian mites of the genus Androlaelaps Berlese s. lat. (Acari: MesostigmataL Bull.
Brit. Mus. (Nat. Hist.) Zool. 10(1):104 pp, [LAELAPIDAE]
Tipton, V.J. (1960) The genus Laelaps with a review of the Laelaptinae and a new SUbfamily Alphalae'
laptinae IAcarina: Laelaptidae). Univ. Calif. Pub!. Ent. 16(6):233·356 + 25 plates.
Vltzthum, H .G. (1931). Resultats scientifiques du voyage aux I ndes Orientales Neerlandaises de LL.AA.

I Belgique. Acarinen. Mem. Mus. Hist. Nat. Belgique IH.S.) 3(5):1.55. [PACHYLAELAPIDAE]
Womersley, H. (lm;6l. On some new Acarina - Mesostigmata from Australia, New Zealand and New
Guinea. Linn. Soc. Jour,·Zool. 42(288):505·599. [LAELAPONYSSIDAE]

I
I
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74
I
Superfamily Sejoidea I
DIAGNOSIS: With a single epigynial shield with one, three or several pairs of epigynial
setae; sternal shield divided behind sternals II; with one to several dorsal I
shields plus marginal platelets; tarsi I with or without apoteles. Male
genital aperture within sternal shield; male chelicerae unmodified for
I
•
sperm transfer.

The Sejoidea includes three recognized families, of which the SEJIDAE is the
best known. Sejids occur commonly in forest humus and litter in the tropics, but a few
species are found also in temperate North America and Europe. Zuluacarus termitophilus
(Trag.) is found scavenging in termite nests. One species was discovered on a rat in Puerto
Rico (Fox 1947), although no ectoparasitic relationship was established. Only three species
I
of MICROGYNIIDAE are described, with one being found in Northwestern United States
and two occurring in Scandinavia. Their habits are unknown, although their mouthparts
do not appear to be well adapted for a predaceous existence. Like the SEJIDAE,
I
microgyniids occur in humus and litter, or under the bark of rotting tree stumps. The
UROPODELLIDAE also are found in forest litter, where they may feed on fungi or organic
I
debris. Uropodellids have been collected in North and South America.
I
I
Useful References

Athias . Henrio!, C. (1960). Contribution aux Mesostigmates d'Algerie (Parasitiformes: Uroaspidae,

Veigaiidael. Acarologia 2(2): 15g·174. [SEJI DAEj
Balogh, J. (1938). Systematische Studien uber eine neue Milbengattung: Willmannia gen. nov. Zoo I. Anz.
123:259-265. [SEJIDAEj
I
Camin, J.H. (1955). UropodeJHdae, a new family of mesostigmatid mites based on Uropodella laciniata
Berlese, 188B (Acarina: Liraaspina). Bull. Chicago Acad. Sci. 10(5):65-81.
Gamin, J.H. and F.E. Gorirossi (1955). A revision of the suborder Mesostigmata based on new interpre-
I
I
tations of comparative morphological data. Chicago Acad. Sci. Spec. Pub!. 11 :70 pp.
Fox, J. (1947). Seven new mites from rats in Puerto Rico. Ann. Ent. Soc. Amer. 40:598-603.
[SEJIDAEj
Krantz, G,W. (1961). A re-evaJuation of the Microgynioidea, with a description of a new species of
Microgynium (Acarina: Mesostigmata). Acarologia 3(1):1-10.
Sellnick, M. (1953). Ichthyostomatogaster nyleni, eine neue Acaride aus Schweden. Ent. Tidsk.
I
74(1·2):24-37. [UROPODELLIDAEj
Tragardh, I. (1942). Microgyniina. a new group of Mesostigmata. Ent. Tidsk. 63(3-4):120-133. [MICRO-
GYNllDAEj
I
I
•
Superfamily Trachytoidea

DIAGNOSIS: With a single epigynial shield, without genital setae; sternal shield entire,
fused with endopodals and (in some cases) the metasternals; with two
dorsal shields (the posterior one a small pygidial plate) and usually with
I
•
marginal platelets or shields. Tritosternal base broader than long; tarsi I
with or without apoteles. Male genital aperture within sternal shield, male
chelicerae unmodified for sperm transfer.

I
•
II 75
II The Trachytoidea is an assemblage of species found in forest and treehole litter, moss,
under tree bark, in ant nests and on insects throughout most of the world. The superfamily
II consists of three families, the DISCOURELLIDAE, the TRACHYTIDAE and the POLY·
ASPI DAE. The discourellids are a holarctic group, with most of the known species being
European in distribution (Johnston 1961). Deutonymphs of the families TRACHYTIDAE
II and PO LY ASPI DAE often are found attached to insects. Apionoseius deutonymphs
(family TRACHYTIDAE), for example, have been recovered from trogine and geotrupine

II scarabs, while deutonymphs of Polyaspis (family PO LY ASPI DAE) are found frequently on
passalid beetles in neotropical regions. Polyaspid nymphs also have been recovered from
ant nests. Anal pedicels similar to those seen on phoretic deutonymphs of U ROPODI DAE

11 (Superfamily Uropodoidea) have been observed in the POLYASPIDAE.

Adult trachytoid mites apparently feed on organic detritus or fungi. Caminella

II peraphora (TRACHYTIDAE) is virtually semi-aquatic in habitat, being found only in wet

moss anchored in streams (Krantz and Ainscough 1960). Its feeding habits are unknown.

:1
:.
Useful References

Gamin, J.H. (1953). A revision of the cohort Trachytina Tra"gardh, 1938, with the description of
Dyscritaspis whf3:rtoni, a new genus and species of polyaspid mites from treeholes. Bull. Chicago

II Acad. Sci. 9(171: 335·385.

Hirschmann, W. (1961). Gangsystematik der Parasitiformes. Teil 4. Die Gattung Trichouropoda
Berlese 1916. Cheliceren und System der Uropodiden. Schriftenreihe fur vergleichende Milbenkunde:
41 pp. + plates.

I Johnston, D.E. (1961). A review of the lower uropodoid mites (former Thinozerconoidea, Protodiny-
choidea and Trachytoidea) with notes on the classification of the Uropodina. Acarologia 3(4):
522·545.

I Krantz, G.W. and B.D. Ainscough (1960), Caminella peraphora, a new genus and species of mite from
Oregon (Acarina: Trachytidae). Ann. Ent. Soc. Arney. 53(11 :27·34.
Tragardh, I. (1941). Further contributions towards the comparative morphology of the Mesostigmata.

I III. On the Polyaspididae Berl. Zoo I. Bidrag Fran Uppsala Bd. 20:345-357. [POL YASPIDAE]

I Superfamily Uropodoidea

DIAGNOSIS: With a single epigynial shield, without genital setae; sternal shield entire,
I typically fused with endopodal and ventral shields to form a ring; with one
or two dorsal shields (the posterior one a small pygidial plate), with or
without marginal shields; tarsi I with or without apoteles. Tritosternal
I base usually partially covered by coxae I, not broader than long. Male
genital aperture within sternal shield, male chelicerae unmodified for

I sperm transfer.

The Uropodoidea is a large cosmopolitan group consisting of fungivores (Radinovsky

I and Krantz 1961), insect associates (Elzinga and Rettenmeyer 1966), nidicoles (Berlese

•
1904) and occasional predators. Deutonymphs of the family U ROPODI DAE often attach to
the cuticle of insects by means of an anal pedicel (Fig. 9), leaving the carrier only after
molting to the adult stage. Adult uropodids occur commonly in forest litter and soil
detritus, while others are myrmecophilous or termitophilous (Urojanetia, Trachyuropoda,

I
I
76
I
Dentidinyehus and Urobove/la, for example). Urodiseella philoetena (Trouessart) attaches
to the tibial spur of legs J of its ant host and feeds on the debris which the ant cleans from
I
itself. Uropodid nest scavengers probably feed on fungi or detritus. Mycetophagous
uropodids also occur as part of the stored products and dung arthropod communities I
(Leiodinyehus, Trematura and Fuseuropoda spp.). One species of UROPODIDAE
(Fuseuropoda agitans (Banks)) often occurs in great numbers in commercial fishworm
beds, competing with the worms for food and providing a source of concern for producers
I
(Stone and Ogles 1953).

The peculiar species of the families COXEQUESOMIDAE, PLANODISCIDAE and

I
CI RCOCYLLIBAN IDAE are neotropical myrmecophiles which are associated with army
ants, sometimes in rather bizarre ways (Sell nick 1926). Coxequesoma eollegianorum
Sellnick, for example, is a capsulate form which covers the terminal antennal segments of
I
the ant host, presenting a smooth hard exterior to potential enemies. Members of the
families DINYCHIDAE, THINOZERCONIDAE, EUTRACHYTIDAE and METAGYNEL-
LI DAE are free-living species with no apparent associations with insects. A Prodinyehus
I
species (family DINYCHIDAE) has been observed to feed on slow-moving microarthropods.
This may occur among related uropodoid groups also, although the structure of the I
chelicerae points to a non-predatory mode of life.
I
Useful References

Berlese. A. (1904). Acari mirmecofili. Redia 1:299·474 + plates.

I
Camin, J.H. (1953). Metagynellidae, a new family of uropodine mite, with the description of Metagvnella
parvula, a new species from tree holes. Bull. Chicago Acad. Sci. 9(18):391-409.
Elzinga, R.J. and C.W. Rettenmeyer (1966). A neotype and new species of Planodiscus (Acarina:
I
Uropodina) found on doryline ants. Acarologia 8(2) :191-199.
Halbert. J.N. (19151. Acarinida: ii . Terrestrial and marine Acarina.
45-136. [THINOZERCONIDAE]
Proc. Royal Irish Acad. 31(39ii): I
Hirschmann, W. (1961). Gangsystematik der Parasitiformes. Teil 4. Die Gattung Trichouropoda
Berlese 1916. Cheliceren und System der Uropodiden. Schriftenreihe fur vergleichende Milbenkunde:
41 pp. + plates.
I
I
Johnston, D.E. (1961). A review of the lower uropodoid mites (former Thinozerconoidea, Protodiny·
choidea and Trachytoidea) with notes on the classification of the Uropodina. Acarologia 3(4):
522-545.
Krantz, G.W. (1969). The mites of Quintana Roo. l. A new species of Eutrachytes from Yucatan
Peninsula, with observations on the classification of the genus. Ann. Ent. Soc. Amer. 62(1 ):62-70.
Radinovsky. S. and G.W. Krantz (19611. The biology and ecology of granary mites of the Pacific
I
Northwest. II. Techniques for laboratory observation and rearing. Ann. Ent. Soc. Amer. 54(4):
512-518. [UROPODIDAE]
Sellnick, M. (1926). Alguns novos acaros (Uropodidae) myrmecophilos e termitophilos. Arch. Mus. Rio de
Janeiro 26:29-56. [COXEQUESOMIDAE, PLANODISCIDAE, CIRCOCYLLIBANIDAEj
I
Stone, P. and G.D. Ogles (1953). Uropoda agitans, a mite pest in commercial fishworm beds. Jour. Econ.
Ent.46:711. [UROPODIDAE] I
Tra·gardh. J. (1943). Zur Kenntnis der Prodinychidae (Acarina). Ark. Zoo I. 34A(21) 1-29.
Tragardh. J. (1944). Zur Systematik der Uropodiden. Ent. Tidskr. 65:173-186.
I
•
I
I
II 77

II Superfamily Protodinychoidea

11 DIAGNOSIS: With a single epigynial shield, without genital setae; sternal shield entire,
fused with endopodal, peritremal and parapodal shields and surrounding the
epigynial shield anteriorly; vertical shield of female free from median
11 dorsal shield, marginal shields absent; tarsi I with apoteles. Tritosternal
base twice as long as wide, not covered by coxae I. Male genital aperture

II within sternal shield, male chelicerae unmodified for sperm transfer.

Asinglefamilv, the PROTODINYCHIDAE, comprises this superfamily. Protodinychus,

11 the type and only genus, contains only one described species, although another is known
from North America (Johnston 1961). P. punctatus Evans was found in flood water
debris in England, while nymphs of the undescribed American species were taken from a

11 beetle associated with the habitat of the adults (beaver lodges). Their feeding habits are
unknown.

'I Useful References

!'.I
Evans, G.O. (1957). An introduction to the British Mesostigmata (Acarina) with keys to families and
genera. Linn. Soc. Jour. ~ Zool. 43(291):203~259.

Johnston, D.E. (1961). A review of the lower uropodoid mites (former Thinozerconoidea, Protodiny-
choidea. and Trachytoideal with notes on the classification of the Uropodina. Acarologia
3(4) :522~545.

I Superfamily Diarthrophalloidea

I DIAGNOSIS: With a single epigynial shield, without genital setae; sternal shield entire,
without lateral endopodal extensions, fused with metasternals; with one or

I two dorsal shields, marginal shields absent. Tritosternal base various, flanked
by pair of setae. Adanal setae long, often more than 1/2 length of body,
Tarsi I without apotele. Male genital aperture within sternal shield, male

I chelicerae unmodified for sperm transfer.

The Diarthrophalloidea is represented by a single family, the DIARTHROPHALLIDAE.

I The family is considered to comprise four genera, all of which are intimately associated
with beetles of the family Passalidae in New Guinea, Australia (Womersley 1961), and
throughout much of the neotropical realm. Diarthrophallid mites usually are found
I under the elytra of the beetle host either as larvae, nymphs or adults. Their constant
association with their beetle carrier may indicate a parasitic relationship. Diarthrophallus

I quercus (Pearse and Wharton) is the only known species of the family from North America
(H unter and Glover 1968c).

I
•
Useful References

Hunter, P.E. and Sandra Glover (1968a). The genus Passalobia Lombardini, 1926, with description of a
new species (Acarina: Diarthrophallidae). Proc. Ent. Soc. Wash. 70(1) :38-42.

I
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78
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Hunter, P,E, and Sandra Glover (1968b). The genus Brachvtremella Tragardh, 1946, with descriptions of
three new species (Acarina: Diarthrophallidael. Proc. Ent. Soc. Wash. 70(2): 114·125.
Hunter, P.E. and Sandra Glover (1968c). The genus Diarthrophallus Tragardh 1946 (Acarina: Diarthro·
phallidae). Proc. Ent. Soc. Wash. 70(3):193·197.
Tra'gardh, I. (1946). Diarthrophallina, a new group of Mesostigmata, found on passalid beetles. Ent.
Medd.24:369-394.
Womersley, H. (1961). On the family Diarthrophailidae (Acarina - Mesostigmata-Monogynaspida) with
particular reference to the genus Passalobia Lombardini 1926. Trans. Royal Soc. S. Australia
I
84:27-44.
I
Superfamily Cercomegistoidea

DIAGNOSIS: With two elongate latigynial and a reduced mesogynial shield covering
I
female genital aperture; sternal shield entire or divided; with one or two
dorsal shields, usually exhibiting hypertrichy. Tritosternal laciniae fused I
for at least 1/2 their length. Epistome without keel;' rarely with apoteles
on tarsi I. Male genital aperture within sternal shield, male chelicerae
unmodified for sperm transfer.
I
The Cercomegistoidea is a small group of trigynaspid mites contained in a single
family, the CERCOMEGISTIDAE. Members of the group have been found on ferns in the
I
southwest Pacific (Vitzthum 1935), or associated with bark beetles under bark of pine
trees in Australasia and in the New World. An undescribed species was collected from an ant
nest in India. The habits of the CERCOMEGISTIDAE are unknown, but it is probable that
I
these mites are fungivorous or predaceous on other small arthropods.
I
Useful References
I
Berlese, A. (1914). Acari nuovi, Manipulus IX. Redia 10:145-146.
Kinn, D.N. (1967).
9(3) :488-496.
A new species of Cercomegistus (Acari: Mesostigmata) from California. Acarologia I
Vitzthum, H.G. (1935). Terrestrische Acarinen von dem Marquesas. Bernice P. Bishop Mus. Bull.
142:64-99.
I
Superfamily Antennophoroidea I
DIAGNOSIS: Latigynial shields reduced, or expanded and fused, usually with distinct
mesogynial shield; sternal shield weakly sclerotized, entire or fragmented;
sternals I may be on jugular shields or free in integument; dorsal shield
I
entire. Tritosternal laciniae divided their entire length; epistome with
central keel; without apoteles on tarsi I. Male genital aperture within
sternal shield, male chelicerae unmodified for sperm transfer. Male and
I
female chelicerae with filamentous excrescences.
I
The superfamily Antennophoroidea contains three families, most of the species
of which are associated in the adult stage with arthropods. Members of the family

•
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II 79

11 ANTENNOPHORI DAE often are found in ant or bee nests in both temperate and tropical
climates of the New World and Europe (Berlese 1904). Species of the genus Antennophorus
11 attach to the gulal' region of the head, or to the body, of the ant host (Banks 1915). A.
foreli Berlese, as well as A. grandis Berlese (Donisthorpe 1927), have been observed to
stroke the ant's mouthparts with their first pair of legs, feeding on the droplet of fluid
II disgorged by the «nt. Members of the genus Physalozercon may also occur in ant nests.

The family PARAMEGISTIDAE is comprised of species associated with insects and

11 myriapods. One g,enus, however, is found on reptiles (Ophiomegistus). Species of Echino-
megistus and Antennomegistus often are found with beetles, while members of the genera

11 Paramegistus and Neomegistus may be observed on myriapods (Tragardh 1907). Unlike

most of the Trigynaspida, representatives of the PARAMEGISTIDAE commonly are
collected in the Old World tropics.
il The PARANTENNULIDAE comprises three genera which are associated primarily
with millipedes in both the Old and New Worlds. A species of Micromegistus has been
I found on a carabid beetle and in carrion at two localities in north central United States.
Their feeding habits are unknown, although it seems likely that parantennulids may be

I capable of deriving food directly from their hosts in a manner similar to that seen in
mrymecophilous ANTENNOPHORI DAE.

II Useful References

I Banks, N. (19151. The Acarina or Mites. U.S.D.A. Rept. 108:86.

Berlese. A. (19041. Acari mirmecofili. Redia 1:299-474 + plates. [ANTENNOPHORIDAE]
Camin, J.H. and F.E. Gorirossi (1955). A revision of the suborder Mesostigmata based on new interpreta-

I tions of comparative morphological data. Chicago Acad. Sci. Spec. Publ. 11 :70 pp.
Donisthorpe, H. St. J.K. (1927). The Guests of British Ants. London:244 pp. + 16 plates.
Tragardh. I. (19061. Neue Acariden aus Natal und Zululand. Zool. Anz. 30:870-877. [PARAMEGI-
I STIDAE]
Tragardh, I. (1907). Description of two myriopodophilous genera of Antennophoridae with notes on
their development and biology. Ark. Zool. 3(281:1-33 + plate. [PARAMEGISTIDAE]

I Tragikdh, I. (1943). Further contributions towards the comparative morphology of the Mesostigmata
(Acarina). The Antennophoridae and the Megisthanidae. Ark. Zool. 34A(20): 1-1 O.

I Superfamily Megisthanoidea

I DIAGNOSIS: Latigynial and mesogynial shields fused, with entire or divided sternogynials;
sternal shield entire or divided behind sternal setae I, fused with meta-
sternals; with jugular shields or fleshy projections bearing sternal setae I;
I dorsal shield entire. Tritosternal laciniae divided, epistome with keel; tarsi I
without apoteles. Male genital aperture within sternal shield, male chelicerae

I unmodified for sperm transfer. Male and female chelicerae with tree-like
excrescences.

I
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80

Two families are relegated to the superfamily Megisthanoidea-the MEGISTHANIDAE

and the HOPLOMEGISTIDAE. Both are associated primarily with passalid beetles. 1
from which they may derive sustenance via the glandular secretions of their host. Megis-
thanus floridanus Banks is a common species on Popilius disjunctus III., the passalid beetle
found throughout the southeastern United States. Other species of the genus are found
••
clinging to passalid and dung beetles throughout much of the world, including the neo-
tropical and australasian realms (Stoll 1886, Womersley 1937).
I
Members of the HOPLOMEGISTIDAE also utilize passalid beetles, and are common
in the neotropics (Stoll 1886). Their feeding habits are unknown.
I
Useful References
I
Johnston, D.E. (1960). Laelaptid mites associated with hermit crabs (Paguridea). The genera Aspidilaelaps
and Cyclothorax (Acarina - Mesostigmata). Acarologia 2(4):446. [MEGiSTHANIDAE]
I
Stoll. O. (18861. Arachnida Acaridea. Bioi. Centrali - Americana: 55 pp. + xvi + plates.
THANIDAE, HOPLOMEGISTIDAE]
Triigardh, I. (1937). Zur Svstematik der Mesostigmata. Ark. Zoo I. 298(11):1-8. [HOPLOMEGISTIDAE]
IMEGIS-
I
TrEigardh, I. (1943). Further contributions towards the comparative morphology of the Mesostigmata
(Acarinal. Ark. Zoo I. 34A(20):1·10. [MEGISTHANIDAEj I
Vitzthum, H.G. (1937). Acari in dar Kiemenh6hle von Birgus latro (Crustacea, MacruraL Zeitschr.
Parasit.9(5):638-647. [MEGISTHANIDAE]
Womersley, H. (19371. Australian Acari of the genus Megisthanus Thorell. Trans. Roy. Soc. So. Austral.
61:175-180.
I
I
Superfamily Celaenopsoidea

DIAGNOSIS: With distinct latigynials, either fused or freei mesogynial shield present or
I
absent without sternogynialsi sternal shield entire, metasternals free or
fused with each otheri dorsal shield entire. Tritosternal laciniae normali
epistome with keeli without apoteles on tarsi I. Male genital aperture at
I
anterior margin of sternal shield,· male chelicerae unmodified for sperm
transfer.
scences.
Male and female chelicerae with tree-like or brush-like excre- I
The Celaenopsoidea contains the families DIPLOGYNIIDAE, SCHIZOGYNIIDAE, I
EUZERCONIDAE and CELAENOPSIDAE, most species of which are found in association
with insects or myriapods. Members of the DI PLOGYN II DAE are common on passalid and
histerid beetles in the New World and in Ceylon, and have been found under bark (Tnjgardh
I
1950) and on a variety of dung beetles from Central Africa. Adults of Ophiocelaeno
sellnicki Johnston and Fain are found on snakes (Johnston and Fain 1964). Other
collections include species from treeholes in north central United States, associates of
I
1Cy c/othorax carcinicola von Frauenfeld, ostensibly a member of the MEGISTHANIDAE found in
I
association with coenobitid crabs, is probably a member of the monogynaspid family LAELAPIDAE
(Johnston 19601. I
I
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II
81
II termites in Oregon (Krantz 1958), and a symbiont from a palm weevil in Nicaragua (Hicks
1958). The feeding habits of diplogyniids have not been studied.
II Species of SCH IZOGYN II DAE occur with passalid beetles in South Africa (Tragardh

il 1950, Ryke 1957), India (Sellnick 1954), and with bark beetles in western United States
(Kinn 1966). An undescribed species has been collected in Central America. Several species
of CE LAENOPSI DAE are known to occur with bark beetles in Europe and Africa (Tragardh
il 1950, Kinn 1968), one with ants in Tasmania (Tragardh 1951) and others in association

i.
with bark beetles in southern U.S.A. The celaenopsids comprise a primarily holarctic
family which, according to Tragardh, may be a relict group which has survived in the
temperate climates of Europe [and of North America] in the galleries of their bark beetle
associates.

I Species of the family EUZE RCON I DAE are common associates of passalid beetles and
my ria pods in the Western Hemisphere and Africa. They are known to occur in the East
Indies and the Philippines, and probably will be found to be world-wide in distribution.
I Observations on Euzercon latus (Banks), an associate of the horned passalus beetle of east-
ern United States, indicate that immature E. latus may be scavengers or fungivores in

I decaying wood and litter (Hunter and Davis 1965). Only adults are found on the beetle
host. Hunter and Davis observed adults to feed on Collembola eggs and on inactive
Collembola. They conclude that E. latus is not an active predator in the adult stage.

I Useful References

I Funk, R.C. (1964). An investigation of the Euzerconidae (Mesostigmata: Celaenopsoidea) based on the
procedures of numerical taxonomy. Acarologia 6(fasc. h.s.): 127-132.
Hicks, EA. (19581. A new genus and species of diplogyniid from Nicaragua (Order Acarina, Family

I Diplogyniidae). Iowa State Call. Jour. Sci. 33(2):103-110.

Hunter. P.E. and R. Davis (1965). Mites associated with the passalus beetle. III. Life stages and
observations on the biology of Euzercon latus (Banks) (Acarina: Euzerconidael. Acarologia
7(11:30-42.
I Johnston, D.E. and A. Fain (1964).
associated with snakes (Acari
Ophiocelaeno sel/nicki, a new genus and species of Diplogyniidae
Mesostigmata). Bull. Am. Soc. Roy. d'Ent. Belg. 100(6):79·91.

I Kinn, D.N. (1966). A new genus and species of Schizogyniidae from North America with a key to the
genera. Acarologia 8(41 :576-586.
Kinn, D.N. (1968). A new species of Pleuronectocelaeno (Acarina: Celaenopsidae) associated with bark
beetles in North and Central America. Acarologia 10(21:191-205.
I Krantz, G.W. (19581. Lobogyniella tragardhi, a new genus and species of diplogyniid mite associated
with dampwood termites in Oregon (Acarina: Diplogyniidael. Proc. Ent. Soc. Wash. 60(3):127,131.

I Ryke, P.A.J. (1957). Mixogynium proteae, a new genus and species of Celaenopsoidea (Mesostigmata:
Acarina) from South Africa. Ann. Mag. Nat. Hist. 10:579-584. [SCHIZOGYNIIDAE]
Samsinak. K. (19571. Die mitteleuropaischen Arten der Familie Diplogyniidae (Acari). Acta Soc. Ent.
Cechoslov.54("1):1·6.

I Sell nick, M. (1954). Indogynium lindbergi nov. gen. nov. spec., eine neue Acaride aus Indien. Ent.
Tidskr.75:285-291. [SCHIZOGYNIIDAE]
Triigardh, I. (1941). Contributions towards the comparative morphology of phylogeny of the Mesostig-
I mata. IV. On the Celaenopsidae and Euzerconidae. Ent. Tidskr. 62: 169·176.

•
Tragardh, I. (19501. Studies on the Celaenopsidae, Diplogyniidae and Schizogyniidae (Acarina). Ark.
Zool. ser. 2,1(:251:361-451.
Tragardh, I. (1951). Brachycelaenopsis, a new genus of Celaenopsidae (Acarina) from Tasmania.
Ent. Tidskr. 72(1-21:60-64.

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82
I
Superfamily Fedrizzioidea I
DIAGNOSIS: Latigynials distinct or reduced to narrow rims adjacent to enlarged
sternogynial shield, mesogynial shield distinct or reduced to remnant;
I
sternal and metasternals fused, with free or fused jugular shields carrying
sternal setae I; dorsal shield entire. Tritosternal laciniae divided; epistome
with keel; tarsi I without apoteles. Male genital aperture within sternal
I
shield, male chelicerae unmodified for sperm transfer. Male and female
chelicerae with filamentous excrescences. I
Members of the two families of the Fedrizzioidea are tropical in distribution, and are
found commonly on beetles of the family Passalidae. This may be either a purely phoretic I
association, with no feeding taking place on the beetle, or an association in which the
mites feed on glandular secretions of the beetle hosts. Non-phoretic phases may be
predaceous or fungivorous. Representatives of the family FEDRIZZIIDAE are reported I
from passalid species in Central and South America and in the southwest Pacific. Some
species have not been found in direct contact with a passalid host (Neofedrizzia vitzthumi
(Oud.) and N. camini Womersley, for example), but rather in a locale which might well be
I
frequented by passalids (rotting logs, fungi etc.) (Womersley 1959a). The three recognized
genera of KLI NCKOWSTROEMII DAE apparently are restricted to the neotropical realm
(Womersley 1959b), where they associate with passalid beetles.
I
Usefu I References
I
Camin, J.H. and F.E. Gorirossi (1955). A revision of the suborder Mesostigmata based on new interpreta-
tions of comparative morphological data. Chicago Acad. Sci. Spec. Pub!. 11 :70 pp.
I
Hunter, P.E. and L. Butler (1966). New Klinckowstroemia mites from Costa Rican passalid beetles
(Acarina: Klinckowstroemiidael. Jour. Georgia Ent. Soc. 1(41 :24·30.
Sellnick, M. (1938). Eine neue Milbengattung aus Slid-und Mittelamerika. Arb. Morph. Taxon. Ent.
I
5:184-186. [KLINCKOWSTROEMIIDAE]
Tragardh, I. (19371. Zur Systematik der Mesostigmata. Arkiv. f. Zool. 29B(11 ):1-8. I
Triigardh, l. (1946). Outlines of a new classification of t~e Mesostigmata (Acarina) based on comparative
morphological data. Kung!. Fysiografiska Sa'lIskapets Handl. N.F. 57(4):1-37.
Turk, F.A. (19481. Insecticolous Acari from Trinidad, B.W.1. Proc. Zoo!. Soc. London 118(1) :82-125,
[KUNCKOWSTROEMIIDAE]
I
Womersley, H. (1959a). Some Acarina from Australia and New Guinea paraphagic upon millipedes and
cockroaches and on beetles of the family Passalidae. Pt. 2. FedrizzHdae. Trans. Royal Soc. So.
Australia 82: 11-54.
I
Womersley, H. (1959b). Klinckowstroemiella helleri (Ouds.. 1929) nov. comb. for Fedrizzia helleri Ouds.,
1929 (Acarina· Klinckowstroemiidae). Zoo!' Meded. 34(19):281·288. I
I
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II
83
il Suborder Mesostigmata

:. (Plates 15 to 52, pp. 95-132)

KEYTOTHE FAMILIES

I 1. One primary genital cover (epigynial shield) well developed and functional or, if

I reduced and non-functional, usually represented by remnant flanked by or bearing

a pair of genital setae (Fig. 16). Legs I with or without apoteles (legs II-IV with
apoteles in both supercohorts). Chelicerae without filamentous or tree-like
I excrescences on movable digit (Plate 16-1) (one or two arthrodial brushes may
occur at the base of the movable digit) Supercohort MONOGYNASPI DA 2

I Three primary genital covers (two latigynials and one mesogynial shield) functional
or variously coalesced or reduced (Fig. 17); when coalesced, without outwardly
opening, trapdoor-like genital shield (17c); when reduced a sternogynial shield always

I present. Legs I usually without apoteles. Chelicerae with filamentous, brush-like or

tree-like excrescences on movable digit (Plate 16-2); occasionally with additional
excrescences at base of digit Supercohort TRIGYNASPJDA '" 48

I 2. Epigynial shield well developed or reduced, with one or more pairs of setae
(epigynial shield reduced in many parasitic forms, lost in some endoparasites of
I vertebrates); epigynial setae may flank the shield in species where the shield is
narrowed (Fi£!. 16f). Hypostomal setae in a subtriangular pattern or in a straight
line. Male genital aperture within sternal shield, (Plate 16-3) or at its anterior
I margin (Plate 16-4) 3
Epigynial shield well developed and without genital setae (the subgenus Dyscritaspis

I (Fig. 16d) and the genera Dipolyaspis and Protodinychus each have a pair of setae
on the epigyr ial shield, but these are thought to be the metasternals and can be
distinguished from true genital setae in that they are accompanied by two pairs of
I pores, one of which is the 3rd sternal pores). Hypostomal setae in a straight line,
never in a triangular pattern. Male genital aperture within sternal shield 37

I 3. Epigynial shield with one, three, or several pairs of genital setae (Plate 16-5,6,7)'
Sternal shield divided behind sternal setae II. With one to several dorsal shields

I plus marginal platelets (Plate 18). Male genital aperture between coxae II, III or IV.
Male chelicerae unmodified Cohort SEJINA, Superfamily SEJOIDEA 4
Epigynial shield with one pair of setae (Fig. 16a) or, if coalesced with ventral
I shield (Fig. 1Gb), with only one pair of setae in the podosomal region (if not on the
shield, the genital setae may be distinguished by the pair of pores associated with
them (Plate 26-1)). Sternal shield entire, although presternal fragments may be
I present (Plates 19-1, 22-10). Usually with one or two dorsal shields and no marginal
platelets. Male genital aperture at anterior margin of sternal shield, or within it.

I Male chelicerae mayor may not bear a spermatodactyl (Plate 5-1, p. 15)

4. With one pair of genital setae; male genital aperture behind coxae II .
7

I ...................................... (Plate 17) Family MICROGYNIIDAE

With more than one pair of genital setae; male genital aperture between coxae II
.................................................................. 5
I
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 I
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I
Fig. 16. Representative epigynial and genitiventral shields of the supercohort Monogynaspida. Arrows
I
•
indicate possible evolutionary trends in expansion or reduction of shield size from a simple
truncate type (16a).

I
II
85
II
II
!I
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I
I Fig. 17. Representative genital shields and associated structures in the supercohort Trigynaspida. Arrows
indicate possible evolutionary trends in expansion, fusion or loss of various components, with the

I diplogyniid genital apparatus considered the basic type (17a), The mesogynial shield is stippled
for clarity.

I
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86
I
5. Epigynial shield large, covering portions of sternal shield II; with one to four dorsal I
shields 6
Epigynial shield not covering sternal shield II; with two to seven dorsal shields
· (Plate 18) Family SEJIDAE
I
6. Both sexes with a pair of large median subequal dorsal shields and a pair of smaller
lateral shields; female with large metapodal shields; male with holoventral shield
I
· (Plate 17) Family UROPODELLIDAE
Female with a large podonotal shield and a small pygidial shield; male with entire
I
dorsal shield; metapodal shields of female strongly reduced; male with a separate
ventrianal shield. . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Asternolaelaps Berlese 1 I
7. Sternal shield of female fused with endopodal, parapodal and peritremal shields and
partially encircling epigynial shield (Plate 42-2)
Sternal shield of female various, but not as above
· :
Cohort UROPODINA (pars)
.
Cohort GAMASINA, Superfamily PARASITOIDEA 8
42

8. Stigmata located dorsolaterally in a distinctive tuberculate-reticulate dorsal shield;

•
I
peritremes absent. Tarsi I with specialized clubbed sensory setae, without claws.
Free-living (Plate 19) Family EPICRIIDAE
I
Stigmata usually located ventrally or lateroventrally (located dorsally in certain
parasitic families), peritremes present or absent. Dorsal shield variously ornamented I
but not as above. Tarsi I without clubbed setae as above 9

9. Epigynial shield triangular and flanked by large discrete metasternal shields. Pal pal
I
apotele three-tined. Male with apophyses on legs II. Free-living .
· (Plate 20 [see also 10-1]) Family PARASITIDAE
Epigynial and metasternal shields, when present, not as above. Palpal apotele
I
various. Male with or without apophyses on legs II

10. Hypostome with distal, usually mustache-like, fringe. Palpal apotele with three tines
10
I
and with associated inflated hyaline, membranous expansion. Epigynial and ventral
elements usually separated by incomplete suture. Dorsal shield partly or completely I
divided subequally. Free-living... (Plate 21 [see also 4-4,16-4]) Family VEIGAIIDAE
Hypostome with or without distal fringe, but not mustache-like. Palpal apotele with
two or three tines, without membranous expansion. Epigynial and ventral shields
I
not as above. Dorsal shield divided or entire 11
I
11. With one or two plumose or filamentous arthrodial processes, or with an arthrodial
brush and an adjacent coronet of setae at the base of the movable digit of the
chelicera. Palpal apotele with three well-developed tines. . . . . . . . . . . . . . . . . . .. 12
Arthrodial process at base of movable chelicera I digit, when present, rarely forming
a distinct brush. Pal pal apotele two- or three-tined (when three-tined, the most
proximal element often is reduced) 13
•
I
l A genus of the family ICHTHYOSTOMATOGASTERIDAE, which has not been figured in the
accompanying plates. The reader is referred to Johnston (1968) for illustrations.
I
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II 87
II 12. Peritreme generally looped proximally, joining the stigma posteriorly; legs I usually
without apoteles; epigynial shield with a pair of well developed lateral accessory
il sclerites; laterodistal elements of pretarsi II-IV (posterior paradactyli) divided or
serrate distally, or membranous proximally. Free-living or insect associates .
· (Plate 22 [see also 3-1, 3-2,8-4, 15-5] ) Family MACROCHELI DAE
I Peritreme normal, joining the stigma anteriorly; accessory sclerites weak or absent;
legs I with or without apoteles; laterodistal elements of pretarsi II-IV setate, not

I divided distally. Free-living ..... (Plate 22 [see also 5-1]) Family PARHOLASPIDAE

13. Sternal shield complete, much longer than wide, reaching to middle of coxae IV;
I peritremal, parapodal and metapodal shields contiguous or fused, extending
posteriorly beyond coxae IV. Free-living or associated with insects .
· " , (Plate 23) Family PACHYLAELAPIDAE
I Sternal shield complete or fragmented, mayor may not be longer than wide;
sternal elements rarely reaching middle of coxae IV (e.g., HETEROZERCONIDAE).

I Peritremal and metapodal shields separate 14

14. Sternal shield over six times wider than long at its widest point. Epigynial and
I ventral shields forming an insensibly fused membrane, anal shield separate.
Opisthosoma of female considerably broader than long, with a fringe of spatulate
setae. Parasites of snakes (Plate 23) Family OMENTOLAELAPIDAE
I Sternal shield dimensions, condition of ventral shields and shape of opisthosoma
various, never as above 15

I 15. Venter of opisthosoma with a pair of well-developed discs or suckers. Associated

with snakes and arthropods , (Plate 24) Family HETEROZERCONIDAE

I Venter without discs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

16. Apotele of pal pal tarsus usually with two tines (if three-tined, then corniculi may be
I divided distally) '
Apotele of pal pal tarsus with three tines; corniculi undivided distally
20
17

I 17. Chelae chelate· dentate or, if fixed digit reduced, with movable digit dentate 18
Chelae simple and finger-like. Associated with insects .
I · " (Plate 23) Family LAELAPONYSSIDAE

I 18. Tarsi I without claws and empodium. Dorsal shield entire (if abbreviated, then with
mesonotal scutella) 19
Tarsi I usually with claws and empodium, or with claws (when claws are absent the
I dorsal shield is divided). A heterogenous aggregation of free-living species .
.. . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... (Plate 25) Family RHODACARIDAE 1

I
I lComprises the families CYRTOLAELAPIDAE, HALOLAELAPIDAE and RHODACARIDAE of
Johnston (1968).

I
I
88
I
19. Tarsus I terminating in one or two long whip-like setae. Epigynial shield truncate, I
with epigynial setae inserted on shield. With a well-developed ventrianal shield.
Male genital aperture at anterior edge of sternal region. Free-living
· ,."
.
(Plate 24 [see also 15-7]) Family PODOCINIDAE
I
Tarsus I terminating in a series of short simple hairs. Epigynial shield rounded
posteriorly, epigynial setae adjacent to shield. With an anal shield. Male genital
aperture within sternal region. Free-living .
I
· (Plate 26 [see also 15-3, 16·1]) Family ARCTACARIDAE
I
20. Epigynial shield truncate posteriorly (Fig. 16a) or fused with a ventrianal shield
(Fig. 16g), rarely rounded (if sides are parallel, shield may be slightly convex
posteriorly. If shield is rounded posteriorly, then anal shield is not triangular in
I
shape) 21
Epigynial shield rounded or pointed posteriorly (Fig. 16j), occasionally greatly
reduced (Fig. 16k) or lacking. Usually with a separate triangular anal shield.
I
Parasites of mammals, snakes, birds or insects, or phoretically associated with
arthropods 26 I
21. Sternal shield of female with two pairs of setae (III-IV off the shield). Palpal
apotele 2· or 3-tined. Corniculi may be divided distally or otherwise modified in the
I
distal portion (Ameroseius). Free-living or associated with insects .
........................................ (Plate 27) Family AMEROSEIIDAE
Sternal shield with 0-4 pairs of setae. Corniculi not modified as above 22
I
22. Metasternal setae on separate platelets or inserted free in integument adjacent to I
sternal shield. Sternal shield with two or three pairs of setae. Dorsal shield entire,
incised or divided medially
Metasternal setae inserted on the posterolateral angles of the sternal shield; shield
23
I
with four (rarely three) pairs of setae. Dorsal shield divided medially. Free-
living or associated with insects (Plate 28) Family DIGAMASELLIDAE I
23. With fewer than 20 pairs of dorsal setae; with 0-3 pairs of sternal setae
With more than 20 pairs of dorsal setae; with three pairs of sternal setae
24
25
I
24. Fixed cheliceral digit normally produced; anal opening subterminal. Predators,
usually found on shrubs or trees, or in some cases, fungus or plant feeders .
I
.................................... (Plates 28, 29) Family PHYTOSEIIDAE
Fixed chelicera I digit reduced; anal opening terminal. Parasites of insects
·
.
(Plates 29, 30) Family OTOPHEIDOMENIDAE
I
25. Dorsal shield divided medially. With a transverse row of four large "pores" on the I
posterior edge of opisthonotal shield. Peritremes greatly reduced in length. With a
ventrianal shield. Male genital aperture within sternal region. Free-living .
I
•
· (Plate 30 [see also 16-3]) Family ZERCONIDAE

I
I
I 89

I Dorsal shield variously produced. Without distinct opisthonotal pores as above.

Peritremes normally produced. With a ventrianal or an anal shield, the latter

I rarely being triangular. Male genital aperture at anterior edge of sternal region.
Free-living, or phoretically associated with insects or birds .
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. (Plate 31) Family ASCI DAE

I 26. Chelicerae lar!}e, hooked, set in an obvious camerostome. Bat parasites .

· (Plate 32) Family SPELAEORHYNCHIDAE
I Without camerostome or hooked chelicerae 27

I 27. Peritremes absent or greatly reduced.

birdS or snakes
Respiratory tract parasites of mammals,
28
Peritremes may be shortened, but always distinct. External parasites of vertebrates
I or invertebrates, or free-living 30

28. Epigynial shield lacking or rudimentary (distinct in Zumptiella bakeri); sternal

I shield generally present. Stigmata ventral. Respiratory tract parasites of mammals.
.. . . .. .. .. . . .. .. . . . . . . . . . . .. .. . . .. .. (Plate 33) Family HALARACHNIDAE

I Epigynial shield present, sometimes reduced. Stigmata dorsal or lateroventral ..... 29

29. Sternal and epigynial shields well developed but often weakly sci erotized; sternal
I setae weak or absent. Stigmata lateral or lateroventral. Respiratory tract parasites
of snakes (Plate 34) Family ENTONYSSIDAE

I Epigynial shield present, but reduced; sternal shield reduced or lacking, but with
distinct sternal setae in podosomal region. Stigmata dorsal. Respiratory tract
parasites of birds (Plates 34, 35) Family RHINONYSSI DAE

I 30. Tritosternum absent or vestigial (if present, then legs I extremely stout and
peritremes extending anteriorly to middle of coxae I) 31

I Tritosternum well developed, with laciniae 32

I 31. Legs I greatlv enlarged, considerably thicker than legs II-IV. Stigmata ventral.
Parasites of edentates (Plate 35) Family DASYPONYSSI DAE
Legs I-I V of comparable thickness. Stigmata usually dorsal. Parasites of bats

I ......................... (Plate 36 [see also 15-10]) Family SPINTURNICIDAE

32. Chelicerae weakly developed; stylettiform, edentate, or with poorly defined

I transparent te'9th. Corniculi often indistinct
Chelicerae strong, dentate. Corniculi well defined
33
35

I 33. Chelicerae attenuated, stylettiform; palpal trochanter without ventral spur 34

Chelicerae not stylettiform; chelae normally developed, edentate; palpal trochanter
I with ventral spur. Parasites of mammals, birds and reptiles .
· . . . . .. . . . .. . . . . . . . . .. . . . . . . . . . .. (Plates 36, 37) Family MACRONYSSIDAE

I
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I
90
I
34. Distal cheliceral segment elongate, far exceeding basal segment in length; movable
digit minute. Parasites of rodents and birds .
I
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. (Plate 37) Family DERMANYSSIDAE1
Distal cheliceral segment normally developed, considerably shorter than greatly
I
elongated basal segment (Plate 4-6). Palpal trochanter without spur. Idiosoma
attenuate terminally. Parasites of porcupines and snakes
· (Plate 38 [see also 4-6]) Family HYSTRICHONYSSIDAE
. I
35. Corniculi elongate and barbed distally; chelicerae with large, often recurved
teeth. Ventral seta of coxae II, I-II, or I-III spur-like. Parasites of snakes .
I
................................... (Plate 38) Family IXODORHYNCHIDAE
Corniculi, chelicerae and coxal setae not as above. Parasitic on vertebrates,
I
associated with invertebrates, or free-living 36

36. Central element of epistome typically long and tongue-like. Epigynial shield
I
almost parallel-sided. Peritremal shields often extend beyond coxae IV. With only
one anterolateral seta on genu I and on tibia I (Plate 39-1). Commonly associated
with insects or crustaceans ..... (Plates 38,39 [see also 15-2]) Family EVIPHIDIDAE
I
Central portion of epistome not as above. Epigynial shield drop-shaped (when
posterior border of shield extends to the anal shield, then posterior portion may I
assume flattened (Fig. 16c, p. 84) or invaginated aspect; sternal, epigynial and/or
dorsal shields may exhibit hypertrichy. Peritremal shield usually not extending
beyond coxae IV. Genu and tibia I each with two anterolateral setae (Plate 39-6).
I
Free-living, associated with invertebrates, or parasitic on birds or mammals
· (Plates 39, 40) Family LAELAPIDAE I
37. Sternal shield without lateral, intercoxal projections. Adanal setae long, often
more than one-half the length of the body. Associated with passalid beetles
Cohort DIARTHROPHALLINA Superfamily DIARTHROPHALLOIDEA
.
.
I
................................. (Plate 41) FamilyDIARTHROPHALLIDAE
Sternal shield insensibly fused with endopodals and projecting laterally between
I
coxae. Adanal setae normal. Free-living or insect associates .
· Cohort UROPODINA (pars) 38 I
38. Coxae I usually contiguous and covering portions of tritosternal base (Plate 41-3),
Fovae pedales (Plate 45-1) present or absent. (If coxae I are widely separated, not
covering tritosternal base, then fovae pedales or peritreme-bearing humeral
I
projections are present.) Superfamily UROPODOIDEA 43
Coxae I widely separated. Fovae pedales lacking (Plate 41-4) . . . . . . . . . . . . . . . .. 39
I
I
1The familial concept of the DERMANNYSSIDAE has been expanded by Evans and Till (19661, to
include the LAELAPIDAE (s. lat.l. MACRONYSSIDAE, RHINONYSSIDAE, ENTONYSSIDAE, HALA·
I
RACHNIDAE, IXODORHYNCHIDAE and HYSTRICHONYSSIDAE.

I
I
I
I 91

I 39. Tritosternal base twice as long as wide. Epistome short, broadly triangular with
large, irregular teeth. Dorsal marginal shields absent, vertical shield of female

I separated from median dorsal shield

.. . . . . .. .. . '" '"
Superfamily PROTODINYCHOIDEA
(Plate 42) Family PROTODINYCHIDAE
.

I Tritosternal base broader than long. Usually without peritreme-bearing humeral

projections (if humeral projections are present, then the vertex setae are absent).
· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Superfamily TRACHYTOI DEA ... 40

I 40. Palpi 4-segmented (tibia and tarsus fused); corniculi more than twice as long as
broad and re<lching to or beyond level of distal margins of pal pal femur. Legs I

I without claws. . . . . . . . . . . . . . . . . . . . . . . . . . . .. (Plate 42) Family POLYASPI DAE

Palpi 5-segmellted; corniculi less than twice as long as broad and/or not reaching to
level of distal margins of palpal femur. Legs I with or without claws 41
I 41. Epigynial shield extending anteriorly to margin of sternal shield. Idiosoma ovoid

I · (Plate 43) Family DISCOURELLIDAE

Epigynial shield not extending anteriorly beyond level of sternal setae II. Idiosoma
pyriform or ovoid (Plate 43 [see also 9-1,2]) Family TRACHYTI DAE
I 42. Tritosternal base broader than long Superfamily TRACHYTOIDEA .
· (Plate 42-3) Polyaspis (Dyscritaspis) of Family POLYASPIDAE
I Tritosternal base longer than broad
·
Superfamily PROTODINYCHOIDEA .....
" .. (Plate 42) Family PROTODINYCHIDAE

I 43. Jugular shields present. Epigynial shield a tongue-like flap. Dorsal marginal setae
not borne on platelets (Plate 44) Family THINOZERCONIDAE
I Jugular shields absent. Epigynial shield variable, not as above.
setae on platelets or elongate marginal shields
Dorsal marginal
44

I 44. Fovae pedales usually lacking. Peritreme-bearing humeral projections often

present and prominent, or absent (if fovae pedales present, humeral projections

I also present) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 45
Fovae pedales present. Peritreme-bearing humeral projections absent or very
small " 46
I 45. Humeral pro}9ctions absent. Epigynial shield not extending anteriorly to margin of

I sternal shield (Plate 44) Family DINYCHIDAE

Humeral projections present. Epigynial shield extending anteriorly to margin of
sternal shield. . . . . . . . . . (Plate 45 [see also 11-1,2,3] ) Family EUTRACHYTI DAE
I 46. Metapodal shiields fused with ventral shield or separated by a fine metapodal line ... 47

I Metapodal shields completely separated from ventral shield by a membranous

suture (Plates 45,46 [see also 15-1] )
· ..... Families COXEQUESOMI DAE-PLANODISCIDAE-CI RCOCYLLIBAN IDAE 1
.

I 1These three families, all associated with army ants (Dorylinae), have many similarities and are diffi-
cult to separate. Thev may constitute a single family.

I
I
 I
92

47. Epigynial shield displaced posteriorly to a position behind coxae IV. Sternal pores
I
III also behind coxae IV ,. (Plate 46) Family METAGYNELLIDAE
Epigynial shield located between coxae II-IV. Sternal pores III in region of coxae
III-IV (Plates 46, 47 [see also 10-2]) Family UROPODIDAE l
I
48. Latigynial shields elongate, overlapping reduced mesogynial shield. With two
subequal dorsal shields or, if a single dorsal shield, at least a faint line of fusion
I
present. Usually with a pelage of body setae. Tritosternal laciniae fused as one for
more than one-half the length, sometimes separated only at tips or not at all. I
Epistome with anterior projections or serrations, lacking keel. Pretarsal elements
rarely present. Commonly associated with arthropods ....Cohort CERCOMEGISTINA
....................................... Superfamily CERCOMEGISTOIDEA
I
" (Plate 48 [see also 15-12]) Family CERCOMEGISTIDAE
Genital shields variable. With one dorsal shield. Body setae moderate in number.
Tritosternum with two distinct laciniae fused less than one-half their length,
I
usually less than one-quarter. Epistome simple or slightly serrate, with distinct
ventral keel (Plate 15-6). Pretarsal elements never present . I
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Cohort ANTENNOPHOR INA .. .49

49. Sternogynial shield(s) present and distinct, clearly bearing sternal pores III (Plate
I
48-5) " , , , 50
No distinct sternogynial shield; sternal pores III on sternal or metasternal shield or
indiscernible 54
I
50. Ventral, anal and metapodal shields contiguous or fused; venter with depressions I
accommodating folded legs and gnathosoma. Tetartosternum (Plate 48-5) often
present (~ jugulars separate from sternal, but fused with each other) in female and
in male. Chelicerae with filamentous excrescences. Turtle-like .
I
.. . .. .. .. .. . . , Superfamily FEDRIZZIOIDEA 51
Ventral and metapodal shields separated by at least a narrow band of soft integu-
ment, anal shield separate from, or fused with, ventral shield. First sternal setae on
I
51.
sternal shield, jugular shields or in soft integument, but not on a tetartosternum ... 52

Latigynial, mesogynial and sternogynial shields well developed (Fig. 17d, p. 85);
I
vaginal sclerites present, but reduced. Male genital aperture oval, much wider
than long, located between coxae III. Associated with insects . I
. . .. .. . .. ..... . .. . . (Plate 48 [see also 15-8J) Family KLiNCKOWSTROEMIIDAE
Latigynial and mesogynial shields lacking or represented only by a narrow band
bordering genital aperture; sternogynial shield enlarged (Fig. 17g, p. 85); vaginal
I
sclerites lost or completely fused with sternogynial shield. Male genital aperture
round or only slightly wider than long, located between coxae II and III. Associated
with insects " (Plate 49) Family FEDRIZZIIDAE
I
l The families TRACHYUROPODIDAE, OPLITIDAE. UROACTINIIDAE and UROPODIDAE of
I
Johnston (1968) are herein considered to be a single family.
I
I
I
I 93

I 52. Latigynial and mesogynial shields coalesced. Chelicerae with tree-like excrescences.
Palps 5-segmented. Associated with passalid beetles and other insects .
I · '" Superfamily MEGISTHANOIDEA
Latigynial and mesogynial shields free and functional (Fig. 17e, p. 85), hinged or
53

I fused with ventral shield. Chelicerae with filamentous excrescences.

distal segmem. Associated with arthropods and reptiles

Palps
4-segmented, tibia and tarsus fused so that apotele appears to arise from middle of
.

I .... Superfamily ANTENNOPHOROIDEA ... (Plate 49) Family PARAMEGISTIDAE

53. Well developed jugular shields; sternal and fused latigynial shields independent of
I each other and of ventral shield; fusion of latigynial shields indicated by medial
indentation on anterior margin and often on posterior margin (Fig. 17h, p. 85);
sternogynial shield divided; vaginal sclerites well developed, with hammer-like
I heads (Plate 49) Family HOPLOMEG ISTIDAE
Without jugular shields; sternal setae I on fleshy projections flanking tritosternum;

I sternal and 'fused latigynial shields coalesced around genital aperture and lati-
gynials not indented at point of fusion; sternogynial shield large, well developed,
entire or divided (Fig. 17i, p. 85); vaginal sclerites reduced, without heads, but often

I with thickened arms (Plate 50) Family MEGISTHANIDAE

54. Sternal setae I always on sternal shield; vaginal sclerites well developed, usually with
I heads (Plate !i1-1). Chelicerae strong, with dendritic or brush-like excrescences and
with large proximal tooth on the movable digit. Male genital aperture under free
anterior edge of sternal shield. Associated with insects, especially passalid beetles.
I ·." Superfamily CELAENOPSOIDEA
Sternal setae I free in soft integument, on independent jugular shields, or on
55

I weakly sci erotized sternal shield; vaginal sclerites reduced, without heads and with
a bow-shaped base-piece. Chelicerae tapered, with filamentous excrescences, often
edentate or with numerous minute teeth. Male genital aperture within sternal
I shield , , , Superfamily ANTENNOPHOROIDEA 58

55. Latigynial shields well developed, free medially, hinged or fused with ventral shield
I posteriorly; mesogynial shield well-developed, reduced or lost '
Latigynial shields partially or entirely fused medially; mesogynial shield never
" 56

I well developed 57

56. Latigynial shields not usually extending posteriorly beyond hind margins of coxae
I III and never to middle of coxae IV, hinged to ventral shield; mesogynial shield
usually hinged, sometimes reduced and fused with ventral shield or absent; with or
without free metasternal shields (Fig. 17a, p. 85) .
I · (Plate 50 [see also 5-2, 15-9, 16-2J) Family DIPLOGYNIIDAE
Latigynial shields elongate, often extending posteriorly beyond coxae IV; fused, not

I hinged to ventral shield, mesogynial shield reduced and fused with ventral shield.
Associated with snakes and with beetles (Plate 50) Family SCH IZOGYN II DAE

I
I
I
94
I
57. Latigynial and mesogynial shields fused with ventral shield and with each other, I
except along anterior margins (Fig. 17b, p. 85) .
........................... (Plate 52 [see also 15-6]) Family EUZERCONIDAE
Latigynial shields entirely fused with ventral shield and with each other, except for
I
a shallow indentation anteriorly (Fig. 17c, p. 85); no remnant of mesogynial shield
...................................... (Plate 51) Family CELAENOPSIDAE I
58. Latigynial shields mostly membranous, unhinged.
teeth. Pedipalpal tibiae and tarsi distinct
Chelicerae usually with some
59 I
Latigynial shields well sclerotized, hinged or fused with ventral shield along
posterior margins. Chelicerae edentate. Pedipalpal tibiae and tarsi insensibly fused.
Associated with arthropods and reptiles (Plate 49) Family PARAMEG ISTI DAE
I
59. Latigynial shields sclerotized along anterior edges; mesogynial shield reduced,
unhinged, and almost entirely coalesced with the latigynials. Chelicerae with
I
minute teeth. Associated with insects ..... (Plate 51) Family ANTENNOPHOR IDAE
Latigynial shields! sometimes with a sci erotized seta-bearing patch; mesogynial shield
I
hypertrophied, hinged and often bearing setae (Fig. 17f, p. 85). Chelicerae with a
few distal teeth. Associated with insects and myriapods .
. .. .. . . . .. . . .. . .. . .. .. .. . . . .. .. .. .. (Plate 52) Family PARANTENNULIDAE
I
I
I
I
I
I
I
I
I
I
I
I
I
!I PLATE 15
95

I
I
I
15-1
I
I
I
I
I 15-5
I 15-6

I
I
I
I
I
I
I JL
I Hi-9 15-10 15-12

I 15-' to 15·6; epistomes of selected mesostigmatid mites. 15-'; family PLANODISCI DAE, Planodiscus burchelli Elzinga and
Rettenmeyer (Panama): 15-2; family EVIPH I 01 DAE, Eviphis stefaninianus (Berlesel (Congo): 15-3; family
AHCTACARI DAE, Arctacarus sp. (Oregon, USA): 15-4; family LAELAPI DAE, Haemogamasus pan tiger (Berlesel (Oregon,
USA): 15-5; family MACROCHELIDAE, Macrocheles montanus Willmann (Germany): 15-6; family EUZERCONIDAE,

I Euzercon sp. (Mexico)

15-7 to 15·9; genital shields of selected mesostigmatid mites. 15-7; family PODGe IN IOAE, Podocinum sagax Berlese (Italy):
15-8; family KLiNCKOWSTROEMIIDAE, 'genus (Brazil): 15-g;family DIPLOGYNIIDAE, 'genus (Ohio, USA)
15-10 to 15-12; tritosterna of selected mesostigmatid mites. 15·10; family SPI NTU RN ICIDAE, Periglischrus sp. (Mexico): 15-11;

I family UROPODIDAE, Neodiscopoma sp. (Oregon, USA): 15-12; family CERCOMEGISTIDAE, Cercomegistus sp.
(Oregon, USA)
 PLATE 16 I
96
I
I
I
I
•
I
I
16-3
I
I
I
16-5 I
I
I
~~J~ I
i'
Ii/if iI'
I
~~ ~ ~~~
16-4

16-6
~ "
I(}V
"
I';;
/
16-7
I
I
I
16·1;
UroPod~1I16'5;familySh~~~n~:
~~~);I:ternitigenital
family ARCTACARIDAE
Lobogyniella tragardhi
SEJIDAE S'. 16·4; family VEIGAIIDA~le;, 16·3; family ZE'RCONiDDAIPLOGYNIlDAE,
~~~:gon, USAI,gCh~li~;r~l~fc~:~cera
,Arctacarus sp. {Ore on .
I
16'~s sternitig~"
of female: 16·2 famil
US asp. (Ohio USA) .' ejussp. (Oregon US ,velgalasp (W h' E,Zercon (0
A), epigynial shi.eld ,eplgynial shield: 16.]', f amlly MICROGYNIIDA
AI, epigynial shield: . ,amily
flnQt,on,UROPODELLI
USAj, nita I reg.on,
shield
E, Mlcrogyn;um"mCfsum Krantz
DAE, (Oregon,
I
I PLATE 17
I
I X
17-2

I
I
I
I
I 17-5
!
I
I
I
I
I
I
I
•
I
I
I
I
I
 I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
18.1 to 18-5; family SEJIDAE. 18-1; Sejus sp. (Oregon, USA), dorsum of female: 18-1; ?genus (Africa) dorsum of female:
I
18-3; Sejus sp. (Oregon, USA), dorsum of male: 18-4; types of dorsal shield setae: 18-5; Sejus sp. (Oregon, USA).
venter of female
I
I PLATE 19
99
I
I cJJ1(~1..",..;>4~

_/TI~·
I
I ;' J \ 't,
I
.....
~

I o.r---...o
0--

I o ~

I ~f3t~ legs I elongate

narrow '

I
I ::;) ad setae terminally

I
I
I
I
I l",.:H_+-genital
aperture

I
I extends to or beyond
coxae IV 19-5
I
I 19-1 to 19-4:
19-6- famIly
- EPICRIIDA
,tuberculate -
dorsal ornamentation:
E, Eplcriu5 19- 5,: vent~r
sp. 19-1- sternltigenital ra·'
of female" 19·2;
glon tarsus. I19.6:
of male" of fadorsum of f~~~:s
rna Ie: 19-3" t of palpal apoteles:

I
 PLATE 20 I
100
I
I
I
20-2
I
=-~--",.--_sternal pore I
sternal shield I
~
l"l,,=4\\-l!--sternal pore II

~~E:jt'~=~rmetasterna,
sternal pore" I
shield I
li.._M----epigynial shield triangular I

~
/~~," -+ ventrianal shield I
Y ----" '\ r~ I
~/<> ~\ r~
~~)~ I
20-1 \ ~/
I
I
I
I
I
I
I
I
20-1 to 20-4; family PARASITIDAE_ 20-1; venter of female (Oregon, USA): 20-2; palpal apotele: 20-3; leg II of male (Oregon,
USA): 20-4; Poecilochirus necrophori Vitzthum (Oregon, USA), venter of deutonymph
I
I
I PLATE 21
101

I
I
I
I
I
I
~I.o?f---I,-I-- epigynial shield
I
--,f-.--H-- ventral shield
I (fused to epigynial, or free)

21-2
I 21-1

I I
I
I
I
I
I
I
I
I 21-4 21-5

I 21-1 to 21-4; family VEIGAll DAE, Veiga;a sp. 21-1; venter of female (Oregon, USA): 21-2; subcapitulum of female: 21-3;
epistome of female: 21-4,5; dorsal shield types

I
102
PLATE 22 I
I
22-3 ••
I
peritreme
usually joins
stigma
posteriorly r
I
I
I
22-2
I
22-4
I
I
I
22-6
I
~ I
r' H

' / \
I
I
~~
22-8

~
/
22-7
: : :brushlike
process
r±J.................. .,
22-9
I
I
22-1 to 22-5; family MACROCHELIDAE. 22-1; Lordocheles ryke; Krantz (Congo), venter of female: 22-2; Haloee/aeno me/is;
Krantz (Brazill. venter of female: 22-3; palpal tarsus: 22-4; Holostaspella bifoliata (Tragardh) (Oregon, USA). chelicera
of female: 22·5; Macrocheles sp. (Trinidad), tarsus III
22·6 to 22·10; family PARHOLASPIDAE. 22-6; pal pal tarsus: 22-7; Parholaspulus lobatus Krantz 10regon, USA), chelicera of
I
female: 22-8; Neoparholaspulus coalescens Krantz (Louisiana, USA), epistome: 22-9; Calha/aspis berlesei Krantz
(Maryland, USA) epistome: 22-10; Parholaspu!us parvilobatus Krantz (Oregon, USA), venter of female
I
I PLATE 23
103
I
I
I
I
I
sternal spatulate setae
I shield
elongate
lateral shields

I fused

I
I 23-1
I 23-3

I
I dorsal, ventral shields
weakly sclerotized

I
I chelicerae edentate

I
I
I 23-5

I
I 23-1; family PACHYLAELAPI DAE, Pachylae/aps sp. 10regon, USA); venter of female
23-2 and 3; family OMENTOLAELAPI DAE. Omentolae/aps mehe/yae Fain (Congol. 23-2; venter of female lafter Fain 1961):
23-3; dorsum of female (after Fain 1961)
23-4 to 23-7; family LAELAPONYSSI DAE, Laelaponyssus mitis Womersley (Australia). 23-4; palpal tarsus: 23-5; chelicera of

I female: 23-6; dorsum of female: 23-7; venter of female

 PLATE 24 I
104
I
\~
/{f-\~
r
'\ \ 1 \
T
24-'
- w i t h one or two whiplike
setae on tarsus ,
•
I
1 \ 1,,/ I
I
I
I
I
I
I
I
I
I
I
I
24-4
•
I
I
24-1 to 24-3;· ,family PODOCINI DAE, Podocinum sp. (Kansas, USA). 24-1; dorsum of female: 24-2; venter of female:
24-3; ornamentation of dorsal shield
I
24-4; family HETEROZERCONIDAE, Heterozercon 'p_ (Florida, USA); venter of female
I
I PLATE 25
105
I
I
I
I
I
I
I
I
I
I
(), 25-6

I
I
I
I 7--+-+Hl---tubulus

I annu[atus

I
I 25-5
I
I 25-1 to 25-5; family RHODACARIDAE. 25·1; Rhodacarus sp. (Oregon, USA), venter of female: 25-2; Rhodacarus sp., dorsum
of female: 25-3; Gamasellus sp. (Norway), dorsum of female: 25-4; Gamasellus sp., venter of female: 25-5; Gamasiphis sp.

I (India), venter of female: 25-6; palpal tarsus of rhodacarid

 PLATE 26
I
106
I
26-1 I
26-2
I
I
•
I
~~~.4I---I--metasternal setae
free
--+"':>'::=+4+-..genital seta
I
+::\:-::p.~c:::d::!genjtaJ pore
epignial shield
without setae
I
I
I
I
I
I
I
I
coxa II

I
mesonotal
scutella
I
I
26-1 to 26-6; family ARCTACARI DAE, Arctacarus sp. (Oregon, USA). 26-1; venter of female: 26-2; palpal tarsus: 26·3;
I
chelicera of female: 26-4; pilus dentilis of femare chelicera: 26-5; sternitigenital region of male: 26-6; dorsum of female

I
I PLATE 27
107
I
I
I
I
~
I 27-2
27-3
I
I
I
I
I
I
•
I \ •epigynial may be

I 1 )/ \ 'ftruncate and

:f~:~".'""
•
'.
I ! \

27-5
I
I
•
27-1 to 27-5; family AMEROSEII DAE. 27-1; Ameroseius sp. (Oregon, USA), venter of female: 27-2; typical dorsal seta of
Ameroseius: 27-3; Ameroseius sp., subcapitulum: 27-4; Ameroseius sp., dorsum of female: 27·5; Kleemannia sp.
(Oregon, USA); epigynial and ventrianal shields of female
 PLATE 28 I
108
I
epistome often
trifid 28-3
I
I
I
I
I
I
I
I
I
I
28-4 I
)) \~ I
pilus dentilis

L ~
co:::::. -==:::r c::=~
I
I
~
,/r/\.:/'-:.I\-'\ I
28-7
I
28-5
Q2S-6
"-..:1/ I
28.1 to 28-4; family DIGAMASE LLI DAE, Digamasellus sp. (Oregon, USA). 28-1; dorsum of female: 28-2; venter of female:
28-3; epistome of female: 28-4; chelicera of female
28-5 to 28-7; family PHYTOSEIIDAE. 28-5; Amblyseius sp. (Oregon, USA), chelicera: 28-6; Iphiseius degenerans (Berlese)
I
(Italy), epigynial and ventrianal region of female (after Chant 1959): 28-7; chaetotactic diagram of genu and tibia I of a
phytoseiid mite (after Evans 1963)
I
I PLATE 29
109

I 29-2/\
I
I LF
I
I
I
I
I
I
I
I
I
I
I
d-
coxa III

I
I
It,..
..............

I 29-4

I
I 29-1 to 29-3; family PHYTOSEII DAE. 29-1; Amblyseius ,p_ (Oregon, USA), venter of female: 29-2; palpal tarsus of generalized
phytoseiid mite: 29-3; Typhlodromus tiliae Oudemans (Oregon, USA), dorsum of female
29-4 to 29-7; family OTOPHEIDOMENIDAE. 29-4; Hemipteroseius indicus (Krantz and Khot) (Indial. spermatheca: 29-5; dorsal
shield of Nabiseius: 29-6; dorsal shield of Otopheidomenis: 29-7; dorsal shield of Hemipteroseius

I
110
PLATE 30 I
loo---- with or without trito'temum ---:-=.....-::--'"
often with only one pair of sternal setae ~
I
on shield, or none

30-2 ! '\ I
---. I
~~

I
I
I
J! \ \ I
I
""n/
1-1 ~0-3 I
30-5
_ -,.7'
!&'-"~
"""r
<1":
I
...-"
~~
,/
<-....
'\''C::
I
~--:J •

I I
•

'\
~ I
/' I' I
r /' '" '\ I
/ I
I
I
I
dorsal pores

I
30-1 to 30-3; family OTOPHEI DOMENIDAE. 30-1; Hemipteroseius indicus, venter of female: 30-2; H. indicus, chelicera:
30-3; Otopheidomenis sp., venter of female
30-4 to 30-6; family ZERCONIDAE. 30·4;Zercon sp. (Oregon, USA), venter of female: 3D-5;Zercon sp., venter of male:
I
30-6;Zercon sp. (Norway), dorsum of female
I
I
111
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I 31-1 to 31-6; family ASCIOAE. 31-1; B/attisocius tarsalis (Berlese) (Oregon, USA). dorsum of female: 31-2; Asca sp. (Oregon,
USA), dorsum of female: 31-3; chaeta tactic diagram of genu and tibia I of certain ascid mites (after Evans 1963):
31-4; Lasioseius garambae Krantz (Congo). venter of female: 31·5; palpal tarsus of ascid mite: 31-6; Blattisocius tarsalis,

I venter of female
 PLATE 32 I
112
I
chelicerae adapted for
grasping
I
I
I
I
I
I
without tarsal apoteles
I
I
palp-----\\.
camerostome----~~~~~~
I
hypostome --,.....~--l-\:;:'1r;:
I
sternal shield reduced--~/!-,~=I___,I,_..¥.Jr_
I
peritreme reduced ---11M
I
I
I
32-3
I
genital aperture posterior,
I
without shields ---~-7'----,..'~::'f!~ •.

anal orening terminal

I
32-1 to 32-3; family SPELAEORHYNCHIDAE, genus Spelaeorhynchus (South America). 32·1; dorsum of temare: 32-2, chelicera
of female: 32-3; venter of female
I
I
I PLATE 33
113

I apotele, when present, 2-tined

I 33-1
33-2
I
I stigmata lateroventral
or \lentral

I
-.+-Y>r'b'r...._st.rnal shi.ld without .ndopodal

I extensions

I
I
I
I dorsal shi.ld
entire

I
I 33-3
I opisthosoma elongate,
filled with tracheae

I
I
I
I 33-5

I 33-4
~m:;('[7t!~.:...-. .plgynial shi.ld
',:.\ absent

I 33·1 to 33·6, family HALARACHNiDAE. 33·1, Orthohalarachne attenuata {Banks} (North Pacific), v.nter of f.mal.: 33·2; O.
attenuate, anterodorsal aspect of female: 33-3; Halarachne americana Banks, sternal and genital region of female (after
N.w.ll 1947); 33·4; Pneumonyssoides sp.• palpal tarsus: 33·5, P. caninvm Chandl.r & Ruh., sternal and genital region

I of f.male (after Bak.r et al. (958): 33·6; P. caninurn, dorsum

 PLATE 34 I
114

dorsal shield
I
entire 34-2
34-1
corniculi distinct
I
..,...-I-""::;F-tritosternal laciniae
weak or absent
fixed cheliceral digit
sometimes not developed I
I
I
I
I
I
34-3 I
I
I
I
I
I
I
I
I
I
34-1 to 34-3; family ENTONYSSIDAE, genus Ophiopneumicola (Washington. USA). 34-1; venter of female: 34·2; dorsum of
female: 34-3; chelicera of female
I
34-4 and 34-5; family RHINONYSSIDAE. 34-4; Neonvssuscolumbae Crossley (Texas, USA), venter of female: 34-5; Cas
angrensis (Castro) (Texas, USA), venter of female
I
I PLATE 35
115

I
I
I
I
I
I
I
I
tritosternum
present "'tl~ )
35-6

I sternal shield r···;""'·~

weakly
sClerotized~
i /'
~.,

'\.
I
I
I
I
I dorsal shield
divIded or entire

I
I
I .j> If
/ no sternal shield \

I
I 35·1 to 35-5; family RH INONYSSIDAE. 35-1; dorsum of Ptilonyssoides: 35-2; dorsum of Cas: 35-3; dorsum of Lar;nyssus:
35-4; dorsum of Neonyssus: 35·5; dorsum of Rhinoecius
35-6 to 35-8; family DASYPONYSSI DAE. 35-6; Manitherionyssus heterotarsus Vitzthuml (Africa), anteroventral aspect of female

I (after Strandtmann and Wharton 1958): 35-7; Dasyponyssus (after Baker et at. 1958): 35-8; Dasyponyssus, dorsal sh ield
(diagrammatic)
 PLATE 36 I
116

I
I
I
I
I
I
I
I
I
I
I
36-4
I
I

a 36-7
I
I
I
I
I
36-1 to 36-7; family SPINTURNICIDAE. 36-1; Periglischrus sp. (Mexico), venter of female: Spinturnix sp. (Indiana, USA), venter
of male: 36·3;Spinturnix sp.t dorsum of male: 36-4 to 36-7; tritosternal bases of various spinturnicid species (after Rudnick
1960)
I
36-8; family MACRONYSSI DAE, Ornithonyssus sylviarum (C. & F.J (Oregon, USA), venter of female
I
I PLATE 37
117
I
I
I
I
I
I
I
I 37-4

•
"chelicerae attenuate,
stylettiform

I
I dorsal shield entire or
fragmented posteriorly

:::7-5
I
I
I
I
I
I
37-7
I
I 37-' to 37-3; family MACRONYSSIOAE. 37·' ;Steatonyssus sp. (California, USA), venter of female: 37-2; Steatonyssus sp.,
dorsum of female: 37-3; Ornithonyssus sylviarum, dorsum of female
37-4 to 37~6; family DERMANYSSIDAE, Dermanyssus triscutatus Krantz (Alaska, USA). 37-4; chelicera of female: 37-5; dorsum
I of female: 37·6; venter of female: 37-7; D. gallinae (DeGeerl (Oregon, USA), dorsum of female
 PLATE 38 I
118

38-1
38-2 I
I
I
38-3 I
coxal spur"--I+-=+\---./€J
I
I
I
I

-t--+- anal shield

•
I
I
I
I
I
I
peritremal shield
extends beyond
I
coxa IV

I
I
38.1; family HYSTRICHONYSSIOAE, Hystrichonyssus turneri Keegan et al., dorsum
38.2 to 38-4; family IXODORHYNCHIDAE, Ixodorhynchus !iponyssoides Ewing. 38·2; venter of female: 38-3; chelicera of
I
female: 38-4; dorsum of female
38-5; family EVIPHI DI DAE, Eviphis stefaninianus (Berlese) (Congo). venter of female (after Shoemake and Krantz 1966)
I
I PLATE 39
119

I
I 39-2 ~394
I
I
I
I
I
I
I 39-1
I
I
I
I
I
I 39-6

I
I
I 39-7

I 39-1 to 39-5; family EVIPHIDIDAE. 39·1; chaetotactic diagram of genu and tibia I of an eviphidid mite (after Evans 1963): 39-2;
Eviphis stefaninianus, epistorne (after Shoemake and Krantz 1966): 39·3; Scarabaspis rvke; Shoemake and Krantz (Congo),
epistorne (after Shoemake and Krantz lHooJ: ~1:l-4; palpal tarsus ot eviphidid mite: 39-5; Eviphis sp. (Africa), dorsum and
venter of female
I 39-6 to 39-8; family LAELAPIDAE. 39-6; chaetotactic diagram of genu and tibia I of a laelapid mite (after Evans 1963): 39-7;
Haemogamasus sp. (Oregon, USA), epistorne: 39-8; Androlaelaps sp. (Oregon, USA), epistorne and related structures
 PLATE 40 I
120

40-2
I
pilus dent ins may be
expanded
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
40-1 to 40-5; family LAELAPIDAE. 40-1; Androlaelaps fahrenholzi (Berlese) (Oregon. USA), chelicera of femaie:
40-2; Hirstionyssus sp. (Oregon, USA), chelicera of female: 40-3; Hirstionyssus sp., venter of female: 40-4; Laelaps sp.
I
(Brazil), venter of female: 40-5; Hypoaspis sp. (Oregon, USA), venter of female
I
I PLATE 41
121
I 41-2
I 41-1

I
I
I +.-'l;.'-'!5:!:=i'-'-......_ epigynial
shield

I
I
I
I
I
I
I
I 41-4
~a
I
I -v-
coxae I widely
separated
_

-
I -+_+-__ fovae pedales
41-3
I
"'=--_---,{-- metapodal line

I
I 41·1 and 41·2; family DIARTH ROPHALLIDAE. 41·1; Diarthrophallus sp. (Mexico), venter of female: 41·2; D. quercus (Pease
and Wharton), dorsum of female
41-3; venter of Uropodoidea (diagrammatic) showing typical coxal and tritosternal characters: 41-4; venter of Trachytoidea

I (diagrammatic) showing typical coxal and tritosternal characters

 PLATE 42 I
122
I
~~~~~_., ~ I
I
dorsal shield
generalfusionofs~ields
around coxae and
epigynial shield
. - ( \ " '•"
~
//-
_
I
I
1:
«_",..lon~ epigynial shield W i t h - - - - V• '
setae (?metasternals) ~. •~ I
broader than

/ 1
42-2
I
I
perigenital rim
•
I
I
I
I
I
I
I
I
42-4 I
42-1 and 42-2; family PROTODINYCHI DAE. 42-1; Protodinychus punctatus Evans (England), vertical shield (after Evans 1957):
42-2; Protodinychus sp. (Canada), anteroventral aspect of female (after Johnston 1961)
42-3 to 42-5; family POL YASPI DAE. 42-3; Dyscritaspis sp. (Oregon, USA), sternitigenital region of female: 42-4; Dyscritasp;s sp.,
I
anterodorsal aspect of female: 42·5; Polyaspis sp. (Brazill, venter of female
I
I PLATE 43
123
I 43-2=---__...,--..... epigynial shield extends to

• r.,. anterior edge of sternal shield

43-1 'e
j ') ! \'\\7
\'\'1
marginal shields./' ~
entire I .... r' \' / all
43-3

)/()(~ J\~"\r(
I
I
• ~ I J\) ••
I )).L==.h((
I
I
I
I tectostracal
"wing"

I metasternal shield

I
I
I
I
I
I
I
I 43·1 and 43·2; family DISCOURELLIDAE, 43-1; Discourella sp. (North Americal. dorsum of female (after Johnston 1961):
43-2; Discourella sp., sternitigenital region (after Johnston 1961)
43-3 to 43-6; family TRACHYTIOAE. 43-3; Caminelfa peraphora Krantz and AinscQugh (Oregon, USA), chaetotactic diagram of

I
genu and tibia I: 43-4; Trachytes sp. (Oregon, USA), dorsal shields and tectostracal wing: 43-5; Polyaspinus sp. (Oregon,
USA), venter of female: 43-6; C. peraphora, dorsum of female
 PLATE 44 I
124
I
I
I
I
I
epigynial
shield
tongue-like I
I
44-1
I
I
I
I
I
I
I
I
I
I
I
44·1 and 44-2; family THINOZERCONI DAE. 44-1; Thinozercon michael; Halbert (Ireland), anteroventral aspect of female {after
Tragardh1941l: 44·2; T. michaeli, dorsum of female (after Halbert 19151
I
44.3 to 44·5; family DINYCHIDAE, Phyllodinychus sp. (Oregon, USA). 44·3; venter of female: 44-4; Chelicera of female:
44-5; dorsum of female
I
I PLATE 45
125
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
•
I 45-1; family EUTRACHYTI DAE, Eutrachytes maya Krantz (Mexico), venter of female
45-2 and 45-3; family CI RCOCYLLlBANIOAE, Circocylliba sp. (Panama). 45-2; dorsolateral ornamentation and setation:
45-3; venter of female

I 45·4; family COXEQUESOMIDAE, Coxequesoma sp. (Panama), venter of male

 PLATE 46 I
126
I
46-2
I
I
I
I
I
46-1 I
I
-+
I
metapodal-_ _
shield

I
46-3
I
I
I
I
1\1 .1
/
I 0 • \

.011 I i
'\
I
/ r rIo
V~\ r
1 ' ""'<
;" \y I
:\ I
~~ I
4&5
I
46-1;
46·2;
46·3;
family COXEQUESOMIDAE, Coxequesoma sp. (Panama), chelicera of male
family PLANODISCIDAE, Planodiscus burchelli Elzinga and Rettenmeyer (Panama), venter of female
family METAGYNELLIDAE, Metagynella sp. (South Africa). venter of female
I
46-4 and 46-5; family UROPODIDAE. 46·4; Leonardie/la sp. (Congo), dorsum of female: 46-5; ?Uropoda sp. (Congo), venter
of female I
I PLATE 47
127

I
47-1
I
dorsal and marginal
shields fused

I with or without

I scalloping

I
I
I
I J~ marginal shield free,
entire, abbreviated posteriorly

I
I scI erotized dorsomedial furrow

I
marginal shield

I
I
I
I 47-3

I
I •
(Q)
I 47-5

I 47-' to 47-5; family UROPODIDAE. 47-1; Urodiaspis sp. (Oregon, USA), dorsum: 47-2; Fuscuropoda agitans (Banks) (Maryland,
USA), dorsum: 47·3; Phaulodiaspis sp. (Congo), dorsum: 47-4; ?genus (Congo), dorsum: 47·5; chaetotactic diagram of
genu and tibia I of a uropodid mite

I
 PLATE 48 I
128

I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
48-3
I
48-1 to 48-4; family CE RCOMEGISTIDAE, Cercomegjstus sp. (Oregon, USA). 48-1; venter of female: 48-2; chelicera of female:
48-3; dorsum of female: 48-4; epistome
48-5 and 48-6; family KLlNCKOWSTROEMIIDAE. Klinckowstroemia sp. {BrazilL venter of female: 48-6; Klinckowstroemia sp.,
dorsum of female with detail of dorsolateral ornamentation
•
I
I PLATE 49
129

I H--1~--1~--laciniae separate throughout length

I
_III--_:"'-_tetartosternum

I !!I)";/-.IJ.-----\-- sternal shield

I ...-_...,--\-_ stern09ynial shield

~ _ _---I,-- remnant of latigynial shield

I ~---I--remnant of mesogynial shield

I
I
I
I
I
I
I
I
I latigynial
shield
mesogynial
shield
I
I
I
I 49-4

I 49-1;
49-2;
49-3;
family KLiNCKOWSTROEMIIOAE, Klinckowstroemia sp. (Brazill, chelicera of female
family FEDRIZZIIOAE, Fedrizzia sp. (New Guinea), venter of female
family PARAMEGISTIDAE, Echinomegistus wheeleri (Wasmann) (Kansas, USA), venter of female

I 49·4; family HOPLOMEGISTIDAE (Brazil), venter of female

 PLATE 50 I
130

50-1
I
I
presternal setae on
fleshy lobes - ----=",.,""
I
I

~
sternogynial entire
or divided

fused latigynials.
I
50-3

I
I
dendritic excrescence
I
I
I
50-5
I
I
I
I
I
I
\ I
.! \
r
o

~I
• .' • 0

I
mesogynial
remnant
I
50-1 and 50-2; farnily MEGISTHANIDAE, Megisthanus floridanus Banks (Florida, USA). 50-1; venter of fernale: 50-2; chelicera
of female
I
50-3 and 50-4; farnily DIPLOGYN II DAE (Ohio, USA). 50-3; chelicera of fernale: 50-4; venter of fernale
50-5; farnily SCHIZOGYNIIDAE, Schizogvnium sp. (Panarnal. venter of fernale
I
I PLATE 51
131

I
I
I
I
I
I
I
I
I sternal setae I free in integument,
on fragment of sternal shield or
I on tetartosternum

I
I
I sternal shield ---------,""-~:"""""--cIH1+

peritreme - - - - - - - \ - ' l l ! \ \
I
latigynial shield -----+-he=:--'r--zP~:"-::;f-'<
I
I
ventrianal shield """'~----+

I
51-3
I
I 51-1; family CELAENOPSIOAE (Colorado, USA), venter of female
51-2 and 51-3; family ANTENNOPHOR IDAE, Antennophorus sp. (British Columbia). 51-2; chelicera of female: 51-3; venter
of female

I
 PLATE 52 I
132

I
I
metasternal
shield
I
I
I
I
I
I
52-1
I
sternal setae in platelets
I
unsclerotized sternal entity I
I
~--'<-;rtt""-Iatigynial shield
I
I
I
mesogynial shield often
with setae
I
anal shield ---~"'-':""""'p-'f-t::
I
52-3
I
52·1; family EUZERCONIDAE, Euzercon sp. (Arkansas, USA). venter of female
52.2 and 52·3; family PARANTENNULIDAE. 52-2; Micromegistus bakeri Triigardh (Ohio, USA), venter of male: 52·3; M. bakeri
I
(Kansas, USA), venter of female -

I
I 133

I NOTES

I
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I
135
I ORDER PARASITIFORMES

I Suborder Metastigmata

I The Metastigmata, or ticks, have often been considered to be distinct from other mites
on the basis of their great size (adults may range from 2,000 to over 30,000 J1 in length).

I Yet there is an obvious affinity with other mite groups, particularly with the Parasitiformes.
Ticks are ectoparasitic in all postembryonic stages, feeding primarily on the blood of
mammals, reptiles and birds. The hypostome of the tick is modified into a holdfast organ

I armed with retrorse teeth (Fig. 18), with which the tick may anchor itself to its host.
Other important features include:

I
I
I
I
I
I
I
I
I Fig. 18. Gnathosoma of an ixodid tick (Congo) showing the median
hypostome armed with retrorse teeth and the anteromedian

I chelicerae.

1. Lack of an apotele on the pal pal tarsus, the tarsus itself often being reduced
I (Fig. 18).

2. A peritreme in the form of a stigmal plate surrounding each of the stigmata which
I are located laterad of, or posterior to, coxae IV (Fig. Ga, p. 20).

3. Chelicerae with a series of external harpoon-like excrescenses, an internal

I stylettiform protuberance, and a hook-like dorsal process (Fig. 18).

I
I
136
I
4. A sensory "capsule" and adjacent pit on the dorsum of tarsus I comprising the I
Haller's organ (Plate 53-3). Portions of this organ may serve as olfactory centers
and as hygroreceptors.
I
5. Lack of an anteriorly produced epistome.

6. Lack of a tritosternum.
I
The Metastigmata are cosmopolitan in distribution and play an important role in the
transmission of many diseases of man and animals. Three families are recognized, including
I
one represented by a single intermediate species (the NUTTALLlELLlDAE).
I
Superfamily Ixodoidea
I
DIAGNOSIS: Weakly sclerotized but with thick leathery cuticle, with or without a
dorsal shield; gnathosoma terminal or ventral, hypostome armed with
retrorse teeth; palpi simple, telescoped or normal. With a sensory pit, or
I
Haller's organ, on dorsum of tarsus I; all tarsi with apoteles.

The Ixodoidea, or ticks, are hematophagous ectoparasites of vertebrate animals

I
throughout most of the world (Aponomma ecinctum (Neumann) (IXODIDAE) occurs on a
beetle, but such associations are quite rare). The superfamily contains three families, of
which one, the NUTTALLlELLIDAE, is monotypic.
I
The IXODIDAE, or hard ticks, comprises approximately 700 species in 9-12 genera. I
Ixodids generally attach to different host species in their different active life stages (larva,
nymph and adult) and feed only once on each host. Some, however, remain attached to a
single host (Boophilus annulatus (Say), or detach from the original host only before the I
adult molt (Rhipicephalus evertsi Neum.). Attachment to the host in the IXODIDAE, as
well as in the ARGASIDAE, is accomplished through piercing the skin of the host with the
chelicerae, and anchoring by inserting the barbed hypostome into the wound (Arthur 1957).
I
Subcutaneous infestations of Ixodes or Hyalomma (IXODIDAE) have been observed in
some hosts. Attraction and subsequent attachment of an ixodid to its host is brought about
through receipt of chemosensory and temperature stimuli from the host. Haller's organ,
I
located on the dorsum of tarsus I in all ticks, is of prime importance as a receiver for these
stimuli (Plate 53-3). Chance of exposure to possible contact with a host is enhanced by the
tick's habit of climbing to the tops of grass blades or twigs on animal runways (Lees 1946).
I
Desiccation may occur after some hours in this exposed situation but many genera are
highly resistant to rapid water loss (i.e., Dermacentor and Hyalomma) (Balashov 1960). I
Ticks which are unsuccessful in finding a host regain moisture by moving down to ground
level for short periods of time. Males of several species of the genus Ixodes apparently are
non-feeders, a phenomenon similar to that in certain ARGASI DAE.
I
Members of the family ARGASIDAE are particularly evident as nest inhabitants and
parasites of birds and small mammals in semitropical and tropical situations (Cooley and
I
Kohls 1944). Unlike the ixodids, the argasids feed intermittently, and often on the same
host. Argasids generally do not move out of the nesting areas of their hosts, and so I
I
I
I 137

I rarely have an opportunity to attack more than one animal species or, in some cases, more
than one individual. Argasids are extremely resistant to water loss and commonly are found

I in dry arid situations (species of the genera Ornithodoros and Argas for example). Some
feed only in larval and nymphal stadia (Otobius megnini (Duges)), while others feed in all
but the larval stage (Ornithodoros savignyi (Audouin)). There are approximately 100
I recognized argasid species grouped into four genera.

Ixodid and argasid ticks cause injury to their hosts through exsanguination, secondary
I infection at sites of attachment, and through the transmission of disease organisms
(Philip 1963). Because of their sedentary habits, argasid ticks are considerably less impor-

I tant than the more unrestricted ixodids in disease transmission.

The family NUTTALLlELLIDAE is represented by a single species found under a

I stone in Africa. Its host relationships are unknown (Bedford 1931).

I Useful References

Arthur, D.R. (1957), The capitulum and feeding mechanism of Dermacentor parumapertus. Parasito!.

I 47:169-184. [IXODIDAE]
Arthur. D.R. (19601. Ticks. A Monograph of the Ixodoidea. Part 5. On the genera Dermacentor.
Anocentor. Cosmiomma. Boophilus and Margaropus. Cambridge Univ. Press: 251 pp. [IXODIDAE]

I Balashov, 1.5. (1960). Water balance and the behavior of Hyalomma asiaticum in desert areas.
Paraz. i Paraz. Bolezni 29(31:313-320. [IXODIDAE]
Med.

Bedford, G.A.H. (1931). Nuttalliella namaqua, a new genus and species of tick. Parasitol.23:230-232.

I Clifford, C.M., G.M. Kohls and D.E. Sonenshine (19641. The systematics of the sUbfamily Ornitho-
dorinae (Acarina: Argasidael. I. The genera and sUbgenera. Ann. Ent. Soc. America 57(41:429-437.
Cooley, R.A. (1946). The genera Boophilus. Rhipicephalus and Haemaphysalis (Ixodidae) of the New
I World. N.1. H. Bul. 187:54 pp.
Cooley. R.A. and G.M. Kohls. (19441. The Argasidae of North America, Central America, and Cuba.
Amer. MidI. Nat. Monograph 1:152 pp.

I Cooley, R.A. and G.M. Kohls (1945). The genus Ixodes in North America. N. I. H. Bul. 184:246 pp.
Filippova, N.A. (1961). Larvae and nymphs of the sUbfamily Ornithodorinae (Ixodoidea. Argasidael in
the fauna of the Soviet Union. Parazit. Sborn. Zool. Inst. Akad. Nauk. SSSR 20:148-184.
I George, J.E. (1963). Responses of Haemaphysalis leporispalustris to light.
Cornell Univ. Press, Ithaca 1:425-430.
Advances in Acarology,

I Hoogstraal, H. (1956). African Ixodidae. Ticks of the Sudan. Bur. Med. Surg., U.S. Navy 1:1101 pp.
Hoogstraal, H. (1966L Ticks in relation to human diseases caused by viruses. Ann. Rev. Ent.11:261-308.
Hoogstraal, H. (1967). Ticks in relation to human diseases caused by Rickettsia species. Ann. Rev. Ent.

I 12:377-420.
Lees, A.D. (1946).
Lees, A.D. (19481.
The water balance in Ixodes ricinus (L.) and certain other ticks. Parasito!. 37:1-20.
The sensory physiology of the sheep tick. Ixodes ricinus L. Jour. Exp. BioI. 25:145-

I 207.
Nuttall, G.H.F., C. Warburton, W.F. Cooper and L.E. Robinson (1908-1926). Ticks. A Monograph of the
Ixodoidea. Parts 1A. The Argasidae. Cambridge Univ. Press.

I Philip. C.B. (1963l. Ticks as purveyors of animal ailments: A review of pertinent data and of recent
contributions. Advances in Acarology, Cornell Univ. Press. Ithaca 1:285-325.
Pospelova - Shtrom, M.V. (1969). On the system of classification of ticks of the family Argasidae Can.,

I 1890. Acarologia 11 (1): 1-22.

I
I
138
I
Ricketts, H.T. (1907). A summary of investigations of the nature and means of transmission of Rocky
Mountain spotted fever. Trans. Chicago Path. Soc. 7:73-82.
I
Sonenshine, D.E., C.M. Clifford and G.M. Kohls (1962). The identification of larvae of the genus Argas
(Acarina: Argasidae). Acarologia 4(2): 193-214. I
I
I
I
I
I
I
Suborder Metastigmata
I
(Plates 53 and 54)
I
KEYTOTHE FAMILIES
I
1. Dorsal shield (scutum) present in all stages; gnathosoma anterior and visible from
above 2 I
Without dorsal shield; gnathosoma inferior, not visible from above , ..
............... , , , (Plate 54) Family ARGASIDAE
I
2. Sclerotization of dorsal shield weak; palpal tarsus normal, terminal .
............... , , , Family NUTTALLlELLIDAE
Dorsal shield strongly sclerotizedin all stages; palpal tarsus reduced, often inserted
I
on ventral apex of tibia .. (Plates 53, 54 [see also Fig. 18, p. 135J) Family IXODIDAE
I
I
I
I
I
I
I PLATE 53
139

I
I
I
I
I
I
I
I adanal shield ----I+_

I 53-'
I
I
I 53-2

I
I
I
I
I
I
I 53·1 to 53·3; family IXODIDAE. 53-1; Ixodes sp. (Oregon, USA), venter of male: 53·2; Ixodes sp" dorsum of male: 53-3; Ixodes
t;Jffin;s Neumann (Mexico), Halter's organ

I
 PLATE 54 I
140
I
I
I
I
I
I
I
I
I
I
IlJ6\li;::s;=~===-p!:,a'P
U hypostome I
1trti8=--\\--+--genital area
I
m»---+-_S~r;;:1
I
I
I
I
I
54-3 I
54-1 and 54·2; family IXODI DAE, Amblyomma cajennense (Fabr.l (Mexico). 54-1; dorsum of larva: 54-2; dorsum offemale
54-3 and 54·4; family ARGASIDAE. 54-3; Argas sp.. venter of female: 54-4; dorsum of larva (diagrammatic) illustrating rhedian
I
position of dorsal scutum

I
I 141

I ORDER ACARI FORMES

I Suborder Prostigmata

I The tremendous diversity of morphological characteristics in the Prostigmata suggests

that the suborder is in reality a composite of several subordinal entities. The Endeostigmata,
for example, might well be considered an independent suborder, as might the extraordinary
I water mites of the Supercohort Hydrachnellae. The great number of exceptions and
alternate features listed in existing descriptions of the Prostigmata, or in key separations of

I the Prostigmata from other suborders, amply illustrate the "waste basket" aspect of the
group.

I The lack of distinguishing features which apply to the Prostigmata as a whole often
necessitates identification of the suborder through elimination; i.e., identification based on
lack of a particular structure or combination of structures possessed by other acarine

I sUborders.

Terrestrial, aquatic and marine mites existing as predators, phytophages and parasites
I are included in the Prostigmata. Certain parasitic species may not exceed 100 J1 in length,
while others may attain lengths of 10,000 J1 or more. Many prostigmatid species have a
pair of stigmata at the bases of the chelicerae (Fig. 7b, p. 20) or, as in the Supercohort
I Parasitengona, between the chelicera I bases (Fig. 20). Others may have propodosomal
stigmata (Fig. 6bl. while a substantial number of forms have no discrete stigmatal openings

I of any kind (the Halacaroidea and Eriophyoidea, for example). Similar diversity exists in
the structure of the chelicerae (stylettiform (Plate 4-7). chelate (Plate 61-1) or reduced
(Fig. 19)) and the palps (simple (Fig. 19). fang-like (Plate 57-1) or with a palptibial claw

I (Fig. 20)). Although the median, or empodial, element of the ambulacra of legs II-III may
be claw-like or sucker-like in certain exceptional families, empodia II-III of the Prostigmata
generally are pad-like, membranous, or rayed (Plate 5-3,4,5,6). Typically, the Prostigmata

I are weakly sclerotized forms when compared to the parasitiform mites. A distinct sternal
shield does not occur and genital shields, where present, do not exceed two in number.
Two or three pairs of discs or "suckers" mayor may not flank the genital opening (plate
I 56-1). Ocelli may be present (Plate 62-1) or absent.

Unlike the majority of suborders, sexual dimorphism in the Prostigmata is often so

I obscure that it is difficult or impossible to separate males from females. Differences
between late nymphal instars and adults may be confined to differences in the number of

I genital discs, or in size.

The Prostigmata is cosmopolitan in distribution, and virtually unlimited in habitat.

I Their great morphological variety is reflected in the necessity for five supercohort categories
to contain the approximately 60 families of terrestrial Prostigmata, and 53 families of
water mites comprising the Supercohort Hydrachnellae.

I The Hydrachnellae is an apparently polyphyletic group (Fig. 21) containing highly

diverse species from aquatic and semi-aquatic habitats. Specialization for life in restricted
I niches (i.e., the surface of submerged rocks, surface film on wet substrates, stream

I
I
142
I
tibial claw
I
I
I
I
I
I
I
I
Fig. 19. Gnathosoma of a tydeoid mite. Fig. 20. Gnathosoma of Trombidium sp. (Wyoming) showing
I
the palpal thumb-claw complex.

bottoms, hot springs, etc:} and under more or less constant environmental conditions has
I
resulted in distinctive and occasionally startling modifications in morphology. While the
genera of Hydrachnellae are quite distinct, most attempts at a familial classification have I
been something less than satisfactory. Relationships between genera are difficult to
establish, owing to the almost complete reliance of many specialists on unstable exoskeletal
structure. Viets (1961) lists 53 families of water mites which are relegated to 12 super-
I
families. These categories were established primarily on exoskeletal structure of adults.
Mitchell (1957, 1964) has considered structural and behavioral features of hydrachnellid
larvae also, and proposes the erection of four superfamilies based on larval morphology and
I
behavior as correlated with life history.

Mitchell's superfamilies are keyed in the following pages, and illustrations of represent-
I
ative larval and adult types are given.
I
Useful References
I
Cunliffe, F. (1955). A proposed classification of the trombidiforme mites. Proc. Ent. Soc. Wash.
57(5) :209-218.
Mitchell. R.D. (1957). Major evoiutionary lines in water mites. Syst. ZooI. 6(3): 137-148. I
Mitchell, R.D. (1964). An approach to the classification of water mites. Acarologia 6 (fasc. h.s.):75-79_
I
I
I
I
143

I
I
Suborder Supercohort Cohort Superfamily
I Endeostigmata - - - - Pachygnathina - - - - - PachygnathoidBa

I Heterostigmata---- Tarsonemina - - - - - Tarsonemoidea

I Eupodoidea

Tydeoidea
Eupodostigmatina

I Bdelloidea

I Halacaroidea

Stomatostigmatina--- Labidostommoidea

I Promata - - - - - " \ Caeculoidea

I Prostigmata
Raphignathoidea

lolinoidea

I i Eleutherogonina I Tetranychoidea

I L Tetrapodilina------- ~--- . . Eriophyoidea

Anystoidea

I Cheyletoidea

I Hydrovolzioidea

Hydryphantoidea
-------------=="'1
I Hydrachnellae
Hydrachnoidea

I __ _ _ _ _ _ _ _ _ _ _ _ _[
Hygrobatoidea

Erythraeoidea

I Parasitengona
Trombidioidea

I Fig.21. Dendrogram illustrating possible relationships within the order ACARIFORMES: suborder
Prostigmata of the subclass ACAR I.
I
I
I
144
I
Schweizer, J. and C. Bader (1963). Die Landmilben der Schweiz (Mittelland, Jura und Alpen). Trombidi-
formes Reuter. Mem. Soc. Helvet. Sci. Nat. 84(2):209·378 + vi.
I
Viets, K.O. (1961). Die Familien· Namen der Milben des Slisswassers. Eine Revision nach den Regeln der
ICZN. Senck, Bioi. 42(1·2):123-130. I
Superfamily Pachygnathoidea I
DIAGNOSIS: Soft-bodied, occasionally showing signs of segmentation; chelicerae variable
but always with opposed chelae; palpi simple; stigmatal openings absent. I
With 7 or 2 pairs of pseudostigmatic organs on propodosoma; tarsi with or
without apoteles. With 2-3 pairs of genital discs.
I
The Pachygnathoidea is a cosmopolitan assemblage comprising seven families, all of
which commonly are found as free-living forms in litter, humus or moss. While it is
convenient to group the various members of the superfamily in a single taxon on the basis
I
of particular characteristics (see key to families), there are strong indications that certain of
the families may in fact have affinities with the suborder Cryptostigmata rather than with
the Prostigmata. The family PACHYGNATHIDAE, for example, might easily be grouped
I
with the primitive Oribatei Inferiores (page 293), while the ubiquitous family NANOR-
CHESTIDAE shares more structural similarities with the eleutherogonine Prostigmata. I
The PACHYGNATHIDAE commonly inhabit forest litter, but species are known also
from the littoral zone. They are found commonly in North America and Europe, and have I
also been collected in South America (Thor and Willmann 1941) and Japan (Shiba 1969).
Their feeding habits are unknown. The fossil species Protacarus crani Hirst, considered by
some authors to be a member of the superfamily Eupodoidea, probably is a pachygnathid
I
(Hirst 1923).

The A LI CO RHAG I J DAE is a small family composed of a single genus and less than a
I
half dozen species, all of which are inhabitants of moss or leaf mold. These tiny (ca. 300/.1)
mites appear to be unsuited as predators, and may feed on vegetable debris or mold. They
are recorded from Europe and from Japan. Members of the family LORDAL YCHIDAE
I
are found commonly in moss and lichens in North America, Europe and Japan, and
species of the lordalychid genus Hybalicus have been recovered from humus in Java (Thor I
and Willmann 1941). The monotypic family SPHAEROLICHIDAE comprises species
which live in dry vegetable debris in Europe, North America and Japan. Few species have
been identified (Grandjean 1939, Thor and Willmann 1941), and the feeding habits of the I
family are unknown.

Members of the family NANORCHESTIDAE are found throughout the world in moss,
I
humus and soil. A species of Speleorchestes has been taken from ant nests in the Pacific
Northwest, where it may be associated with fungi. Nanorchestes antarcticus Strandtmann
is common to the Antarctic (Strandtmann 1964), and is one of the most southern terres-
I
trial arthropods known. It has also been collected in subalpine habitats in Japan (Shiba
1969). N. amphibius is a littoral species which is found living in the minute cracks
between rocks in the intertidal region in Europe. Specimens of what may be N. amphibius
I
have been found in rock cracks in the Oregon intertidal zone.
I
I
I
I 145

I The TERPNACAR IDAE comprises three described genera, the members of which occur

•
I
in dry situations in vegetable debris. One undescribed species from the Pacific Northwest
occurs in wood dust generated by the activities of subterranean termites. The family
PEDICULOCHELIDAE, once felt to be a primitive group of the Astigmata, is considered
by modern workers to be a pachygnathoid family. Paralychus raulti (Lavoipierre) has been
found associated with bees in South Africa (Lavoipierre 1946). Other pediculochelids
have been collected from chickens in the Philippines and, probably accidentally, from a rat
in Florida.
I
I Useful References

Grandjean, F. (1939). Quelques genres d'acariens appartenant au groupe des Endeostigmata. Ann.

I Sci. Zool., Ser. 11:1-122.

TERPNACARIDAEJ
[ALICORHAGIIDAE, LORDALYCHIDAE, SPHAEROLICHIDAE,

Grandiean, F. (1942)_ Quelques genres d'acariens appartenant au groupe des Endeostigmata (2 Ser_).
Premiere partie. Ann. Sci. Nat. Zool., Ser.l1(4):85-135. [PACHYGNATHIDAEJ
I Grandjean, F. (1943). Quelques genres d'acariens appartenant au groupe des Endeostigmata (2 Ser.),
Deuxieme partie. Ann. Sci. Nat. Zool" Ser. 11 (5):1-59. [PACHYGNATHIDAEj
Hirst, S. (1923). On some arachnid remains from the Old Red Sandstone (Rhynie Chert 8ed, Aberdeen'
I shire). Ann. Mag. Nat. Hist. 12(9):455-474. [PACHYGNATHIDAEj
Lavoipierre, M. (1946). A new acarine parasite of bees. Nature 158(4004) :130. [PEDICULOCHELIDAEj

I Shiba. M. (1969). Taxonomic investigations on free-living mites in the subalpine forest on Shiga Heights
IBP area. IL Prostigmata. Bull. Nat. Sci. Mus. 12(1) :65-115.
Strandtmann, R.W. (1964). Insects of Campbell Island. Prostigmata: Eupodidae, Penthalodidae,
Rhagidiidae, Nanorchestidae, Tydeidae, Ereynetidae. Pac. Insects Mono.: 148-156.
I Thor, S. and C. Willmann (1941). Acarina. Prostigmata 6·11 (Eupodidae, Penthalodidae, Penthaleidae,
Rhagidiidae, Pachygnathidae, Cunaxidae). Das Tierreich 71a: 1·186 + xxxvi.

I Womersley, H. (1944). Australian Acarina, families Alycidae and Nanorchestidae. Trans. Roy. Soc. So.
Austral. 68( 11: 133·143.

I Superfamily Tarsonemoidea

I DIAGNOSIS: Soft-bodied or sclerotized, often showing signs of segmentation; chelicerae

stylet-like, inserted in an inconspicuous gnathosoma; palpi simple, minute.
Stigmatal openings absent in males, opening posterolaterad of gnathosoma
I in female. Pseudostigmatic organs present or absent; tarsi usually with

•
claws; number of legs variable, from 1 to 4 pairs.

The Tarsonemoidea contains approximately 350 species grouped into five families.
Representatives of these families are found virtually throughout the world as insect
associates, fungivores or plant feeders. Most of the known species are small in size
I (200-400 iLl and lightly sclerotized, although species of more than one tarsonemoid
family exhibit heavy sclerotization.

I The PYEMOTI DAE, one of the larger families of the Tarsonemoidea, includes over
100 species of which several are of agricultural or medical significance (Cross 1965).

I Siteroptes cerealium Kirchner is a known vector of Fusarium poae, the fungal pathogen
which causes carnation bud rot (Cooper 1940). S. cerealium may also be implicated in

I
I
146
I
silver top of grasses in North America and Europe although Hardison (1959) presents I
evidence to the contrary. Pyemotes ventricosus (Newp.), commonly a parasite of larval and
pupal Lepidoptera, often is a part of the arthropod fauna of grain storages. P. ventricosus
may attack grain handlers, causing a dermatitis which can be quite severe (Nixon 1944).
I
Species of Pyemotes attack and generally kill immature Homoptera, Coleoptera, Diptera
and Hymenoptera, as well as Lepidoptera. Certain members of the pyemotid genus
Acarophenax may be beneficial in that they parasitize graminivorous beetles of the genera
I
Tribolium and Cryptolestes (Newstead and Duvall 1918, Cross and Krantz 1964).

The TARSONEMIDAE includes phytophagous, fungivorous and insectophilous mites

I
grouped in 10-12 genera. Although the family is primarily tropical or subtropical (Beer
1954). many species are found in both the nearctic and palaearctic regions. Species of the
genera Steneotarsonemus and Hemitarsonemus are phytophagous, with the cyclamen mite
I
S. pallidus (Banks). and Hemitarsonemus latus (Banks). being of prime economic importance
(Moznette 1917, Gadd 1946). Other members of the genus Steneotarsonemus feed I
primarily on monocotyledonous hosts. Two species of Steneotarsonemus utilize the galls
of eriophyid mites (superfamily Eriophyoidea). forcing the eriophyids to abandon the galls
(Beer 1963).
I
Mites of the genus Iponemus prey on the eggs of bark beetles and live as commensals
in the beetle galleries (Lindquist and Bedard 1961, Lindquist 1969). The adults are
I
phoretic on the beetles and are easily carried from gallery to gallery. Other tarsonemids
are associated with bees (Lindquist 1968) or with beetles (Beer 1958). The majority of the
species of the genus Tarsonemus are fungivorous or algivorous.
I
All of the known members of the family PODAPOLIPIDAE are arthropod
parasites which usually illustrate rather bizarre reductions in number of appendages.
I
Podapolipids pass through a hexapod larval and a larviform adult stage. Females later
molt to sac-like reproductive forms which often exhibit reduction in leg number. I
Podapolipus, for example, has only one pair of legs when mature, while Eutarsopolipus
has three pairs. Locustacarus trachealis Ewing infests the tracheae and air sacs of grass-
hoppers, probably piercing the tracheal walls to feed (Wehrle and Welch 1925). Podapolipus
I
grassi Berlese feeds externally on grasshoppers, while Tetrapolipus and Eutarsopolipus are
found under the elytra of beetles. I
The CROTALOMORPHIDAE is a manuscript name for an unusual species of
tarsonemoid mite found under the elytra of the beetle Agonoderus pallipes in Kansas. I
Useful References

Beer, R.E. (954). A revision of the Tarsonemidae of the western hemisphere. Univ. Kansas Sci. Bull.
I
36.2(16):1091-1387.
Beer, R.E. (1958). The genus Tarsonemella Hirst, with description of a new species (Acarina, Tarso-
nemidae). Jour. Kans. Ent. Soc. 31(2):188-192.
I
Beer, R.E. (1963). Social parasitism in the Tarsonemidae, with description of a new species of tarsonemid
mite involved. Ann. Ent. Soc. Amer. 56(2):153-160.
Beer, R.E. and A. Nucifora (1965). Revisione dei generi della famiglia Tarsonemidae (Acarina). Boll.
I
Zoo I. Agr. Bachic., Ser. 2, 7:19·43.
Cooper, K.W. (1940). Relations of Pediculopsis graminum and Fusarium poae to bud rot of carnation.
Phytopath.30(0):853-859. [PYEMOTIDAE]
I
I
I
I 147

I
I
•
I
I
I ~genus
. ~
no. legs
if
Dorslpes 3 4
I Locustacarus ~3 3
I Tarsopolipus./ ~ 4
Eutarso~ipus 3
I . .~.,tap1IlpUS 2-3
Podapolipus 1-3
I
I Fig. 22. Chart indicating degree of leg reduction in selected genera of the family PODAPOLIPIDAE. The
larviform female of Locustacarus is figured on the left, and the reproductive female of
Podapolipus is shown on the upper right.

I Cross, EA. (19651. The generic reiationships of the family Pyemotidae (Acarina: Trombidiformes).
Univ. Kansas Sci. Bull. 45(21 :29-275.

I Cross, EA. and G.W. Krantz (19641. Two new species of the genus Acarophenax Newstead and Duvall
1918 (Acarina: Pyemotidae). Acarologia 6(21:287·295.
Gadd, C.H. (1946). Observations on the yellow tea mite Hemitarsonemuslatus (Banks) Ewing. Bull. Ent.

I Res. 37:157-162. [TARSONEMIDAEj

Hardison, J.R. (1959). Evidence against Fusarium poae and Siteroptesgraminum as causal agents of silver
top of grasses. Mycologia 51(51 :712-728. [PYEMOTI DAEI

I Karafiat, H. (19591. Systematik und Okologie der Scutacariden.

Acarina 1(4) :627-712.
Karl, E. (1965a).
Beitr. Syst. Okol. mitteleurop.

Untersuchungen zur Morphologie und Okologie von Tarsonemiden gartnerischer

I Kulturpflanzen. I. Tarsonemus pallidus Banks. BioI. Zbl. 84:47-80.

Karl, E. (1965b). Untersuchungen zur Morphologie und Okologie von Tarsonemiden gartnerischer
Kulturpflanzen, 11. Hemitarsonemus latus (Banks), Tarsonemus confusus Ewing, T. talpae
Schaarschmidt, T. setiter Ewing, T. smithi Ewing und Tarsonemoides belemnitoides Weis-Fogh.
I BioI. Zbl. 84:331-357.
Krczal, H. (1959). Systematik und Okoiogie der Pyemotiden. Beitr. Syst. Okol. mitteleurop. Acarina
1(3) :385·625.

I Lindquist, E.E. (1969). Review of holarctic tarsonemid mites (Acarina: Prostigmata) parasitizing eggs of
ipine bark beetles. Mem. Ent. Soc. Canada 60:111 pp.

I
I
148
I
I
•
Lindquist, E.E. and W.D. Bedard (1961). Biology and taxonomy of mites of the genus Tarsonemoides
(Acarina: Tarsonemidae) parasitizing eggs of bark beetles of the genus Ips. Can Ent. 83:982-999.
Newstead, R. and H.M. Duvall (1918). Bionomic, morphological and economic report of acarids of
stored 9rain and flour. Roy. Soc. Rept. Grain Pests (War) Cmttee. No. 2:48 pp. [PYEMOTIDAE]
Nixon, J.W. (1944). Cheese 'itch' and 'itchy' cargoes in reference to workmen's compensation. Proc. Roy.
Soc. Med. London 87:405-410. [PYEMOTIDAE]
Regenfuss, H. (1968). Untersuchungen zur Morphologie, Systematik und Okologie der Podapolipidae
(Acarina, Tarsonemini) Zeitschr. wissenschaftl. Zoo I. 177(3/4): 183·282.
I
Rennie, J. (1921). Isle of Wight disease in hive bees-Acarine disease; the organism associated with the
disease· Tarsonemus woodi. sp. n. Trans. Roy. Soc. Edinburgh 52:768·779. [TARSONEMIDAE] I
Schaarschmitt, L. (1959). Systematik und Okologie der Tarsonemiden. Beitr. Syst. Okol. mitteleurop.
Acarine 1(5) :713-823.
Volkonsky, M. (1940). Podapolipus diander n. sp. acarien heterostygmate parasite du criquet migrateur
(Locustamigratoria L.). Arch.lnst. Pasteur Alger. 18:321-340. [PODAPOLIPIDAEJ
I
Wehrle, L.P. and P.S. Welch (1925). The occurrence of mites in the tracheal system of certain Orthoptera.
Ann. Ent. Soc. Amer. 28(1 ):35·44. [PODAPOLIPIDAE] I
Superfamily Eriophyoidea 1 I
DIAGNOSIS: Soft-bodied or sc/erotized, annulate worm-like forms; chelicerae stylet-like,
palpi minute, simple. Without conspicuous tracheal system. With only 2
I
pairs of legs, inserted anteriorly; apoteles present. Genital opening
transverse. I
The Eriophyoidea includes a number of 4-legged elongate, annulate phytophagous
mites which show strong morphological and behavioral affinities for the Tetranychoidea
(page 156). The eriophyoids are considered here as comprising a single family, the
I
ERIOPHYIDAE.
I
Eriophyids are referred to as blister mites, rust mites, bud mites or gall mites,
depending on the type of injury they cause. They are found almost exclusively on perennial
plants, and illustrate a high degree of host specificity. Generally, the feeding injury I
occasioned by eriophyids is mild, although some species may cause noticeable damage to
their hosts. I
The ERIOPHYIDAE may be divided into subfamilial groups on the basis of habitat,
and certain morphological traits. For example, fusiform eriophyid mites which are found
wandering on the leaf or bud surfaces of their hosts are classified as rust mites or leaf
I
vagrants. These mites have the gnathosoma partially covered by an anterior extension of
the dorsal shield. Included here are the subfamilies Sierraphytoptinae and the Phyllo-
coptinae (Keifer 1952). Soft-bodied worm-like eriophyids which are found within buds, in
I
blisters or in galls are referred to as bud or gall mites. These mites have no dorsal shield
extension, and the entire gnathosoma is visible. Bud and gall mites are grouped in the sub- I
families Phytoptinae, Eriophyinae and Aberoptinae (Keifer 1952, 1966). Economically
important species are found primarily in the Phytoptinae and Eriophyinae.
I
Leaf injury caused by eriophyid mites may be of several types, and may involve the
surface tissue or the leaf mesophyll (Keifer 1952). Vasates cornutus (Banks), a rust mite
1See footnote, page 174.
I
I
I
I
149

I found on peach, causes a browning or silvering of the leaf surface. Aceria granati (Can.)
brings about edgerolling or folding of pomegranate leaves, while A. brachytarsus (K.)
I causes pocketing of leaf tissue, or purse galls, on its walnut host. Stunting of leaves is a
symptom produced on Euonymus by Eriophyes convolvens (Nal.). Another species of
Eriophyes, E. pyri (Pgst.}, is a blister mite which invades the mesophyllous leaf tissue of
I pears and causes serious injury to the host. Gall-forming mites, such as Aceria fraxinivorus
K. on ash, cause a type of tissue pocketing in which the pocket is virtually closed, and in

I which the abnormal growth of leaf hairs forms a protective mat for the eriophyid inhabi-
tants. The formation of hairy patches or erinea on the underside of grape leaves infested
by Eriophyes vitis (Pgst.l constitute "open galls," in which pocketing of leaf tissue is

I minimal.

Bud injury may be brought about by surface feeding, or by gall formation. The
I citrus bud mite, Aceria sheldoni (Ewing}, is an important surface feeder which causes
severe fruit and leaf malformation in California coastal fruit orchards (Boyce and Korsmeier
1941). The filbert mite, Phytoptus avellanae Nal., forms galls in terminal filbert buds and
I prevents fu rther growth.

I In addition to leaf and bud injury, some eriophyids cause "witches broom" (adventi-
tious twig development), flower galls, shortening of internodes, or secondary development of
leaf hair. Aceria tulipae (K.l, a species found in bulb storages, feeds between tissue layers of

I onion, tulip or garlic bulbs and brings about eventual deterioration.

Eriophyids are the only mites which have been reported to transmit plant viruses

I (Slykhuis 1963, Nault et al. 1967). Aceria tulipae transmits kernal red streak of corn,
wheat spot and wheat streak mosaic viruses, while A. ficus (Cotte) transmits fig mosaic.
Other examples of plant viruses transmitted by eriophyid mites include currant reversion
I (Phytoptus ribis (Westw.) peach mosaic (Eriophyes insidiosus Keifer and Wilson) and
ryegrass mosaic (Abacarus hystrix (Nal.)).

I Mites of the subfamily Phyllocoptinae produce, in addition to the ordinary females, a

special overwintering female called a deutogyne. Eggs laid in the spring by deutogynes

I produce ordinary summer females. Detailed information on deuterogyny is given by

Keifer (1942, 1952).

I Useful References

I Boyce, A.M. and R.B. Korsmeier (1941). The citrus bud mite, Eriophyes sheldoni Ewing. Jour. Ecan. Ent.
34(61 :745-756.
Hodgkiss. H.E. (1930). The Eriophyidae of New York. II. The maple mites. Cornell Univ. Agr. Exp. Sta.

I Tech. Bull. 163:45 pp.

Keifer. H.H. (1942). Eriophyid studies XII. Bull. Calif. Dept. Agr. 31:117-129.
Keifer, H.H. (1946). A review of North American economic eriophyid mites. Jour. Ecan. Ent. 39(5):563-

I 570.

•
Keifer, H.H. (1952). The eriophyid mites of California (Acarina: Eriophyidael. Bull. Calif. Insect Surv.
2(11:123 pp .
Keifer, H.H. (1964). Eriophyid studies B-l1. Bull. Calif_ Dept. Agr.:20 pp.
Keifer, H.H. (1966), Eriophyid studies B-18. Bull. Calif. Dept. Agr.:20 pp.

I
I
150
I
Nault, L.R., M.L. Briones, L.E. Williams and B.D. Barry (1967). Relation of the wheat curl mite to kernel
red streak of corn. Phytopath. 57(9) :986-989.
I
Slykhuis, J.T. (1963). Mite transmission of plant viruses. Advances in Acarology 1:326-340.
I
Superfamily Eupodoidea
I
DIAGNOSIS: Soft-bodied, with a tubercle on anterior portion of propodosoma bearing a
pair of setae; movable cheliceral digit stylet-like or chelate and opposed to
movable digit; palpi simple. Tracheal system present, opening at base of
I
chelicerae. With or without apoteles on tarsi I, with 1 or more rhagidial
organs on tarsus I lying flush with tarsus; with 2 or 3 pairs of genital discs. I
Four families are included in the Eupodoidea-the EUPODIDAE. RHAGIDIIDAE,
PENTHALODIDAE and PENTHALEIDAE. Eupodoids may be predaceous, fungivorous, I
phytophagous or parasitic, and are world-wide in distribution.

The EUPODIDAE comprises a group of species which, for the most part, are confined
I
to damp soil, humus or moss in temperate climates. The several known species of the
genus Eupodes have well developed femora IV (Thor and Willmann 1941) and are accom-
plished jumpers. Members of the genus Linopodes are fungivores which are often found
I
under stones or debris in open situations. When threatened, Linopodes is capable of rapid
backward movement, an ability which no doubt serves as a defensive response. Species of
Linopodes may be encountered in mushroom houses, where they cause economic injury
I
through feeding on the root system of the mushroom host. Benoinyssus najae Fain, the sole
representative of the eupodid subfamily Benoinyssinae, was found in the nasal fossae of the
cobra Naja melanoleuca in Ruanda-Urundi (Fain 1958).
I
Members of the predaceous family RHAGIDIIDAE are soft-bodied, fast mites found I
under stones or debris in essentially the same habitats as many of the EUPODIDAE. Their
strongly developed chelate chelicerae are excellently su ited for their predatory existence.
Rhagidiids are common to the holarctic regions, although species have been collected in
I
South Africa as well (Meyer and Ryke 1960). The often colorful, delicately ornamented
species of the family PENTHALODIDAE also may be predaceous (Baker 1946), and
generally are collected in moss or leaf mold throughout much of the world.
I
The winter grain mite, or blue oat mite, Penthaleus major (Duges) is one of the
economically important species of the phytophagous family PENTHALEIDAE. This
I
cosmopolitan species attacks a wide variety of host plants including peas (Campbell 1941),
grain (Chada 1956) and clover (Swan 1934). Halotydeus destructor (Tucker), the red-
legged earth mite or black sand mite, is a common pest of vegetable and legume crops in
I
South Africa (Meyer and Ryke 1960) as well as in Australia. Species of both Halotydeus
and Penthaleus have been collected from non-plant habitats (Thor and Willmann 1941), so I
it appears likely that at least some species are not phytophagous.
I
Useful References

Baker, E.W. (1946). New species of North and Central American mites of the family Penthaleidae
I
(Acarina). Jour. Wash. Acad. Sci. 36(121:421-425. [PENTHALODIDAE)
I
I
I 151

I Campbell, R.E. (1941). Further notes on the blue oat or pea mite Penthaleus major (Duges) in California.
Calif. Dept. Agr. Bull. 30:312-314. [PENTHALEIDAE]

I Chada, H.L. (1956). Biology of the winter grain mite and its control in small grains. Jour. Econ. Ent.
49(4) :515-520.
Fain, A. (1958). Un nouvel acarien Trombidiforme parasitant les fosses nasales d'un Serpent au Ruanda-

I Urundi. Rev. Zool. Bot. Afr. 57(1-2):177-1B3. [EUPODIDAE]

Meyer, M.K.P. and P.A.J. Ryke (1960). Mites of the superfamily Eupodoidea (Acarina, Prostigmata)
associated with South African plants. So. Afr. Jour. Agr. Sci. 3(4):481-496.

I Solomon, M.E. (1937). Behaviour of the red-legged earthmite, Halotydeus destructor, in relation to
environmental conditions. Jour. Anim. Ecol. 6(2):340-36"
Swan, D.C. (1934). The blue oat mite (Penthaleus major Duges). Jour. Agr. So. Austral. 38:365-367.

I Thor, S. (1934). Einzelne neue, besonders Norwegische Eupodes Arten mit einigen alteren Arten
vergleichen. Zool. Anz.105:201-215. [EUPODIDAE]
Thor, S. and C. Willmann (1941). Acarina,71a. Eupodidae, Penthalodidae, Penthaleidae, Rhagidiidae,

I Pachygnathidae, Cunaxidae. Das Tierreich 71a: 1-186 + xxxvi.

I Superfamily Tydeoidea

DIAGNOSIS: Soft-bodied,' cheliceral bases usually fused, movable digit stylet-like; palpi

I simple. Stigmata, when distinct, opening at base of chelicerae. Idiosoma

with one or two pairs of pseudostigmatic organs. With apoteles on legs;
a
one or several erect sensory rods on tarsus I; with 2 or pairs of genital discs.

I The superfamily Tydeoidea is a grouping of four families, of which at least three

include predatory forms. The superfamily is cosmopolitan in distribution.

I The elongate, secondarily segmented members of the family PARATYDEIDAE occur

in soil and litter in Europe and in North and Central America (Baker 1949, 1950) where

I they apparently prey on other minute animals. Collections at hand include species from a
bird nest in Utah, and from moss in Oregon. The NEMATALYCIDAE is represented by
elongate worm-like species found in coastal sand or in pasture soil (Strenzke 1954, Cunliffe
I 1956). It is included in the Tydeoidea only provisionally. The feeding habits of nemataly-
cids are unknown.

I While some members of the EREYNETI DAE are predators in soil or vegetable debris,
others may be intimately associated with terrestrial gastropods, or found in the respiratory

I system of birds, mammals or amphibians. Ricardoella limacum (Schrank), the slug mite, as
well as other members of this genus, feed on the mucous secretions of slugs and snails.
Members of the ereynetid subfamily Speleognathinae inhabit the mucosa of the nasal

I passages of a wide variety of birds and mammals throughout the world (Clark 1960, Fain
1963). Clark notes that Boydaia sturnellae Clark, a North American speleognathine,
appears to be capable of feeding on whole blood in the turbinates of its meadowlark host.
I Nasal parasites of frogs and toads comprise the third ereynitid subfamily, Lawrencarinae
(Fain 1956). The adaptation to nasal passages as a preferred habitat, therefore, is apparent
throughout most of the family.
I The TYDEI DAE is a large cosmopolitan family of weakly sclerotized mites which

I may be predators or plant feeders or, conceivably, both predators and phytophages (Baker
1965, Brickhill 1958). Some species are found in stored products where they prey on
other mites. A species of tydeid in Oregon granaries is, in turn, the preferred prey of a

I
I
152

species of Eupalopsis (superfamily Raphignathoideal. Other mites generally are ignored by

this sedentary prostigmatid. Another tydeid, Tydeus molestus (Moniez) may attack man and
•I
animals, causing irritation to the skin. The habitat for the majority of known tydeids is the
bark or leaves of woody plants, although specimens have been collected from bird nests, I
forest litter, and from insects.
I
Useful References

Baker, E.W. (1949). Paratydeidae, a new family of mites (Acarina). Proc. Ent. Soc. Wash. 51(3): 119-122.
I
Baker, E.W. (1950). Further notes on the family Paratydeidae (Acarina) with a description of another new
genus and species. Jour. Wash. Acad. Sci. 40(9) :289-291.
Baker, E.W. (1965). A review of the genera of the family Tydeidae (Acarina). Advances in Acarology
I
2:95-133.
Brickhill, C.D. (1958). Biological studies of two species of tydeid mites from California. Hilgardia
27(20) :601-620.
I
Clark, G.M. (1960). Three new nasal mites (Acarina: Speleognathidae) from the grey squirrel, the
common grackle, and the meadowlark in the United States. Proc. Helminth. Soc. Wash. 27(1):
103-110. [EREYNETIDAE]
I
Cunliffe, F. (1956). A new species of Nematalycus Strenzke with notes on the family (Acarina.
Nematalycidae). Proc. Ent. Soc. Wash. 58(6):353-355.
Fain, A. (1956). Nouvelles observations sur ies acariens de la famille Speieognathidae parasites des fosses
I
nasales chez les batraciens, les oiseaux et les mammiferes. Ann. Parasitol. 31:644-662. [EREY-
NETIDAE]
Fain, A. (1963). Chaetotaxie et classification des Speleognathinae (Acarina: Trombidiformes). Bull.lnst.
I
Roy. Sci. Nat. Belg. 39:1-80. [EREYNETIDAE]
Fleschner, CA. and K.Y. Arakawa (1953). The mite Tydeus califomicus on citrus and avocado leaves.
Jour. Econ. Ent. 45: 1092. I
Meyer, M.K.P. and P.A.J. Ryke (1959). New species of mites of the families Tydeidae and Labidostom-
midae (Acarina: Prostigmata) collected from South African plants. Acarologia 1(4) :408-420.
Strenzke, K. (1954). Nematalycus nematoides n. gen. n. sp. (Acarina Trombidiformes) aus dem
I
grundwasser der Algerischen Keste. Vie et Milieu 4(4) :638-647.
Thor, S. (1933). Acarina. Tydeidae, Ereynetidae. Das Tierreich 60:1-82 + xi.
Turk, F.A. and S.M. Phillips (1946). A monograph of the slug mite-Ricardoella limacum (Schrank).
I
Proc. Zool. Soc. London 115(3,4):448-472. [EREYNETIDAE]
I
Superfamily Bdelloidea

DIAGNOSIS: Soft-bodied; cheliceral bases free, moving scissors-like aver gnathosoma,

I
movable cheliceral digit small; rostrum elongate, palpi simple or raptorial.
Stigmata open at base of chelicerae. Usually with twa pairs of pseudo-
stigmatic organs on propodosoma; eyes present. With apoteles on legs;
I
with several erect sensory rods an tarsus I; genital area with 3, 2 or 0
pairs of discs. I
Two free-living families are included in the superfamily Bdelloidea-the BDELLJDAE
and the CUNAXIDAE. Both families are cosmopolitan in distribution.
I
The BDELLJDAE is a large family of active red, reddish-brown or green mites which
prey on small arthropods or arthropod eggs (Atyeo 1960). Some are of great size (3000-
I
I
I
I
153
I 4000 Ill. but most species measure between 700-1200 II. Bdellids, or "snout mites:' are
found in a variety of environments, being common in dry exposed desert situations, on
I rock faces or debris in the intertidal zone, or in cool moist forest habitats. Certain species
may be important in regulating populations of spider mites (Snetsinger 1956) or of
springtails (Womersley 1933a). Odontoscirus virgu!atus (C. & F.) has been found associ-
I ated with the phytophagous prostigmatid mite Ha!otydeus destructor (Tucker) in South
Africa (Womersley 1933b, Meyer and Ryke 19591. but its effect on Ha!otydeus populations

I is not known. Similarly, substantial numbers of Biscirus uncinatus (Kramer) were recov-
ered from balsam fir infested with balsam woolly aphid in Canada. It is possible that
aphids are utilized for food by this large bdellid species. Bdella !ignico!a Can. may be

I recovered from arthropod-infested stored grain and flour (Hughes 1961) where it preys on
other mites.

I The members of the family CUNAXIDAE are small (ca. 450-550 II) red or reddish-
brown species which prey on other arthropods in dry or damp humus, leaves, moss or
straw. The effect of cunaxids on prey populations is not clear, although Ewing (1917)
I refers to Cunaxoides parvus (Ewing) as an enemy of the oyster-shell scale. Muma (1961)
reports the presence of species of Cunaxa and Cunaxoides on Florida citrus trees. Cunaxids
often are found on the bark of fruit trees infested with spider mites in the Pacific Northwest,
I but their effect on spider mite populations has not been measured. Occasional records of
cunaxids in stored products also exist (Hughes 1961).

I There is little doubt that, on the basis of striking morphological similarity, the
CUNAXI DAE is closely related to the marine superfamily Halacaroidea (page 1551. and that

I the HALACARIDAE may be a direct offshoot of a cunaxid ancestral stock resembling the
modern genus Bonzia (Atyeo 1958).

I Useful References

I Atyeo, W.T. (1958).

[BDELLIDAEI
The genus Bonzia in the New World. Jour. Kansas Ent. Soc. 31(2):173-177.

Atyeo, W.T. (1960). A revision of the mite family Bdellidae in North and Central America. Univ.
I Kansas Sci. Bull. 40(8) :345-499.
Atyeo, W.T. (1963). The Bdellidae (Acarina) of the Australian Realm. Bull. Univ. Nebraska State Mus.
4(8): 113-21 O.

I Baker, E.W. and A. Hoffmann (1948).

5(3-41 :229-273.
Acaros de la familia Cunaxidae. Anales Esc. Nac. Ciencias BioI.

Ewing, H.E. (1917). New Acarina. Part II. Descriptions of new species and varieties from Iowa,

I Missouri, Illinois. Indiana, and Ohio. Bull. Amer. Mus. Nat. Hist. 37(2):149-172. [CUNAXIDAEj

••
Grandjean, F. (19381. Observations sur les Bdelles (Acariens). Ann. Soc. Ent. France 107:1-24.
Hughes, A.M. (19611. The Mites of Stored Food. Minis, Agr .. Fish. and Food Tech. Bull. 9:287 pp. + vi.
Meyer, M.K.P. and P.A.J. Ryke (1959). Cunaxoidea (Acarina: Prostlgmata) occurring on plants in
South Africa. Ann. Mag. Nat. Hist. Ser. 13,2:369-384.
Muma, M.H. (1961). Mites associated with citrus in Florida. Univ. Fla, Agr. Exp. Sta. Bull. 640:39 pp.
Snetsinger, R. (1956). Biology of Bdella depressa, a predaceous mite. Jour. Econ. Ent. 49(6):745-746.
Thor, S. (19311. Bdellidae, Nicoletiellidae, Cryptognathidae. Das Tierreich 56:1-65 + xiii.

I Thor, S. and C. Willmann (19411. Acarina. Prostigmata 6-11 (Eupodidae. Penthalodidae, Penthaleidae,
Rhagidiidae, Pachygnathidae, CunaxidaeL Oas Tierreich 71a: 1-186 + xxxvi.

I
I
154
I
Womersley, H. (1933a). A possible biological control of the clover springtail or lucerne fiea (Sminthurus
viridis L.) In Western Australia. Jour. Austral. Council Sci. Ind. Res. 6(2):83·91. [8DELLIDAE]
I
Womersley, H. (1933b). Dn some Acarina from Australia and South Africa.
Austral. 57:108·112. [8DELLIDAE]
Trans. Roy. Soc. So.
I
Superfamily Labidostommoidea I
DIAGNOSIS: Heavily sclerotized forms; cheliceral bases free, movable and fixed digits
opposed; palpi simple. Stigmata open at base of chelicerae. With two pairs
I
of pseudostigmatic organs on the idiosoma; lateral eyes present, along with
a pair of conspicuous lens-like protuberances. With apoteles on legs; with
two erect sensory rods on tarsus I; genital area covered by shields in both
I
sexes.

The Labldostommoidea is represented by a single cosmopolitan family, the LABIDO-

I
STOMMIDAE, which is comprised of a small number of heavily armored ornamented preda-
tory species. Labidostommids are moderately large (500-200011), distinctive forms which I
differ radically from related prostigmatid families in their morphology (Grandjean 1942).
They are found in humus, lichens, soil or moss where they subsist on other small arthropods.
Specimens also have been taken from caves in Europe. Labidostomma pacifica (Ewing), a
I
common labidostommid in the Pacific Northwest, stalks its prey with the well developed
first pair of legs held as pincers. Legs II-IV are utilized for locomotion. I
Useful References I
Grandjean, F. (1942). Observations sur les Labidostommidae.
14(2):118-125; (3):185·192; (5):319-326; (6):414-418.
8ull. Mus. Hist. Nat. Paris Ser. 2,

Greenberg, 8. (1952). New Labidostommidae with keys to the New World species (Acarina). Jour. N.Y.
I
Ent. Soc. 60:195·209.
Shiba, M. (1969). Taxonomic investigations on free-living mites in the SUbalpine forest on Shiga Heights
18PArea. II. Prostigmata. 8ull. Nat. Sci. Mus. 12(1):65·115.
I
Thor, S. (1931). 8deliidae, Nicoletiellidae, Cryptognathldae. Das Tierreich 56:66-77.
I
Superfamily Caeculoidea
I
DIAGNOSIS: Heavily sclerotized forms, with several dorsal shields; chelicerae short, thick,
with strong sickle-shaped movable chelae; palps with thumb-claw complex.
Stigmata open at the base of chelicerae; lateral eyes present. With apoteles I
on legs, legs I with strong internal spine-like setae; tarsus and tibia I each
with a pit containing a sensory seta; without genital discs. I
A single family, the CAECU LI DAE, comprises the superfamily Caeculoidea. Caeculids
are large (1000-300011) heavily armored, wrinkled, slow-moving predators which appear to
prefer dry hot open habitats to more protected sites. They are commonly encountered in
I
desert situations under rocks, where they apparently lie in wait for their more agile
I
I
I
I 155

I arthropod prey. The inner faces of legs I are armed with strong sclerotized spines which
serve to capture or hold the prey. An undescribed species of caeculid is found in sagebrush

I litter in central Oregon, where it preys on other prostigmatid mites found in the same
habitat.

I Useful References

I Andre, M. (1935). Notes sur Ie genre Caeculus Dufour (Acariens) avec descriptions d'especes nouvelles
africaines. Bull. Soc. Hist. Nat. Afr. Nord 26:79·127.
a
Coineau, Y. (1963). Contribution I'etude des Caeculidae. Premiere serie: Developement postlarvaire de

I Allocaeculus cata/anus Franz 1954,-Premiere partie: la chaetotaxie du corps. Acarologia 5(2):189-

212.
Coineau, Y. (1966), Contribution a I'Eltude des Caeculidae. Deuxieme serie: Developement postlarvaire

I de Microcaeculus hispanicus Franz 1952. Acarologia 8(2):23-44.

Franz, H. (1952). Revision der Caeculidae Berlese 1894 (Acari). Bonner Zoo I. Beitr. 2(1-2):91·124.
Grandjean, F. (19441. Observations sur les Acariens du genre Caeculus. Arch. Sci. Phys. Nat., 5, 26:33-46.

I Mulaik, S. (1945). New mites in the family Caeculldae. Bull. Univ. Utah 35(171:1·23.

I Superfamily Halacaroidea

I DIAGNOSIS: Primarily marine; with sclerotized dorsal and ventral shields; movable digit of
chelicera reduced to a membranous structure,' palpi simple or raptorial,
opposed to elements of rostrum in one group (Lohmannellinaei. Stigmatal
I openings apparently absent; eyes present. Apoteles present on all legs; with
three pairs of genital discs.

I The marine acarine superfamily Halacaroidea includes a single family, the HALA-
CARIDAE, which is an assemblage of over 300 species of mites from a wide range of
habitats. Most of the known species have been collected from intertidal, arenicolous,
I cavernicolous or shallow subtidal situations (Newell 1947), but some have been found at
depths exceeding 4000 meters (Newell 1967). Members of several genera occur in brackish

I or fresh water, and some have evolved as external or internal parasites of marine inverte-
brates (Viets 1931, 1938). The members of the subfamily Rhombognathinae are thought
to be algivorous, although feeding in these forms has not been studied extensively.

I Morphologically, species of the Lohmannellinae and the Simognathinae seem well suited to
predation. Thus, the HALACARIDAE contains species which appear to have occupied all
of the major marine habitats. Despite their great diversity in habitat, however, the

I halacarids present an unusual degree of morphological uniformity.

While the HALACARIDAE inhabit oceans and seas throughout the world, certain
I genera appear to be dominant at different latitudes. For example, the genera Meta-
rhombognathus and Arhodeoporus appear to be confined to temperate seas, while Copid-
ognathus is common in tropical waters. Others are found principally in fresh or brackish
I water (Astacopsiphagus, Soldanellonyx) or in sand (Actacarus).

I
I
I
156
I
Useful References I
Andre, M. (1946). Halacariens marins. Off. Centro Faun. 46:1-152.
Newell, I.M. (1947). A systematic and ecological study of the Halacaridae of eastern North America. Bull.
Bingham Ocean. Coil. 10(3): 1·232.
I
Newell, I.M. (1953). The natural classification of the Rhombognathinae (Acari, Halacaridael, Syst. Zool.
2(3):119·135. I
Newell, I.M. (1967). Abyssal Halacaridae (Acari) from the southeast Pacific. Pac. Ins. 9(4):693-708.
Viets, K. (1931). Uber die an Krebskiemen parasitierende Susswassermilbe Astacocroton Haswell, 1922.
Zool. Anz. 97(3·4) :85-93. I
Viets, K. (19361. Spinnentiere oder Arachnoidea, Vii. Wassermilben oder Hydracarina (Hydrachnellae
und HalacaridaeJ. Tier. Deutschl. 31, 32:1-574 + x.
Viets, K. (1938). Eine neue, in tiefsee·echiniden schmarotzende Halacaridengattung und Art (Acari).
I
Zeit Parasit. 10(2):210-216.

I
Superfamily lolinoidea
DIAGNOSIS: Soft-bodied; movable cheliceral digit long, whiplike; palp simple, 7-
segmented. Stigmatal openings at base of chelicerae, peritremes indistinct;
I
eyes absent. Apoteles absent on legs I; genital opening transverse.
I
lolina nana Pritchard is the type species of the family IOLINIDAE, which in turn is the
only representative family of the lolinoidea. I. nana is a parasite of tropical roaches
(Pritchard 1956) and is considered by Pritchard to be an intermediate form between the
I
predaceous Raphignathoidea and the phytophagous Tetranychoidea. Fain and Evans (1966)
have shown that I. nana is closely related to species of the tydeoid genus Proctotydeus,
associates of locusts. The lolinoidea may, therefore, be more properly considered a family
I
under the superfamily Tydeoidea (p. 151).

Useful References
I
Fain, A. and G.O. Evans (1966). The genus Proctotydeus Berl. (Acari: lolinidae) with descriptions of two
new species. Ann. Mag. Nat. Hist. 9(131:149·157. I
Pritchard, A.E. 119561. A new superfamily of trombidiform mites with the description of a new family,
genus, and species (Acarina: loiinoidea: lolinidae: lolinanana). Ann. Ent. Soc. Amer. 49(3):204·206.
I
Superfamily Tetranychoidea

DIAGNOSIS: Soft-bodied; movable cheliceral digit long, whiplike; with or without a

I
palpal thumb-claw complex. Stigmatal openings at base of chelicerae,
peritremes well developed; eyes present or absent. Apoteles on all legs, with
tenent hairs on claws, and often on empodia; genital opening transverse.
I
The five families of Tetranychoidea include over 350 phytophagous species, many of I
which are of economic importance. The majority of species are contained in two families-
the TETRANYCHIDAE and the TENUIPALPIDAE.
I
Species of the family TETRANYCH IDAE are found throughout the world on
virtually every major food crop and ornamental plant. Their feeding often results in I
I
I
I 157

I serious injury or death of the host. Tetranychid mites are green, yellow, orange or red in
color, and often are found associated with fine silk webbing which the mites spin from

I glands probably located in the palps (Grandjean 1948).

Grasses and other low-growing plants often are infested by bryobiine tetranychids.
I Bryobia praetiosa Koch is common on grasses in Europe and North America (Pritchard and
Baker 1955). It often invades houses in great numbers, particularly in the fall. Mono-
ceronychus, a grass-inhabiting bryobiine genus, is more or less confined to grass, while
I Petrobia, Aplonobia and Hystrichonychus (among others) may be collected from a
variety of low-growing weeds and food crops. Not all bryobiines are confined to low-

I growing plants. Bryobia rubrioculus (Scheuten) is an important pest of pome and stone
fruit trees in both the Old and New Worlds (Morgan 1960). Unlike the tetranychine spider
mites, many bryobiine tetranychids apparently do not spin webs on their plant hosts

I (Geijskes 1939).

The tetranychid subfamily Tetranychinae includes a number of economically

I important forms, only a few of which can be mentioned here. Tetranychus cinnabarinus
(Boisduval) is a tropical and subtropical pest of cotton and many other plants. One of a
complex of species formerly considered the "two-spotted spider mite," T. cinnabarinus now
I enjoys separate specific status (Boudreaux 1956). T. urticae Koch, the green two-spotted
spider mite, is a widespread major pest of deciduous fruit trees in temperate parts of the
world. The Mcdaniel spider mite, T. mcdanieli McGregor, also is an important fruit tree
I pest, particularly in the Pacific Northwest. The European red mite, Panonychus ulmi
(Koehl. may be found in association with Tetranychus species on deciduous fruit trees.

I P. ulmi is considered a major economic problem in both European and North American
orchards (Groves and Massee 1951). P. citri (McGregor) attacks citrus trees and other
plants (Pritchard and Baker 1955) in the southwestern United States, South Africa and

I Asia. It has been highly injurious in California orchards, particularly in coastal and
intermediate citrus-growing areas (Jeppson 1963).

I The timothy mite, Oligonychus pratensis (Banks), has proven to be an important pest of
grasses and grains in the Pacific Northwest. While species of the pratensis group are common
on grasses, grains, and other low-growing plants, the majority of known Oligonychus species
I are found in trees and shrubs. Members of the ununguis group, for example, are usually
collected on coniferous hosts. O. ununguis (Jacobi), the spruce tree spider mite, is a major

I pest which severely injured over 800,000 acres of fir trees in Montana and Idaho during
1957 (U.S.F.S. 1958). The southern red mite, O. ilicis (McGregor) is injurious on holly
and rhododendron in New York state (Weidhaas and Reeves 1963). Other species of

I Oligonychus feed on deciduous trees (0. gambellii Tuttle and Baker, and O. propetes
Pritchard and Baker, for example), or on shrubs.

I Members of the genus Eotetranychus feed primarily on trees and shrubs, with some
species being considered as economically injurious pests. E. willamettei (McGregor) causes
serious injury on grapes in California, and heavy infestations of E. coryli (Reck) have been
I found on maple in western Oregon. Other species of the genus are important on fruit and
ornamental trees (Pritchard and Baker 1955).

I The false spider mites, or TENUIPALPIDAE, are small, somewhat flattened red or

I
I
158
I
green phytophagous mites which appear to be best adapted to subtropical or tropical
climates. Feeding usually occurs on the leaves of the host (Pritchard and Baker 1958),
I
although some species are collected on bark, in leaf sheaths of grass, or from plant galls.
Members of the genera Brevipalpus and Tenuipalpus are of particular importance as plant
pests.
I
Brevipalpus californicus (Banks) is a cosmopolitan pest of citrus (Baker and Pritchard I
1960). as is B. obovatus Donnadieu. Grapes, ornamentals, ivy and berry crops are some
of the other piant hosts of this genus. Tenuipalpes pacificus Baker is an important feeder
on greenhouse orchids (Geijskes 1939), as is T. granati on grapes (Sayed 1942). The
I
elongate bright red members of the genus Dolichotetranychus are found in grasses (Pritchard
and Baker 1958). I
Larvacarus transitans (Ewing). a gall-forming tenuipalpid, has reduced palpi and only 3
pairs of legs in the adult stage, as is the case in the genus Phytoptipalpus. There is a strong
indication in these reduced forms of a relationship with the elongate, unusual phytophagous
I
mites of the superfamily Eriophyoidea (page 148).
I
The families TUCKERE LLI DAE, ALLOCHAETOPHOR IDAE and L1NOTETRANI DAE
are monogeneric groups containing few species. A species of Tuckerella has been collected
from the roots of plants in California, but other species of this genus are aerial forms. I
Allochaetophora californica McGregor feeds on Bermuda grass, while Linotetranus
cylindricus Berlese, the type species of the genus, was collected in moss. L. protractulus
Athias-Henriot was recovered from soil and litter in Algeria (Athias-Henriot 1961).
I
Useful References
I
Athias-Henriot, C. (1961). Nouveaux acar)ens phytophages d'Algerie (Actinotrichida, Tetranychoidea:
Tetranychidae, Linotetranidael. Ann. Ecole Nat. d'Agr. d'Alger 3(3):1-10.
I
Baker, E.W. and A.E. Pritchard (1953). The family categories of tetranychoid mites, with a review of the
new families Linotetranidae and Tuckerellidae. Ann. Ent. Soc. Amer. 46(2) :243-258.
Baker, E.W. and A.E. Pritchard (1960). The tetranychoid mites of Africa. Hilgardia 29(11):455-574.
I
Blauvelt, W.E. (1945). The internal morphology of the common red spider mite (Tetranvchus telarius
Linn,). Cornell Univ. Agr. Exp. Sta. Mem. 270:3-46.
Boudreaux, H.B. (1956), Revision of the two-spotted spider mite (Acarina, Tetranychidae) complex,
I
Tetranvchus telarius (Linnaeus). Ann. Ent. Soc. Amer. 49(1) :43-48.
Geijskes, D.C. (1939). Beitrage zur Kenntnis dar europaischen Spinnmilben (Acari, Tetranychidae),
mit besonderer Berucksichtigung dar niederlandischen Arten. Meded. Landbouwhoogeschool
I
42(4): 1-68.
Groves, J.R. and A.M. Massee (19511. A synopsis of the world Iiterature on the fru it tree red spider mite
Metatetranvchus ulmi (C.L. Koch, 1835) and its predators. Commonw. Inst. Ent., London: 180 pp.
I
Jeppson, L.R. (1963). Interrelationships of weather and acaricides with citrus mite infestations.
Advances in Acarology 1:9-13. ITETRANYCHIDAEJ
McGregor, E.A. (1950l. Mites of the family Tetranychidae. Amer. MidI. Nat. 44(2):257·420.
I
Morgan, C.V.G. (1960). Anatomical characters distinguishing Bryobia arborea M. & A. and B. praetiosa
Koch (A,arina: Tetranychidae) from various areas of the world. Can. Ent. 92(8):595-604.
Pritchard, A.E. and E.W. Baker (19551. A revision of the spider mite family Tetranychidae. Pac. Coast
I
Ent. Soc. Mem. Ser. 2:472 pp.
Pritchard, A.E. and E.W. Baker (1958). The false spider mites (Acarina: Tenuipalpidael. Univ. Calif.
Publ. Ent. 14(3): 175-274.
I
I
I
I 159

I Sayed, M.T. (1942). Contribution to the knowiedge of the Acarina of Egypt: II. The genus Tenuipalpus
Donnadleu (Tetranychidael. Soc. Fouad 1er Ent. Bull. 26:93·113.

I Tuttle, D.M. and E.W. Baker (1968). Spider mites of the southwestern United States and a revision of the
family Tetranychidae. Un;v. Arizona Press, Tucson: 143 pp. + vii.
U.S.D.A. Forest Service (1958). Forest insect conditions in 1957: a status report. Div. Forest Ins. Res.:

I 20 pp.
Walnstein, B.A. (19601. Tetranychoid mites of Kazakhstan (with revision of the famiiyl Kazakh. Akad.
Sel·sk. Nauk. Nauch.·issled. Inst. Zash. Rast. Trudy 5:1-276.

I Weidhaas, J.A. and R.M. Reeves (1963). The occurrence and importance of tetranychid and eriophyid
mites on woody plants in New York. Advances in Acarology 1:25-29.

I Superfamily Raphignathoidea

I DIAGNOSIS: Soft-bodied, but with various numbers of dorsal shields; cheliceral bases
usually fused, movable cheliceral digit stylet-like or elongate; with or
without a palpal thumb-claw complex. Stigmatal openings at base of
I chelicerae, peritremes well developed__ eyes present or absent. Apoteles
usually present on all legs, tenent hairs present on claws (when claws are

I present).

The eight families currently included in the Raphignathoidea are made up of small

I (350-500 Ill, generally red or reddish-brown species which are predaceous in moss, soil or
lichens but which are also to be found feeding on aerial phytophagous forms in pomaceous,
stone and citrus fruit trees.

I The family POMERANTZII DAE is represented by a single species Pamerantzia charlesi

Baker (1949) which, because of its unusual morphology, is somewhat difficult to include in
I the Raphlgnathoidea (Summers 1966a). The delicately ornamented members af the
family CRYPTOGNATHIDAE comprise a well-defined group, the members of which have
an elongated gnathosoma covered dorsally by a sheath (Summers and Chaudri 1965).
I Cryptognathids are particularly common in leaf mold and moss. An eiongate gnathosoma
alsa is found in the EUPALOPSELLIDAE, but no ornamental gnathosomal sheath is

I present. The eupalopsellid Exatharhis caudata Summers is found in association with citrus
(Summers 1960a), although most eupalopsellids are collected in moss or leaf mold.

I The elongate, rather uncommon species of the family TARSOCHEYLIDAE are

predators in forest iitter in Europe and North America (Atyeo and Baker 1964). One species
of Hoplacheylus, however, was found beneath the elytra of a passalid beetle in the Congo

I (Cooreman 1951).

The family STIGMAEIDAE is a large cosmopolitan group of genera which often are
I characterized by particular combinations of dorsal shields (Summers 1966b). The genus
Ledermuelleria, a strongly sclerotized and ornamented genus, is essentially confined to soil
or litter habitats (Summers and Price 1961) while members of the lightly sci erotized
I genera Zetzellia and Agistemus are commonly found as aerial predators (Summers 1960b,
Gonzalez-R. 1965). Summersiella ancydactyla Gonzalez-R., a predator in leaf cavities in

I New Zealand, is practically unsclerotized (Gonzalez-R. 1967). Zetzellia mali (Ewing) has
been observed to feed on spider mites on both po me and stone fruit trees throughout the
United States and in Europe, and is regarded as a beneficial species.

I
I
160
I
Mites of the family RAPHIGNATHIOAE are often found as predators on and under I
tree bark (Atyeo et al. 1961), One species has been recovered from stored grain both in
New York state and in Oregon, Summers (1966a) includes only the type genus, Raphig-
nathus, in the family,
I
The CALIGONELLIOAE consists of fewer than 20 described species in six genera, the
majority of which are collected in leaf and grass litter (Summers and Schlinger 1955),
I
Molothrognathus fulgidus Summers and Schlinger has been found under the bark of prune
and almond trees in California. An unidentified caligonellid species was collected from
cherry tree twigs in the Pacific Northwest. The stilt-legged NEOPHYLLOBIIOAE, once
I
considered a part of the CALIGONELLIOAE, now constitutes a separate family.
Species of Neophyllobius may be found in leaf mold or on tree bark (McGregor 1950)
where they prey on other arthropods. One species has been recovered from sagebrush
I
litter in the eastern Oregon desert. Camerobia australis Southcott, an Australian inhabi-
tant of eucalyptus bark, considered by Southcott (1956) and Summers (1966a) to I
represent a separate family of Raphignathoidea on the basis of the presence of a camero-
stome, has been included in the NEOPHYLLOBIIOAE in this manual. C, australis has
obvious close affinities with Neophyllobius, making a generic rather than a familial
I
separation seem more realistic,
I
Useful References

Atyeo, W.T. and E.W. Baker (1964), Tarsocheylidae, a new family of prostigmatic mites (Acarina). Bull.
I
Univ. Nebraska State Mus. 4(11):243-256.
Atyeo, W.T" E.W. Baker and D.A. Crossley, Jr. (1961). The genus Raphignathus Duges (Acarina,
Raphignathidae) in the United States with notes on the Old World species. Acar%gia 3(1):14-20.
I
Baker, E.W. (1949), Pomerantziidae, a new family of mites. Jour. Wash. Acad. Sci. 30(8):269·271.
Cooreman, J, (1951), Notes et observations sur les Acariens.
27(1):4-7. [TARSOCHEYLIDAEJ
(IV). Bul/. Inst. Roy. Sci. Nat. Belg. I
Gonzalez-Rodriguez, R.H. (1965). A taxonomic study of the genera Mediolata, Zetzellia and Agistemus
(Acarina: Stigmaeidae). Univ. Calif. Pub/. Ent. 41:1-64.
Gonzalez-Rodriguez, R.H. (1967). Summersiel/a, a new stigmaeid mite from New Zealand (Acarina:
I
Prostigmata). Pan-Pac. Ent. 43(3):236-239. [STIGMAEIDAEJ
McGregor, EA, (1950). Mites of the genus Neophyl/obius. Bull. So. Calif. Acad. Sci. 49(2):55·70. I
Southcott, R.V. (1956). Description of a new Australian raphignathoid mite, with remarks on the
classification of the Trombidiformes (Acarina). Proc. Linn. Soc. N.S.W. 81(3):306-312. [NEO-
PHYLLOBII DAEJ
Summers, F.M. (1960a). Eupalopsis and eupalopsellid mites (Acarina: Stigmaeidae, Eupalopsellidae).
I
Florida Ent. 43(3): 119-138.
Summers, F.M. (1960b). Several stigmaeid mites formerly included in Mediolata redescribed in Zetzellia
Ouds. and Agistemus, new genus (Acarina). Proc. Ent. Soc. Wash. 62(4) :233-247.
I
Summers, F,M. (1966a). Key to families of the Raphignathoidea (Acarina). Acarologia 8(2):226-229.
Summers, F.M. (1966b). Genera of the mite family Stigmaeidae Oudemans (Acarina). Acarologia
8(2) :230-250.
I
Summers, F.M. and W.M. Chaudri (1965). New species of the genus Cryptognathus Kramer (Acarina:
Cryptognathidae). Hilgardia 36(7):313-326 + plates.
Summers, F.M. and D.W. Price (1961). New and redescribed species of Ledermuel/eria from North
I
America (Acarina: Stigmaeidae). Hilgardia 31 (1 0) :369·382 + plates.
Summers, F.M. and E./. Schlinger (1955). Mites of the family Caligonellidae (Acarina).
23(12):539-561.
Hilgardia I
I
I
I 161

I Superfamily Anystoidea

I DIAGNOSIS: Soft-bodied; cheliceral bases free, moving scissors-like over gnathosoma,

movable digit reduced; with a well developed palpal thumb-claw complex.
Stigmatal openings and peritremes distinct at base of chelicerae; eyes
I usually present. Apoteles absent on tarsi I of certain species, with or
without tenent hairs on claws,' with 3, 2 or a pairs of genital discs_

I Both predaceous and parasitic species are found among the four families which make
up the superfamily Anystoidea. The cosmopolitan family ANYSTI DAE is a predatory

I group, as are the less common TENERIFFIIDAE and PSEUDOCHEYLIDAE. The family
PTERYGOSOMIDAE is an exclusively ectoparasitic group.

I Members of the ANYSTI DAE are moderately large (550-1500 11) red fast-moving
mites found on a variety of plants. Species also have been recovered from under boards in
meadows, from leaf litter, bird nests and soil. A species of Tarsotomus is a regular
I inhabitant of farm storage bins in Oregon. Anystis species commonly run on open surfaces
in a whirlagig pattern, and may be difficult to capture.

I Like the ANYSTIDAE, the TENERIFFIIDAE are predaceous and move rapidly when
disturbed. Teneriffiid species have been collected in habitats ranging from intertidal sand

I banks to the undersides of rocks in the Tyrol at altitudes exceeding 4000 feet (Eller and
Strandtmann 1963, Irk 1939). Specimens also have been collected from dry barren sandy
deserts in sagebrush litter in company with other prostigmatid mites. The predaceous

I PSEUDOCHEYLIDAE are elongate, soft mites which may be found under tree bark or in
moss (Tragardh 1906). They are collected only rarely.

I Members of the family PTERYGOSOMIDAE are external parasites of lizards,

attaching under scales or between the toes of the host (Hirst 1925). The broad short
species of the genus Pterygosoma are common on agamid lizards in North and East Africa,
I and in India (Jack 1961). Gekkonid and iguanid lizards are parasitized by species of
Geckobia and Geckobiella (Lawrence 1953, Davidson 1958). Geckobiella texana (Banks)
has been recovered from iguanids of the genus Sceloporus in Texas and in California,
I while Geckobia species are common on geckos of the genus Hemidactylus in southeast
Asia. Specimens of Hirstiella bakeri Cunliffe have been found on Ctenosaura hemilopha in

I the San Diego Zoo, California.

Members of the pterygosomid genus Pimeliaphilus apparently are confined to

I arthropods. Two species attach to the conjunctival tissues of scorpions, where they feed on
the blood of their hosts (Beer 1960, Cunliffe 1949). P. podapolipophagus is an ectopara-
site of cockroaches (Cunliffe 1952), often proving troublesome in laboratory rearing
I operations. Heavy infestations of this mite may cause roach mortality within a few hours.
Several species of triatomine kissing bugs (family Reduviidae) also are attacked by species of
Pimeliaphilus (Newell and Ryckman 1966). The bugs may be killed by the feeding of large
I mite populations.

I
I
I
 162
I
Useful References I
Beer, R.E. (1960). A new species of Pimeliaphilus (Acarina: Pterygosomidae) parasitic on scorpions, with
discussion of its postembryonic development. Jour. Parasit. 46(4) :433-440. I
Cunliffe, F. (1949). Pimeliaphilus isometri, a new scorpion parasite from Manila, P. I. Proc. Ent. Soc.
Wash. 51:123-124. [PTERYGOSOMIDAE]
Cunliffe, F. (1952). Biology of the cockroach parasite, Pimeliaphilus podapolipophagus Tragardh, with a
discussion of the genera Pimeliaphilus and Hirstiella. Proc. Ent. Soc. Wash. 54:153-169. [PTERY-
I
GOSOMIDAE]
Davidson, J.A. (1958). A new species of lizard mite and a generic key to the family Pterygosomidae
(Acarina, Anystoidea). Proc. Ent. Soc. Wash. 60(2):75-78.
I
Eller, R. and R.W. Strandtmann (1963). Notes on Tenerifflidae (Acarina: Prostigmata). Southwest. Nat.
8(1) :23-31,
Hirst, S. (1925). On the parasitic mites of the suborder Prostigmata (Trombidioidea) found on lizards.
I
Jour. Linn. Soc. London Zool. 36:173-200. [PTERYGOSOMIDAE]
Irk, V. (1939). Drei neue Milbenarten aus dem Tiroler Hochgebirge. Zool. Anz. 128(7·8):216-223.
[TENERIFFIIDAE]
I
Jack, K.M. (1961). New species of Near Eastern agamid scale-mites (Acarina, Pterygosomidae) with
notes on the developmental stages of Geckobia hemidactyli Law., 1936. Parasit. 51:241-256.
Lawrence, R.F. (1935), The prostigmatic mites of South African lizards. Parasit. 27(1) :1-45. [PTERY-
I
GOSOMIDAE]
Lawrence, R.F. (1936). The prostigmatic mites of South African lizards. Parasit.28(1):1-39_ [PTERY·
GOSOMIDAE)
I
Lawrence, R.F. (1953). Two new scale-mite parasites of lizards. Proc. U.S. Nat. Mus. 103(3312):9-18.
[PTERYGOSOMIDAEJ
Meyer, M.K.P. and P.A.J. Ryke (1960). Acarina of the families Anystidae, Pseudocheylidae and
I
Cheyletidae (Prostigmata) found associated with plants in South Africa. Jour. Ent. Soc. So. Afr.
23(1):177-193.
Newell, I. and R.E. Ryckman (1966). Species of Pimeliaphilus (Acari: Pterygosomidae) attacking insects,
I
with particular reference to the species parasitizing Triatominae (Hemiptera: Reduviidae). Hilgardia
37(12):403·436.
Oudemans, A.C. (1936). Neues uber Anystidae (Acari). Archiv. f. Naturg. N.F. 5:364-446.
I
Thor, S. (1911). Eine neue Acarinenfamilie (Teneriffiidae) und zwei neue Gattungen, die eine von
Teniriffa, die andre aus Paraguay. Zoo I. Anz. 38: 171-179. I
Superfamily Cheyletoidea I
DIAGNOSIS: Soft-bodied, but often with weak dorsal shields; cheliceral bases fused with
gnathosoma, movable digit stylet- or whip-like; thumb-claw complex
present Or absent. Stigmatal openings, when distinct, at base of chelicerae,
I
peritremes usually well developed; eyes present or absent. Apoteles various,
but generally present; tenent hairs often present on empodia. I
The Cheyletoidea is a rather heterogenous assemblage of nine families, of which only
the CHEYLETIDAE includes non-parasitic species. Parasitic cheyletoids are found on
I
arthropods, reptiles, birds, and mammals-including man.
I
•
The CHEYLETIDAE is a large family of essentially free-living predators (Baker 1949),
although some exceptions should be cited. Cheyletiella parasitivorax (Megnin), Chela-
caropsis moorei Baker and species of Hemicheyletus are found in the fur of mammals

I
I
I 163

I where they apparently subsist as predators of ectoparasitic arthropods in the same habitat
(Lawrence 1954, VOlgin 1960). Smiley (1965) mentioned that Cheyfetieffa yasguri Smiley

I causes a superficial mange on dogs. The same species subsequently was found as a
hyperparasite of dog-infesting louse flies in Iran (Vercammen-Grandjean and Rak 1968).
Cases of eczema in humans as the result of contact with C. parasitivorax have been
I reported by Olsen and Roth (1947). Neocheyfetiella, Ornithocheyla and Cheletosoma
species are found on birds (Baker 1949, Lawrence 1959) and probably feed on parasites.
Bakericheyfa chanayi (Berlese and Trouessart) is a true hematophagous ectoparasite of
I fringillid birds which constructs "nests" of silk fibers on the skin of its hosts (Furman and
Sousa 1969). Free-living cheyletids have been collected from many habitats including

I mite-infested plants (Baker 1949, Yunker 1961, De Leon 1962), in ant and termite nests,
associated with bark beetles or scale insects, in caves, in bird nests and in bat roosts.
Woodroffe (1956) reports that Acaropsis docta Berlese and a species of Chefetonefla were

I observed to feed ectoparasitically on house-martins in England.

Species of Cheyletus, Cheletomorpha, Cheyfetia and Acaropsis often occur in stored

I grain which is infested by astigmatid grain mites (Hughes 1961), upon which they prey.
Cheyfetus eruditus (Schrank) is considered by some observers to be effective in controlling
grain mite populations (Pulpan and Verner 1965), but somewhat seasonal in its effectiveness
I (Solomon 1946). Cheletomorpha fepidopterorum (Shaw) has been observed to consume an
average of 4 grain mites per day under optimum laboratory conditions (20'C and 80% rh).
The rate of predation for other grain inhabiting cheyletids is not known.
I Mites of the family MYOBII DAE are ectoparasites of rodents, marsupials, bats and

I insectivores. One species, Archemyobia inexpectatus Jameson is associated with an

opossum (Jameson 1955). Feeding on the host by myobiids is more or less confined to the
hair follicle bas~s, and hematophagy does not occur under most circumstances. Jameson,

I however, has observed Myobia simplex Ewing to contain blood after feeding on its host
(1948). Myobia musculi (Schrank) and Radfordia ensifera (Poppe), common myobiid
parasites of laboratory mice and rats, cause a mild dermatitis in their rodent hosts (Baker
I et al. 1956), and apparently ingest blood.

Mites of the family HARPYRHYNCHIDAE are parasitic in or under the cuticle of

I birds (Zumpt 1961), and those of the SYR INGOPH I LI DAE usually infest quills of flight
feathers of domestic fowl and wild birds (Zumpt 1961, Clark 1964). Syringophilids feed

I within the quill, inserting the mouthparts into the tissues at the base and feeding on the
seeping fluids. Several species of Syringaphifus, the only genus of the family, are known
from Europe, North America and Africa.

I The family OPHIOPTIDAE includes nearly circular mites which live in small pits in the
scales of snakes. Sambon (1928) reported that Ophioptes parkeri Sambon formed these

I pits in the scales of its colubrid host, a conclusion also reached by Ewing (1933) for O.
tropicalis on another colubrid species. Fain (1964) and others have described additional
species of "pit mites" from various parts of the world. Their feeding habits are unknown.
I Members of the PSORERGATIDAE are minute (ca. 200 /-,) round mites which infest the
surface layers of skin of their mammal hosts, causing a dermatitic condition which may be
aggravated by rubbing or biting by the host. Psorergates avis Womersley, the sheep itch
I mite, is a major pest of sheep in the United States and in Australia (Baker et al. 1956).

I
I
164
I
P. bos Johnston was found to cause symptoms on a Hereford bull in New Mexico which
were similar to those caused by P. ovis on sheep (Johnston 1964). Other psorergatids occur
I
on murid rodents, monkeys and porcupines (Fain et al. 1966, Johnston op. cit.).
I
The minute, annulate, usually worm-like species of the family DEMODICIDAE are
parasitic in the skin of many kinds of mammals. Mange is not uncommon in domestic
animals attacked by Demodex canis Leydig. Demodectic mange in dogs and cats may be I
caused or aggravated by a Staphylococcus which occurs with the demodicid mite (Hirst
1919). D. equi Railliet may cause pruritis in horses, with concomitant loss of hair.
Subsurface skin pustules containing demodicid mites are found in cattle, sheep and goats.
I
Lemurs are affected by species of Stomatodex and Chinodex (Fain 1960), and Demodex
folliculorum Owen is common in the hair follicles of the forehead and eyebrows of man.
D. folliculorum infestations usually cause no noticeable symptoms in man, although
I
dermatitis of the scalp attributed to this mite has been described (Miskjian 1951).

Species of the highly modified family CLOACAR IDAE are parasitic in the mucosa of
I
the cloaca of turtles (Camin et al. 1967, Fain 1968). Cloacarids are thought to be venereally
transmitted from host to host and are highly host specific. Species of the family HETERO- I
CHEYLIDAE are ectoparasites of arthropods (Tragardh 1950). one species being collected
with regularity from beneath the elytra of the passalid Popilius disjunctus in southeastern
United States.
I
Useful References
I
Baker, E.W. (19491. A review of the mites of the family Cheyletidae in the United States National
Museum. Proc. U.S. Nat. Mus. 99(3238):267-320.
I
Baker, E.W., T.M. Evans, D.J. Gould, W.B. Hull and H.L. Keegan (1956). A Manual of Parasitic Mites of
Medical or Economic Importance. Nat. Pest Cont. Assoc. Tech. Pub!.: 170 pp.
Camln, J.H., W.W. Moss, J.H. Oliver, Jr. and G. Singer (1967). Cloacaridae, a new family of cheyletoid
I
I
mites from the cloaca of aquatic turtles (Acari: Acariformes: Eleutherengonal. Jour. Med. Ent.
4(3) :261·272.
Clark, G.M. (1964). The acarine genus Syringophilus in North American birds. Acarologia 6(1):77-92.
[SYRINGOPHI L1DAE]
DeLeon, D. (1962). Three new genera and seven new species of cheyletids (Acarina: Cheyletidael.
Florida Ent. 45(3):129-137.
I
Ewing, H.E. (1933). A new pit-producing mite from the scales of a South American snake. Jour.
Parasit.20(1):53-56. [OPHIOPTIDAE)
Fain, A. (1960). Les acariens psoriques parasites des chauves-souris. XIII. La famille Demodicidae
I
Nicolet. Acarologia 2(1):80·87.
Fain, A. (1964). Les Ophioptidae acariens parasites des ecailles des serpents (Trombidiformes), Ins!. Roy.
Sci. Nat. Belg. 40(15):1-57.
I
Fain, A., F. Lukoschus and P. Hallmann (1966). Le genre Psorergates chez les murides. Description de
trois especes nouvelles (psorergatidae: TrombidiformesL Acarologia 8(2):251·274.
Fritsch, W. (1958). Die Milbengaltung Syringophilus Heller 1880 (Subordo Trombidiformes, Fam.
I
Myobiidae Megnin 1877). Zool. Jahr. 86(3):227-244. [SYRINGOPHILIDAE]
Furman, D.P. and O.E. Sousa (1969). Morphology and biology of a nest-producing mite, Bakericheyla
chanayi (Acarina: Cheyletidae). Ann. Ent. Soc. Amer. 62(4):858-863. I
Hirst, S. (1919). The genus Demodex Owen. 8rit. Mus. (Nat. Hist.) Studies on Acari 1:1-44. [DEMO-
DICIDAE]
I
I
I
I 165
I Hirst, S. (1922). Mites injurious to domestic animals. Brit. Mus. (Nat. Hist.) Econ. Ser. 13:107 pp.
Hughes, A.M. (19611. The mites of stored food. Min. Agr. Fish. and Food Tech. Bull. 9:287 pp. + vi.
I Jameson, E.W. (1948). Myobiid mites (Acarina: Myobiinae) from shrews (Mammalia: Soricidae) of
eastern North America. Jour. Parasit. 34(4):336-342. [MYOBIIDAE]
Jameson, E.W. (1955). A summary of the genera of Myobiidae (Acarina). Jour. Parasit. 41(4):407-416.
I Johnston, D.E. (1964). Psorergates bas, a new mite parasite of domestic cattle (Acari-Psorergatidae).
Ohio Agr. Exp. Sta. Res. Circ. 129:7 pp.

I Lawrence, R.F. (1954). The known African species of Cheyletidae and Pseudocheylidae (Acarina:
Prostigmata). Ann. Natal. Mus. 13:65-77.
Lawrence, R.F. (1959). New mite parasites of African birds (Myobiidae, Cheyletidae). Parasit. 49(3-4):
416-438.
I Miskjian, H.G. (1951). Demodicidosis (Demodex infestation of the scalp). Arch. Derm. Syph. 63:282-283.
[DEMODICIDAE]

I
Norris, J.D. (1958). Observations on the control of mite infestations in stored wheat by Cheyfetus spp.
(Acarina, Cheyletidae). Ann. Appl. BioI. 46(3) :411·422.
Olsen, S.J. and H. Roth (1947). On the mite Cheyletie/la parasitivorax, occurring on cats, as a facultative
parasite of man. Jour. Parasit. 33:444-445. [CHEYLETIDAE]

I Pulpan, J. and P.H. Verner (1965), Control of tyroglyphoid mites in stored grain by the predatory mite
Chevletuseruditus (Schrank). Can. Jour. Zool. 43:417-432.

I
Sambon, L.W. (1928). Ophioptes parkeri. A new species and genus of Cheyletid inhabiting the scales of
reptiles. Ann. Trap. Med. & Parasit. 22(1):137-142.
Smiley, R.L. (1965). Two new species of the genus Chevletiella (Acarina: Cheyletidae). Proc. Ent. Soc.
Wash. 67(2) :75-80.

I Solomon, M.E. (1946). Tyroglyphid mites in stored products.

33(1):82-97. [CHEYLETIDAE]
Ecological studies. Ann. Appl. BioI.

Southcott, R.V. (1956), Notes on the acarine genus Ophioptes, with a description of a new Australian

I species. Trans. Roy. Soc. So. Australia 79:142-147.

Tragardh, l. (1950). Description of a new species of Heterocheylus Lombardini from Africa, with notes
on 'he classification of the Pseudocheyletidae. Ent. Tidsk. 71(2): 104-11 O. [PSEUDOCH EY LI DAE]

I Vercammen-Grandjean, P.H. and H. Rak (19681. Chevletiel/a vasguri Smiley, 1965, un parasite de Canid"s
aux Etats-Unis et hyperparasite d'Hippoboscide en Iran (Acarina: Cheyletidae). Ann. Parasit. Hum.
et Camp. 43(3) :405-412.

I Volgin, V.I. (1960). On the taxonomy of predatory mites of the family Cheyletidae.
Chevletiella Can. Akad. Nauk Zool. Inst. Parasit. Sborn. 19:237-248.
II. Genus

Woodroffe, G.E. (1956). Some insects and mites associated with bat-roosts, with a discussion of the

I feeding habits of the cheyletids (Acarina)_ Ent. Month. Mag. 92: 138-141.
Yunker, C.E. (1961), The genera Sak, new genus, and Cheletomimus Oudemans, with descriptions of
three new species (Acarina: Cheyletidae). Can. Ent. 93(111: 1023-1035.

I Zumpt, F. (ed.) (1961). The Arthropod Parasites of Vertebrates in Africa South of the Sahara (Ethiopian
Region). Vol. I. (Chelicera'a). Publ. So. Afr. Inst. Med. Res. 50(11) :457 pp.

I Superfamily Erythraeoidea

I DIAGNOSIS; Soft-bodied,' adults and nymphs hypertrichous, with sensory setae in an

anterodorsal crista metopicai movable cheliceral digit long, needle-like,
retractile in cheliceral sheathi thumb-claw complex well developed. Stig-
I matal openings between cheliceral basesi 1-2 pairs of eyes. With apoteles on
all legs, empodia absenti with 2 or a
(rarely 1J pairs of genital discs.

I Larvae heteromorphic, with anal opening present or absenc coxae 1·11

contiguous or separate.

I
I
166
I
The three families of erythraeoid mites, like other members of the Parasitengona, are I
parasitic in the larval stage and predaceous in the nymphal and adult phases. The larvae of
all three famiiies are heteromorphic and parasitize arthropods, with few exceptions.
I
The ERYTHRAEIDAE is a cosmopolitan family of red or reddish-brown species which
may be found in moist or dry habitats in the nymphal and adult stages as predators of
small arthropods. Erythraeid larvae are for the most part parasitic on a variety of insects
I
and other arthropods (Southcott 1961 a). However, larvae of the genus Balaustium have
been observed to feed on pollen (Grandjean 1947), a habit which may be shared by larval
forms of the genus Microsmaris. Larvae of Caeculisoma (Southcott 1961b), Erythraeus
I
(Lawrence 1940) and Charletonia (Southcott 1965) are common parasites of locusts,
while species of Balaustium are found on aphids and on hemipteran insects (Womersley I
1934). Rainbowiaimperator (Hirst 1928) and certain Erythroides species (Southeott 1946)
parasitize Homoptera in Australia. Larvae of the widespread genus Leptus have been found
on spiders (Lawrence 1940), dragonflies, scorpions, butterflies and moths, phalangids and I
dipterans. One record exists of a Leptus species having been reared on a lizard in the South
Pacific area. Another species has been found in great numbers in fescue grass plantings
infested by the thrips Aptinothrips rufus in Oregon. Several thrips were found to be
I
parasitized by the larval mites.

Adult and nymphal erythraeids occur on plants, in humus, or on open ground where
I
they prey on other mites or on small insects. Balaustium sp. was collected from balsam fir
infested with aphids, and may have been feeding on these insects. Ebeling (1934) reports
B. aonidaphagus (Ebeling) as being predaceous on citrus red scale in California. Newell
I
(1963) reports three cases of a Balaustium species attacking man, and an instance of a
Balaustium larva feeding on green leaves of a corn-lily. This would seem to indicate that I
Balaustium spp. are general feeders capable of phytophagy or hematophagy as well as
insect parasitism and predation. Myrmicotrombium brevicristatum Womersley has been
found with ants, but its relationship to the ants is obscure (Southcott 1957).
I
The larvae of SMARIDIIDAE are poorly known, with few observations having been
made on host preferences. Womersley and Southcott (1941) record a species of psocid as
I
the host for the larva of Smaris prominens, but the larvae of Sphaerotarsus leptopilus
Womersley, another smaridiid, refused psocids and other insects offered to them in the
laboratory (Southcott 1960). Adult smaridiids are wide-ranging predators with essentially
I
the same habits as adult erythraeids.

The CAL YPTOSTOMIDAE is a small family of large mites (1500-2500 f.J) which are
I
found on wet stream banks or in moss in Europe (Turk 1945), Japan (Shiba 1969), Africa
and North America. Additional collecting in semi·aquatic habitats should result in I
recovery of these rather rare mites from other parts of the world. Larvae of Calyptostoma
velutinus (Muller) are reported as parasites of tipulid flies in Ehime Prefecture, Japan
(Shiba 1969).
I
Useful References
I
Ebeling, W. (1934). A new predaceous mite from southern California.
[ERYTHRAEIDAEJ
Pan·Pac. Ent. 10(1) :33·34. I
I
I
I 167

I
•
Grandjean, F. (1947). Etude sur les Smarldiidae et quelques autres Erythroides (Acariens). Arch. de Zool.
Expt. Gen. 85(1):1·126 .
Hirst. S. (1928). On some new Australian mites of the families Trombidiidae and Erythraeidae. Ann. Mag.
Nat. Hist. 10, 1(4):563·571.
Lawrence, R.F. (1940). New larvai forms of South African mites from arthropod hosts. Ann. Natal

I Mus. 9(3):401-408. [ERYTHRAEIDAE]

Newell, i.M. (1963). Feeding habits in the genus Balaustium (Acarina, Erythraeidae), with speciai
reference to attacks on man. Jour. Parasit. 49(3):498-502.

I Shiba, M. (1969). Taxonomic investigations on free-living mites in the subalpine forest of Shiga Heights
IBP Area. ii. Prostigmata. Buli. Nat. Sci. Mus. 12(11:65-115. [CALYPTOSTOMIDAE]
Southcott. R.V. (1946). Studies on Australian Erythraeidae (Acarina). Proc. Linn. Soc. N. S. W. 71(1·2):

I 6·48.
Southcott, R.V. (1957). The genus Myrmicotrombium Womersley 1934 (Acarina, Erythraeidae), with
remarks on the systematics of the Erythraeoidea and Trombidioidea. Rec. So. Austral. Mus.
13(1) :91-98.
I Southcott. R.V. (1960). Notes on the genus Sphaerotarsus (Acarina, Smaridiidae). Trans. Roy. Soc. So.
Austrai. 83:149·161.
Southcott, R.V. (1961a). Studies on the systematics of the Erythraeoidea (Acarina), with a critical
I revision of the genera and subfamilies. Austral. Jour. Zool. 9(3) :367·61 O.
Southcott, R.V. (1961b), Notes on the genus Caeculisoma (Acarina: Erythraeidae) with comments on the
biology of the Erythraeoidea. Trans. Roy. Soc. So. Austral. 84: 163·178.
I Southcott. R.V. (1963), The Smaridiidae (Acarina) of North and Central America and some other
countries. Trans. Roy. Soc. So. Austral. 86:159-245.
Southcott, R.V. (1965). Revision of the genus Charletonia Oudemans (Acarina: Erythraeidae), Austral.
I Jour. Zool. 14:687-819.
Turk, F.A. (1945). Studies of Acari, V. - Notes on and descriptions of new and little·known British Acari.
Ann. Mag. Nat. Hist. 12(111:785·820. [CALYPTOSTOMIDAE]

I Womersley, H. (1934). A revision of the trombid and erythraeid mites of Australia with descriptions of
new genera and species. Rec. So. Austral. Mus. 5(2) :179·254.
Womersley, H. and R.V. Southcott (19411. Notes on the Smaridiidae (Acarina) of Australia and New
I Zealand. Trans. Roy. Soc. So. Austral. 65(1) :61·78.

I Superfamily Trombidioidea

DIAGNOSIS: Soft-bodied, adults and nymphs hypertrichous, with 1-2 pairs of sensory
I setae associated with anterodorsal crista metopica; movable cheliceral digit
short, hook-like, non-retractible; thumb-claw complex well developed.

I Stigmatal openings between cheliceral bases; eyes present. With apoteles on

all legs, empodia present or absent; with or without genital discs. Larvae
heteromorphic, with anal opening; coxae I-II usually contiguous.
I Three families comprise the Trombidioidea and, like the Erythraeoidea, all are
parasitic in the larval stage and predaceous on other arthropods as nymphs and adults.
I The TROMBIDIIDAE is a large and complex group of mites, the larvae of which
parasitize insects. The larvae may be found attached to species of Orthoptera, Lepidoptera,
I Hemiptera and Diptera (Thor and Willmann 1947), among others. Larvae of Allothrombium
aphidis (DeGeer) and Microtrombidium sp. have been recovered from aphid species in

I northwestern United States, and A. neapolitanum Oud. was collected from a phalangid in
Italy. Paratrombium egregium Bray. is parasitic on a spider wasp of the genus Pompilus in

I
I
168
I
France, while Hoplothrombium quinquescutatum Ewing was found attached to a cryptostig-
matid mite in the stomach of a North American toadl (Thor and Willmann 1947, Ewing
I
1925). Larvae of Eutrombidium rostratus (Scopoli) are parasites of various Orthoptera
throughout most of the world. The life cycle of this mite has been studied by Severin
(1944).
I
Larvae of the trombidiid genera Dinothrombium and Angelothrombium commonly
feed on locusts, and the adults and nymphs prey on termites (Newell and Tevis 1960). The
I
adults of these two genera are referred to as giant red velvet mites on the basis of their
great size (2,500-13,000 Ill, their color and the extreme hypertrichy of body setae I
exhibited in all species. Newell and Tevis tell of an unusual eruption of what must have
been one or more species of Dinothrombium discovered from the air near Tucson, Arizona,
following a heavy rainstorm. The 5-acre eruption from the interstices of the soil was seen
I
from 1500' altitude as a bright red bloom. Another species of Dinothrombium sent to
Oregon State University was collected in India in great numbers under virtually identical
circumstances.
I
Adults and nymphs of the family TROMBICULIDAE prey on small arthropods, and
are most commonly taken in soil. One species is a commensal of termites in East Africa
I
(Vercammen-Grandjean 1965a). Comparatively few species of trombiculids are known from
the postlarval stages (Thor and Willmann 1947) but the larvae, or chiggers, have been
studied both extensively and intensively by many acarologists. Trombiculid chiggers are
I
parasitic on vertebrates-with few exceptions-and may cause injury through their bite or
through the transmission of disease organisms. Their importance as parasites of man and I
animals helps to account for the enormous bibliography of literature on the subject (see
bibliographies in Fuller 1952, Bakeretal. 1956, Andre 1965, Audy 1968).
I
Every major group of vertebrates is attacked by trombiculid larvae, and the results of
such attacks often create a distinctive symptomatology on the host. The formation of a
feeding tube, or stylostome (Fig. 23) at the site of chigger attachment is characteristic of
I
chigger attack (Cross 1963). Dermatitis caused by
trombiculid mites, or trombidiosis, is a symptom
commonly observed in man as the result of
; ,
;1
I
attacks by chiggers of the groups Trombicula,
Eutrombicula, Acomatacarus, Sch6ngastia,
;:
stylostome----i--J'j: I
Eusch6ngastia and Apolonia (Baker et al. 1956).
Trombicula (Neotrombicula) autumnalis (Shawl,
the harvest mite of Europe, is particularly annoying I
to man but also causes dermatitic symptoms in
domestic animals, rodents and birds. Species of the
subgenus Eutrombicula, genus Trombicula, cause
I
trombidiosis in man in the Western Hemisphere.
These mites also attack other mammals, as well as
birds, reptiles and amphibians. Eutrombicula palp
I
(Eutrombicula) a/freddugesi (Oud.) is the most
common and troublesome chigger in North America
(Williams 1946L while members of the genera
chelicera
I
Fig. 23. Feeding tube, or stylostome,
Acomatacarus and Sch6ngastia are common pests
of man in the Orient and Australasia (Womersley
of a trombiculid larva attached
to the gnathosoma. I
1952, Womersiey and Heaslip 1943).
I
I
I 169

I
•
Apolonia tigipioensis Torres and Braga attack chickens in Brazil, often causing death of
young birds (Torres and Braga 1939). Attacks by this mite on humans also have been
recorded.

Species of Trombicula (Leptotrombidium) are commonly found on rodents in the

I Orient and Australasia, but attack man and other animals with regularity. Three of these
species, T. (Leptotrombidium) de/iense Walch, scute//aris Nagayo et al. and akamushi
(Brumptl serve as the vectors of tsutsugamushi fever, or scrub typhus (Audy 1968). Scrub
I typhus is an infectious disease of the Asiatic-Pacific region which proved to be a major
medical problem for both the Allied and Japanese military in the Pacific theatre of

I operations during World War II.

Some chiggers have been found living in the nasal passages of their animal hosts. For

I example, members of the genus D%isia live as intranasal parasites of bats (Yunker and
Brennan 19621, and Vatacarus species occur in the nasal fossae, trachea or lungs of marine
iguanids and sea snakes (Southcott 1957, Vercammen-Grandjean 1965b). Reptiles and
I amphibians often are found to have sub-surface skin infestations of chiggers of the genera
Hannemania and Endotrombicula. H. hylae (Ewingl is a subcutaneous parasite oftreefrogs
in North America (Ewing 1926). A species of Hannemania recently was found in the lip
I tissue of a hylid frog in Oregon.

I The family JOHNSTONIANIDAE is similar to the TROMBIDIIDAE in habits, being

parasitic on insects as larvae, and predaceous as adults. Johnstonianid larvae are particularly
abundant on insects found living in aquatic and subaquatic situations (Newell 1957). Adults

I have been collected from many habitats, but might well be considered as subaquatic, since
they are usually found near water. Newell postulates that, on the basis of morphology and
habitat, the JOHNSTONIANIDAE may have provided the evolutionary stepping-stones

I both toward strictly terrestrial trombidioids, and toward the aquatic mites of the super-
cohort Hydrachnellae.

I Useful References

I Andre, M. (ed.1 (19651. Acarologia 7 (fasc. supplementaire): 335 pp. [TROMBICULIDAE]

Audy, J.R. (19681. Red mites and Typhus. Univ. London. Athlone Press: 191 pp. + x. [TROMBI-

I
CULIDAE]
Baker. E.W .. T.M. Evans. D.J. Gould, W.B. Hull and H.L. Keegan (19561. A Manual of Parasitic Mites of
Medical or Economic Importance. Nat1. Pest Control Assoc. Tech. Pub!.: 170 pp.

I Brennan, J.M. and E.K. Jones (1959). Keys to the chiggers of North America with synonymic notes and
descriptions of two new genera (Acarina: Trombiculidae). Ann. Ent. Soc. Amer. 52(1):7-16.
Cross, H.F. (1964). Observations on the formation of the feeding tube by Trombicula splendens larvae.
Acarologia 6 (fase. h.s.l: 255-261.
I Crossley, D.A. (1960). Comparative external morphology and taxonomy of nymphs of the Trombiculidae
(Acarinal. Univ. Kansas Sci. Bull. 40(6): 135-321.

I Ewing, H.E. (1925). A contribution to our knowledge of the taxonomy of chiggers (Trombidiidae),
including the descriptions of a new genus, six new species and a new variety. Amer. Jour. Trap.
Med.5:251·265. [TROMBICULIDAE]
Ewing. H.E. (19261. The life history and biology of the Tree-Toad Chigger. Trombicula hylae Ewing.
I Ann. Enl. Soc. Amer.19:261-267.
Feider, Z. {19551. Fauna Repub\icii. Populare Romine. Arachnida 5(1). Acarina; Trombidioidea: 186 pp.

I
I
170
I
I
•
Fuller, H.S. (1952). The mite larvae of the family Trombiculidae in the Oudemans collection: taxonomy
and medical importance. Zoo I. Verhandel. 18:261 pp.
Newell, I.M. (1957). Studies on the Johnstonianidae (Acari, Parasitengona). Pac. Sci. 11:396·466.
Newell, I.M. and L. Tevis, Jr. (1960), Angelothrombium pandorae n. g., n. sp. (Acari, Trombidiidae), and
notes on the biology of the giant red velvet mites. Ann. Ent. Soc, Amer. 53(3):293-304.
Severin, H.C. (1944). The grasshopper mite Eutrombidium trigonum (Hermann) an important enemy of
grasshoppers, So. Dakota Agr. Exp. Sta. Tech, Bull. 3:36 pp. [TROMBIDIIDAE] I
Southcotl, R,V, (1957). On Vatacarus ipoides n. gen., n. sp. (Acarina: Trombidioidea). Trans. Roy. Soc.
So, Austral. 80:165-176. [TROMBICULIDAE]
Thor, S. and C. Willman (1947). Trombidiidae. Das Tierreich 71b:187-541.
I
Torres, S. and W, Braga (1939). Apolonia tigipioensis g. et sp. n. (TrombiculinaeJ parasito de Gallus
gallus domesticus. Bolo Soc. Brasil. Med. Vet. 9:28-34.
Vercammen-Grandjean, P.H. (1965a). Tenotrombicula mintneri n. g" n. sp" an interesting commensal of
I
African termites (Trombiculidae: Acarina). Acarologia 7 (fasc. suppl.): 259-265.
Vercammen-Grandjean, P.H. (1965b). Iguanacarus, a new subgenus of chigger mite from nasal fossae of
the marine iguana in the Galapagos Islands, with a revision of the genus Vatacarus Southcott (Acarina,
Trombiculidae). Acarologia 7 (fasc, suppl.): 266-274.
I
Vercammen-Grandjean, P.H. (1968). Chigger mites of the Far East (Acarina: Trombiculidae & Leeuwen-
hoekiidae), U.S. Army Med. Res. Dev. Command, Washington, D.C. Spec. Study: 135 pp. I
Wharton, G.W, and H.S. Fuller (1952), A manual of the chiggers. Mem: Ent. Soc. Wash. 4:185 pp.
Williams, R.W. (1946). A contribution to our knowledge of the bionomics of the common North
American chigger, Eutrombicula alfreddugesi (Oudemans) with a description of a rapid collecting
method, Amer. Jour. Trap, Med. 26:243-250.
I
Womersley, H. (1952). The scrub-typhus and scrub-itch mites (Trombiculidae, Acarina) of the Asiatic-
Pacific region. Rec. So. Austral. Mus. 10:1-673. I
Womersley, H. and W.G. Heaslip (1943). The Trombiculinae (Acarina) or itch-mites of the Austro-
Malayan and Oriental regions. Trans. Roy. Soc, So. Austral. 67:68-142.
Yunker, C.E. and J.M. Brennan (1962). Endoparasitic chiggers: II. Rediscovery of Doloisia synoti
Oudemans, 1910, with descriptions of a new subgenus and two new species (Acarina: Trombiculidae).
I
Acarologia 4(4) :570·576.
I
Superfamily Hydrovolzioidea
I
DIAGNOSIS: Aquatic; dorsum with series of weakly sclerotized shields (an anterior,
posterior, and a pair of laterals); swimming hairs absent on legs, leg axes not
parallel; weak swimmers. Larvae heteromorphic, with a propodosomal shield
I
flanked by a pair of club-shaped setae; chelicerae in dorso-ventral plane.

A single family, the HYDROVOLZIIDAE, is contained in the Hydrovolzioidea.

I
Hydrovolziids are known primarily from the Asiatic-Pacific region and from Europe, but
species have been identified from North Africa and North America (Mitchell 1954). The
larvae of Hydrovolzia gerhardi Mitchell, for example, parasitize terminal nymphs and
I
adults of the water bug Microvelia americana in Illinois. The nymphal and adult mites are
active predators, H. placophora Monti and H. cancellata Walter are cold water species
usually found at high altitudes or latitudes in Europe (Monti 1905, Walter 1906) and are
I
considered to be glacial relicts.
I
I
I
I
I 171

I Useful Referenoes

I Mitohell, R,D. (1954). A desoription of a water·mite. Hydrovolzia gerhardi new species, with observations
on the life history and eoology. Nat. Hist. Misc. (Chi. Acad. Sel.l 134:1-9
Monti, R. (19051. Genre e specie nouvi di Idrachnidae. Rend. Istit. Lomb. ScI. e Lett. 2, 38:168·176.

I Waiter, C. (1906), Neue Hydrachnidenarten aus der Schweiz. Zoo I. Anz. 30:570-575.

I Superfamily Hydryphantoidea

DIAGNOSIS: Aquatic; usually without distinctive dorsal shields; swimming hairs on legs
I present or absent; leg axes not parallel; weak swimmers. Larvae hetero-
morphic, with median dorsal sc/erite armed with sensory setae; chelicerae

I in typical longitudinal plane; palpi raptorial.

The Hydryphantoidea includes water mites of several families, all of which are

I parasitic on insects in the larval stage, and which show strong morphological affinities in the
larval stage to the Trombidioidea (page 167) (Mitchell 1957, Imamura and Mitchell 1967).
The majority of known larval forms parasitize adult Diptera, although larvae of the genera
I Eylais and Piersigia (families EYLAIDAE and PIERSIGIIDAE) attack aquatic beetles or
bugs. The parasitic habit has been suppressed or lost in some families. Hydryphantoid
nymphs and adults usually are non-swimming or weak-swimming large mites which prey on
I aquatic or semi-aquatic insects. Species of Limnochares (family L1MNOCHARIDAE), Eylais
and Hydryphantes (family HYDRYPHANTIDAE) are commonly found in standing water.
Piersigia limophila Protz is found in rotting vegetation of marshes and swamps (I mamura and
I Mitchell 1967), and may represent an early step toward a truly aquatic environment from a
terrestrial trombidiform progenitor.

I Useful References

I Imamura, T. and R.D. Mitchell (1967). The ecology and life cycle of the water mite, Piersigia limophila
Protz. Ann. Zoo I. Japon. 40(1):37-44. [PIERSIGIIDAE]

I Mitchell, R.D. (1955).

[PIERSiGliDAE]
Two water-mites from Illinois. Trans. Amer. Micro. Soc. 74(4):333-342.

Mitchell, R.D. (1957). Locomotor adaptations of the family Hydryphantidae. Abh. naturw. Ver.

I Bremen 35(1):75-100.
Soar, C.D. and W. Williamson (19251. The British Hydracarina. Ray Society, London 1:216 pp. + x +
plates.

I Stout, V. (1953). Eylais waikawae n. sp. (Hydracarina) and some features of its life history and anatomy.
Trans. Roy. Soc. N. Zeal. 81:389-416. [EYLAIDAE]

I Superfamily Hydrachnoidea

I DIAGNOSIS: Aquatic; soft-bodied, globular, red mites with papillate integument; leg axes
more or less parallel, swimming hairs present on legs, indifferent swimmers.
Larvae heteromorphic, with greatly enlarged mouthparts nearly equalling
I size of idiosoma; palpi raptorial.

I
I
172
I
I
•
A single family, the HYDRACHNIDAE, is included in the Hydrachnoidea. The
hydrachnids are considered primitive forms (Mitchell 1957) whose larvae are parasitic on
nymphal or adult hemipterans, or on adult beetles. The larvae are weak swimmers, but tend
to be found in the upper levels of their aquatic habitat. The feeding habits of the adults are
not clear, although repeated observations of Hvdrachna attaching to plants has led some
workers to conclude that they may be herbivorous. The HYDRACHNIDAE is virtually
cosmopolitan in distribution.
I
Hydrachnid larvae, like those of the hydryphantoid family EYLAIDAE, remain
attached to their insect host until the nymphal development is completed. This nvmphophan
I
stage is greatly enlarged due to the engorgement of the larva, and resembles an elongate
pouch (Fig. 10, p. 47). I
Useful References
I
Mashuda, Y. (19341. Notes on the life history of Hydrachna (Schizohydrachna) nova Marshall. Jour. Sci.
Hiroshima Univ. Ser. B., Div. 1,3:33-43.
I
Mitchell, R.D. (19571. Major evolutionary lines in water mites. Syst. Zool. 6(31:137-148.
Soar. C.D. and W. Williamson (1925). The British Hydracarina. Ray Society, London 1:216 pp. + x
+ plates.
I
Stout, V. (1953). New species of Hydracarina, with a description of two. Trans. Roy. Soc. N. Zeal.
81 :417-466.
I
Superfamily Hygrobatoidea I
DIA GNOSIS: Aquatic; with or without distinctive sclerotization; leg axes more or less
parallel, swimming hairs present or absent, generallv strong swimmers.
Larvae heteromorphic (or suppressed), assuming a variety of shapes and
I
ornamentation; palpi raptorial.

The Hygrobatoidea includes the great majority of the water mite families. Aside from
I
one questionable case in the family AXONOPSIDAE (Viets 1935). hygrobatoid larvae are
either non-parasitic (the UNIONICOLIDAE, for example) or suppressed, so that the first I
active stage in ontogeny is the nymph (e.g., the PIONIDAE) (Lundblad 1927). In the
UNIONICOLIDAE, the nymphal and adult stages are parasitic rather than free-living forms,
and the host range is restricted to molluscs and sponges.
I
Larvae of many hygrobatoid families are found at all depths in a variety of aquatic
situations. Since many have lost parasitic associations, it is not necessary for them to
I
contact aerial insect hosts on the water surface. Suppression of the parasitic habit in these
forms has permitted a greater degree of ecological and morphological diversity in the
Hygrobatoidea than in any other water mite assemblage.
I
I
I
I
I
I 173

I Useful References

I Cassagne-Mejean, F. (19661. Contribution a I'etude des Arrenuridae (Acari, Hydrachnellae) de France.

Acarologia 8, fasc. suppl.: 186 pp.
Lundblad, O. (19271. Die Hydracarinen Schwedens I. Zool. Bidr. Uppsala 11:185-540.

I Mitchell, R.D. (19551. Anatomy, iife history and evoiution of the mites parasitizing mussels.
Michigan Misc. Publ. Zool. 89:27 pp. + plates. [UNIONICOLIDAE]
Univ.

Mitchell, R.D. (19571. Major evoiutionary lines in water mites. Syst. ZooI. 6(31:137-148.

I Munchberg, P. (1954). Zur Kenntnis der an Culiciden schmarotzenden Arrenurus-Larven. Zeit. fur
Parasit.16:298-312. [ARRENURiDAE)
Soar, C.D. and W. Williamson (19271. The British Hydracarina. Ray Society, London 2:215 pp. + viii
I + plates.
Soar, C.D. and W. Williamson (19291. The British Hydracarina. Ray Society. London 3:184 pp. + viii
+ plates.
I Stout, V. (1953). New species of Hydracarina, with a description of two. Trans. Roy. Soc. N. Zeal.
81:417-466. [PiONIDAEj
Viets, K. (1935). Die Wassermilben von Sumatra, Java und Bali nach den Ergebnissen der Deutschen
I Limnologischen Sunda-Expedition. Arch. Hydrobiol. Suppl. Bd. 13:595·738.

I
I
I
I
I
I
I
I
I
I
I
I
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174
I
Suborder Prostigmata I
(Plates 55 to 97, pp. 183-225)

KEYTOTHE FAMILIES
I
(Supercohort and cohort designations are used only where such designations are felt to
I
lend clarity to the key.)
I
1. Found in fresh water (one family, PONTARACHNIDAE, is found in brackish water),
often capable of swimming Supercohort HYDRACHNELLAE
Rarely found in fresh water; non-swimming forms primarily confined to brackish or
60
I
salt water (HALACAR IDAE), free-living on land, or parasites of animals or plants.. 2

2. Body annulate, usually elongate and worm-like (Plate 55-1) .

I
· Supercohort PROMATA (pars) '" 3
Body more or less rounded, not worm-like or annulate. . . . . . . . . . . . . . . . . . . . . .. 5
I
3. With four pairs of legs
With two pairs of legs; with a transverse genital aperture located behind the
, 4 I
second pair of legs. Plant feeders
Superfamily ERIOPHYOIDEA,
Cohort TETRAPODI L1NA,
(Plate 55 [see also 10·6]) Family ERIOPHYIDAE 1 I
4. Female genital aperture between legs IV; genital discs absent. Body and leg
setation absent. Leg segments telescoped. Skin parasites . I
· , Cohort ELEUTHEROGONINA (pars), Superfamily CHEYLETOIDEA (pars),
· (Plate 56) Family DEMODICIDAE
Female genital aperture behind legs IV; with three pairs of genital discs. Free·living
I
in soil or arenicolous Cohort EUPODOSTIGMATINA (pars),
" .. , ., ... " .... Superfamily TYDEOIDEA, (Plate 56) Family NEMATALYCIDAE I
5. Gnathosoma tubular, oriented vertically; palpi 1-segmented and fang-like, without
discernible chelicerae. From cloaca of aquatic turtles . I
· Cohort ELEUTHEROGONINA (pars), Superfamily CHEYLETOIDEA (pars),
· (Plates 56, 57) Family CLOACARIDAE
Gnathosoma not as above; palpi developed as segmented appendages, not fang-like;
I
chelicerae present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 6

6. Gnathosoma usually circular in outline, with minute palpi lying closely appressed
I
laterally; chelicerae tiny, stylet-like (Plate 59-4). With four pairs of legs, or less;
empodium of tarsi II-III commonly a broad membranous pad. Stigmata of female I
opening on propodosoma, behind (and lateral to) gnathosomal base (Plate 58-1);
males without stigmata or tracheae ... " ..... Supercohort HETEROSTIGMATA,
.................. Cohort TARSONEMINA, Superfamily TARSONEMOIDEA ... 7
I
l The Eriophyoidea is considered by Keifer (1964) to comprise three families: the PHYTOPTIDAE.
the RHYNCAPHYTOPTIDAE and the ERIOPHYIDAE.
I
I
I
I 175
I Gnathosoma variously shaped, generally conspicuous and with distinct chelicerae;
palpi usually well developed. Rarely with fewer than four pairs of legs; empodia
I highly variable. Stigmata opening at base of chelicerae 11

7. Both males and females with four pairs of legs; pseudostigmata often present ..... " 8
I Females with one to three pairs of legs; males with three (seldom four) pairs of legs;
pseudostigmata absent. Associated with insects .

I · (Fig. 22, p. 147) Family PODAPOLIPIDAE

8. Podonotal shield forming a broad roof-like cover over gnathosoma. Leg IV ending
I in many whip-like setae which may exceed more proximal setae by twice their
length; tarsi IV may be long and attenuate, bearing claws. Free-living or associated
with insects , (Plate 57) Family SCUTACARIDAE
I Podonotal shield forming a broad roof-like gnathosomal cover only rarely (I.e., the
genus Glyphidomastax). Setae of tarsi IV typically subequal in length to more

I proximal setae, ending in two whip-like setae, or none 9

9. Legs IV of female with claws and membranous empodia; without terminal whip-like
I setae
Legs IV of female without claws or empodia; with two terminal whip-like setae.
10

I Plant feeders or insect associates (Plate 58) Family TARSON EMIDAE

10. Idiosoma of female elongate or oval (ovoid or terminally rounded in gravid individ-

I uals). Tarsi II-IV each with two claws and a membranous empodium, or with a
sucker-like empodium and no claws; tarsus I usually with a single sessile claw (an
exception is the genus Dolichocybel. Associated with insects, or found in plant
I materials " ., , " . (Plate 59 [see also 5-3] 1 Family PYEMOTIDAE
Idiosoma of gravid female lobed laterally, more or less pointed posteriorly. With

I two sessile claws on tarsi I-IV, empodia absent. Associated with insects
· (Plate 60) Family CROTALOMORPHIDAE ms.
.

I 11. With a distinct palpal thumb-claw process (Fig. 20, p. 142)

Without a palpal thumb-claw process, or complex indistinct
38
12

I 12. Gnathosoma elongate; palpi slender, attenuate, with palpal tarsus usually terminal;
thumb-claw, when present, is obsolete; coxae II-II I well separated. Free living .....
· Superfamily RAPHIGNATHOIDEA,
I · (Plate 88 [see also 7-61) Family EUPALOPSELLIDAE
Without above combination of characters 13
I 13. Gnathosoma enclosed within a sheath formed by an anterior extension of the
propodosoma. With two pairs of genital discs. Free-living .
I · ..... Superfamily RAPHIGNATHOIDEA, (Plate 60) Family CRYPTOGNATHIDAE

•
Gnathosoma not enclosed as above. With 0-3 pairs of genital discs 14

I
•
176
I
14. Primarily marine, estuarine or arenicolous species; 3-4 palpal segments; with four I
dorsal and four ventral shields. Predaceous, phytophagous, or parasitic forms
· .. Superfamily HALACAROIDEA, (Plate 61 [see also 4-3]) Family HALACARIDAE
Terrestrial species; with variable numbers of palpal segments and idiosomal shields;
I
15.
discrete ventral shields often absent. 15

Rod-like sensory setae of tarsus I lying flush with tarsus in a specialized membranous
I
depression (Plate 61-1). Anterior portion of propodosoma with a tubercle bearing
a pair of setae Cohort EUPODOSTIGMATINA (pars). I
· Superfamily EUPODOIDEA 16
Rod-like sensory setae of tarsus I, when apparent, erect, arising from a small
circular membranous base, never within a specialized membranous depression (Plate
I
65-2); anterior tubercle may be present or absent

16. With small and sometimes distorted cheliceral shears (Plate 62-3)
19

17
I
With large, opposed cheliceral shears. Free-living ... (Plate 61) Family RHAGIDIIDAE I
17. Soft bodied, with or without roof-like projection over gnathosoma. Free-living or
plant feeders 18
Body sclerotized in a tuberculate pattern, and with a roof-like extension over the
I
gnathosoma. Idiosoma often with a "V" or "Y" suture dorsally. Free-living
· (Plate 62) Family PENTHALODIDAE I
18. Anal aperture ventral; femur IV often greatly enlarged. Free-living predators and
fungivores (Plates 62, 63) Family EUPODIDAE
I
Anal aperture dorsal or terminal; femur IV not enlarged. Plant feeders
·
.
(Plate 63) Family PENTHALEIDAE I
19. Chelicerae whip-like, long.
or insect parasites
Genital aperture transverse (Plate 64-3). Plant feeders
20 I
Chelicerae either opposed, stylet- or scissors-like (if appearing whip-like, then the
genital aperture is longitudinal). Free-living or animal parasites (some members
of the family TYDEIDAE may be plant feeders) 21
I
20. Chelicerae arising from an eversible stylophore (Plate 4-7). Tarsal claws with tenent
hairs; adults occasionally with only three pairs of legs (genus Phytoptipalpus).
I
Plant feeders Superfamily TETRANYCHOIDEA,
I
•
· (Plates 63, 64) Family TENUIPALPIDAE
Chelicerae not arising from an eversible stylophore. Without claws on tarsi I; tarsal
claws II-IV without tenent hairs; adults always with full complement of legs.
Parasites of cockroaches... Superfamily IOLINOIDEA, (Plate 64) FamilY IOLINIDAE

21. Cheliceral bases fused, or if not fused, not capable of a lateral scissors-like motion
over gnathosoma .- 22
I
I"
I
I;
I
177
I Cheliceral bases not fused, the chelicerae moving scissors-like over gnathosoma (Plate

I 65-1); anterior portion of gnathosoma produced into a long "snout." With two
pairs of long sensory setae, or sensilla, on the propodosoma. Free-living
· Superfamily BDELLOIDEA 23
.

I 22. Chelicerae and rostrum fused into a cone. Parasites of vertebrates or arthropods
· Superfamily CHEYLETOIDEA (pars) .., 24
I Chelicerae and rostrum not fused into a cone. Free-living 28

I 23. With two pairs of genital discs (rarely 3 or 0). Palpi extending beyond gnathosoma,
or shorter and approximately equal to the chelae in length; with distal segment
raptorial, adapted for grasping (Plate 65) Family CUNAXIDAE

I With three pairs of genital discs.

distal setae; antenniform ,
Palpi long, often elbowed, usually with strong
(Plate 65) Family BDELLIDAE

I 24. Palpal tibia claw-like, with vague remnants of tarsal thumb. Tarsus I without claws
or empodium; other tarsi with large discoid empodia. Ectoparasites of arthropods
·" , ., (Plate 78 [see also 5-7]) Family HETEROCHEYLIDAE
I Palpal tibia and tarsus not suggesting a thumb-claw process. Tarsi various 25

I 25. All legs normal, ambulatory

Legs I adapted for clasping hairs of mammal hosts
26
(Plate 79) Family MYOBIIDAE

I 26. Palpi distinct; empodia rayed

Palpi foreshortened; empodia pad-like. With a strong hook-like spine on the
27

I venter of femora I-IV. Skin parasites of mammals

·
.
(Plate 80) Family PSORERGATIDAE

I 27. Claws apparently lacking; claw remnant(?) cup-like and with rayed empodium. Body
rounded. Parasites of snakes (Plates 77,78) Family OPHIOPTI DAE

I Claws present; empodia simple and with a double row of tenent hairs. Body
elongate. In quills of birds " (Plate 79) Family SYRINGOPHILIDAE

I 28. Strongly armored mites, generally bright orange or yellow in life. With a pair of
large lens-like structures laterally on the idiosoma in addition to ocelli. Coxae
forming characteristic apodemal patterns. Chelae with opposed digits. Free-living.
I ·
·
Cohort STOMATOSTIGMATINA, Superfamily LABIDOSTOMMOIDEA,
, (Plate 66) Family LABIDOSTOMMIDAE

I Soft-bodied mites, without large lateral lens-like structures or strong apodemal

patterns as above. Chelae with opposed digits, or short and needle-like 29

I 29. Cheliceral digits opposed. Propodosoma with one or two pairs of distinct differen-
tiated pseudostigmata and pseudostigmatic organs (Plate 67-2). Free-living
· Supercohort ENDEOSTIGMATA, Cohort PACHYGNATHINA,
.

I · Superfamily PACHYGNATHOJDEA 30

I
I
 I
178

Fixed cheliceral digit reduced, movable digit short and needle-like (Plate 70-5).
I
Propodosoma with a pair of sensory setae (pseudostigmatic organs) distinguishable
by their length and by type of insertion (Plate 70-4)
· . " .. , ., ., Cohort EUPODOSTIGMATINA (pars), Superfamily TYDEOIDEA
.
36
I
30. Tarsi I-IV each with two claws and empodium 31 I
Some or all of the tarsi lacking true claws, empodia or both 33

31. With two pairs of genital discs. Fixed chela often reduced. With one or two pairs of
I
propodosomal pseudostigmatic organs. Ocelli absent
· (Plate 69) Family LORDALYCHIDAE
.
I
With three pairs of genital discs. (only two pairs on deutonymphs). Fixed chela
normal , , " 32
I
32. With one pair of pseudostigmatic organs. One pair of lateral ocelli present.
· (Plate 69) Family TERPNACARIDAE
With two pairs of pseudostigmatic organs. Ocelli present or absent .
I
......... , , , , (Plate 67) Family PACHYGNATHIDAE

33. Tarsus I with true claws but lacking empodium; tarsi II-IV with claws and empodia.
I
With two pairs of ocelli and two pairs of hair-like pseudostigmatic organs
· " (Plate 67) Family SPHAEROLICHIDAE
.
I
True claws lacking on all legs, but with claw-like or pad-like empodia 34
I
34. Empodia claw-like (Plate 68-5) 35
Empodia pad-like. Ocelli absent; with a pair of club-shaped pseudostigmatic organs
on the propodosoma " .. _. (Plates 69,70) Family PEDICULOCHELIDAE
I
35. With two pairs of pseudostigmatic organs .
I
· " (Plate 68 [see also 8-5,10-7]) Family NANORCHESTIDAE
With one pair of pseudostigmatic organs (Plate 68) Family ALiCORHAGIiDAE I
36. Hysterosoma divided transversely by one or two sutures. Empodia claw-like; tarsus
I with several erect dorsal sensory rods. Elongate forms with distinct peritremes on I
the anterior aspect of the propodosoma. Free-living .
· , ., ., ., ., , , (Plate 70) Family PARATYDEIDAE
Hysterosoma without transverse sutures. Empodia pad- or hair-like; tarsus I with
I
one erect sensory rod. Peritremes indistinct or absent

37. With an ereynetal organ opening in the distal portion of tibia I (Plate 72-1),
37
I
consisting of an internal sac-like structure and a narrow distal "duct" which
terminates at or near the insertion of a hair-like, brush-like or otherwise highlY I
modified seta; with or without genital discs, hysterosomal sensory setae, net-like
pattern on the legs, or distal tarsal concavities on tarsi I-II. Free-living, associated
with molluscs, or parasites in nasal passages of vertebrates . I
· , , , .. (Plates 71,72 [see also 4-5, 5-4]) Family EREYNETIDAE
I
I
I 179

I Without ereynetal organ. Genital discs, hysterosomal sensory setae, net-like

pattern on the legs and tarsal concavities absent. Free-living .
I · (Plates 70, 71) Family TYDEIDAE

38. Body of adults and nymphs densely clothed with setae. Stigmatal openings between
I chelicera I bases (Plate 89-1,2). Larvae usually heteromorphic, parasitic; adu Its and
nymphs free-living predators Supercohort PARASITENGONA 55

I Body setae of adults and nymphs relatively few, arranged in transverse rows.
Stigmatal openings behind cheliceral bases (Plate 74-2). Larvae homeomorphic,
similar to nymphs and adults in habits 39
I 39. Body strongly armored, with several dorsal shields. With several internal setae on
legs I arranged so as to give the latter a rake-like aspect. Chelicerae short, thick, with
I strong sickle-shaped movable chelae and weak fixed chelae. Free-living
· Superfamily CAECULOIDEA, (Plates 72, 73) Family CAECULIDAE
.

I Body not so constructed 40

40. Chelicerae not fused together; hinged at base so as to move scissors-like over the
I gnathosoma (Plate 73-6); palpal tibia with 1-3 claws .
.. .. . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . Superfamily ANYSTOIDEA 41

I Chelicerae completely or partially fused, or contiguous, not capable of scissors-like

motion; with needle-like movable digits _ 44

I 41. Pal pal thumb (tarsus) long and prominent; chelicerae hooked distally. Free-living.
· (Plate 73 [see also 10-6]) Family ANYSTIDAE
Palpal thumb not prominent; chelicerae variously developed 42
I 42. Claws of tarsi I-I I strongly rayed; with three pairs of weakly developed genital discs.

I Free-living (Plate 75) Family TENERIFFIIDAE

Claws of tarsi I-II either absent or, if present, not rayed; genital discs absent 43

I 43. Tarsi with or without claws and/or empodia; claws never with tenent hairs.
Free-living (Plate 74) Family PSEUDOCHEYLIDAE

I Tarsi with claws but without empodia; claws with tenent hairs. Parasites of
lizards or arthropods (Plate 74) Family PTERYGOSOMIDAE

I 44. Cheliceral bases fused with gnathosoma and without indication of suture. Peritreme
present on gnathosoma, often shaped like an arch, or like an "M"
· (Plate 76-4) Superfamily CHEYLETOIDEA (pars)
.
45

I Cheliceral bases closely contiguous or fused with each other, but not generally with
gnathosoma. Peritremes usually present on anterior portion of propodosoma.
Predators or plant feeders 46
I 45. Tarsi II-IV (and usually tarsi I) with claws and empodia. Free-living predators or

I associates of vertebrates (Plate 76 [see also 4-1 J) Family CHEYLETJDAE

I
I
 I
180

Tarsi III-IV without claws or empodia, and ending in long, whip-like setae; tarsi I-II
I
with claws and rayed empodia. Parasites of birds (Plate 77 [see also 7-2]).
· Family HARPYRHYNCHIDAE I
46. Chelicerae long, recurved and whip-like, and arising from a stylophore (Plate 81-4).
Genital aperture transverse. Plant feeders ... Superfamily TETRANYCHOIDEA ... 47 I
Chelicerae not long and whip-like, but rather short and stylet-like (Plate 84-2).
Genital aperture longitudinal. Free-living ...Superfamily RAPHIGNATHOIDEA '" 50
I
47. Ocelli present on propodosoma 48
Ocelli absent. Elongate striate forms with widely separated coxae II-II I
·
.
(Plate 83) Family L1NOTETRANIDAE
I
48. Claws each with a series of comb-like tenent hairs (Plate 83-1), empodia similarly I
ornamented; caudal aspect of idiosoma with a series of flagelliform or dendritic setae.
.................................................................. 49
Each claw terminating in a pair of tenent hairs (Plate 81-2) or bordered with combs
I
of tenent hairs (Plate 81-1 J; empodia claw-like or pad-like (rarely appearing absent),
often divided distally or with a basal divided spur, with or without tenent hairs,
caudal aspect of idiosoma without highly modified setae .
I
· (Plates 81,82 [see also 5-6, 7-2]) Family TETRANYCHIDAE
I
49. Dorsum of hysterosoma with 36 fan-shaped setae. Caudal aspect of idiosoma with a
series of long flagelliform setae... (Plate 83 [see also 5-5]) Family TUCKERELLIDAE
Dorsal setae simple. Caudal aspect of idiosoma with a series of short dendritic setae.
I
·......................... (plates 83,84) Family ALLOCHAETOPHORIDAE

50. With a series of four or five dorsomedian hysterosomal shields arranged end-to-end.
I
Empodia present or absent; when present, without tenent hairs
Dorsal shields various, not as above. Empodia present, with tenent hairs
51
52 I
51. With a pair of clavate pseudostigmatic organs on the propodosoma; with four
dorsomedian hysterosomal shields. With no more than two solenidia on tarsi I;
I
empodia I present or absent. Without genital discs .
· (Plate 84) Family TARSOCHEYLIDAE
Propodosoma without clavate pseudostigmatic organs; with five dorsomedian
I
hysterosomal shields. With more than three solenidia on tarsi I; empodia I-IV
absent. With three pairs of genital discs (Plate 84) Family POMERANTZIIDAE I
52. Coxae II-III contiguous (Plate 85) Family RAPHIGNATHIDAE
Coxae I-II distinctly separated from coxae III-IV. or coxae equally separated from
I
one another

53. Peritremes extend into chelicera I bases

53

54
I
Peritremes confined to the anterior edge of propodosoma
·
.
(Plates 86,87) Family STIGMAEIDAE
I
I
I
I
181
I 54. Dorsal and leg setae ornate, often with raised insertions; phalangid-like, with legs
considerably longer than body (Plate 88) Family NEOPHYLLOBIIDAE 1
I Dorsal and leg setae generally simple. Idiosoma elongate, with legs rarely exceeding
body length (Plates 86, 87) Family CALIGONELLIDAE

I 55. Movable chelae of adults short, curved (Plate 91-1), sometimes toothed distally,
hinged at base. Empodia present or absent. Larvae with urstigmata (Plate 93-2). no
I distinct separation between coxae I-II; anus present; parasites of vertebrates or
arthropods Superfamily TROMBIDIOIDEA 58
Movable chelae of adults long, straight and extrusible (Plate 89-1 l. Empodia absent.
I Larval morphology variable; parasitic on arthropods .
.. . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . .. Superfamily ERYTHRAEOIDEA 56

I 56. Genital discs absent (rarely with one pair). With distinct prodorsal crista metopica
(Plate 89-4). Larvae without urstigmata, coxae I-II not contiguous; anus absent .... 57

I With two pairs of genital discs. Propodosomal sensory setae not inserted in a
distinct crista metopica. Gnathosoma capable of being withdrawn into body.
Larvae with urstigmata; coxae I-II contiguous; anal aperture present .
I · (Plate 90 [see also Fig. 8, p. 24]) Family CALYPTOSTOMIDAE

I 57. Gnathosoma small and narrow, capable of being completely withdrawn into body.
Propodosoma usually elongate anteriorly, with ossiform crista or scutellate shield.
Body setae generally flattened and serrate (Plate 901 Family SMARIDIIDAE

I Gnathosoma large, incapable of being withdrawn into body. Propodosoma not

elongate anteriorly. Generally with an ossiform crista. Body setae simple or
pectinate, and often somewhat broadened .
I · (Plate 89 [see also 10-3]) Family ERYTHRAEIDAE

I 58. With two pairs (rarely one) of propodosomal sensory setae in larvae and adults;
dorsal setae of larvae and adults simple, each borne on an individual platelet.
Paragenital sclerites of adults well developed, with many setae. Urstigmata always

I associated with coxae I. Larvae parasitic on arthropods .

· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. (Plates 91,92) Family JOHNSTON IAN IDAE
Always with a single pair of propodosomal sensory setae; dorsal setae of larvae and
I adults often ornamented, and mayor may not arise from individual platelets.
Paragenital sclerites of adults various, but with fewer setae than found on more

I medial genital sclerites (Plate 91-3). Larval urstigmata associated with coxae I or 11..59

59_ With only one or two setae on tectum. Adults may be figure "8"-shaped. Larvae

I with a single dorsal shield, with a few

posterior to palpal femur (Plate 92-5).
Larvae parasitic on vertebrates
exceptions. Seta on palpal coxa of larva
Urstigmata always associated with coxae I.
(Plates 92, 94) Family TROMBICULlDAE 2 ,3

I 1 Includes the family CAMEROBII DAE (Southcott 1957).

21ncludes the family LEEUWENHOEKIIDAE.

I 3Adult trombiculids are difficult or impossible to separate from adult trombidiids by existing
methods.

I
I
1S2
I
With numerous setae on tectum (when present). Adults not figure "S"-shaped. I
Larvae usually with more than one dorsal shield. Seta on palpal coxa of larva
usually extending anterior to base of palpal femur (Plate 93-4). Urstigmata
associated either with coxae I or with coxae II. Larvae parasitic on arthropods.
I
.. . . . .. . . . . .. (Plates 91,92,93 [see also Fig. 20. p. 142J) Family TROMBIDIIDAE

60. Adults generally inefficient swimmers, although swimming hairs on legs may be
I
present. Larvae parasitic, found on water surface rather than beneath it, with
rounded idiosoma, weak apodemes, and long posterior dorsal setae; sclerotization
weak or absent, leg axes not parallel (Plate 94-5) 61
I
Adults generally efficient swimmers. Larvae parasitic, free-living or suppressed; when
present, found actively swimming on or beneath the surface film, with dorsoventrally I
flattened idiosoma usually protected by sclerites or shields; most body setae often
modified or lost, although long dorsal hairs may be present; leg axes more or less
parallel (Plate 96-2) 62
I
61. Adults without swimming hairs on legs; with an anterior and posterior dorsal shield
flanked by a pair of small lateral shields. Larvae with an anterior dorsal shield
I
flanked by a pair of club-shaped setae; chelicerae arranged in a dorsoventral plane.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. (Plate 94) Superfamily HYDROVOLZIOIDEA I
Adults with or without swimming hairs, commonly without dorsal shields (when
dorsal shields occur, not as above). Larvae of at least the known families with a
median dorsal sclerite armed with sensory setae; club-shaped setae absent; chelicerae
I
arranged in a typical longitudinal plane; palpi raptorial _ .
............................... (Plate 95) Superfamily HYDRYPHANTOIDEA I
62. Adults with swimming hairs; globular, red, large forms with papillate integument.
Larvae parasitic, with greatly enlarged gnathosoma nearly equalling size of idiosoma;
palpi raptorial. Weak swimmers ..... (Plates 95, 96) Superfamily HYDRACHNOIDEA
I
Adults mayor may not have swimming hairs; variable in form and sclerotization.
Larvae usually free-living, but sometimes parasitic or suppressed; larval mouthparts
I
normal in size, less than half the length of the idiosoma; palpi raptorial. Strong
swimmers .. , ..... " (Plates 96, 97 [see also 4-2]) Superfamily HYGROBATOIDEA I
I
I
I
I
I
I
I
I PLATE 55

.55 2
183
I 0

I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I genital flap
transverse

I
I 55-1 to 55-5; family ER IOPHYI DAE, 55-1; Phy top tus avellanae Nalepa. (Oregon, USA), lateral aspect, with detail of annulations
and microtubercles: 55-2; P. avellanae, featherclaw: 55-3; p, avellanae, leg I: 55-4; Rhyncaphytoptus strigatus Keifer
(Maryland, USA), lateral aspect, with detail of leg I: 55-5; R. strigatus, genital area

I
 PLATE 56 I
184
I
I
chelicera
propodosomaJ shield I
ri:'\---\tb':t-- aedeagus points
I
IJI--~-#
forward

I
claw
I

/~ 56-1
I
I
I
I
I
I
I
I
aedeagus
I
I
56-4
56-5
I
I
56-1; family NEMATAL YCIDAE, Nematalycus (diagrammatic), venter of female with detail of chelicera, dorsal seta and apotele
of tarsus IV
56-2 and 56-3; family DEMODICIDAE, Demodex 'p. (Oregon, USA). 56-2; dorsum of male: 56·3; venter of female
I
56-4 and 56-5; family CLOACARIDAE, Cloacarus faini Camin etal. (Kansas, USA), 56-4; dorsum of female: 56·5; dorsum of
male (after Camin etal. 1967)
I
I PLATE 57
185
I
-)~
•
I
I
I
I
I
57·2
I
I
I 57-1

I
I
I
I
I 57-3

I
I
I
I 57-1; family CLOACAR IDAE, Cloacarus faini Camin et al. (Missouri, USA), venter of female with detail of leg I
57-2 to 57-4; family SCUTACARIDAE. 57-2; Scutacarus SP't lateral aspect (diagrammatic) (after Karafiat 1959): 57-3; apotele of

I leg II or III of a scutacarid mite: 57-4; Scutacarus sp. (Oregon, USA), venter
 PLATE 58 I
186
I
pseudostigmatic-__ N, /'-..;........~
organ

tracheal--4-.~
•
I
I
I
I
58-3 I
58-1
I
I
I
\"",W./
~{
~
tarsus IV
I
58-5 I
I
58-7
I

iii ~-X
I
I
I
I
58-1 to 58-6; family TARSONEMIDAE. 58-1; Tarsonemus sp. (Oregon, USA), dorsum of female: 58-2; characteristic apotele of
tarsus II or III of tarsonemid mite: 58-3; ?Tarsonemoides sp. (Oregon, USA), venter of female: 58-4; Steneotarsonemus sp.
I
(Oregon, USA). dorsum of larva: 58-5; leg IV of a saprophytic or fungivorous tarsonemid male: 58·6; leg IV of a phyto-
phagous tarsonemid male: 58-7; male propodosomal patterns of (feft) Steneotarsonemus and (right) Tarsonemus
I
I PLATE 59
187
I
•
I
59-2

I
I
I
I
•
I
59-4
chelicera

I \. 1'(1,. "._
1..\ ; ) •."'- ~ stigma
I ~, ~, . il/ >
I ~y\ J,\1P
I
I ~~~­
'59-~
I
I~
.~ 59-6
1 9<1 to 59-7; family PYEMOTI DAE. 59-1; Siteroptes reniformis Krantz (California, USA), dorsum of female (after Krantz 1957):
59-2; S. cerealium Kirchner (from Krantz 1959): 59-3; Resinacarus sp. (Oregon, USA), tibiotarsus I: 59-4; Pyemotes sp.
(Oregon, USA), dorsum of gnathosoma and portion of propodosoma: 59-5; Resinacarus sp. (Oregon, USA), tibia and

I
tarsus I: 59-6; Pygmephorus sp. (Oregon, USA), terminus of leg 1: 59-7; Acarophenax nidicolus Cross and Krantz (Colorado,
USA), venter of female (after Cross and Krantz 1964)
 PLATE 60 I
188
I
I
I
I
I
I
I
I
I
I
I
I
I
I
60-2
I
•
I
60-3

60.1; family CROTALOMORPHIDAE (manuscript name), Crotalomorpha (Kansas, USA), gravid female
60.2 and 60·3; family CRYPTOGNATHIDAE, Cryptognathussternalis Krantz (Oregon, USA). 60·2; propodosomal hood:
60·3; dorsum (after Krantz 1958)
•
I
I
I PLATE 61
189

~
I
I \ m,,",,_.;.w.'
I
I
61-1

r tz ~n
(,~~

I
I
61-3
I
••
I rostrum >

I
I
I
I genital disc 61-4 61-5

I
I
I
I
I 61-1 to 61-3; family RHAGIDllDAE (Oregon, USA). 61-1; venter of female with detail of rhagidial organs: 61·2; propodosoma:
61·3; palp
61·4 and 61·5; family HALACAR I DAE. 61·4; Agave sp. (Oregon, USAI. dorsum of female with detail of tarsus III: 61·5;

I Lohmannella sp., lateral aspect of rostrum and palpi

 PLATE 62 I
190

62-5 I
••
I
I
I
~I"'~'b¥-- propodo,omal
protuberance
I
~
62-6
•
I
I
•
I
I
I
I
I
I
62-4 anal opening
ventral
62-' to 62-3; family PENTHALODIDAE. Penthalodes ,p_ (Oregon. USA). 62-'; dorsum of female: 62-2; genital valves: 62-3;
•
I
•
chelicera
62-4 to 62-7; family EUPODIDAE_ 62-4; Eupodes sp. (Oregon. USAI. lateral aspect with detail of chelicera: 62·5; Linopodes ,p.
(Oregon, USA), dorsum with detail of apotele of tarsus II: 62-6; ?genus (Oregon, USA), sensory seta of tarsus I: 62-7;
Eupodes 'p. (Oregon. USA), venter
I· PLATE 63

.ff'
191
I
•• 63-1

~)~
63-2

~r
'k,~
"
I ,
63-3

I
I
I anal aperture dorsal or ' "

I 63-4
terminal - e ' "

I
I
I
I
I
I
I
I anal aperture

I
I 63-6

I 63·'; family EUPOOI OAE, Linopodes sp. (Oregon, USA), rostrum

63-2 to 63-5; family PENTHALEI DAE, Penthaleus major (Duges) (Oregon, USA). 63·2; tarsus 1with detail of sensory rod and
apotele: 63-3; genital valves: 63-4; lateral aspect: 63-5; dorsum
63-6 and 63-7; family TENUIPALP1DAE. 63-6; Do!ichotetranychus sp. (Utah, USA), dorsum: 63-7; Tenuipalpus sp. (Florida,
I USA), dorsum
 PLATE 64 I
192
I
6~2A
ocelli~ ~ ••
It i
) I
";,;"~~!.'. I
\'"' \ l I
I
\ I •
I
I
":(l I
,'>:
.~

I
genital aperture
64-4 I
""~~.\, I \ I
'ci'./
"'I ' ' 1/~ '~'\';
'!";;;';\'"
)~1
'.
I
I
I
64-5/f~'~
64-3
I
64-1 and 64-2; family TENU IPALPIOAE, Brevipalpus essig; Baker (Oregon, USA). 64-1; venter of female with detail of palp and
of tarsus II: 64-2; dorsum
I
64-3 to 64-5; family rOLlNIDAE, la/ina nana Pritchard (Massachusetts. USA). 64-3; genital aperture of female (after Pritchard
1956): 64-4; venter of gnathosoma (after Pritchard 1956): 64-5; dorsum with detail of tarsus II (after Pritchard 1956)
I
I PLATE 65
193

I
I
I 65-2
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I an~~~~nf' vanter of f~mal~
65-1 to 65-3; family BDELLIDAE 651-

•
65-4
, US_A).CUNAXIDAE.
_. amlly • anterodorsal
65-4- C aspect of typical bdellid"" 65-2"• t arsus I of bdellid: 65-3; Bdella p
• unaxa sp_ (Oregon. USAL dorsum". 65- 5,_Cunaxoldes _ sp. (AfricaL palp s "
 PLATE 66 I
194
I
•
I
I
66-2 I
I
I
•
I
I
I
~""~~~~4 I
~h" 66-3
I
I

~ Ol~~
genital area
••
I
~t--':---J
anal area

66-5 I
I \ (Oregon, USA)
. .re=,.
66·1; venter of fe male: 66-2; chelicera I
66-1 to 66-5,. familyI of
LAB IDOST0
female. . ; dorsumLabidostOf~"U~~.~·genjtal_anar
6 6MMIDAE, of female. area. 0 f male
66-3; tarsus 4

I
I
195
I
•• 3 pairs genital discs

I 2 pairs pseudostigmatic organs

I
I \ \
,
I '\ ~
, I
I
I ;t
~

J ,
f
'\
67-2

I J ~ ~ ~ \ ~
'i~o 1/ ~ ~')(
I I'..... I-j-l·~r(
I
I
I 67-5
I
I
I
I
I 67·1 and 67-2; family PACHYGNATHIDAE (Oregon. USA!. 67-1; genital area: 67-2; dorsum with detail of tarsus IV and dorsal

•
ornamentation
67-3 to 67-7; family SPHAEROLICHIDAE. Sphaerolichus barbarus Grandjean (after Grandjean 1937). 67-3; anterodorsal aspect:
67·4; chelicera: 67-5; tarsus III: 67-6; tarsus I: 67-7; genital area
 PLATE 68 I
196
I
•
I
I
•
I
I
I
I
I
68-3
I
\-cJ ~o
y--h I
(;~.~'* f) \
o

I", ! /0;\
I
\ ~!~
2 pairs genital
discs
68-4 I
I
I
68-5 68-6
I
I
68-1 to 68-3; family NANO~CHESTIDAE. 68-1; Nanorchestes sp. (Oregon, USA), dorsolateral aspect of female with detail of
I
•
pseudostigmatic apparatus: 68-2; Nanorchestes sp., tibia and tarsus I: 68-3; Speleorchestes sp. (Oregon, USA), body seta
68-4 to 68-6; family ALiCORHAG iIDAE, Alicorhagia sp. (after Grandjean 1939). 68-4; genital area: 68-5; tip of tarsus I:
68-6; chelicera
I PLATE 69
197
I
I 69-'
I
I
I
I
I
I
I
I
I
I
I
I
69-6
I

69~
I
I -..
~
~--~ ... \

\, 69-8

I
I 69-1 to 69·4; family TERPNACARIOAE, Terpnacarus sp. (Oregon, USA). 69-1; dorsum of female with detail of rutellum-. 69-2;

69-5 to
dorsal ornamentation: 69-3; genu, tibia and tarsus I: 69-4; genital area
69~7; family LORDALYCHIDAE, Lordalychus peraltus Grandjean (after Grandjean 1939). 69-5; terminal portion of
tarsuS I: 69·6; genital area: 69-7; chelicera
I 69·8; family PEDICULOCHELIDAE, Pedieulocheles (diagrammatic). genital area
 PLATE 70 I
198
I
chelate-dentate
chelicerae
2 pairs genital
discs

•
I
I
I
claw-like
I
empodium
I
postpedal_";';:"~,,>---
furrow
_ _~----\i'<'.
I
I
•
I
I
70-' I
I
70-6
f rect s e n s o / y
rod
I

~~
I
h----+--peritreme
I

70-1; family PEDICU LOCH Ell DAE, Pedicl1locheles (diagrammatic), dorsum

70-7
\ \ pad like
empod,um

70·2 and 70-5; family PARATYDEI DAE, Paratydeus sp. (Oregon, USA). 70-2; genital area: 70-3; tibia and tarsus I: 70-4; dorsum:
70-5; anterodorsal aspect
•
I
70-6 and 70·7; family TYDEIDAE. 70-6; genital area of typical tydeld mite: 70·7; tarsus I of tydeid mite
I
I PLATE 71
199
I
••
I 71-1
I pseudostigma -----JC-_.i)(

I soft striated _--''----+H-

integument

I
I
I
I
I
I
I
I
I
I
I
71-5
I 71-6 posterior sensillum

I 71-' to 71-3; family TYDEIDAE. 71-1; ParaJorry;a sp. (Oregon, USA), dorsum: 71-2; Coccotydeus sp., sensory seta: 71-3;
Microtydeus sp., sensory seta
71-4 to 71-7; family EREYNETI DAE, Ereynetes sp. (Oregon, USA). 71-4; venter: 71-5; detail of propodasomal sensory area:

I 71-6; genital area: 71-7; dorsum

 PLATE 72 I
200

72-3
I
claws in
72-2 concavity
I
I
I
I
I
ereynetal
nr----I~ organ
I
I
ereynetal organ
I
I
I
...L--I-_ _ palpal tarsus
I
I

~
fixed chela
reduced
I
(H)~ I I
;y ~~ I
72-6 I
I
72.' to 72·4; family EREYNETIDAE. 72-1; Ereynetes sp. (Oregon, USA), tibia and tarsus I: 72·2; Speleognathus sp., dorsum
(diagrammatic); 72-3; Boydaia sturm" WomersJey, tibia and tarsus IV; 72-4; Speleognathus sp., genital area
I
72-5 and 72-6; family CAECULIDAE, Caeculus sp. (Oregon, USA). 72-5; dorsum with detail of dorsal seta: 72-6; gnathosoma
and palp
I
I PLATE 73
201
I 73-2 ~~~
•
I
73-1
~#fr#~
empodium aLike
claw-like or p

I tibia

I
I
I
I
I
I
I
I
I
I
I hel icerae free,
~inged at base

I
73-5
I
I
I
I
 PLATE 74 I
202
I
•
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
7
74·1 to 44-
-, family PTER YGOSOMIDAE.
ofgnathoso ma- 74-3;
YL-rOAE,Pseu 0
I'
G
tarsu~ ~heVlus'PO. dorI:~m
74-4;eckobie/la
74-1; Gecko b- texana
dorsu (diagrammatic)
m
. USA), venter of female: 74·2; G. texana
Banks (Califor~ra, f
I
74-5; dorsumpSEUDOCHE
I
family
I PLATE 75
203

I
•
I
I
I palpal genu

I
I ocellus~~
75-1
I
I
I
I
I
I
I 3 pairs of discs

I r1 J r

I
I ~.~,,~."
•
I
75-2

!
I
75-1 and 75-2; family TENER I FFII DAE (Oregon, USA). 75-1 ;1 dorsum with detail of tarsal claws and of pal pal thumb-claw process:

•
75·2; genital and anal area
,
i

I
 PLATE 76 I
204

tenent hairs on
I
.m~,"m)'
I
I
76-3
I
I
I
I
I
I
76-4
I
opisthonotal (hysterosomal) shield
I
I
76-2
aedeagus
points
I
anteriorly

I
I
I
I
I
76-1 to 76-6; family CHEYLETIDAE. 76-1; Cheyletus malaccensis Oudemans (Oregon, USA), palpsl thumb-claw process:
76-2; C. malaccensis, dorsum with detail of tarsus I: 76·3; C. malaccensis, apoteJe of tarsus I: 76-4; Acaropsis docta
Berlese (Oregon, USA), dorsomedian aspect of gnathosoma: 76-5; A. docta, aedeagal area of male: 76-6; Cheyletiella
I
parasitivorax (Megnin) (Oregon, USA), aedeagal area of male
I
I 205
I
I --
I
•
I
I 77-1

I
I
I
I
I
I

~ '0--:::::,
genital aperture ant .
or posterior enor
I
I r".,;) .
I
I
I 77f2
i !
~ ~)O;"
I n . ~ 7
L ,om'" '" . 7-3
I arrangeme~tY'~fPH
palp PYRHYNCHID
77-2 and 77 3 fam I
emales
AE, Harpyrhynchus s
IOPTIofDAE 77 2 ,OphlO
A froph/optes (antenorl t OPh/O:;~ '
ptes pcol(fa'lforn,a,
-ndber Radf USAI d orsum
(Indlal, d etalis of apot e Ie of tarsus I and of the
with of
dorsum
I FainFf;;61~641. 77 3, genital
I I (postenor) (ad aptedmale
from(after
 PLATE 78 I
206
I
I
78-1
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
78-1; family OPHIOPTIDAE, Ophioptessouthcotti Fain (Thailand), venter of male (after Fain 1964)
78·2 and 78-3; family HETEROCHEYLIDAE, Heterocheylus sp. (Georgia, USA). 78·2; venter of female: 78-3; dorsum of
I
gnathosoma and propodosoma
I
I PLATE 79

I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I '19·1 and 79·2; family MYOBII DAE. 79·1; Protomyobia sp. (ciregon, USA), dorsum of male: 79·2; Radfordia sp. 10regon, USA),
leg I of female ;

I 79·3 and 79-4; family SYRINGOPHI L1DAE, Syringophilus sp.j (Delaware, USA), 79·3; gnathosoma of female: 79-4; venter of
female !
 PLATE 80 I
208
I
I
iql!\f\~P'-~;::::::;;=----,I----pal pal segments telescoped ventrally

I
I
femora with a ventral spur
I
coxae radially arranged
I
~L.-

I
I
I
I
I
I
I
I
I
I
I
I
80.1 and 80-2; family PSORERGATI DAE. 80-1; Psorergates simplex Tyrell, venter of female: 80-2; P. (PsorobiaJ bas Johnston
(New Mexico, USA), dorsum of femare (after Johnston 1964)
I
I
I PLATE 81
209
I claws with tenent
~hairs
I ~:""-~",,"f-- chelicera

I 81-1 rostrurr!-Ho-.",..--/-c---,,. 1/\

i
whip-like

I stylophore-f-'-lc------J--!-,;)1

I
I peritreme--~--\'...j,~:s;:;::::Y'~~,Lil'l\
\.(hRHJ

I
81-4
I
I
I
I oce\lus-'~~I---I!J
:il
f-\ ~,l~dorsal\!l.,.~~
propodosomal seta

";r e \ - - dorsocentral hysterosomal seta

AJ
I ~17 (~;Dj
L \
I /"J~ \ [\ ".=,..,.'""~_..,...
humeral seta

I
I
~/\ /J .i-~~-- sacral Seta

I 'V
l~f\\i
)/'J
I 81-5 ~1-6
~
I
I
I
clunalfseta

-----
81-1 to 81-6; family TETRANYCHIOAE. 81-1; apotele of retr~nycopsis: 81-2; apotele of Petrobia: 81-3; apotele of Tetranychus:
81-4; Tetranychuspacificus McGregor (Oregon, USA), arl~terodorsal aspect of female: 81·5; Tetranychus sp. (Oregon, USA).
dorsum of idiosoma: 81-6; Tetranychus sp., aedeagu5 of, male
 PLATE 82 I
210
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
82.1; family TETRANYCHIDAE , T-etranychus sp. (Oregon, USA), venter
I
I
I PLATE 83

I I 211

I genital

I
I
I
I ocellus---=-",,-l.I <

I
I
I
I
I
I
I
I
I
I ~
I
I
~-- 83-3

I 83-1 and 83-2; family TUCKERELLJDAE, Tuckerella sp. (Hatvaii, USA). 83-1; dorsum of female with detail of tarsus I: 83-2;
genital-anal area of female I
83-3; family ALLOCHAETOPHORIDAE, Allochaetophora qalifornica McGregor (California, USA), caudal aspect of nymph

I 83-4 and 83-5; family LINOTETRAN IOAE. Linotetranus sp_ j 83A; genital-anal area of female: 83-5; dorsum of female
 PLATE 84 I
212
I
I
84-4
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
84-1; family ALLOCHAETOPHORI DAE, Allochaetophora californica, dorsum of nymph (after McGregor 1950)
84-2 and 84·3; free and fused cheliceral bases in raphignathoid mites
84-4; family TARSOCHEYLl DAE, Hop/achey/us sp. (?California, USA), dorsum of female with detail of apoteJe of tarsus I
I
84-5 and 84-6; family POMERANTZIIDAE, Pomerantzia charles; Baker (Georgia, USA), dorsum of female (adapted from Baker
1949): 84-6;P. charlesi, genital-anal area (adapted from Baker 1949) I
I 213

I
I
I
I
I
I
I
I
I cheliceral bases fused

, ~~' . peritreme

I
?,},~k~~~~
19- /" L ~-;;~

I ·:.;' .; u:· 8
(t.g '~JI~
ocellus .
",i
I /;!TV'~ "'.
,.,(.:,,'A
~_; 85-2

I h' ~~ r~
I /7.•,./. '.. ~ \. '~. "'=
/
M~r
~ \ \. f
~- -
\ -l"'==
0

I \ \
, ..
.~ ~ t'\
I \~\J'
J!~fl'" ,~"'~
).
\.. '

I 1.!1\
85-3 .' i "
I lr~
~" "
I 85·' to dorsum
85·3; family
RAPHIGNATHID;~
RAPHIGNATHID
of female . h detaL!. of AE.
palpal85-';
tibiatarsus 1 of tyj't~al/aPhignathid
and tarsus
'
!i n 0 the terminusmite:
of tarsusI: 8,f.;,gnathus
85·2' Ra sp. (Oregon, USA)
I'' coxal arrangement in the '

I ,
 PLATE 86 I
214

I
86-~
86-1 palpal tibia

polpal genu

pal pal femur ~

. ;.l
Ii(

"",.
Ii -- palpal
tarsus I
I
I
I
dr--+-peritremes in
I
cheliceral base
I
I
I
I
I
I
I
I
I
I
I
I
86-1 and 86·2; family STIGMAEIDAE (Oregon. USA). 86-1; dorsum with detail of terminal portion of tarsus I: 86·2; venter
86-3; family CALIGONELLIDAE, Coptocheles sp. (Oregon, USA), gnathosoma and anterior portion of propodosoma
I
I
 I
I PLATE/87
215
I !

87·1
I
I
I
I
coxae I-II separate f
I coxae III.IV rom

I
I 87-2
I
I
I
I ,;

I dorsum may be
ornamented

I ('

I setae Sim~le

I \
t\
, \

I 1Jl~ 1-

I A,~\~I! \ '
,ii\:. !
I 87-1 and 87 2
STIGMAEI6~ONELLIDAE
87-3', f amily
-; family CA Ledermuell~ri;Ptocheles , r
E,
, :
C sp. (Oregon {O!:; egon,
sp. USt USA)~ith
, dorsum 87 ~1etall
; coxal arranget~~~~_c~7-2; dorsum
of palpal
; l aw process

I
i
 PLATE 88 I
216
I
88-1
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
88-2 I
I
88-1 and 88-2; family NEOPHYLLOBIIDAE. 88-1; Neophyllobius sp. (Oregon, USA). dorsum: 88-2; Camerobia australis
Southcott (Australia), body seta (after Southcott 1956)
I
88-3; family EUPALOPSELLIDAE, Eupalopsellus sp_ 10regon, USA), dorsum with detail of palp

I
I PLATE 89
217
I 89-1

I ,i
stigmata opell~ between
cheliceral bases

I I

I
I , peritreme
89-2

I if
:
l

:':'~:'
'
,1L-chelicerae
:\ \
", retractile
III i;\
I ItI

til ~ ~
\:lui
II I

I
I
I
I
I
I
I
I
I
I
I 89-4

I 89-1 to 89-4; family ERYTH RAEIDAE. 89-1; rostral area of 1'r ythraeid mite: 89-2; peritremes and stigmata of an erythraeid
mite: 89-3; Leptus sp. (Oregon, USA), dorsum of larva with detail of apotele of tarsus I: 89-4; ?Balaustium sp. (Oregon,
USA), dorsum with details of terminal portion of tarsU! I and of body seta
I
I
 PLATE 90
I
218
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
90-1 to 90-3; family SMARI DII DAE. 90-1; Fessonia sp. (Oregon, USA), anteromedian portion of propodosoma: 90·2; scuteJJate
shield with sensory setae (diagrammatic): 90-3; Smaris sp. (Oregon, USA), venter with detail of parpal thumb·claw process
I
and dorsal seta
90-4 and 90-5; family CALYPTOSTOMI DAE, Calyptostoma sp. (California, USA). 90-4; dorsum: 90-5; genital-anal area I
I
219

I
I
I '-4-~'f4-- 3 pairs of
genital discs

I
I
I
I
I
I
I
I
I
I
I '\~.:~~rt:
"
II \,

'~~~~
I '\'0{
\ I

\-
I
'
t
paragenital Sc1 ites

I with fewer set- ethan

on medial scler:tes

I 91-4

I 91-1 and 91-2; family JOHNSTONIANIDAE. 91-1; dorsum wilth detail of setal platelets: 91-2; Dip!othrombium sp., genital area
~fumale i
91·3 and 91·4; family TROMBIDIIDAE, Trombidium sp. (Ore~on. USA). 91·3; genital area of female: 91-4; dorsum of female
I with detail of setae '
 PLATE 92 I
220
92-2 92-3 ~
urstigma-~~
I
claw A- I
I

f~'1V.
t I

1 \
\
I
I
92-1 i I
I
I
I
I
I
I
empodium
prernt
I
I
I
I
,
'"
palpal coxal seta-{~.<~~-::~~_ 92·6
I
92-5 /
I
I
92-1 to 92-3; family JOHNSTON IAN I DAE. 92-1; dorsum of larva: 92·2 and 92-3; apoteles characteristic of johnstonianid species
92-4; family TROMBIDII DAE. dorsum of engorged larva
I
92-5 and 92-6; family TROMBICULlDAE_ 92-5; Sasacarus sp. (Nevada. USA). palp of larva: 92-6; Trombicula sp. (Korea). dorsum
of larva
I
I PLATE! 93
I 221

I j

I
I 93-4 ~
I urstigma palpal coxal seta

I
I
I
I
I
I
I
I
I
I
I
I 93-6
I
I
I 93-1 to 93-4; family TROMBIDII DAE, Alfothrombium ,p. IvVaShington, USA). 93-1; tarsus 1: 93-2; venter of larva: 93-3; dorsum
of larva: 93-4; palp !
93-5 to 93-7; dorsal shield patterns of trombidiid larvae :'

I I
 PLATE 94 I
222
I
I
I
I
I

@
a,~
I
I \
I
/I:~
anal~----=--\ 94-2 I
~
aperture

Q; 94-3
I
I
I
I

~
I

,,~~ ~Iil ~.
I
(Or
vA
\t~~
r/1r 'Y. ""~
\/r~I~~y
I
I
I
'%;\
94-4 \7M LI DAE Euschongasoa
94 1 and 94-2, faml Iy TROMBICU, sp_ 10reg
EA Hydrovo!zJa sp_ on
94-5
, USA).
943
."'''M''.",.,
of ad u It
94-1,
, venter
~"."''',",_,"''
944, dorsum of adult 94-5", dorsum of
I
I
I~~V; superfamily HYD~O:~aL~~7elrDMlt~hell 1957)
94 3 to larva'with detail of gnat as
I
I PLATE 195
! 223
I
95-'
I
I
I
I
I 95-2

I
I
~
I
I
I
I
I 77J~
I
I
}OW ~ ~
@)
"At
I ~
~
r
I ~
95-4
I
I 95-1 to 95-4; superfamily HYDRYPHANTOIOEA. 95-1; Thva~ barbigera Viets (Michigan, USA), anterodorsal portion of idiosoma:
95-2; Eylais sp., eye plate of adult: 95-3; T. barbigera, d~orsum of larva (after Mitchell 1957): 95-4; T. barbigera, venter of
adult i
I 95-5; superfamily HYDRACHNOIDEA, Hydrachna sp., dorsup, of adult

i
 PLATE 96 I
224
I
96·1 I
gnathosoma
enlarged
I
I
I
I
I
I
I
I
I
I
I
I
I
I
96-4 I
I
I
I
 I
I PLATEI97
:1 225

I
97/.1
I
I
I
I
I
I
~_-I±::=-

I genital acetabula

I
I
I
I
I
I
I
I
I 97-2 !

I
I 97·1 and 97·2; superfamily HYGROBATOiDEA. 97·1; Pionl;psis 'p. (Oregon, USA), venter of femaie: 97·2; Hvgrobates sp.
(Michigan, USA), venter of male '

I
 I
226

NOTES
I
I
I
I
I
I
I
I
I
I
I
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I
I
I
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I
I
I
I
I 227
I
I
I
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I
228
I
ORDER ACARIFORMES I
Suborder Astigmata
I
The Astigmata is a fairly homogenous assemblage of slow-moving, weakly sclerotized
mites which range in size from 200 to 1500 iJ.. With few exceptions, the astigmatids have
I
evolved to a non-predaceous existence and may be considered a highly advanced group.
Respiration is apparently integumental throughout the suborder, although trachea-like
ducts have been observed in some species. These ducts may open into the genital vestibule.
I
A supracoxal gland may be found dorsad of trochanter I, the gland emptYing into the
podocephalic canal (Fig. 24). Other important identifying characteristics include: I
1. Fusion of leg coxae with venter, the position of the coxal fields often being
demarcated by subintegumental apodemes (Plate 98-1). I
2. Presence of a terminal (rarely dorsal) bursa copulatrix in females (Plate 8-1)
which often is extrusible. Males usually have an aedeagus (Plate 7-4,5).
I
3. A pair of laterodorsal opisthonotal "glands" in the majority of families (Plate
102-4) (absent in the SARCOPTIDAE and others).
I
4. A single-segmented palp which usually consists of two false segments, or
podomeres (Plate 100-3) [exceptions include the families ANOETIDAE (Plate
I
5.
98-2) and CHIRORHYNCHOBIIDAE (Plate 123-1)J.

Chelicerae often chelate-dentate (Fig. 25), with a paraxial seta on the fixed digit.
I
Rutella are absent.
I
6. Absence of true claws. The empodium may be claw- or sucker-like (Figs. 26,27).

7. Absence of discrete sternal or anal shields. The genital region may be covered by
I
a pair of lateral paragynial flaps and a posterior epigynial flap (Fig. 28). Two
pairs of genital discs may occur in adults (Plate 99-2). I
8. Two or three rod-like sensory setae, or solenidia, on tarsus , (Plate 119-3).
Solenidia also may occur on genua and tibiae. I
Astigmatid mites are cosmopolitan and have achieved success as fungivores, sapro-
phages, predators, graminivores and parasites. The latter category includes feather and I
skin mites of birds, skin parasites of mammals, gill inhabitants of Crustacea, and respiratory
and visceral parasites of a variety of animal hosts. Approximately 40 families are included
in the Astigmata, and these are grouped in two supercohorts (Fig. 29).
I
I
I
I
I
I
229
I
I
paraxial seta

I
I
I
I
I Fig. 24. Anterolateral aspect of an astigmatid mite
(ACARIDAE),
Fig. 25. Chelate-dentate chelicera of an
ast;gmatid mite.

I
I
I
I
condylophore
I
I
I
,~~
I ~SOlenidion w1

I
I Fig. 26. Tarsus II of an astigmatid mite (ACARIDAE) Fig. 27. Tarsus 11 of an astigmatid mite

I with a claw-like empodium. (PSOROPTIDAEI with a sucker-

like empodium.

I
I
 I
230

/
/
I
/

I
I
I
I
I
I

I
I
I
~
I \
I I
I
I
I
I
I
pregenital sclerite~ I
I
I
Fig. 28. Genital regions of various astigmatid mites: ACARIDAE
I
(upper left), CHORTOG LYPHIDAE (upper right),
AUDYCOPTIDAE (lower left), SARCOPTIDAE (lower
right). I
I
I
I
I
Supercohort Superfamily I
A 'd'
can la .
r= Anoetoidea
L[ Canestrinoidea I
Astigmata ~ Acaroidea
I
~
[ Listrophoroidea
Psoroptidia Ewingoidea
Psoroptoidea
Analgoidea
Sarcoptoidea
Cytod itoidea
I
Fig. 29. Dendrogram illustrating possible relationships within the Order ACARI FORMES:
Astigmata of the subclass ACARI.
suborder I
I
I
I
I I
I
231

Useful References I
I Andre, M. (19491. Ordre des Acariens. Traite de ZOOlogielpariS 4:794-892.
DUbinin, V.B. (1953). Fauna of the U.S.S.R. Arachnoid. a 6(61. Analgesoidea Pt. II, Epidermoptidaeand
Freyanldae. Publ. Zool.lnst. Acad. SCI, U.S.S.R., M scow: 411 pp,
I Vltzthum, H.G. (1942).
Book 5(6) :801-912.
Acarina. Bronn's Klassen und Ordnungen des Tierreichs, Leipzig: 5, Sect. 4,
,
tLe
I Yunker, C.E. (1955). A proposed classification of
Helminth. Soc. Wash. 22(2):98-105 I'
Acaridiae (Acarina: Sarcoptiformes).

Zakhvatkin, A.A. (1952). Division of the Acarina int%rders and their position in the system of the
Proc.

Chelicerata. Mag. Parasit. Moscow 14:5-46. .

I
Superfamily Anbetoidea
I ,i
DIAGNOSIS: Soft-bodied; with prodorsal shield! sejugal furrow present; movable cheliceral
digit reduced, palpi highly mOdif~'ed as "strainers." With an empodial claw
I and sessile caruncle on each leg. !Female genital opening transverse, flanked
by 2 pairs of ring-like structures. r Male with aedeagal apparatus between or

I behind coxae I V; ringlike strucljures present. Hypopal stage commonly

encountered. '

The Anoetoidea contains a single family, tt~e ANOETI DAE or slime mites. Anoetids
I are virtually world-wide in distribution, living in highly organic wet substrates. The
deutonymphal or hypopal stage is commonly fo~~nd as a phoretic associate of millipedes or
I insects although it may be encountered crawlil g actively with other stages on the food
substrate (Hughes and Jackson 1958). The adul s probably feed on microorganisms in their
semiaquatic habitats, although Histiosoma mu chiei Hughes and Jackson and H. berghi
I Jensen have been observed to feed on eggs ,of earthworms and leeches respectively.
!
I The adaptation of the ANOETI DAE to Isemiaquatic habitats reaches a climax in
species such as Histiosoma nigrellii H. and J. an~ H. cyrtandrae (Vitzthum), both of which
live completely submerged in water (Hughes~'and Jackson 1958). Both species were

I observed to feed in debris among aquarium ,Iants. All stages of the anoetid Anoetus
gibsoni Hunter and Hunter (including eggs) wer collected from below the surface of liquid
in a pitcher plant (Hunter and Hunter 1964). ther members of the genus also are found
I in this habitat (Hirst 1928, Nesbitt 1954). MOle typical, however, are substrates in which
the mites live beneath a thin water film. Defaying vegetables, or insect cultures which
have gone out of condition are common s~urces of burgeoning anoetid populations.
I i

I Useful References

Hirst, S. (1928). A new Tyroglyphid mite (Zwickia nJpenthesiana-sp. n.) from the pitchers of Nepenthes

I ampul/aria. Jour. Malay Brit. Asiat. Soc. 6: 19-221

N.S.(1):5-198.
I

Hughes, R.D. and e.G. Jackson (19581. A review bf the Anoetidae (Acari).
'
Virginia Jour, Sci. 9,

Anoet~s
I Hunter, P.E. and C.A. Hunter (1964).
Proc. Ent. Soc. Wash. 66(1):39-46.
A new mite from pitcher plants (Acarina: Aneotidae).

I
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232
I
Nesbitt, H.H.J. (1954). A new mite, Zwickia gibsoni n. sp. from the pitchers of Sarracenia purpurea.
Can. Ent. 86:192-197.
I
Scheucher, R. (1959). Systematik und Okologie der deutschen Anoetinen. Beitr. Syst. Oko!. mitteleurop.
Acarina 1(2) :233-384. I
Superfamily Acaroidea I
DIAGNOSIS: Soft-bodied; often with a prodorsal shield, sejugal furrow present; chelicerae
chelate-dentate; palpi simple. Apoteles present, consisting of a fleshy
I
stlaked or sessile caruncle and empodial claw. Female genital opening
longitudinal, resembling an inverted "Y," with 2 pairs of genital discs; male
with anal, and often with tarsal, suckers. Hypopal stage commonly
I
encountered.

The twelve families comprising the Acaroidea are a somewhat heterogenous group of
I
taxa which include both free-living and phoretic species found throughout most of the
world. Some are of economic importance as contaminants of stored or cured products. I
The family ACARIDAE is a large assemblage of saprophagous, graminivorous,
fungivorousand phytophagous species which may be found from extremely wet to fairly dry
I
habitats, generally feeding on organic debris of plants or animals. Species of the genus
Acarus, however, feed directly on stored grain and often cause considerable injury to the
commodity (Solomon 1946, Griffiths 1964). Tyrophagus putrescentiae (Schrank) is a
I
common contaminant of stored products which, because of its astonishing fecundity, is a
cause for concern in laboratory insect cultures. T. dimidiatus (Hermann) is reported as a
crop pest in greenhouses (Van den Bruel 1940), while Rhizoglyphus echinopus (F. & R.)
I
attacks bulb tissue both in the field and in storage (Garman 1937). Primary damage to lily
bulbs in Oregon may be caused by a fungus which causes deterioration of bulb tissue and
invites secondary attack by R. echinopus. It is possible, however, that the mite may cause
I
primary as well as secondary injury.
I
Although species of Caloglyphus may occur in food storages, most species are soil or
litter forms. C. berlesei Michael is often found in stored products (Hughes 1961) but is
equally prevalent as a saprophage in soil. Species of the acarid genera Suidasia, Thyreophagus,
I
Lardoglyphus and Aleuroglyphus also occur as contaminants of stored products. A species
of Tyrophagus has been implicated as the causal agent in cases of human intestinal and
urogenital acariasis. Acarid hypopi occur primarily on insects, although other animals may
I
serve as carriers (Zakhvatkin 1941, Samsinak 1965). Hypopi of the acarid genera Horstia,
Lackerbaueria and Tortonia are common on bees and wasps (Baker 1962). I
Acaroid species of the families GLYCYPHAGI DAE, CHORTOG LYPHI DAE, CTENO-
GLYPHIDAE, CARPOGLYPHIDAE and LABIDOPHORIDAE also occur in stored pro- I
ducts (Hughes 1961), but are not necessarily confined to this habitat. Grammolichus and
Sclerolichus spp. (GLYCYPHAGIDAE) are collected from the nests of rodents (Fain
1968a), as are species of Ctenoglyphus (family CTENOGLYPHIDAE) (Fain 1967).
I
Glycyphagus domesticus (DeGeer) is an intermediate host of a rodent tapeworm (Joyeux
and Baer 1945), and is also a causative agent of "grocers itch" of food handlers. Species of I
I
I
I i 233

I ctenoglyphids, chortoglyphids, and 9IYCYPhagid<] may be found in the dry dust of hay or
fodder in warehouses and farm feed storages. G ycyphagid hypopi usually occur on insects

I or birds or in rodent nests, but inert G'YCYPhagrS hypopi are carried by air currents from
one point to another inside the protonymphal skin (Fig. 30). Carpoglyphus lac tis (L.)
I
I I

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Fig. 30. Inert hypopus of G1vcvphagus destructor (Sellrank) within the protonymphal skin (left) and free

I (right). !

(family CARPOGLYPHIDAE) is an inhabitant of dried fruit, milk products and decaying

I food materials. Wine, caramel and flour also IraVe been reported as satisfactory substrates
for these species (Hughes 1961). ! ,
I Gohieria fusca Oud. (family LABIDOP'HORIDAE) is a common contaminant of
stored grains and feed. on which it may ca'use perceptible damage (Zakhvatkin 1941 l.

I Other labidophorids are free-living inhabitants t,f rodent nests (Drummond 1957, Rupes and
Whitaker 1968), although adults of the labic!ophorid Xenocastor fedjushini Zakhv. have

I
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234
I
been found living on their host (a beaver) rather than in the nest (Zakhvatkin 1941). Unlike I
the majority of labidophorid adults, labidophorid hypopi usually are found attached to the
hairs of their rodent hosts. Hypopi of Labidophorus sciurinus (Koch) and L. soricis Oud.
are found on mice, squirrels and shrews in North America and Russia. Other species of this
I
genus are carried by moles, water rats and muskrats (Turk and Turk 1959, Zakhvatkin
1941). Dermacarus ondatrae Whitaker, D. hypudei (Koch) and D, heptneri (Zakhv.) have
been collected from North American rodents (Rupes and Whitaker 1968). In addition to
I
labidophorids, mole and field mice nests in Europe and North America often harbor adults
of Fusacarus laminipes Michael (family FUSACARIDAE) (Michael 1903). The feeding
habits of fusacarids have not been studied.
I
It has been found that many birds, as well as certain rodents, may harbor subcutaneous
infestations of hypopi. A hypopal species identified as Falculifer rostratus (Buchholz), a
I
dermoglyphid, by Robin and Megnin (1877) was found subcutaneously in pigeons. Fain
and Bafort (1966) postulated, on the basis of chaetotaxy, that the hypopus considered to I
be Falculifer is in fact a free-living nest inhabitant in the adult stage rather than a feather
mite. The family Hypodectidae (=HYPODERIDAE Murray 1877) was erected to contain
this mite and other related forms found in the tissues of birds in the hypopal stage.
I
Characteristically, hypopi of HYPODERIDAE are elongate forms which achieve considerable
size with increasing age of the host (Fain 1967b). Hypopi of Hypodectes propus (Nitzsch),
for example, increased in average size from approximately 200 /l to over 500 /l in pigeons
I
from two weeks to two months old.

Hypopi of the genus Rodentopus resemble those of the HYPODERIDAE in many

I
ways, including choice of habitat (R. muris Fain was taken from a skin lesion of its host
(Fain 1965). although its host was a rodent rather than a bird). The subfamily Rodento- I
pinae, which also includes other hypopal associates of rodents, has been considered a part
of the GLYCYPHAGIDAE (Fain 1967c). Recently Fain described a species of HYPOD-
ERIDAE which was collected from a rodent in Africa (Fain 1968b). It was made the I
representative of a new hypoderid subfamily, the Muridectinae, on the strength of a
number of chaetotactic characters including the presence of vertical external setae on
the propodosoma. Rodentopus shares this and other characters, for which reason the
I
muridectine hypoderids and the rodentopine glicyphagids are herein considered as one
assemblage. They are treated as part of the LABIDOPHORIDAE with which they share
many morphological and habital characters. The HYPODERIDAE is considered a separate
I
family, with affinities with the GLYCYPHAG IDAE.

Bees provide the habitat and transport mechanism for members of the family
I
CHAETODACTYLIDAE. Adult Chaetodactylus, Sennertia and related genera are found
primarily in the nests of wood-boring xylocopid and solitary megachilid and osmid bees in I
both tropical and temperate climates (Zakhvatkin 1941, Krombein 1962). HYQopi
utilize these bees to migrate into new nesting sites. According to Popovici-Baznosanu
(1913), Chaetodactylus osmiae (Dufour) may be truly parasitic on Osmia bees. Krombein I
(1962a) observed C. krombeini Baker to feed on eggs and larvae of Osmia.

Members of the SAPROGLYPHIDAE are unsclerotized opaque species which may be

I
fungivorous or saprophytic, and which may be found associated with insects. Czenspinskia
lordi Nesbitt is common in filbert buds and in apple trees where it apparently feeds on
fungi or vegetable matter (Nesbitt 1946). Vegetable material also provides a substrate for
I
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I
235
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I )
I
I Fig, 31, Comp oSlte
, dorsum
hypopus of Neott' :nd venter of the
Iii hypopus of o~~~ and venter of the
Fig. 32. Composite d
ILABIDOPHORI;1topus sciuri Fain
bassani (Monta uj '(' ges (Pelecanectes) i
ladapted from ~ , HYPODERIDAEj
I am 19671.
;i,1
Fain 1965). Ej (adapted from

I
 236
I
the saproglyphid Oulenzia arboricola (Oud.) (Oudemans 1928). Species of Ensliniella and
Winterschmidtia spp. both are found on or with insects, while adults of Calvolia heterocoma
I
Michael apparently are associated with rodents. Other species of Calvolia are found in
granaries (C. romanovae Zakhv. and C. kamenski Zakhv.) or on decaying plants (c. I
hebeclinii Sicher). Hypopi of Calvolia occur on bees, beetles and flies in many parts of the
world. All stages of C. furnissi Woodring are found in bark beetle galleries (Woodring 1966).
I
Hypopi of the saproglyphid genus Monobiacarus are common on bees and wasps
(Krombein 1961). Vidia cooremani Baker is found in the hypopal stage with a solitary
wasp in eastern United States (Baker 1964). Other Vidia species are found associated with
I
wood-boring Hymenoptera. Kennethiella trisetosa, a saproglyphid, is transmitted venereally
from males to females of the wasp Ancistrocerus antilope (Cooper 1955). I
The family ROSENSTEINIIDAE, once considered a part of the CANESTRINIIDAE,
is a group of small (300-400 ;.d weakly sci erotized, sometimes scaly mites which are found
either on bats (Ah and Hunter 1968) or in bat caves (Strandtmann 1962). Nycteriglyphus
I
species are common on bats in the Far East, Europe and Africa. N. bifolum Strandtmann
is known from Texas, and Mydopholeus capillus McDaniel and Baker occurs on a free-tailed I
bat in Mexico. Species occurring on bats commonly are found on the wing membranes of
the host, as well as on the body proper. The feeding habits of rosensteiniids have not been
studied.
I
Species of the family HYADESII DAE are found in tide pools, submerged algae and in
mussel beds in coastal areas of Europe, (Halbert 1915, Michael 1901) and the Americas.
I
Hyadesia algivorans (Michael) is found in algae bathed by fresh water but within reach of
sea spray. An undescribed species was taken in red coralline algae on the Caribbean coast
of southeast Mexico where the mites were perpetually submerged. H. uncifer Megnin, on
I
the other hand, is considered semiaquatic (Andre 1931).

Useful References
I
Ah, H.S. and P.E. Hunter (1968). Nycteriglyphus vespertilia n. sp. a new acarid mite associated with bats
from Korea (Acarina: Rosensteiniidae). Aearologia 10(2):269-275.
I
Andre, M. (1931). Sur Ie genre Hyadesia Megnin 1889 (Sarcoptides hydrophiles). Bull. Mus. d'Hist. Nat.
Paris Ser. 2, 3(6):496-506. [HYADESliDAEj
Baker, E.W. (1962). Some Acaridae from bees and wasps (Acarina). Proe. Ent. Soc. Wash. 64(1):1-10.
I
[ACARIDAE]
Baker, E.W. (1964), Vidia cooremani, a new species of Saproglyphidae from a crabronine wasp (Acarina).
Ent. News 75(2) :43-46.
I
Cooper, K.W. (1955). Venereal transmission of mites by wasps, and some evolutionary problems arising
from the remarkable association of Enslin/ella trisetosa with the wasp Ancistrocerus anti/ope.
Trans. Amer. Ent. Soc. 80:119-174. [SAPROGLYPHIDAE]
I
Drummond, R.O. (1957). Observations on fluctuations of acarine populations from nests of Peromyscus
leucapus. Ecol. Monogr. 27:137-152. [LABIDOPHORIDAEj
Fain, A. (1963). Les Tyroglyphides commensaux des Chauves-souris Insectivores. Description de cinq
I
especes nouvelles. Rev. Zool. Bot. Afr. 67:33-58. [ROSENSTEINIIDAE]
Fain, A. (1965). Un nouveau type d'hypope, parasite cuticole de rongeurs Africains (Acari: Sarcoptiformes)
Zeit. fUr Parasit. 26:82-90. [LABIDOPHDRIDAEj
I
Fain, A. {1967aJ, Acariens nidicoles et detriticoles d'Afrique au Sud du Sahara. II. Ctenoglyphus
schoutedeni sp. n., vivant dans les nids de murides au Congo (Glycyphagidae: Sarcoptiformes).
Rev. Zoo I. Bot. Afr. 75(1-2):162-170. [CTENOGLYPHIDAE]
I
I
I
I
237
I Fain, A. (1967b). Les hypopes parasites des tissus celluI ires des oiseaux (HYPodecticae: Sarcoptiformesl.
Bull. Inst. Roy. Sci. Nat. Belg. 43(4):1-139. [Hype DERIDAE]

I Fain, A. (1967c). 1 phoretique sur des rongeurs et des marsupiaux

Nouveaux hypo pes vivant en aSSOCia tion
(Acarina: Glycyphagidae). Acarologia 9(2) :415-43 . [LABIDOPHORIDAE]

I
Fain, A. (1968a). Acariens nidicoles et detriticoles en

1 frique au Sud du Sahara. IiI. Esp"ces et genres

nouveaux dans les sous·familles Labidophorinae et ( rammolichinae (Glycyphagidae: Sarcoptiformes).
Acarologia 10(1) :86·110. [LABIDOPHORIDAE, . LYCYPHAGIDAE]
Fain, A. (1968b). Un hypope de la famille !-iypoderidbe Murray 1877 vivant sous la peau d'un rongeur
j
I (Hypoderidae: Sarcoptiformes). Acarologia 10(1): 11-115. [LABIDOPHOR 1DAEl
Fain, A. and J. Bafort (1967).
(Nitzsch) acarien nidicole
Cycle evolutif et morphologie de Hypodectes (Hypodectoides) propus
adeutonymphe paraSl~e tissulaire des pigeons. Bull. Acad. Roy. Belg.

I 53(5) :501·533.
Garman, P. (1937). A study of the bulb mite (Rhizogiy· hus hyacinthi Banks). Connecticut Agr. Exp. Sta.
Bull. 402:888-904 + plates.
If., 1758 (Acaridae, Acarina).
I Griffiths, D.A. (1964). A revision of the genus Acarus
Hist.) ZooI. 11 (6) :415-464 + plate. :
Bull. Brit. Mus. (Nat.

Hughes, A.M. (1961). The Mites of Stored Food. Ministry Ag. Fish and Food Tech. Bull. 9:287 pp. + vi.

I Joyeaux, C. and G. Baer (1945). Morphologie, evo1ution et position systematique de Catenotaenia

pusilla (Goeze, 1782). Cestode parasite de Rongeurs.
PHAGIDAE] ~
Rev. Suisse Zool. 52(2):13·51. [GLYCY·

I Knulle, W. (1959). Morphologische und Entwicklungsge chichtliche untersuchungen zum phylogenetischen

System der Acari: Acariformes Zachv. II. Acari iae: Acaridae. Mitt. ZooI. Mus. Berlin 35(2):347-
417.

I Krombein, K.V. (1961). Some symbiotic relations betreen saproglyphid mites and solitary vespid wasps
(Acarina, Saproglyphidae and Hymenoptera, V spidae).
Krombein, K.V. (1962a).
Jour. Wash. Acad. Sci. (Oct.):89·92.
Natural history of PIUmn[ers Island, Maryland XVI. Biological notes on
Chaetodactylus krombeini Baker, a parasitic mite of the megachilid bee, Osmia rOsmiaJ Iignaria Say
I (Acarina: Chaetodactylidae). Proc. BioI. Soc. wasl' 75:237-250.
Krombein, K.V. (1962b). Biological notes on acarid m tes associated with solitary wood-nesting wasps and
bees (Acarina: Acaridae). Proc. Ent. Soc. Wash. 6 (1): 11-19.

I McDaniel, Jr. B. and E.W. Baker (1962).

Fieldiana-Zool. 44(16): 127-131.
A new 6enus of Rosensteiniidae (Acarina) from Mexico.
.1
Michael, A.D. (1901). British Tyroglyphidae. Ray 10c" London 1:291 pp. + xiii + plates. [GLYCY-
I PHAGIDAE]
Michael, A.D. (1903). British Tyroglyphidae. Ray
,
~oc.,
London 2:183 pp. + vii + plates. [CHORTO-
GLYPH I DAE, FUSACAR IDAE, CHAETODACT LI DAE, GLYCYPHAGI DAE, CARPOG LYPH IDAE,

I ACARIDAE] ,
Nesbitt, H.H.J. (19461. Three new mites from Nova SI!cotian apple trees. Can. Ent. 78:15-22. [SAPRO-
GLYPHIDAEJ

I Oudemans, A.C. (1928). Acarologische aanteekeninger XCI. Lenziidae n. fam. Ent. Ber. 7(161) :327, 328.
[SAPROGLYPHIDAE]
Popovici Baznosanu, A. (1913). Etude biologique ur l'Acarien Trichotarsus osmiae Duf. Arch. Zool.

I Exp. Paris 52:32,41. [CHAETODACTYLIDAE]

Robertson, P. (1959). Revision of the genus Tyroph gus with a discussion on its taxonomic position in
the Acarina. Austral. Jour. ZooI. 7(2) :146-181. ACARIDAE]

I Robin, C. and M.P. Megnin (1877).

13:209-656 + plates. [HYPODERIDAEJ
Memoire sur J les Sarcoptides plumicoles.
'
Jour. Anat. Physio!.

Rupes, V. and J.O. Whitaker, Jr. (19681. Mites of title subfamily Labidophorinae (Acaridae, Acarina) in

I North America. Acarologia 10(3):493-499. [LA IDOPHORIDAE]

Samsinak, K. (1965). Termitophile Milben aus der V
294. [ACARIDAE]
China, 2. Acaridoidea. Reichenbachia 5(34):291-

I Solomon, M.E. (1~46), Tyroglyphid mites in stored prOducts. Nature and amount of damage to wheat.
Ann. Appl. BIOI. 33(3) :280-289. !

I
I
 I
238

Strandtmann, R.W. (1962). Nycteriglyphus bifolum n. sp., a new cavernicolous mite associated with bats
(Chiroptera) (Acarina: Glycyphagidae). Acarologia 4(4):623·631. [ROSENSTEINIIDAE]
I
Turk, .F. and F. Turk (1959). Systematik und Okoiogle der Tyroglyphiden Mitteluropas.
Okol. mitteleurop. Acarina 1(1) :3·231.
Beitr. Syst.
I
Van den Bruel, W.E. (19401. Un ravageur de I'epinard d'hiver: Tyroglyphus dimidiatus Herm, (fongior
Gervl. Bull. Inst. Agron. Gembloux 9(1-4):81-99. [ACARIDAE]
Woodring, J.P. (1966). North American Tyroglyphidae (Acari): I. New species of Calvolia and Nanacarus,
with keys to the species. Proc. Louisiana Acad. Sci. 29:76-84. [SAPROGLYPHIDAE]
I
Zakhvatkin, A.A. (1941). Fauna of U.S.S.R. Arachnoidea 6(11, Tyroglyphoidea (Acari).
Acad. Sci. U.S.S.R. N. S. 28 (translated by Amer. Inst. BioI. Sci.: 573 pp. + v).
Zool. Inst.
I
Superfamily Canestrinioidea
I
DIAGNOSIS: Soft-bodied; with a distinct prodorsal shield, sejugal furrow present or
absent; palpi simple. Apotele represented by a sucker-like stalked caruncle
I
with or without a minute empodial claw. Female genital opening longi-
tudinal, resembling an inverted "Y," genital discs present. Hypopal stage
infrequently encountered (HEMISARCOPTlDAE).
I
Three families are included here in the Canestrinioidea-the CANESTR IN II DAE, the
L1NOBIIDAE and the HEMISARCOPTIDAE. All are associated with insects.
I
Members of the family CANESTRINIIDAE are weakly sclerotized, moderately large I
(400-600 Il) mites which, because of long dorsal and leg setae, often appear quite hairy.
Canestrinilds generaliy favor protected areas beneath the elytra of beetles (Cooreman 1955),
but some have been coliected from gular or coxal cavities on beetle hosts. Most of the
I
described canestriniid species are tropical or subtropical in distribution, although both
Percanestrinia saetolata Cooreman and Photia chrysocarabi Cooreman were found on
carabid beetles in temperate climates (Cooreman 1950). Species of CANESTRINIIDAE are
I
known from Europe, Africa, Asia and South America primarily from lucanid, carabid,
passalid, and scarabaeid beetles. The feeding habits of canestriniids are unknown, but their
intimate association with their beetle hosts indicates a parasitic or commensal association.
I
Linobia coccinellae (Scopoli), an associate of a Mexican chrysomelid beetle, represents the
family LI NOBII DAE, a taxon closely related to the canestriniids. I
Hemisarcoptes malus (Shimer), one of three described species of the family HEMI-
SARCOPTIDAE, is a tiny (300 Il) unsclerotized mite which feeds upon scale insects and I
their eggs in North America and in Europe (Tothill 1918, Andre 1942). H. malus appears to
exert a significant degree of control on various scale species, although most observations on
natural suppression of scale populations by this species have been confined to the oyster-
I
she Ii scale, Lepidosaphes ulmi L. H. coccophagus Meyer preys on scales in South Africa.
The hypopus of H. cooremani (Thomas) was coliected from under the elytra of a coccineliid
beetle (Thomas 1961), on which it may travel to scale-infested habitats.
I
Useful References
I
Andre, M. (1942). Sur I'Hemisarcoptes malus Shimer (=coccisugus Lignieresl (Acariens). Bull. Mus. d' Hist.
Nat. Paris Ser. 2,14(3):173-180. [HEMISARCOPTIDAE]
I
I
I
I
239

I Cooreman, J. (1950). Etude de quelques Canestriniides (Acari) vivant sur des Chrysomelidae et sur des
Carabidae (Insecta, Coleoptera). Bull. Inst. Roy. Sci. Nat. Belg. 26(33):1-36.

I Cooreman, J. (1954).
33(13) :83-90.
Acariens Canestriniidae de la COllEjction A.C. Oudemans, a Leiden. Zool. Meded.

Cooreman, J. (1955). Acari. Exploration du Pare Na ional Albert. I. Mission G.F. de Witte, Fasc.

I 85:3-43. [CANESTRINIIDAE]
Thomas, H .A. (1961). Vidia (Coleovidia) cooremani, a n w subgenus and new species and notes on the life
history (Acarina: Saproglyphidae). Ann. Ent. Soc. Amer. 54(3):461-463. [HEMISARCOPTIDAE]

I Tothill, J.D. (1918). The predaceous mites, Hemisarcoptes malus Shimer, and its relation to the natural
control of the oyster-shell scale, Lepidosaphes ulmi L. Agr. Gazette Canada 5(3) :234·239.
Turk, F.A. (19481. Insecticolous Acari from Trinidad., B.W.1. Proc. Zoo I. Soc. London 118:82-128.

I [CANESTRINIIDAE] !

I Superfamily Listr~Phoroidea
DIAGNOSIS: Soft-bodied; strongly striated, US~aIIY
with distinct prodorsal or podonotal
I shield; mouthparts, legs I-II, IIJ.lV or anterior coxal region modified for
grasping hairs; empodial claws on Ilegs I-II absent. Female genital opening an
inverted "V" or nearly transverse, ~enital discs reduced or absent.
I ,
Listrophoroid mites are parasites of mammals, and are grouped here in four families-

I the L1STROPHOR IDAE, MYOCOPTI DAE, RHjfrNCOPTI DAE and CH I RORHYNCHOBI-

IDAE. McDaniel (1968a) recognizes two additi nal families, the LABIDOCARPIDAE and
ATOPOMELIDAE, which have been include under the L1STROPHORIDAE in this

I discussion.

Members of the L1STROPHORIDAE are/found attached to hairs of a variety of

I mammals, inclUding rodents, bats and carnivo es throughout the world.' Lemuroecius
cheirogalei Fain and Lemuroptes primarius LjWrenCe are parasites of lemurs in Africa
(Fain 1968, Lawrence 1958), while members f the atopomeline genus Austrochirus are
I found primarily on Australian marsupials (Do I'row 1958). Species of the genera Listro-
phorus, Listrophoroides, and Chirodiscoides (ar~ong others) are more or less restricted to

I rodents. Many of the chirodiscine listrOPhoridS'jhowever, are parasitic on bats (Pinichpongse

1963). The chirodiscine Schizocoptes conjugat s Lawrence is an exception, being found on
an African mole. Listrophorids probably feed 0 the sebaceous secretions which accumulate

I at the bases of the hairs to which the mites adhere.

The MYOCOPTI DAE are cosmopolitan darasites which are common inhabitants of
rodent fur. One species of Chrysocoptes La~lrence parasitizes an insectivore (M. cDaniel
I 1968), but the mite may be an analgoid (page r 43) rather than a listrophoroid. Myocoptes
musculinus (Koch) produces a mangy condi ion in laboratory mice which can cause
I problems in animals which have low resistance (Baker et al. 1956). Myocoptes
ondatrae Lukoschus and Rouwet is a related sdecies found on Ondatra zibethica in Europe
(Lukoschus and Rouwet 1968) and possibly in ~orth America.
I 1Chirodiscus amplexans (Trouessart and NeumaJ ) was collected from a bird, and was considered a

I possible member of the Analgoidea (page 2431. MtDaniel (1968b) has established its identity as a
listrophorid, and suggests that it is a mammal parasitr which was fou~d in accidental association with a
bird.

I
I
240
I
The family RHYNCOPTJDAE is a distinctive monogeneric group of four Iistrophoroid I
mite species which parasitize monkeys in South America and Africa (Fain 1965) and
porcupines in South Africa (Lawrence 1956). While the specific location of Rhyncoptes
recurvidens Lawrence on its porcupine host is not known, the species of Rhyncoptes from
I
monkeys are found embedded in the hair follicles of their hosts. Rhyncoptids may produce
mange, but such an association has not been illustrated. I
Chirorhynchobia urodermae Fain, the sole species of the family CH I RORHYNCHOBI-
IDAE, is a parasite of a South American bat (Fain 1967, 1968). Only one specimen of this
bizarre mite has been collected.
I
Useful References
I
Baker, E.W., T.M. Evans, D.J. Gould, W.B. Hull and H.L. Keegan (1956). A Manual of Parasitic Mites of
Medical or Economic Importance. Natl. Pest Control Assoc. Tech. Publ.: 170 pp. [MYOCOPTIDAE,
I
L1STROPHORIDAE]
Damrow. R. (195B). A summary of the Atopomelinae (Acarina, Listrophoridael. Proc. Linn. Soc. N. S. W.
83:40·54.
I
Fain, A. (1965). A review of the family Rhyncoptidae Lawrence parasitic on porcupines and monkeys.
Advances in Acarology 2: 135-159.
Fain, A. (1967). Diagnoses d'Acariens Sarcoptiformes nouveaux. Rev. Zool. Bot. Afr. 75(3·4):378-382.
I
[CHI RORHYNCHOBIIDAE]
Fain. A. (1968). Notes sur trois Acariens remarquables (Sarcoptiformes).
[CHIRORHYNCHOBIIDAE]
Acarologia 10(2):276-291- I
Lawrence, R.F. (1956). Studies on South African fur mites (Trombidiformes and Sarcoptiformes). Ann.
Natal Mus. 13:337-375. [RHYNCOPTIDAE]
Lawrence, R.F. (1958). Studies on the Iistrophorid fur-mites of Madagascar iemurs. Mem. Inst. Sci.
I
Madagascar 12A:113-125. [LABIDOPHORIDAE]
Lukoschus, F.S. and J.G.J.H. Rouwet (19681. Mycoptes ondatrae spec. nov., ein neuer parasit von Ondatra
zibethica L. (Listrophoridae: Sarcoptiformes). Acarologia 10(3):483·492. [MYOCOPTIDAE]
I
McDaniel. B. (1968a). The superfamily Listrophoroidea and the establishment of some new families
(Listrophoroidea: Acarina). Acarologia 10(3):477·482. [L1STROPHORIDAE. MYOCOPTIDAE,
RHYNOCOPTIDAE]
I
McDaniel, B. (1968b). The genus Chirodiscus Trouessart and Neumann with lectotype designation of
C. amp/exans Trouessart and Neumann (Listrophoroidea: Atopomelidae). Acarologia 10(4):653-656.
[L1STROPHOR IDAE]
I
Pinichpongse. S. (1963). A review of the Chirodiscinae with descriptions of new taxa (Acarina: Listro·
phoridae). Acarologia 5(1) :81-91, 5(2) :266-278, 5(3) :397~404, 5(41 :620-627.
Watson, D.P. (1960). On the aduit and immature stages of Myocoptes musculinus (Koch) with notes on its
I
biology and classification. Acarologia 2(3):335-344. [MYOCOPTIDAEJ
Zumpt, F. (ed.) (1961). The Arthropod Parasites of Vertebrates in Africa South of the Sahara (Ethiopian
Region). Vol. I. (Chelicerata). Publ. So. Afr. Inst. Med. Res. 50(11):457 pp.
I
Superfamily Ewingoidea
I
DIAGNOSIS: Soft-bodied,' without distinct dorsal shield or sejugal furrow; chelicerae
narrowed, divided terminally; palps simple. With empodial claws on tarsi
I
1-1/; legs /II-I V enlarged, with large hook-like empodial claws. Female
genital opening longitudinal, genital discs indistinct. I
I
I
I 241

I A single unusual species, Ewingia coenOiitae Pearse, is included in the family

EWI NG IDAE, the only suprageneric taxon of 1he Ewingoidea. E. coenobitae attaches
I to the gills of
rather rare, E.
pagurid crabs in Florida (Pearse' 929) and the Antilles. Once considered
coenobitae has been collected in large numbers from Coenobita clypeatus
in Florida. I
I i

I Useful References

Pearse, A.S. (1929). Two new mites from the gills of iarld crabs. Carnegie Inst. Wash. Pub!. 391 :225·230.

I
Superfamily PsOroptoidea
I DIAGNOSIS: Soft-bodied, but commonly with l~eaklY sclerotized podonotal, opisthonotal
and lateral shields; without sejug'll furrow; chelicerae chelate or stylet-like,
I weakly developed; palps simple. ~poteles, when present, each consisting of
a stalked terminal or subterminal sucker; tarsi III-I V normal or terminating
in long whip-like setae, with or ithout empodial sucker. Female genital
I opening an inverted "V," "Y" or "U" shape; males usually with anal
suckers, often with enlarged legs II I.

I Five of the six families of Psoroptoidea arel associated with mammals as true parasites.
The sixth famiiy, PYROGLYPHIDAE, assumes a variety of habits and habitats, none of

I which are known to be parasitic. Some psor ptoid species are economically important
pests.
1
The family PSO ROPTI DAE includes sever~1 species and species varieties which may
I cause injury to their mammal hosts. PsoroPlles equi (Hering) and the varieties of this
species parasitize horses, cattle, sheep, goats, el~' rabbit and mountain sheep (Baker et al.
I 1956), often causing a serious mange condition on the body or head of the host. Psoroptic
mange in sheep may be fatal. Dogs, cats and 0 her small carnivores often are attacked by
the ear mite, Otodectes cynotis (Hering). Fever and depression often result from heavy
I infestations of the ears. Chorioptic mange of sleep, goats, horses and rabbits results from
skin infestations of Chorioptes bovis (Gerlac). Although not usually as serious as

I psoroptic mange, heavy infestations of Chorioptes may cause intense irritation in affected

I
animals (Hirst 1922). Chorioptic mange symptoms are considerably more restricted than
those of psoroptic mange, being confined primJrily to the lower portions of the legs and to

I the root of the tail. Sweatman (1957) synOny;riZed the several species and varieties of the
e. bovis "complex" on the basis of his discover of non-specificity in these morphologically
identical forms. e. erewi Lavoipierre, a parasit of the African duiker, is a distinct species.
I Species of the psoroptid genus Paracoroptes arasitize monkeys and gorillas in Africa
(Zumpt 1961) while Psoralges libertus Trouess- rt appears to cause mange symptoms in its
edentate host, the tamandua, in South Americ . Other species of the psoroptid subfamily
I Psoralginae also are found on edentates (Fonsec11954, Fain 1965).

South American edentates of the familJ Bradypodidae serve as hosts for Lobalges
I trouessarti Fonseca 1954, and Edentalges cifoloePi Fain, representatives of the family
LOBALGIDAE (Fain 1965a). Lobalgids live in the skin of their hosts.

I
I
 I
242

The families YUNKERACARIDAE and LEMURNYSSIDAE include species which are I

found in the respiratory passages of mammals. Yunkeracarus muris Fain and Y. faini
Hyland and Clark parasitize rodents in Africa and in North America (Fain 1957, Hyland and
Clark 1959). The yunkeracarid Sciuracarus paraxeri Fain has been collected from the nasal
I
fossae of a sciurid rodent in the Transvaal (Fain 1964b). Lemurnyssus galagoensis Fain
(family LEMURNYSSIDAE) is an inhabitant of the nasal fossae of a lemur in Africa (Fain
1957), while lemurnyssids of the genus Mortelmansia infest the nasal passages of monkeys
I
in South America (Fain 1959).

Species of Audycoptes and Saimiroptes (family AUDYCOPTIDAE) are found in the

I
lip tissues of South American squirrel monkeys (Lavoipierre 1964, Fain 1968). where they
feed on sebaceous materials around sinus·hair and normal hair follicles. I
Approximately 16 species of PYROG L YPH IDAE have been described, and these are
grouped into two subfamilies-the Pyroglyphinae and the Dermatophagoidinae (Fain 1967a). I
Pyroglyphine pyroglyphids are found in the nests of birds (Bontiella bouilloni Fain is an
example) or rodents (Pyroglyphus mor/ani Cunliffe). Others are found living in high
protein substrates such as cottonseed cakes (Euroglyphus maynei (Cooreman)) or fish meal
I
(Pyroglyphus (Hughesiella) africanus (Hughes)) in many parts of the world. Euroglyphus
(Gymnoglyphus) longior Trouessart) is a common contaminant of grain storages (Hughes
1961). Dermatophagoidine pyroglyphids are, with some exceptions, found in house dust,
I
and one species, Dermatophagoides pteronyssinus (Trouessart) has been impl icated as the
causal agent of house dust allergy in the Netherlands (Spieksma and Spieksma . Boezeman
1967). D. pteronyssinus also is found in Africa, Brazil, India and North America (Fain
I
1967b). and was the causal agent of a scalp condition in man reported by Traver (1951).
Species of Dermatophagoides are common in bird nests as feeders on detritus, and are I
occasionally encountered on the feathers of their "hosts" (Gaud 1968). Malayoglyphus
intermedius Fain, Cunnington and $pieksma is an inhabitant of house dust in the Asiatic·
Pacific region (1969). Other species of the subfamily are found in bird or mammal nests
I
or in stored products. The genera Hullia, Onychalges, Paralgopsis and Paramealia are bird
associates (Gaud 1968) with Onychalges and Paralgopsis having been collected from bird
feathers with enough regularity to suggest a possible parasitic association.
I
Useful References
I
Baker, E.W., T.M. Evans, D.J. Gould, W.B. Hull and H.L. Keegan (19561. A Manual of Parasitic Mites of
Medical or Economic Importance. Nat!. Pes! Control Assoc. Tech. Pub!.: 170 pp.
I
Fain, A. (957). Notes sur /'acariase des voies respiratoires chez I'homme et les animaux. Description de
deux nouveaux Acariens chez un lemurien et des rongeurs. Ann. Soc. Beige Med. Trap. 37(4) :469-
482. [LEMURNYSSIDAE, YUNKERACARIDAE)
I
Fain, A. (1.959). Deux nouveaux acariens nasicoles chez un singe platyrhinien Sa/mir! sciurea (L.). Bull.
Soc. Roy. Zool. Anvers 12:3·12. [LEMURNYSSIDAE)
Fain, A. (1964a). Les Lemurnyssidae parasites nasicoles des Lorisidae africains et des Cebidae sud-
I
amekicains. Description d'une espece nouvelle (Acarina: Sarcoptiformes). Ann. Soc. BeIge Med.
Trop. 44(3) :453-458.
Fain, A. (1964b). Chaetotaxie et classification des Gastronyssidae avec description d'un nouveau genre
I
parasite nasicole d'un Ecureuil sudafricain (Acarina: Sarcoptiformes). Rev. Zool. Bot. Afr.
70(1·2):40·52. [YUNKERACARIDAE)
I
I
I
I
243
I Fain, A. (1965al. Les Acariens producteurs de gaie che les Edentes et les Marsuplaux (Psoroptldae et
Lobalgidae: Sarcoptlformes). Buli. Inst. Roy. Sci. Ni' Belg. 41 (171 :1·41.
I Fain, A. (1965b). Les Acarlens nldlcoles et detritlcoles de la famille Pyroglyphldae Cunliffe (Sarcopti·
formes). Rev. Zooi. Bot. Afr. 72(3-41 :267-288.
Fain, A. (1967a). Deux nouvelles especes de Dermatoph goidinae rattachement de cetta sous·famille aux
I Pyroglyphidae (Sarcoptlformes). Acarologla 9(4) :87 ·88L
Fain, A. (1967b). Le genre Dermatophagoides 80gdanov 1864 son importance dans les allergies
respiratolres et cutanees chez I'homme (Psoroptidte: Sarcoptiformes). Acarologia 9(1 I: 179-226.

I [PYROGLYPHIDAEJ
Fain, A. (1968). Notes sur trois acarlens remarquable
[AUDYCOPTIDAE]
(Sarcoptiformes). Acarologia 10(2):276·291.

I Fain, A., A.M. Cunnington and F.Th.M. Spleksma (1969)1. Malayoglyphus intermedIUS n.g" n. sp.• a new
mite from house dust In Singapore and DJakart~ (Pyroglyphldae' Sarcoptiformes). Acarologla
11(1) 121·126

I da Fonseca, F. (1954). Notas de Acarologla XXXIX. SI tematlca e fllogenese de Psoralgldae Oudemans.

Sarcoptlformes parafaglstas de mamlferos com
Butantan 26:92-167. [PSOROPTIDAE. LOBALGldAEj
orphologla de Acan plumicolas. Mem. Inst.

Gaud, J. (1968), Acarlens de la sous-famille des Der~atophagoidinae (Psoroptidael recoltes dans les
I plumages d'oiseaux. Acarologla 10(2) :292-312. [PVROGLYPHIDAE]
Hirst, S. (19221. Mites injurious to domestic anlmals Brit. Mus. (Nat. Hlst.) Econ. Ser. 13:107 pp.

I Hughes, A.M. (1961 I. The Mites of Stored Food. Minlstl y Agr. Fish. and Food Tech. 8ui. 9:287 pp. + vi.
1
Hyland, K.E., Jr. and D.T. Clark (1959). The occurre ce of the genus Yunkeracarus in North America
(Acarina: Epldermoptidae). Acarologia 1(3):365-36H. [YUNKERACARIDAE]
a~d female of Chorioptes crewei Lavolpierre 1958
I Lavoiplerre, M.M.J. (1959). A description of the male
(Acarina: Psoroptidae), together with some rema ks on the family Psoroptidae and a key to the
genera contained In the family. Acarologia 1(3) :35 -364.

I Lavoiplerre, M.M.J. (1964). A new family of Acari es belonging to the Suborder Sarcoptiformes
parasitic in the hair follicles of Primates. AnniNatai. Mus. 16:191·208. [AUDYCOPTIDAE]
Spieksma, F.Th.M. and M.I.A. Spieksma-Boezeman (196 I. The mite fauna of house dust with particular

I reference to the house-dust mite Dermatophago;d s pteronyssinus (Trouessart, 1897) (Psoroptidae:

Sarcoptiformes). Acarologia 9(11 :226-241. [PYRClGLYPHIDAEj
Sweatman, G.K. (1957). Life history, non·specificity ~nd revision of the genus Chorioptes, a parasitic
mite of herbivores. Can. Jour. Zool. 35:641-689. [rSOROPTIDAEJ
I Traver, J. (19511. Unusual scalp dermatitis In humans crUSed by the mite Dermatophagoides. Proc. Ent.
Soc. Wash. 53(11:1·25.
Zumpt, F. (ed.1 (1961). The Arthropod Parasites of Ve tebrates in Africa South of the Sahara (Ethiopian
I Regionl. Vol. I. (Chellcerata). Publ. So. Afr. Inst. Med. Res. 50(11) :457 pp.

Superfamily Atlgoidea

I DIAGNOSIS' Soft·bodied but commonly with ieaklY sclerotized podonotal, opisthonotal

and lateral shields/ without sei~gal furrow/ chelicerae generally chelate,
I weakly developed, fixed digit ma( be reduced/ palps simple. Apoteles each
consisting of a stalked empodial sucker with or without a minute claw,
inserted terminally or subtermi1ally on tarsi. Female genital opening an
I inverted "U" or "V" shape, or ,transverse/ males with anal suckers, often
with enlarged legs III. I
I Six families of Analgoidea are recognized n this treatment, of which all but one are
external forms inhabiting the feathers or skin of their bird hosts. The TURBINOPTIDAE
I
I are respiratory parasites of birds.

The ANALGIDAE Includes no fewer than fwen tv genera of weakly sclerotized feather

I i
I
244
I
mites, representatives of which may be found adhering to the feathers of a wide variety of I
bird species throughout the world (Zumpt 1961). Dubinin (1951) observed that analgids,
as well as other analgoid feather mites, may show distinct preferences for types of feathers
infested, and for particular locales on the feathers. Species of Analges, for example, favor
I
the basal portions of the quill feathers while Mesalges prefers the primary feathers.
Megninia cubitalis (Megnin) and M. ginglymura (Megnin), along with other species of the
genus, infest the feathers of domestic fowl, parakeets and pigeons. M. cubitalis is a
I
common parasite of chickens in Europe and Africa (Zumpt 1961). It is not, however,
considered to be economically important. I
Mites of the families FREYANIDAE and DERMOGLYPHIDAE are feather mites
which generally exhibit strong sclerotization and/or distinctive ventral apodemes, and I
which are common primarily on wild birds. Like the ANALGIDAE, the many species of
FREYAN IDAE are relegated to a large number of genera (Dubin in 1953, Gaud and
Mouchet 1957), the majority of which favor sites on the leading edges of the wings of
I
large birds. Cormorants, ibis and spoonbills are typical hosts.

Species of several genera of the DERMOG LYPHI DAE are found within the feather
I
quills of hosts, rather than on the feather surfaces. Syringobia, Dermoglyphus and
Oxyalges are typical dermoglyphid quill invaders. Dermoglyphus elongatus (Megnin)
commonly invades the quills of domestic fowl, with infestations of canary feathers also
I
having been noted (Baker et al. 1956). Feather surface parasites include species of
Pterophagus and Falculifer. Pterophagus species are particularly common on feathers of
doves, as are the known species of Falculifer (Zumpt 1961).
I
The family PROCTOPHYLLODIDAE is a large, extremely common family of I
feather mites, the members of which are most often encountered as associates of passeriform
birds. The number of described species may be illustrated by the example of one genus,
Proctophyllodes, in which 123 species are included in the latest generic review (Atyeo and
I
Braasch 1966). Like some dermoglyphids, proctophyllodids are found between the
feather ribs, either remaining motionless or moving within the confines of the interstice, or
crossing between interstitial grooves in a crab-like fashion. Proctophyllodid mites apparently
I
are scavengers and inflict no noticeable injury on their hosts, even under conditions of
massive infestation. I
Mites of the family TURBINOPTIDAE live in the nasal fossae of birds (Fain 1957) in
both the Old and New Worlds. Although they may be found in large numbers in infested
birds, there is no record of harmful effects occasioned by these infestations.
I
The EPIDERMOPTIDAE are usually to be encountered on the skin of domestic I
birds (Zumpt 1961). Epidermoptes bilobatus Rivolta, a common skin parasite of domestic
fowl, has also been found under the skin pellicule (Fain and Evans 1963), and was reported
as causing pityriasis in chickens (Baker et al. 1956). Similarly, an infestation of E.
I
odontophori Fain and Evans on an African bird was accompanied by a generalized
superficial mange on the body of the host. Rivoltasia bifurcata (Rivolta) attacks domestic
fowl, while R. dermicola is found on the European sparrow. Infestations of R. bifurcata
I
in chickens cause intense itching and scurfiness, particularly around the head.
I
I
I
I
I I 245

•
Useful References I

Alyeo, W.T. {19661. A new genus and six new species ol feather mites primarily from Tyranni (Acarina:
Proctophyllodidae). Jour. Kansas Ent. Soc. 39(3) :1~l1-492.
Atyeo, W.T. (1967). Two feather mite genera with polymorphic males (Analgoidea: Proctophyllodidael.

I Jour. Kansas Ent. Soc. 40(41 :465-471. I

Atyeo, W.T. and N.L. Braasch (1966). The feathet mite genus Proctophyllodes (Sarcoptlformes:
Proctophyllodidae). Bull. Univ. Nebraska State Mu,. 5:1·354.

I I
Atyeo, W.T. and J. Gaud (1966). The chaetotaxy of ,arcoptiform feather mites (Acarina: Analgoidea).
Jour. Kansas Ent. Soc. 39(21 :337·346.
Baker, E.W., T.M. Evans, D.J. Gould, W.B. Hull and H.l. Keegan (1956). A Manual of Parasitic Mites of

I Medical or Economic Importance. Natl. Pest ContTI Assoc. Tech. Publ.: 170 pp.
Boyd, E.M. (1949). A new genus and species of mi~e from the nasal cavity of the ring-billed gull,
(Acarina, Epidermoptidael. Jour. Parasit. 35(3):295-300. [TURBINOPTIDAEJ
Dubinin, W.B. (1951). Feather mites (Analgesoidea). P~rt I. Introduction to their study. Fauna U.S.S.R.
I 6(5):1·363. :
i
Dubinin. W.B. (1953). Feather mites (Analgesoidea). Part II. Epidermoptidae and Freyanidae. Fauna
!
I U.S.S.R. 6(6):1·411.
Dubinin, W.B. (1956). Feather mites (Analgesoidea). ~art III. Pteroliehidae. Fauna U.S.S.R. 6(7): 1-814.
[DERMOGLYPHIDAE] i
Fain, A. (1957), Les Acariens des families EPidermOP~~idae et Rhinonyssidae parasites des fosses nasales
I d'oiseaux au Ruanda·Urundi et au Congo beige.: Ann. Mus. Roy. Congo beige Ser. 8, 60:176 pp.
+ xi. [EPIDERMOPTIDAE, TURBINOPTIDAEJ [
Fain, A. (1965). A review of the family EPidermoPtid~ie Trouessart parasitic on the skin of birds. Parts
I I-II. Verhandl. Konin. Vlaam. Acad. Wetensch. Be g. 27(841:1·176; 1-144.
Fain, A. and G.O. Evans (1963). On three mites of He genus Epidermoptes Rivolta (Acari). Ann. Mag.
Nat. Hist. Ser. 13,6:595·608. .
I Gaud, J. and J. Mouchet (1957). Acariens Plumie~les (Analgesoideal des oiseaux du Cameroun. I.
Proetophyllodidae. Ann. Parasit. Hum. Compo 32l5-6) :491-546.
Poppe, E. (196.7). Die Begattung bei den Vogelhilben Pterodectes Robin (Analgesoidea. Acaril.
I Z. Morph. Okol. Tiere 59:1-32. [PROCTOPHYL\LODIDAEJ
Radford, C.D. (1958). The host· parasite relationships of the feather mites (Acarina: Analgesoidea).
Rev. Brasil. Ent. 8:107·170. [
I Zumpt, F. (ed.1 (1961). The Arthropod Parasites of tertebrates in Africa South of the Sahara (Ethiopian
Region). Vol. I. (Chelieeratal. Publ. So. Afr. In~t. Med. Res. 50(11) :457 pp.

I Superfamily S!arcoptoidea

DIAGNOSIS: Soft·bodied, with prodorsal sfiield and anterolateral apodemes sometimes

I present, commonly rounded cjr sac-like in shape, without sejugal furrow;
chelicerae reduced, chelate, parips simple. Legs telescoped, terminating in
empodial suckers or a series of terminal spine-like setae; legs I V occasionally
I absent. Female genital opening a transverse slit.

maml~als
I Skin parasites of both birds and
families are recognized, of which two are of
are found in the Sarcoptoidea.
k~own economic importance.
:
Four

The SARCOPTI DAE live in or on the s~in of mammals of many types, including man.
I Sarcoptes scabiei (DeGeer) invades the skin ~1 man and other animals, and causes sarcoptic
mange or scabies (Mellanby 1943). Scabies if: highly contagious in crowded conditions and

I is recognized by accompanying scabby, itch'ng lesions which may persist for some time.
Other races-or possibly distinct species-of ,Sarcoptes cause scabies in domestic animals,
!
I
I
246
I
monkeys and camels (Baker et al. 1956), producing symptoms varying in severity. Sarcoptic
mange in cattle or sheep occasionally results in death of the host animal. Notoedres cati
I
(Hering), the common mange mite of cats and rodents, causes face mange which in rabbits
may spread to the hind quarters as well. N. alepis (Railliet and Lucet) attacks rats, as does I
N. muris (Megnin). Sarcoptid parasites of bats are found in the genera Notoedres,
Chirnyssus and Nycteridocoptes (Fain 1959a, 1959b, 1965). Species of Notoedres are
found in the skin of their hosts, and are more or less cosmopolitan. The latter two genera
I
are primarily tropical and may be collected from the buccal mucosa, with species of
Nycteridocoptes also occurring on the ears and on the wing membranes of their hosts. I
Mites of the family KNEMIDOCOPTIDAE are skin inhabitants of birds which
produce various skin disorders in their hosts through their feeding. The scaly-leg mite,
Knemidocoptes mutans (Robin and Lanquetin) causes a condition on the legs of galliform
I
birds which may finally cripple the host. Chickens are commonly attacked, as are pheasants
and turkeys. The depluming mite, Neocnemidocoptes laevis gallinae (Railliet) burrows into I
the skin at the base of the feathers of chickens and other galliform birds, bringing about
intense irritation and feather pulling by the host bird (Baker et al. 1956). Pigeons are
attacked in similar fashion by N. laevis laevis (Railliet) (Fain 1967). Various species of I
Knemidocoptes attack specific areas in passeriform birds, causing varying degrees of
irritation to their hosts. K. jamaicensis Turk burrows into the legs of its host (Turk 1950)
while K. rossor (Ehlers), a similar species, tunnels at the base of the bill. K. pilae Lavoi-
I
pierre and Griffiths attacks psittaciform birds and may precipitate a scaly-leg condition
(Lavoipierre and Griffiths 1951). The same species is found on other parts of the bird also. I
The TEINOCOPTIDAE are skin parasites of bats which generally show considerable
reduction in development of legs III and/or IV (Fain 1962a). Teinocoptes and Chirobia
species are found on or in the skin of the wing of Megachiroptera, while Bakerocoptes
I
cynopteri Fain, the type and only species of the genus, is found in subcutaneous pouches
(in all stages) of the Malaysian bat Cynopterus brachyotis angulatus. I
A single species represents the family EVANSACARIDAE-Evansacarus lari Fain
(1962b) found in the skin of the head of Larus canus (a gull) in Israel. E. lari may
I
consitute a subfamily of the KNEMIDOCOPTIDAE (Fain 1967).
I
Useful References

Baker, E.W., T.M. Evans, D.J. Gould, W.B. Hull and H.L. Keegan (1956). A Manual of Parasitic Mites of
I
Medical or Economic Importance. Nat!. Pest Control Assoc. Tech. Pub!.: 170 pp.
DUbinin, W.B. (1953). Feather mites (Analgesoidea). Part II. Epidermoptidae and Freyanidae. Fauna
U.S.S.R. 6(6):1-411. [KNEMIDOCOPTIDAEj
I
Fain, A.((1959a). Les Acariens psoriques parasites des Chauve-souris: II. Chirnyssus myoticola n.g., n.
sp. parasite du murin Myotis myotis (Borkh) en Belgique. Acarologia 1(1): 119-123. [SARCOP-
TJDAEj
I
Fain, A. (195gb). Les Acariens psoriques parasites des Chauves-sQuris VI. Le genre Prosopodectes
Canestrini 1897 est composite et do it tomber en synonymie de Notoedres Railliet 1893. Acarologia
1(3):324-353. [SARCOPTIDAEj I
Fain, A. (1962a). Les Acariens psoriques parasites des Chauves-sQuris XXII I.-Un nouveau genre hexapode
a taus les stades du developpement (Teinocoptidae: Sarcoptiformes). Bull. Ann. Soc. Roy. d'Ent.
Belg. 98(28) :404-412. I
I
I
I I
[ 247

I Fain, A. (1962b). Un acarien remarquable combinantjes caracteres de plusieurs families: Evansacarus

lari n.g.. n. sp. (Evansacaridae nov. fam.: Sarco tiformes). Bull. Ann. Soc. Roy. d'Ent. Belg.

I 98(9):125·145.
Fain, A. (1964). Le developpement postembryonnaire c ez les Acaridiae parasites cutanes des mammiferes
et des oiseaux (Acarina: Sarcoptiformes). Bull. Classe Sci., Acad. Roy. Belg. Ser. 5, 50:19-34.

I Fain, A. (1965). Notes sur Ie genre Notoedres Railliet, 1893 (Sarcoptidae: Sarcoptiformes). Acarologia
7(2) :321-342. :
Fain, A. (1967). Les acariens de la famille Knemid(~koPtidae producteurs de gale chez les oiseaux

I (Sarcoptiformes). Acta Zoo I. Path.Antwerp. 45:3-115. [KNEMIDOCOPTIDAE, EVANSACARIDAEJ

Fain, A. (1968). Etude de la variabilite de Sarcoptes scaiiei avec une revision des Sarcoptidae. Acta Zool.
Path. 47:196 pp.
Lavoipierre, M.J.J. and R.B. Griffiths (1951). A prelirrinary note on a new species of Cnemidocoptes
I (Acarina) causing scaly-leg in a Budgerigar (Mel0l'jsittacus undulatus) in Great Britain. Ann. Trap.
Med. Parasit. 45(3-4) :253-254. I
Mellanby, K. (1943). Scabies. Oxford War Manuals, Uni~. Press: 81 pp.

I Turk, F.A. (1950). A new species of parasitic mite, :Cnemidocoptes jamaicensis, a causative agent of
scaly-leg in Turdus aurantiacus. Parasit.40:60-62. I

I Superfamily CY}Oditoidea

DIAGNOSIS: Soft-bodied, but occasionally wi~h weakly sclerotized podonotal, opistho-

I notal and/or ventral shields, wiih few body setae; idiosoma rounded or
elongate, without sejugal furrol!'; chelicerae chelate or greatly reduced,
palps simple. Apoteles present or absent, generally consisting of a stalked
I empodial caruncle. Female ger!ital opening a longitudinal slit, a narrow
inverted "V," or transverse. '

I The four families included in the Cytodit~idea

are subdermal, respiratory or visceral
parasites of domestic fowl, bats or rodents. ITwo of these families are represented by
species which are of veterinary importance. '
I Cytodites nudus (Vizioli), a representative lof the family CYTODITI DAE, invades the

I peritoneum of chickens, canaries and other ['birdS (Baker et al. 1956), with primary
infestations occurring in the air sacs. Death of infested hosts may occur when large
numbers of mites are present. Cytodites bal' ksi (Wellman and Wherry) was recovered

I from the lungs of a ground squirrel (Wellman ard Wherry 1910). Other species have been
taken from the nasal fossae of a variety of birdS (Fain and Bafort 1964). The fowl cyst
mite, Laminosioptes cysticola (Vizioli) (family LAMINOSIOPTIDAE) is a subcutaneous
I parasite in many domestic birds, and is fou1d in the skin of the neck, breast, flank,
vent and thighs. Small cysts develop around ~he mites following their death, and these
often are visible as small lumps under the skin ;of the infested bird (Lindquist and Belding
I 1949). Heavy infestations of L. cysticola may cause death. A second species, L. hymenop-
terus Jones and Gaud, is found on the skin ard feathers of the eastern crow (Jones and
Gaud 1962). i
I i
The PNEUMOCOPTIDAE are species whf"Ch inhabit the respiratory tract of rodents.

I A male prairie dog which had died of acute b onchopneumonia was found to be infested
by Pneumocoptes penrosei (Weidman) (Weid an 1917). P. jellisoni occupies the lung
tissue of a species of Peromyscus (Baker 195~), but no damage to lung tissue has been

I noted. '

I
I
248
I
The GASTRONYSSI DAE is a group of four genera which are found in the stomach, I
intestines, nasal cavities or eyes of bats. Gastronyssus is an African genus represented by
G. bakeri, a stomach parasite of fruit bats (Fain 1955). Rodhainyssus yunkeri Fain occurs
in the nasal passages of its bat hosts (Fain 1956) while species of Opsonyssus and Mycter-
I
onyssus invade the nasal cavities and eyes of Mega- and Microchiroptera.
I
Useful References

Baker, E.W.. (1951). Pneumocoptes, a new genus of lung·inhabiting mite from rodents (Acarina:
I
Epidermotidae). Jour. Parasit. 37(6):583·586. [PNEUMOCOPTIDAEj
Baker, E.W" T.M. Evans, D.J. Gould. W.B. Hull and H.L. Keegan (1956). A Manual of Parasitic Mites of
Medical or Economic Importance. Natl. Pest Control Assoc. Tech. Publ.: 170 pp.
I
Fain, A. (1955). Un acarien remarquable vivant dans j'estomac d'une chave-souris: Gastronyssus baked
n.g., n. sp. Ann. Soc. Beige Med. Trap. 35(6) :681-688.
Fain, A. (1956). Una nouvelle famille d'acariens endoparasites des chauves-souris: Gastronyssidae fam.
I
nov. Ann. Soc. Beige Med. Trap. 36(1):87·98.
Fai~. A. (1960). Revision du genre Cytodites (Megnin) et description de deux especes et un genre
nouveaux dans la famille Cytoditidae Oudemans. Acarologia 2(2) :238·249.
I
Fain, A. and J. Bafort (1964). Les Acariens de la famille Cytoditidae (Sarcoptiformes). Description de
sept especes nouvelles. Acarologia 6(3) :504·528.
Jones, J., Jr. and J. Gaud (1962). The description of Laminosioptes hymenopterus n. sp. (Sarcoptiformes)
I
from the American crow. Acar%gia 4(3):391·395.
Lindquist, W.D. and R.C. Belding (1949). A report on the subcutaneous or flesh mite of chickens.
Michigan State Call. Vet. 10:20-21. [LAMINOSIOPTIDAEj
I
Weidman, F.D. (1917L Cyto!eichus penrosei, a new arachnoid parasite found in the diseased lungs of a
prairie dog. Cynomys ludovicianus. Jour. Parasit. 3:82·89. [CYTODITIDAEj
Wellman. C. and W.B. Wherry (1910). Some new internal parasites of the California Ground Squirrel
I
(Otospermophilus beecheyi). Parasit.3:417·422. [CYTODITIDAEj
Zumpt. F. (ed.) (1961). The Arthropod Parasites of Vertebrates in Africa South of the Sahara (Ethiopian
Region). Vol. I. (Chelicerata). Publ. So. Afr. Inst. Med. Res. 50(11):457 pp.
I
I
I
I
I
I
I
I
I
I
I 249

I Suborder Ast/gmata
I
(Plates 98 to 127, ~p. 256-285)
I KEYTOTHE F~MILIES
I I

1. Free-living or associated with insects or ver1tebrates; usually soft-bodied. With two

I pairs of distinct genital discs or ring-like strGctures (may be small as in Plate 101-4).
Caruncle (Fig. 26, p. 229) and empodial!c1aw present on at least some of the
tarsi (sometimes minute but seldom absent).! Supercohort ACARIDIA 2
I Parasitic on birds, mammals, crustaceans 01 insects, rarely nidicolous or free-living;
soft-bodied or with sclerotized plates. Genital discs greatly reduced or absent.

I Caruncles present or absent (if present, a pearing sucker-like (Fig. 27, p. 229); if
absent, terminus of tarsi with claw-like setae or spines (Plate 121-4)) .

I 2.
1
. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . " ..... Supercohort PSOROPTlDIA

Female genital aperture a transverse slit; wi~h two pairs of large ring-like structures,
16

generally not adjacent to genital opening. I' Caruncles sessile, reduced. Palpi with
I terminal segment generally oriented laterall and with two distal flagella. Free-living
in moist habitats Superfamily ANOI TOI DEA, (Plate 98) Family ANOETI DAE

I Female genital silt an inverted "Y" (Plate 102-2); genital discs large or small, always
closely associated with the longitudinal opel/ling. Caruncles on pretarsi not sessile or,
if sessile, not noticeably reduced. Palpi vari?us, but not as above 3

I or associated with, insects "

i
3. Caruncles sucker-like although weak claw mby be present (Plate 100-2). Found on,
1 .. Superfamily CANESTRINIOIDEA 4
I Caruncles rarely sucker-like, empodial claws distinct. Free-living or associated with
vertebrates Superfamily ACAROIDEA 1 6

I 4. Tarsi with narrow sucker-like caruncles; caws absent. Genital and anal openings
contiguous in female and nearly contiguOL s in male, at posterior end of idiosoma.

I Predators of scale insects ,. (Plate 99) Family HEMISARCOPTIDAE

Tarsi with large stalked caruncles, c1awJ small or distinct. Genital and anal
openings not contiguous. Found on insects!. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 5
I 5. Chelicerae without fixed digit, movable di9if well developed, dentate
I

.
.......................................... (Plate 100) Family L1NOBIIDAE
I Fixed cheliceral digit present but sometimet reduced
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 'j' ..
.
(Plate 100) Family CANESTRINIIDAE

I 6. With a bilobed membranous pseUdorutella'fprocess on the mediodistal aspect of the

subcapitulum. With two setae (one media and one lateral) at the base of the fixed
I cheliceral digit. Associated with bats (Plate 101) Family ROSENSTE IN II DAE

I 'The HYPODERIDAE, which occur in connectiJe tissue of birds and rodents while in the hypopal
stage, are free-living as adults and are considered a partlOf the Acaroidea. Through an oversight, the adults
have not been included in this key. 1

I
I
 I
250

Bilobed subcapitular appendage absent. With only one seta on the median aspect of
I
the chel icera. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 7

7. Female genital aperture covered by a pair of lateral lightly sclerotized shields

I
forming a crescent. Dorsal body setae minute. Males with tarsal and anal suckers...
· (Plate 101 [see also Fig. 28, p. 230]) Family CHORTOGLYPHIDAE I
Female genital aperture with or without sclerotized shields; if shields are present,
not crescent-shaped. Body setae various. Males with or without tarsal and/or anal
suckers " 8
I
8. Empodial claw connected to pretarsus by a pair of sci erotized rods, or condylophores
(Plate 100-2). Males usually with tarsal and anal suckers. Weakly sclerotized mites
I
living in a wide variety of habitats (Plates 102, 103 [see
also 10-5,11-4, Fig. 24, p. 229; Fig. 26, p. 229; Fig. 28, p. 230J) Family ACARIDAE' I
With at most a single tendon connecting the empodial claw to the pretarsus (Plate
110-1). Males without tarsal or anal suckers 9
I
9. Tarsi I-II of both sexes in the shape of a large claw; pretarsi I-I J strongly stalked,
joining tarsus proximad of the distal extremity of the tarsal segment. Marine tidal
forms, found on algae , ," (Plate 104) Family HYADESIIDAE
I
Tarsi not as above; pretarsi often stalked, but always attached to the distal extremity
of the tarsus. Non-marine mites 10 I
10. Idiosoma fusiform, convex. Integument smooth, thick, with all but one pair of the
idiosomal setae minute. Dark brown mites commonly found in nests of rodents.
I
· " , " (Plate 103) Family FUSACARIDAE
Idiosoma not as above. Integument may be thick, but with more than one pair of
macrosetae, or with platelets dorsally 11
I
11. Dorsal and lateral setae coarsely bipectinate or squamose. Integument thick, rough,
wrinkled, or covered with warts. Commonly found in hay or stored products.
I
· (Plate 107) Family CTENOG LYPHIDAE
Setae various, but not as above. Integument pigmented or smooth 12
I
12. Integument strongly pigmented, brownish, occasionally spined. Legs I-II often
compressed laterally or ornamented with longitudinal keels. Genital sclerites well
I
developed, often forming a circum genital ring. A heterogenous group
.. .. .. .. . .. . . . . . , (Plates 104,105 [see also Fig. 32]) Family LABIDOPHORJDAE
.
I
Integument soft, colorless or slightly pigmented, smooth, wrinkled or with fine
projections. Legs rarely ornamented or compressed. Genital sclerites variously
developed 13
I
13. Idiosoma without a transverse sejugal furrow between propodosoma and hystero-
soma 14
I
1Includes the LARDOGLYPHIDAE. I
I
I
I
251
I i
Idiosoma divided by a sejugal furrow, be·tween propodosoma and hysterosoma.
Small weakly developed mites similar to AC1RIDAE .
I · (Plat (105,106) Family SAPROGLYPHIDAE 1

I 14. Claws distinctly larger than caruncles, often )lalf the length of the tarsus. Associated
with Hymenoptera (Plates '108, 109) Family CHAETODACTYLIDAE
Claws small or well developed, but no large1r than associated caruncle in either case.
I Commonly found in stored grain, dried fruitlor milk products
i
15

15. Coxal apodemes I-II of female thickenedl fused to sternum and bordering the
I anterior edge of the genital shield. Tarsi n?ver more than twice the length of the
adjacent tibia. Found in dried fruit, miljk products and other processed foods.
· 1. (Plate 106) Family CARPOG LYPH IDAE
I Coxal apodemes I-II of female various, but I' ot as above. Tarsi often three times the
length of the adjacent tibiae. Found in tored grain and grain products, and in

I rodent or bird nests

............. (Plates 109, 110 [see also 7[4, 7-5, 8-1]) Family GLYCYPHAGIDAE 2
.

I
I 16. Female genital aperture a simple transverseflit without genital apodemes. Vertical
setae usually present. Females (and most ales) rounded or sac-like (Plate 126-2)
legs of females often greatly reduced; fem les rarely elongate (when elongate, legs
I IV are absent) with or without terminal empodial suckers. Skin parasites of

I
vertebrates j

I Superfamily SARCOPTOIDEA 37
Female genital aperture various, but never a simple transverse slit; genital apodemes
present (Plate 113-1), sometimes reduced (TURBINOPTIDAE and LEMURNYS-

;:"oP~~i)~1 ~~i~:~r~a ~~~~ga~~ ~~ o.va~~,.Ie~s. ~ t r~~~ce~: .~~th t.e~~.i~~I. ~r. ~~~t~~mi~~117

I
17. Some of the legs, the coxal regions of legs II-II, or portions of the mouthparts (Plate
I 124) strongly modified for clasping host .. ~
Without modifications as above i
33
18

I
I

18. Female genital aperture a longitudinal slilt, or a slightly expanded inverted "V"
(Plate 121-1) (if aperture appears trans erse, it is located between coxae II).
I Subdermal, visceral and lung parasites of m, mmals and birds .
·
1 Superfamily CYTODITOIDEA 30

I Female genital aperture assuming an inverted "U," "Y," or "V" shape, or transverse
(if transverse, the genital aperture is betwe~r coxae III or IV) 19

19. Parasitic on or in mammals, or on hiPPO~_OSCid or mallophagan insect parasites of

I mammals, occasionally free-living or nid iCOiOUS (species of nest-inhabiting PYRO-
GLYPHIDAE may be found on the feather- of birds) .
I · Superfamily PSOROPTOIDEA 20

I
l1ncludes ENSL! N IELL! DAE. OULENZII DAE,
21ncludes HERICIIDAE. 1 INTERSCHM IDTIIDAE and CZENS?I NSKII DAE.

I
I
 I
252

Parasitic on birds Superfamily ANALGOIDEA 25 I

20. Gnathosoma elongate, extending nearly to a point level with the tips of tibiae I.
Ambulacra I-II sucker-like; tarsi III-IV terminating in a spur-like extension and a
I
long seta. Follicle mites .
.................. (Plate 111 [see also Fig. 28, p. 230] ) Family AUDYCOPTI DAE
Gnathosoma not extending to the tips of tibiae I; tarsi III-IV similar to tarsi I-II,
I
and/or with more than one long terminal seta 21
I
21. Chelicerae stylet-like, fixed digit absent. Female with bursa opening dorsally,
female genital aperture an inverted "V"; male without anal or tarsal suckers.
Tarsi I-IV with terminal suckers. In' respiratory passages of lemurs .
I
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Plate 110) Family LEMURNYSSIDAE
Chelicerae dentate, fixed digit present 22 I
22. Integumentary striae of soft cuticle of opisthosoma formed into "scales." With at
most one pair of elongate terminal opisthosomal setae (d5). In respiratory I
passages of rodents (Plate 111) Family YUNKERACARIDAE
Integumentary striae of opisthosoma not scale-like. With two pairs of elongate
terminal opisthosomal setae (d5 and 15 both may be elongate) or terminal setae
I
short or absent _ 23

23. Coxal apodemes I fused in both sexes, forming a median keel. Opisthosoma of
I
female bilobed, setae d5 and 15 elongate; legs III-IV well developed but without
whip-like setae. Skin parasites of South American edentates
........................................ (Plate 114) Family LOBALGIDAE
. I
Coxal apodemes separated in both sexes or, if touching, without a median keel.
Opisthosoma of female generally not bilobed. Legs 111-1 V variously developed ..... 24
I
24_ With a rigid bifurcate or trifurcate epistomal extension of the propodosoma.
Legs III-IV of female inserted ventrally, normally developed and without long
I
terminal setae. Males with or without anal suckers. Nidicoles in nests of mammals
or birds, or free-living (Plate 112) Family PYROG LYPH IDAE I
Epistomal portion of propodosoma truncate or somewhat rounded. Legs III-IV of
female inserted laterally or ventrally (when inserted laterally, often reduced and
with long whip-like terminal setae). Males with anal suckers. Skin parasites of
I
mammals , (Plates 112, 113) Family PSOROPTIDAE 1

25. Legs arising from margins of body; coxae III-IV visible from above. Coxal apodemes
I
variously developed but only rarely fused 26
Legs I I I-IV inserted ventrally, generally not projecting far beyond margin of body. I
Coxal apodemes (particularly III-IV) often tending to fuse together toward the
medial aspect of idiosoma, resulting in closed coxal fields. Apoteles large, either
oval or heart-shaped. Feather mites (Plates 114. 115) Family FREYANI DAE
I
1Includes the family PSORALGIDAE. I
I
I
I 253
i
I 26. Legs III-IV more or less equally developed ill both sexes (in Syringobia, legs IV may
be thickened, but the coxal apodemes do no1 form plates) 27
I Legs III of males enlarged, much longer and thicker than legs III of female.
Transparent "spurs" and short spinose •etae on tarsi, tibiae and genua I-II.

I Coxal apodemes III-IV of males forming coxrl "plates." Feather mites

• • • • • • • • • • • • • • • • • • • . • • • • , •••••••••• 1.
.
(Plates 116,117) Family ANALGIDAE
I
I 27. Caudal aspect of females (and most males)!distinctl y bilobate (in some genera, the
posterior lobes may be fused to form a pOlsterior tail-like process); posterior lobes
with two pairs of strong and often highly modified setae. Vertical setae absent.
I Idiosoma often elongate. Feather mites 1

........................... (Plates 117, 118) Family PROCTOPHYLLODIDAE

.

I Cauda of females broadly rounded, rarely ircised; posterior aspect of males various.
Vertical setae present or absent ,
i
28

I 28. Tarsi I-II markedly shorter than tibiae I-II! usually equipped with claw-like distal
setae. Genital opening of female transversJ or resembling an inverted "U" or "V."
Males often bilobate posteriorly. Respirato~y parasites of birds .
I · ~' (Plate 118) Family TURBI NOPTIDAE
Tarsi I-II various, may be short but not mar edly shorter than tibiae; genital opening

I not as above. Males mayor may not exl, ibit bilobed condition of opisthosoma.
Externa I parasites i 29

I 29. With thin, colorless, finely striated integunlient. Body shields weakly developed or
absent. Apoteles of tarsi bell-shaped or rounded, tarsi often claw-like distally.
Vertical setae absent. Skin parasites of bird ... (Plate 119) Family EPI DE RMOPTI DAE

I Integument thick, strongly sclerotized. iBOdY shields generally well developed.

Apoteles various, usually attached distally to a gradually tapering tarsus. Feather
(Plat s 119, 120) Family DERMOGLYPHIDAE 1
I mites

30. All tarsi ending in apoteles j. . . . . . . . • . . . . . . . . . . . . . . . . . . . . . . . 31

I Tarsi without apoteles (eXception-Mycterf'nyssus has stalked caruncles on tarsi 111-

IV), with distal spine-like setae. Parasites in stomachs, nasal cavities or the eyes of
bats (Plate 121) Family GASTRONYSSIDAE
I i
31. Legs IV inserted on posterior fourth ofl idiosoma. Femur and genu II without
dorsal coalescence '1' 32
I Legs IV inserted anterior to that position dtscribed above. Femur and genu I-II each
coalesced dorsally. Internal parasites of dO,mestic fowl .

I · / .. (Plate 121) Family LAMINOSIOPTIDAE

32. Leg coxae forming large anterior and postdrior coalesced ventral sclerites. Tarsi each
I with a long seta. Parasitic in lungs of roder~ts
· i ..
(Plate 122) Family PNEUMOCOPTIDAE
.

I 1Includes the families PTEROLICHIDAE, SYRI~GOBIIDAEand FALCULI FERIDAE .

•
I
254
I
Coxal plates. not greatly enlarged or coalesced. Without long tarsal setae. Internal I
parasites of fowl. ; (Plate 121) Family CYTODITI DAE

33. Gnathosoma, legs I-II, III-IV and/or anterior coxal region modified for grasping
I
hairs of host; tarsi I-II without empodial claws. External parasites of mammals.
· Superfamily L1STROPHOROIDEA 34
Legs III-IV modified for grasping gills or hairs of host; tarsi I-II with large empodial
I
claws. External parasites of shore crabs
·
,
, .Superfamily EWINGOIDEA, (Plate 122) Family EWJNGIDAE
.
I
34. Legs I-II adapted for clasping hairs; legs III-IV reduced, with whip-like terminal
setae. Chelicerae with recurved teeth. Parasites of porcupines . I
.. . . .. . . . .. . . . . . .. .. .. .. . .. . .. .... .. .. (Plate 123) Family RHYNCOPTIDAE
Tarsi without whip-like terminal setae. Gnathosoma, the anterior coxal regions, or
any of the legs may be adapted for clasping hairs of host animal. Mammal parasites...35
I
35. Gnathosoma greatly enlarged, strongly sclerotized, modified for grasping host; palpi
pointed, without podomeric segmentation, forming a clamp-like apparatus with the
I
dentate chelicerae. Parasites of South American bats
· (Plate 123) Family CHI RORHYNCHOBIIDAE
.
I
Gnathosoma may have adaptation for grasping, but not developed as above. Fur
mites 36 I
36. Palpi normally developed; legs III-IV highly modified for clasping hairs. Quite often
oval in shape. Fur mites of rodents (Plates 123, 124) Family MYOCOPTIDAE I
Palpi, legs I-II and/or coxal fields of legs I-II modified for clasping hairs. Tarsi III-IV
each may be armed with a curved claw and associated spurs. Elongate species
found in fur of mammals (Plates 124, 125) Family L1STROPHORIDAE 1
I
37. Prodorsum (propodosoma) with strong elongate lateral apodemes which flank or
encircle the prodorsal sclerite. Parasites of birds 38
I
Prodorsum without elongate apodemes. Parasites of mammals 39
I
38. Legs IV of female absent; idiosoma elongate, narrowed posteriorly. Legs IV,of male
normally developed; idiosoma not elongated. One species from a gull
.. ----
--~-------_.-- .... - (Laridae) .... 2
.................................... (Plate 125) Family EVANSACARIDAE
I
Legs IV of female present, similar to legs III; idiosoma rounded. Parasites of fowl
and small domestic birds (Plates 125, 126) Family KNEMIDOCOPTIDAE I
39. Legs IV each reduced to a minute papillary structure, or absent; idiosoma rounded
or somewhat elongate. Skin parasites of bats... (Plate 126) Family TEINOCOPTIDAE I
Legs IV normally developed, similar to legs III; idiosoma rounded. Parasites of
mammals (Plate 127 [see also Fig. 28, p. 230] ) Family SARCOPTI DAE I
1Includes the families LABIDDCARPIDAE and ATOPOMELIDAE of McDaniel (1968).
2The EVANSACAR IDAE may constitute a subfamily of the family KNEM IDOCOPTI DAE. I
I
I
I 255

I KEY TO DEUTONYMPHS (HYPOPI) OF SOME FAMILIES OF ASTIGMATA 1

I ,. V"", " ,,'''h~,m, wl<h w wl<h'"' '",k~'" bOf 0_ wi<h ,,,,,.,,. . . . . . . . .. 2

I Venter of opisthosoma with claspers.. (Plate 1105-1) Family LABIDOPHORIDAE s. str.

2. Sucker plate reduced or absent (genital JCkers may be modified for clasping).
Gnathosomal subcapitulum and palpal entiti~s absent. . . . . . . . . . . . . . . . . . . . . . .. 3
I Sucker plate we.1I developed (Plate 108-3ll; gnathosomal elements distinct (Plate
103-2) but occasionally reduced '"1"' . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 5
I 3. Mobile, elongate forms, found in connectivetissues or skin of birds or mammals. .. 4

I Inert, rounded forms often found in and aro nd grain storages

· "
.
(Fig. 30,l p. 233) Family GLYCYPHAGIDAE (pars)

I 4. Vertical external setae present. Found in Sk~' n of South African rodents

· (Fig. 32, p. 235) Family LABIDOPHORIDAE, Subfamily RODENTOPINAE
.

I
I Vertical external setae absent. Found in co nective tissue of birds
· ,
.
, (Fig. 31, p. 235) Family HYPODERIDAE

I~ss
I 5. All legs mor.e or equally developed; Illegs III-IV often directed posteriorly or
laterally, tarsI and tibiae III-IV short ..... , . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 6
Legs III-IV more slender than I-II, directe~ forward; tarsi and tibiae III-IV long and
I slender \ (Plate 98-4) Family ANOETIDAE

I 6. Dorsum strongly ornamented with reticulations, striations or large pits, usually with
separate and distinct propodosomal and hjsterosomal shields; tarsi I-III often with
fleshy pretarsi and greatly enlarged hook- ike claws. Ventral apodemes generally

I robust, forming complex coxal field pa terns. Associated with Hymenoptera.

· (Plates 108, 109) Family CHAETODACTYLIDAE
Dorsum typically weakly sclerotized, withf.lo ut two distinct shields; pretarsi mayor
I may not be elongate, but never with enla ged hook-like claws. Ventral apodemes
generally distinct but not as above " . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 7
i
I 7. Pretarsi long, stalk-like, bearing an empodibl claw. Eyes usually present (Vidia is an
exception) (/Plate 105-2,3) Family SAPROGLYPHIDAE

I Empodial claws sessile, pretarsi small or ab~ent; eyes usually absent. . . . . . . . . . . .. 8

8. Gnathosoma well developed, terminal sen[IOrY organ (seta w) elongate, exceeding

I length of subcapitulum. Sternal setae ofter modified into discs :
.................................. (Plates 11-4, 103-1,2) Family ACARIDAE
.

I Gnathosoma weakly developed, seta w

Sternal setae normal
(~onsiderably
j •.•••.•
shorter than subcapitulum.
Family G LYCYPHAG IDAE (pars)

I 1 1n addition to those families cited in this key, hypJPi are known to occur in the CARPOGL YPH IDAE,
FUSACARIDAE and HEMISARCOPTIDAE. .

I
I
 PLATE 98 I
256
I
palp with distal
flageiia I
I
I
~I---+---{,,--trochanter J 98-2
I
-+--+--+-- coxal field I
,F'I-++---apodeme
I
,---:a;;;==~~;:;";;/;I-.L---!genital
'\{"@~
opening transverse
ring structure
I
I
"=:''1--_ apodeme I
Hr---*"Ir::='*- seta w long I
+--+--Ik--lr-l~ __ gnathosoma
I
I
I
legs III-IV slender,
I
I
I
-I----:-r-_sucker plate
I
98-3 98-4 I
98-1 to 98-4; family ANOETIDAE, Histiosoma sp. (Oregon, USA). 98-1; venter of female: 98·2; palp and portion of chelicera:
I
98·3; tip of tarsus III: 98·4; venter of hypopus

I
I PLATE 99
257
I
I
I
I
I
I
I
I
I
I
I
I
I
I hysttrosoma
J \
I I \
! \
I
I ~\ /A/l
99-3
(\-1 \ 99-4

I
I 99·1 to 99-4; family HEMISARCOPTI DAE, Hemisarcoptes sp. (0' egan, USA). 99-1; posteroventral region of male: 99·2; genital-
ana-I area of female: 99-3; tarsus II of male: 99-4; dorsum of male

I
 PLATE 100 I
258

~ ~fixedCheliceraldigit
I
~
100-1
absent
I
prodorsal shield without
vertical setae
I
I
apotel.. stalked,
sucker-like

I
~\\I -8Jl----vertical seta I
claw small, indistinct
I
I
I
I
I
100-2
I
I
proximal
fixed cheliceral
digit present I
podomere
I
subcapitulum
I
I
100-3
I
100-1; family LINOBIIDAE, Linobia sp., anterodorsal aspect with detail of chelicera (after Baker et al. 1958)
100.2 to 100-4; family CANESTRINIIDAE. 100-2; ?genus (India). tarsus II: 100·3; 'genus (Thailand), gnathosoma with one
I
chelicera removed: 100-4; ?genus (India), dorsum of female
I
 I PLATE 01
259
I
•
101-2

I ~ 101-3

•
I
I
I
I
I
I
•
I
I
•
I
genital discs small-.~.J.4:;;e:=..--'--<l

I 101-4

I
•
I i
101-'1 to 101-3; family ROSENSTEINIIDAE, Nycteriglyphus vespe tilio Ah and Hunter (Korea). venter of female with diagram-

:.
i
matic detail of chelicera (after Ah and Hunter 1968): 101·2; body seta: 101-3; N. pterophorus Berlese, body seta (after
Zakhvatkin 1941)
101·4; family CHORTOGLYPH IDAE, Chortoglyphus arcuatus (T oup.) (Oregon, USA). venter of female with detail of chelicera
 PLATE 102 I
260

/ I
'O)~ I
(j; \ I
I
I
I
I
102-2
I
102-1 I
I
I
I
I
I
I
I
I
I
I
102-4 I
I
102-1 to 102-6; family ACAR IDAE. 102-1; Lardoglyphus ,p., apotele of tar,us I of female: 102-2; genital region of an acarid
female: 102-3; Caloglyphus 'p. (Oregon, USA). venter of male: 102-4; Tyrophagus ,p. (Oregon, USA), dorsum of female:
I
102-5; Rhizoglyphus echinopus (F. & R.), tarsus IV of male: 102·6; undivided empodiaI claw of an acarid mite
I
I PLATE 103
261
I
•
I seta w

I 103-1 \I1--1'.Jl-
11'.11-_ _ palp
lyrifissure

I -+1
/ - _ - seta elcp
subcapitulum

I
I
I
I
I I '\

I
I
I
I
I ~~~.7
. / / - + ' - ' _ genital discs
anterior to
aperture

I
I
I 103-3

I 103-1 and 103-2; family ACARIDAE, Caloglyphus ,p. (Oregon,

hypopus with detail of gnathosoma
103-1; dorsum of tarsis I of hypopus: 103-2; dorsum of

103-3 and 103·4; family FUSACARIDAE,Fusacaruslaminipes (Oregon. USA). 103-3; genital area of female: 103·4;

I venter of male
 PLATE 104 I
262
I
104-2
•
I
I
I
I
I
I \
I
104·5
I
I
/ff\

I
I
,/ i! \
I:' \
,I
,I
\
\
I
I 'I \

,/ II \
I
I
pregenital sclerite fused to coxal
apodeme

/' '\
I
I
anal~J'
7]\\\
LW
sclerite
I
1\
I
I
104-1; family HYADESII DAE, Hyadesia ,p. (Oregon, USA), dorsum of female
104-2 to 104·5; family LABIDOPHOR IDAE. 104-2; Gohieria fusea (Oudeman,) (Oregon, USA) genital area of female: 104·3;
I
Labidophorus sp. (Oregon, USA), genital area of female: 104-4; G. fusca, genu, tibia and tarsus I of female: 104·5;
Xenoryctes sp., venter of female
I
I PLATE 105

I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
r.
105·1; family LABIDOPHORIDAE, Dermacarus sp. (Oregon, US.!), venter of hypopus
105·2 and 105-3; family SAPROGLYPHIDAE, Calvolia sp. (orego~r USA). 105·2; propodosoma of hypopus: 105·3; venter of
hypopus
 PLATE 106 I
264
I
106-1
without sclerotized
I
condylophore

I
~~
. Ad I
I
I
gland canal:---,I--
I
I
I
I
I
I
+---+_trochanter IV
I
I
I
I
106-2
I
I
106.1; family SAPROGLYPHIDAE, Czenspinskia lord! Nesbitt (Oregon, USA), dorsum of female with detail of apotele of
I
106-2;
tarsus I
family CARPOGLYPHIDAE, Carpoglvphuslactis (L.) (Oregon, USA), venter of female
I,
 I PLATE '107
265
I
I
I
I
I
I
~~~~__
integument
ornamented

I N'!{tI.~~ Positaion of genital

rea

I
I
I
I
I
I
I
I I ~~
I
I
I
I
{j(jCAF1[1
IIJ.V 107-2

,I
II '"" .0010' ' '-.
' family CTEN
7-2; C. palmifer IF_ 2~LYPHIDAE. a ter ' Hughes
.J, dorsal seta 107-1'
(f II'"
19 !' ~~ plumiger (Kochl 10
CtenOglVP~
regen, USA), dorsum of f
,m.'.,
II
 PLATE 108 I
266
I
108·1
108-2 I
I
I
I
I
'~---'t--'ll---I--- position of
aedeagus I
I
I
108-3
I
I
I
I
I
I
ventral apodemes'£'---+-
I
robust

sucker plate -----.::~~~~S~~

I
I
I
•
108.1 to 108·3; family CHAETODACTYLIDAE. 108·1 ;Sennertia ,p. (Congo), dorsum of male: 108·2; Sennertia 'p., female
genital area: 108-3; Chaetodacty!us osmiae (Dujardin) (Oregon, USA), venter of hypopus
I PLATE ~09 267
I
I claws enlarged

I 109·1

I
I
I
I
I podonotal shield

I
I opisthonotal shield

I
I
I
I
I
I I
I 'I
\ I

I I
I

'('i
109·4

I ~~~);~Ot~~G1g~r£~~;:D~~'~~~:~~~~~a:~~~y ~~~~um of hypopus

109·1 to tarsus
109-3;I famil y CHAETODACT
109.4; family 109·2; Horstia sp. (M ~,~).
. "",,""
I
 PLATE 110 I
268
I
110-1
I
I
unsclerotized
tendon
~
~
I
I
I
body setae often
pectinate
I
\~~J I
I
I
I
I
I
I
I
~:,v:~~::~;m----+-...,hll
palp----/-
110-4
Q
, I
I
110-3
I
110-1; family GLYCYPHAGIDAE, Glycyphagus domesticus (DeGeer! (Oregon, USA), venter of female with detail of apotele of
tarsus J
110-2 to 110-4; family LEMURNYSSIDAE, Lemurnyssus galagoensis Fain (Ruanda-UrundD. 110-2; dorsum of female (after Fain
I
1957): 110-3; gnathosoma of female (after Fain 1957): 110-4; genital area of female (after Fain 1957)
I
I PLATE 11
269
I 111-2
I 111-1
Y,...il=:::lW--_' gnathosoma elongate

I
I
I
I
I
I
I
I
I
I
I
I
I
I scale-like
ornamentation
----"ir-;':

I 111-3

I
I 111-1 and 111-2; family AUDYCOPTIDAE. 111-1; saimiriOPtesp~radOXUS Fain (South America), venter of female (after Fain
19681: 111-2; ?Audycoptes sp. (Pennsylvania. USA), med oventral aspect of female
111-3 and 111-4; family YUNKERACARIDAE, Sciuracarus parax ri Fain (Transvaal). 111-3; dorsum of female lafter Fain 1964):

I
111-4; posteroventral aspect of female (after Fain 1964)

I
 PLATE 112 I
270
I
112·1
I
I
I
I
I
I
I
I
I
HL....,k,ll--::::.. coxal apodemes
separate I
I
I
J I
J
1 I
I
J
I
112-3 112-4
I
112·1 to 112·3; family PYROGL YPHIDAE. 112·1 ; Euroglvphus (Gvmnoglvphus) longiorTrouessart (England). posteroventral
aspect of female: 112·2; E. long;or, dorsum of female: 112-3; Dermatophagoides pteronyssinus Trouessart (Oregon, USA).
I
venter of female
112·4; family PSOROPTIDAE, Caparinia tripi/is (Michael) (New Zealand). dorsum of female
I
I PLATE '113
I
271

I
I
I
I
I
I
I
I
I
I
I l~
113-2
I
I
I
I
I
I
I r~ r "'

I \, \ . _ , <CO'"
1
13·1 and 113-2; fa mily
(Colorado.
dorsum ofDA~~
USA).PSOROPTI m
fI
1 13·1; Chorioptes ,p. ( regan, USAI. venter of female: 113-2; Chorioptes

I
 PLATE 114 I
272

114-1
I
I
I
l~'---I'oI~~--aedeagus
I
empodia sucker-like
I
I
I
I
I
I
I
I
I
I
I
opisthosoma biJobed
I
114-2 I
114-3
I
114·1; family FREYANIDAE, Kramerel/a sp. (Texas, USA), venter of male
114-2 and 114·3; family LOBALGIDAE, Lobalges trouessarti Fonseca (Brazil). 114-2; posteroventral aspect of male: 114·3; com-
I
posite dorsum and venter of female (after Fain 1965)
I
I PLATE 15
273

I
I
I
I
N
I ......Lil
I
I
I
I
I
I
I
I
I ...
......
Lil

I
I
I
I
I
I
274
PLATE 116 I
~ sucker·like empodia
I
f?-~=--- spurs on legs I-II
I
I
I
I
. _ _ - - 4 - - - pregenital sclerite I
;;i4,--~""'-=::-:=-- genital opening an inverted "V"

I
I
I
I
116-1 I
I
I
I
I
I
I
I
116-1 and 116-2; family ANALGIDAE, Ana/ges sp. (Oregon, USA), 116·1; venter of female: 116-2; dorsum of female
I
I
I PLATE 117
275
I
I
I
I
I
I
I
I
I
I
I
I
I
'i
I legs III enlarged, with
enlarge<! emPOdial claw
anal sucker

I 117-3
I
I terminus of---J-~'-­
opisthosoma with
ornate flanges

I 117·1; family ANALGIDAE,Analges ,p. (Oregon, USA!. vejlter of male

117.Z and 117·3; family PROCTOPHYLLODIDAE. 117-2; ~ ouessartia ,p. (Nigeria!. dorsum of female: 117·3; Troue.sartia 'p.

I
(England!. posteroventrat region of male
 PLATE 118 I
276
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
118-1; family PROCTOPHYLLODIDAE, Trouessartia sp. (Nigeria), venter of female
118-2 to 118-4; family TURBINOPTIDAE. 118·2;Schoutedenocoptes (Kansas, USA), venter of female: 118·3; Turbinoptes
I
I
strandtmanni Boyd (Texas, USA), genital area of female (after Boyd 1949): 118-4; Schoutedenocoptes sp., tarsus I
I PLATE 119
277
I
I
I
I
I
I
I
I
I
I
I
I
119-4
I
I
I
I 119-3
I
I
I 119-1 to 119-3; family EPIDERMOPTIDAE. 119-1; Strelkoviafarus 'p. (Bechuanalandl, dorsum of female: 119-2; Epidermoptes
bilobatus Rivolta (England), composite of female
bilobatus, tarsus I (after Fain and Evans 1963)
dorSUI and venter (adapted from Fain and Evans 1963): 119-3; E.

I 119-4 and 119-5; family DERMOGLYPHIDAE,Falculifersp_ 1 9-4; chelicera of female: 119-5; chelicera of male
 PLATE 120 I
278
I
120-1
empodial suckers terminal
I
I
I
I
chelicera palp I
I
I
I
I
I
I
I
I
I
I
120-2 I
I
120-1;
. DERMOGL YPHIDA E. 120·1; Syringobla
tamll Y of male
dors um
~/
. sp . (Nebraska, USA), venter 0 Ilemale.. 120-2; Pterolichus sp.
(Mexico), I
I
I PLATE
279
I
I
I
~
}~£ @~."
aedeagus~
I 121-3

I
I
I
I elongate
or oval

I
I
I
I
I
I
I
I
I
I
Gastronyss~s bakeri Fain (Ruanda-Urundi), venter of female (after Fain 1959):
I 121·1 to 121·4; family GASTRONYSSIDAE. 121-1;
121-2; Rodha;nyssus yunker; Fain (Central Africa), dorS'al profile (adapted from Fain 1959): 121-3; G. bakeri, aedeagal area
of male: 121-4; Gastronvssus Sp., tarsus I I
121-5; family CYTODITIDAE, Cytodites nudus (Vizioli) venter of female (adapted from Hirst 1922)

I 121·6; family LAMINOSIOPTIDAE, Laminosioptes cysticola (Vizioli). venter of female (adapted from Hirst 1922)

I
 PLATE 122 I
280
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
pretarsus claw-like
I
122-1; family PNEUMOCOPTIDAE, Pneumocoptes jellisoni Baker (Idaho, USA), composite dorsum and venter of female (adapted
from Baker 1951)
I
122-2 to 122-4; family EWINGIDAE, Ewingia coenobitae Pearse (Florida, USA). 122-2; tarsus I: 122-3; posteroventral region of
male: 122-4; venter of female
I
I 281

I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
123-4 123-5
I
•
I
123-' and 123-2; family CH I RORHYNCHOBII DAE, ChirorhynCtbia urodermae Fain, venter of female: 123-2; tarsus I (after Fain
19681
123·3; family RHYNCOPTI DAE, Rhyncoptes anastosi Fain (So th America), composite of dorsum and venter of female (adapted
from Fain 19651
123-4 and 123-5; family MYOCOPTI DAE. 123-4; Myocoptes mUrU'inus (Kochl (Oregon, USA), genital area of female: 123-5;
Trichoecius sp., terminal portion of leg III I
282
PLATE 124 I
I
124-1 I
124-3 I
I
I
I
I
I
legs III-IV
modified for
clasping hairs
I
I
I
I
I
I
I
I
I
124-2
I
124-1 and 124-2; family LlSTROPHORIDAE. 124-1; Listrophorus sp. (Oregon, USA), venter of male: 124-2; Listrophorus
sp. (Oregon, USA), lateral aspect of female
I
124-3; family MYOCOPTIDAE, Myocoptes musculinus (Koch) (Oregon, USA), dorsum of female
I
I PLATE f25
283

I I
I

•• I
I A
legs I-II modified
for clasping hairs
Ii
I 125-1 j
I I
terminal suckers aJsent

•
/ in female

! coxal fields I-II modified

to aid in clasping
i

I
•• '--r I.

- sclerotized apodeme

I
I
I
••
125_!JX~
I
I 125-5 125-6
I
125-1 and 125-2; family L1STROPHOR IDAE. 125-1; Alabidoca,Jusrecurvus (Womersley) (Australia), anterolateral aspect of

I female (after Pinichpongse 1963): 125-2; Campylochirus t~aViae (Hirst), anteroventral region of female (after Hirst, 1922)
125-3 and 125-4; family EVANSACARIOAE, Evansacarus far; Fai (Israel). 125-3; dorsum of female: 125-4; anterodorsal region
of male (after Fain 1962)
125-5 and 125-6; family KNEMIDOCOPTIDAE, Knemidocoptes n utans (Robin & Lanquetin). 125-5; dorsum of leg I of female

I (after Fain and Elsen 1967): 125-6; dorsum of leg I of ml'e (after Fain and Elsen 1967)
 PLATE 126 I
284
I
•
I
I
I
I
I
I
•
I
I
I
I
•
I
leg IV
reduced or
I - l e g III
I
absent
I
126-3 126-4 I
126-1 and 126·2; family KNEMIDOCOPTIDAE, Knemidocoptes mutans. 126-1; anterodorsal region of male: 126-2; composite
dorsum and venter of female
126-3 and 126-4; family TEINOCOPTIDAE. 126-3; Teinocoptes epomophori Rodhain (Congo). venter of male (after Rodhain
1923): 126-4; Chirobia congolensis Fain (Congo), venter of female (after Fain 1959)
•
I
I PLATE
285

I
I
••
J"--_stalked empodial sucker
on legs 1-11

I r--'--=-\----- genital aperture transverse

I
•
I ~~"=",,J--p....- Iegs IV normally developed

I
prodorsal shield

I reduced

anal aperture ---t-.:::>'i'o=:=--lJ

I
•
I
127-1

I
I
I
I
I
I 127-1 and 127-2; family SARCOPTIOAE, Sarcoptes scabiei (OeGrerl (Oregon, USA). 127-1; venter of female: 127-2; dorsum of

I female

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 I
286

NOTES
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I
287
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•
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288
I
ORDER ACARI FORMES I
Suborder Cryptostigmata
I
The Cryptostigmata, or "beetle mites," comprise a cosmopolitan group of approxi-
mately 5000 species which have been relegated to some 500 genera. The majority of species
I
are slow-moving, strongly sci erotized forms which range in size from 200 to 1300 11.
Respiration apparently is carried on through tracheal ducts (Fig. 7c, p. 20) which open into
the acetabular cavities of the legs or through brachytracheae. The latter open into the leg
I
segments themselves or into the prodorsal pseudostigmata. Additional features of impor-
tance are: I
1. Simple palpi, without claws, each composed of 3-5 segments (Fig. 33).
I
2. Well-developed subcapitular rutella (Fig. 33).

3. Chelicerae usually chelate-dentate (Fig. 33).

I
4. Tarsi with 1-3 tarsal claws.
134-2).
Empodium, when present, always claw-like (Plate I
5. An ovipositor in females (Plate 10-8) and aedeagal sclerites in maies. I
6.

7.
Generally with a pair of prodorsal pseudostigmatic organs (Plate 130-1).

Genital and anal openings protected by discrete shields. Genital discs are present
I
(Fig. 34).
I
Cryptostigmatid mites are primarily fungivorous, algivorous or saprophagous, and are
particularly common inhabitants of forest humus and soil.
I
The plethora of morphological characters available for comparative studies of the
Cryptostigmata usually makes specific diagnoses relatively simple. At the same time,
however, there appears to be a tendency to "upgrade" the classification of the group, so that
I
species often become monotypic genera and, on the basis of specific characters, genera are
set up as families. While this topheavy arrangement allows for easy inclusion of newly
discovered species, it appears to encourage a similar expansion of the higher categories as
I
well. Such expansion is reflected in the large number of superfamilies (27) recognized in
this manual, and by the number of familial categories (114) listed in the accompanying
incomplete chart.
I
The cryptostigmatid superfamilies are arranged in three supercohorts, the largest and I
most complex of which is the Oribatei Superiores. Cohortal separation of the Oribatei
Superiores into the Pycnonoticina and the Poronoticina is complicated by the fact that, in
I
•
some cases, a superfamily may fit into either cohort (the Ameronothroidea, for example).
Also, certain superfamilial diagnoses presently used by specialists in the Cryptostigmata are
so broad as to make their definitions contiguous or overlapping. Thus it is not practical to
attempt a key separation of the Ameronothroidea and the Passalozetoidea, nor may the

I
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I 289
I
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•
-+---"--I-_genital disc

I
I Fig. 33. Venter of the gnathosoma of
Ctenacarus sp. (Oregon, USA).

I
I i
FJ 34. Genital and aggential shields of Brachythonius

I
sp. (Oregon, USA), illustrating the position of
the three pairs of genital discs.

I Carabodoidea be clearly distinguished from thf1e Cepheoidea. Similar difficulties occur at the
family level. The OPPIIDAE and AUTOGNET IDAE, for example, are difficult to separate,

I as are most of the families of the superfamil Oribatuloidea. Hopefully, a redefinition of

generic relationships eventually will be attem ted, possibly through comparative studies of
immature stages. Such a system has been devised by Grandjean (1954, 1969) but its

I complexity probably would limit its usefuln ss in an introductory text. For reasons of
simplification, only superfamilies based on adult Cryptostigmata will be included in the keys
and illustrations to follow. Keys to most of the presently recognized families and genera
I appear in two works by 8alogh, both of whi, h were used in writing the keys which appear

•
below. Balogh (1965) stresses that the system hich he followed in erecting his classification
is based solely on adults, and that it is artifi ial and somewhat provisional for that reason .
The classificatory works of Grandjean shoul, , therefore, be consulted for an ontogenetic
approach to cryptostigmatid classification.

I
I
 SUPERCOHORT COHORT SUPERFAMILY FAMILY

~
Archeonothridae
Palaeacaridae
PALAEACARI Palaeacaroidea --------1 ~:~~~~~~;~ae N
to
Adelphacaridae a
Aphelacaridae
Parhypochthonoidea C r;::~~gg~~tt~~~i~~::
~~~~~~~~~~~?:e

-1
Brachychthoniidae
Haplochthoniidae
Cosmochthoniidae
Hypochthonoidea Heterochthoniidae

'"~".oo. 0_,_'""{
Sphaerochthoniidae
Atopochthoniidae
Pterochthoniidae
Phyllochthoniidae
Protoplorphoridae
ORISATEIINFERJORES
Mesoplopharoidea C ~~~~~~~~~~~~~~:~
Phthiracaridae
E
Phthiracaroidea - - - - - - - - Euphthiracaridae
Oribotritiidae

~
Mixonomata Grandjean 1969 ( Perlohmanniidae
EpHohmanniidae
Perlohmannoidea ------~ Eulohmanniidae
Lohmanniidae
Collohmanniidae
Nothridae

~
Holonothridae
Camisildae
Nothroidea - - - - - - - - - Trhypocthoniidae
Malaconothridae
Nanhermanniidae
Hermanniidae
Hermannielloidea C ~I:~~:~~~~~~ae
Liodoidea Neoliodldae
Plateremaeidae

CRYPTOSTIGMATA
Plateremaeoidea - - - - - - -
E ~~~~~~:~~:ae
Licnobelbidae
Damaeoidea - - - - - - - - - Damaeidae tBelbidae)
Cepheoidea Cepheidae
Zetorchestoidea C ~~~~Z~i:~dae
~':g~:~aa:idae

--i
Amerobelbidae
Eremulidae
Staurobatidae
Eremobelbidae
Eremaeoidea Damaeolidae
Basilobelbidae
Heterobelbidae
Ameridae
Ctenobelbidae

---------------------
--------------------- PYCNONOTICINA

~
Peloppiidae
liacaridae
liacaroidea ---------1 ;~~~~~~f;ae
Tenuialidae
Ceratoppiidae

~
Carabodidae
Charassobatidae
Carabodoidea ----------1 ~~~~~:g~:j~::
Eutegaeidae
_ _ _ _ _ _ _ _ ?Tokunocepheidae

ORIBATEI SUPERIORES :::~~~:aZ-tetO;deafjf[~~~~te

Rhynchoribatidae
Dampfiellidae
Qtocepheidae
Eremellidae
Trizetidae
Hydrozetoidea C ~~~~~:~~~~~

~
Ameronothridae
Podacaridae
r Ameronothroidea - - - - - - . . . . . , ~r;;~~e~~~a:eidae
.I Selenoribatidae
------''--FF,ortti"y niid66
Passalozetidae
Passalozetoidea
E
-------+ Scutoverticidae
Licneremaeidae

·l eo"",o","" _-
Pelopoidea

Oribatelloidea ---------1
C ~~~~~:;~~~dae
E
Oribatellidae
Achipteriidae
Tegoribatidae
Microzetoidea - - - - - - - - - Microzetidae

~
Ceratozetidae
Mycobatidae
Ceratozetoidea ---------1 Chamobatidae
Euzetidae
Mochlozetidae
Galumnidae
Galumnoidea --------+_
E Parakalumnidae
Epactozetidae

g~:~~~:~o;ee

----f
Haplozetidae
Oripodidae
Oribatuloidea Zetomotrichidae
Chaunoproetidae
Neotrichozetidae
Stelechobatidae
Sellnickiidae

Fig. 35. Dendrogram illustrating possible relationships within the order ACARIFORMES: suborder Cryptostigmata of the subclass ACARI.
'"
CD
292
I
Useful References I
Balogh, J. (1961). Identification keys of world oribatid (Acaril families and genera. Acta Zool. Acad. Sci.
Hung. 7(3-4) :243-344. I
Balogh, J. (1965). A synopsis of the world oribatid (Acari) genera. Acta Zoo I. Acad. Sci. Hung.
11(1·2):5·99.
Grandjean, F. (1951). Les relations chronologiques entre ontogeneses et phylogeneses d'apres les petits
characteres discontinus des Acariens. Bull. BioI. Fr. Belg. 85(3) :269-292.
I
Grand]ean, F. (1954). Essai de classification des Oribates (Acariens). Bull. Soc. Zool. France 78:421-446.
Grandjean, F. (1969), Considerations sur Ie classement des Dribates leur division en 6 groupes majeurs. I.
I
Les affinites de Col/ohmannia gigantea Selin. 1922. Acarologia 11(1) :127-153.
Hammen, L. van der (1952). The Oribatei (Acari) of the Netherlands. Zool. Verhand. Leiden 17:1·139.
Hammen, L. van der (1959). Berlese's primitive oribatid mites. Zool. Verhand. Leiden 40:1-93.
I
Knul/e, W. (1957). Morphologische und Entwicklungsgeschictliche untersuchungen 20m phylogenetischen
System der Acari: Acariformes Zachv. I. Dribatei: Malaconothridae. Mitt. ZooI. Mus. Berlin
33(1) :97·213.
I
Sel/nick, M. (1960). Formenkreis: Hornmilben. Oribatai [sic]. Die Tierw. Mitteleur. 3(4):44-134.
Trave, J. (1964). Importance des stases immatures des Dribates en systematique et en ec%gie. Acarologia
fasc. h. s. 1964:47·53.
I
Wool/ey, T.A. (1958). A preliminary account ofthe phylogeny of the Oribatei (Acarina: Sarcoptiformes).
Proc. 10th Int. Congo Ent. 1:867·873.
Woolley, T.A. and E.W. Baker (1958). A key to the superfamilies and principal families of the Oribatei
I
(Sarcoptiformes: Acarina). Ent. News 69:85·92.
I
SUPERCOHORT PALAEACARI

DIAGNOSIS: Soft-bodied, with or without transverse sutures; gnathosoma visible from

I
above. With a pair of filiform, fusiform or claviform pseudostigmatic
organs. Genital shields weak, 3 pairs of genital discs present. Femora of
legs divided; tarsus I with 4 sensory solenidia. Tarsi of larvae and nymphs
I
not always monodactylous.
I
The Pa/aeacari includes a single superfamily, PALAEACAROIDEA, and three families.
On the basis of species representing two of these families (the PALAEACAR IDAE and
ACARONYCHIDAE), Tragardh erected a new suborder, Pa/aeacariformes (1932), pointing
I
out the affinities of this group with both the Prostigmata and the Cryptostigmata. Grandjean
subsequently included the Palaeacariformes as a superfamily in the Cryptostigmata (1932),
an emendation which has been followed only by some workers. In a recent revision of the
I
major cryptostigmatid categories, Grandjean (1969) erected the supergroup Palaeosomata in
which he raised certain palaeacaroid families to superfamilies (CTENACARIDAE, ARCHE-
ONOTHRIDAE), with the third superfamily, the PALAEACAROIDEA, represented only
I
by the family PALAEACARIDAE.
I
Unlike the higher Cryptostigmata, members of the Palaeacari are soft-bodied, weakly
sclerotized species which superficially resemble some of the free-living Astigmata rather
than the strongly sci erotized Oribatei. Tragardh (1932) felt that palaeacaroids are
I
phytophagous or saprophagous, probably feeding on mosses, lichens or decayed leaves.
Acaronychus tragardhi Grandjean was collected from moss in Colorado (Woolley 1960). A
species of Aphelacarus (family CTENACARJDAE) has been collected in stored grain and
I
I
I
I
293
I in the nest of a subterranean termite in Oregor. Palaeacarus appalachicus Jacot and P.
hystricinus Tragardh occur in Europe and the Uni ed States (Johnston 1967), and apparently
I feed on fungal hyphae in forest litter. A. acarinu (Berlese) occurs in northern and southern
Europe, while a related species has been col lee ed in South America, Africa and Europe.

I Useful References
I
I Grandjean. F. (1932). Au sujet des Palaeacariformes T)agardh. Bull. Mus. Nat. Hist. Nat. 4(2) :411·426.
Grandjean. F. (1954). Etude sur les Palaeacaroides (Acjriens. Oribates). Mem. Mus. Nat. Hist. Nat. (n.s.)

I A, Zoo I. 7:179·274.
Grandjean, F. (1969). Considerations sur Ie c1assement des Oribates leur division en 6 groupes majeurs. I.
Les affinites de Collohmannia gigantea Selin. (192'). Acarologia 11 (1) :127·153.

I Johnston, D.E. (1967). On the occurrence of two sp~cies of Palaeacarus in the eastern United States
(Acari: Acariformesl. Proc. Ent. Soc. Wash. 69(4) :tl01·302.
Tragardh, 1. (1932). Palaeacariformes, a new suborder ofl Acari. Ark. Zoo I. 24B(2): 1·6.

I Willmann, C. (1931). Moosmilben order Oribatiden (orii' ateil. Tierw. Deutschl. 22:79·200.
Woolley, T.A. (1960). Some interesting aspects of Olibatid ecology (Acarina). Ann. Ent. Soc. Amer.
53(2) :251·253.

I Zakhvatkin, A.A. (1945). On the discovery of Palaeac riformes (Acarina) in the U.S.S.R. Comptes Rend.
Acad. Sci. U.S.S.R. 47(9):673·676.
I

I SUPERCOHORT ORIBA1EI INFERIORES

I DIAGNOSIS: Generally well sclerotized, pty.hoid or aptychoid; gnathosoma not com·

pletely visible from above. With a pair of pseudostigmatic organs usually
present, occasionally minute. I Genital shields variously formed, genital
I discs present. Femora of leg) not divided, genua and tibiae subequal in
size; tarsus I with less than 4 olenidia. Tarsi of immatures always mono-

I dactylous.

Six superfamilies are included in the Or batei Inferiores, with no fewer than thirty

I families being recognized at the present time. [Grandjean (1947) created the Enarthronota to
contain those families comprising the HYPOC~ THONOI DEA and MESOPLOPHOROJ DEA.
fn fater works (1954a, 1969), Grandjean regro ped the families of Enarthronota under new
I superfamilies so that, based on ontogenetic an other considerations, the HYPOCHTHONO-
IDEA and MESOPLOPHOROIDEA indicat d in the accompanying dendrogram are
considered by Grandjean to comprise seven s perfamilies (Grandjean 1969). The PHTH I R-
I ACAROIDEA and PERLOHMANNOIDEA re included by Grandjean under the new
group (?cohort) Mixonomata. These are rec assified under seven superfamilial categories.

I Members of the Oribatei Inferiores are a heterogenous group insofar as adult

characteristics are concerned. The ptychoid groups MESOPLOPHOROIDEA and

I PHTHI RACAROI DEA are flattened laterallyd!and often are strongly sci erotized and tanned.
The aptychoid PARHYPOCHTHONOIDEA, n the other hand, are weakly sclerotized and
without lateral compression. These differ in ~urn from the well-armored often ornamented
I members of the NOTHROIDEA. The heterogrneity of the Oribatei Inferiores is exemplified
by the broad choice of morphological characters which permits inclusion of a family or

I
I
294
I
superfamily within this group (page 301). Similar difficulties, however, exist in defining I
more recent categories such as the Mixonomata (Grandjean 1969).

Parhypochthonoid and hypochthonoid species apparently subsist in leaf litter, soil and
I
humus (Wallwork 1959, 1960). Eniochthonius sp. may feed on fungal spores (Woolley
1960). A species of Parhypochthonius was recovered from a treehole in British Columbia in
association with an ant nest. Pine or fir duff, or moss in cool slightly moist situations, are
I
common habitats for many hypochthonoids.

Mesoplophoroids and phthiracaroids are particularly abundant in woody plant tissues,

I
bark, or decaying tree stumps. A species of Oribotritia is found on fir and cedar needles in
northwestern Oregon. Phthiracarus borealis Triigardh feeds on hemlock needles, as well as
on deciduous leaves (Wallwork 1958). Members of the genus Mesoplophora appear to
I
feed on woody tissues, while Hoplophora and Euphthiracarus spp. are found in twigs of
hemlock and yellow birch where they tunnel and feed (Wallwork 1958). An undescribed I
species of Steganacarus has been taken from holly root crowns in Oregon, where it may
have been feeding on root tissue. Immatures of S. diaphanum Jacot feed on decaying
conifer needle tissue and the nonvascular portion of leaf petioles from deciduous trees
I
(Hartenstein 1962a). Phthiracaroids also occur in soil (Woolley 1960). An undescribed
mesoplophoroid was collected in substantial numbers from an ant nest in a Brazilian rain
forest.
I
Members of the PERLOHMANNOIDEA are primarily litter forms, with some species
having been taken from pasture soils. An unidentified species of LOHMANNII DAE from the
I
Congo was found "avec insecte et arachnide." Others have been collected from coastal
bogs and moss. I
Nymphs of Hermannia sp. (superfamily NOTHROIDEA) feed in lenticels of yellow
birch bark (Wallwork 1958). Related species occur with regularity in forest litter, often at I
high altitudes. Hermannia subglabra Berlese, however, was collected from a salt marsh in
Wales. Twigs of balsam fir in New Brunswick, Canada harbor a species of Camisia, and
a second species is common on fir bark in Oregon. Platynothrus peltifer (Koch) is
I
a fungivorous camisiid in forest soils and litter (Hartenstein 1962b). Nanhermannia
elegantula Berlese may be collected, among other habitats, in saline situations in Europe
(Woolley and Higgins 1958), while other members of this nothroid group occur in moss,
I
litter or treeholes.
I
Useful References

Berlese, A. (1916). Centuria seconda di Acari nouvi. Redia 12:125·177. [NOTHROIDEAJ

I
Berlese, A. (1923). Centuria sesta di Acari nouvi. Redia 15:237-262. [PERLOHMANNOIDEAl
Ewing, H.E. (1917). A synopsis of the genera of beetle mites with special reference to the North
I
American fauna. Ann. Ent. Soc. Amer. 10:116-132. [PHTHIRACAROIDEAl
Grandjean. F. (1932), La famille desProtoplophoridae (Acariens).
[HYPOCHTHONOI DEAl
Bull. Soc. Zoo I. France 57:10·36.
I
Grandjean, F. (1933). Oribates de l'Afrique du nord (Ire serie). Bull. Soc. Hist. Nat. Afr. Nord
24:308-323. [M ESOP LOPHO RO IDEAl
I
I
I
I
295
I Grandjean, F. (1947). Les Enarthronota (Acariens)
I

Pre'~iere serie. Ann. Sci. Nat., Zool. 11, 8:213·248.

I Grandiean,
Grandjean,
Grandiean,
F. (1948). Les Enarthronota (AcarienS)'l(2e serie). Ann. Sci. Nat.. Zool. 11, 10:29·58.
F. (1950a). Les Enarthronota (Acariens). (3e serie). Ann. Sci. Nat.. Zool. 11, 12:85·107.
F. (1950b). Etude sur les Lohmanniidae (0 ibates, Acariens). Arch. Zoo I. Exp. Gen. 87:95·
I
I 162.
Grandjean,
[PERLOHMANNOIDEA]
F. (1954al. Essai de classification des Oribatis (Acariens). Bull. Soc. Zool. France 78:421-446.
Grandiean, F. (1954b). Les Enarthronota (ACariensl.j(4e seriel. Ann. Sci. Nat. Zoo I. 11, 16:311·335.

I Grandjean, F. (1954cl. Posthermannia nematophora n.g., n. sp. (Acarien, Oribate). Rev. Fr. Ent.
21 :298-311. [NOTHROIDEA]

I (n.s.1 A. Zool. 16(31:57-119. I

Grandjean. F. (1958). Per/ohmannia dissimilis (Hewittl) (Acarien, Oribatel. Mem. Mus. Nat. Hist. Nat.

Grandjean, F. (1969). Considerations sur Ie c1assement des Oribates leur division en 6 groupes majeurs. I.
Les affinites de Col/ohmannia gigantea Selin. 1922. Acarologia 11 (11: 127·153.

I Hammen, L. van der (19591. Berlese's primitive o'libatid mites. Zool. Verhand. Leiden 40:1-93.
Hartenstein, R. (1961). Soil Oribatei. I. Feeding specificity among forest soil Oribate; (Acarinal.
Ann. Ent. Soc. Amer. 55(21 :202-206. I

I Hartenstein, R. (1962a). Soil Oribatei VII. Deco~rposition of conifer needles and deciduous leaf
petioles by Steganacarus diaphanum (Acarina: Phtl1liracaridae). Ann. Ent. Soc. Amer. 55(6):713,716.
Hartenstein, R. (1962b). Soil Oribatei V. Investi 9atidns on Platynothrus peltifer (Acarina: Camisiidael.

I Ann. Ent. Soc. Amer. 55(61:709-713.

Jacot, P. (19301. Oribatid mites of the subfamily hthiracarinae of the Northeastern United States.
Proc. Boston Soc. Nat. Hist. 39:209-261.

I Michael, A.D. (18881. British Oribatidae. Ray Society, ondon 2:337·657 + xi + plates.
J
Sell nick. M. (19281. Formenkreis: Hornmilben, Oribatei. Tierw. Mitteleur. 3(9):1·42.
Sellnick, M. and K.-H. Forsslund (1955). Die Ca~isiidae Schwedens (Acar. Oribat.). Ark. Zool.
I 8(2):473-530. [NOTHROIDEA]
Wallwork, J.A. (1958), Notes on the feeding behavior of some forest soil Acarina. Oikos 9(2):260-271.

I Wallwork, J.A. (19591. The distribution and dynami s of some forest soil mites. Ecol. 40(4):557·563.
Willmann, C. (1931). Moosmilben oder Oribatiden (Oril atei). Tierw. Deutschl. 22:79·200.
Woolley, T.A. (1960). Some interesting aspects of a ibatid ecology (Acarinal. Ann. Ent. Soc. Amer.

I 53(2) :251-253.
Woolley, T.A. and H.G. Higgins (19581. A reVISion rOf the family Nanhermannlldae (Acari Oribatei).
Proc 10th Int. Cong Ent. 1913·923. [NOTH OIDEA]

I
SUPERCOHORT ORIBATEI SUPERIORES

I DIAGNOSIS: Well sclerotized, aptychoid; gn8lhosoma not completely visible from above.
Pseudostigmatic organs presentl assuming a variety of shapes. Genital
I shields variously formed, genitar discs present. Femora of legs undivided,
tibiae distinctly longer than ~enua; tarsi I with less than 4 solenidia.
Tarsi of immatures always monoFactYlous.
I Twenty superfamilies and more than seve~ty families comprise the strongly sclerotized

I aptychoid Oribatei Superiores, or "higher Oribatei." On the basis of the presence or absence
of symmetrical pairs of dorsal pores and of dorsolateral "wings" or pteromorphae, the
higher Oribatei may be divided into two sm Iler groups, the cohorts Pycnonoticina and

I Poronoticina.

I
I
 I
296

Cohort Pycnonoticina
I
The Pycnonoticina includes those higher Oribatei which possess neither pteromorphae
nor symmetrical dorsal pores. There are more than forty recognized pycnonoticine
I
families arranged in twelve superfamilies on the basis of adult characters. Grandjean (1954)
has proposed a system of classification of the higher Oribatei in which immature characters
are utilized, including methods of nymphal ecdysis. Immatures in the Pycnonoticina are,
I
however, difficult to categorize, other than to point out general similarities between nymphs
and adults of certain pycnonoticine superfamilies (the L10DOIDEA, EREMAEOIDEA and
ZETORCHESTOIDEA, for example).
I
The habitats of pycnonoticine Oribatei vary from dry sandy areas (Eremaeus spp., I
superfamily EREMAEOIDEA) (Woolley 1960) to aquatic drift in streams (Platyliodes sp.,
superfamily L10DOIDEA). Species of the genus Cepheus (CEPHEOIDEA) are common in
moss and litter (Schweizer 1957), but may be found in punky wood with burrowing
I
arthropods. C. latus (Nic.) was found to be a true wood borer in yellow birch twigs
(Wallwork 1958). C. cepheiformis is a vector of the tapeworms Cittotaenia ctenoides and
C. denticulata (Stunkard 1941) in Germany. Members of the family DAMAE IDAE
I
(DAMAEOIDEA) are litter and moss forms (Schweizer 1957) which may be fungivorous.
Gustavia spp. (ZETORCHESTOIDEA) also are litter forms, but the stylet-like chelicerae of
members of the family GUSTAVIIDAE may indicate a preference for a food substrate which
I
is atypical for the superfamily. Tectocepheus velatus (Mich.) is a widespread species of
CARABODIDAE (CARABODOIDEA) which is common in sandy habitats (Woolley 1960).
Another carabodoid (CARABODIDAE) is a known vector of a tapeworm species, as are
I
members of the litter-inhabiting superfamily L1ACAROIDEA (Allred 1954). Species of
Liacarus, Xenillus and Ceratoppia are Iiacaroid genera which have been implicated as I
tapeworm vectors.

Oppia spp. (OPPIOIDEA) are common inhabitants of pastures, moss or humus

I
(Woolley 1960, Sellnick 1960) but also may be coprophagous in twig galleries (Wallwork
1958). Other members of the superfamily are litter, moss or humus forms (Schweizer
1957, Wallwork 1967). Limnozetes ciliatus (Shrank) and other members of the HYDRO-
I
ZETOIDEA have adapted themselves to semiaquatic situations, and may be found in
floating sphagnum or in very wet moss (Schweizer 1956, Sell nick 1960). I
Cohort Poronoticina I
Poronoticine Cryptostigmata are those which possess a series of pores and porose
areas arranged symmetrically on the dorsal shield (Fig. 36) and which have pteromorphic I
extensions dorsolaterally. Eight superfamilies and over thirty families are recognized in the
Poronoticina on the basis of adult morphology. Using the condition of the nymphal and
larval cuticle as a major characteristic, Grandjean (1954) divided the poronoticines into
I
three groups:

1. Species with regularly folded, deeply striate integument-Plicate forms

I
2. Species in which the cuticle is smooth, but with small sclerites around the I
I
I
I 297
I notogastral setae in which the seta, are usually eccentrically inserted-the
Excentrosclerosae
I 3. Species which have large sclerites 01 both dorsal and ventral surfaces-the
Macrosclerosae.
I interlamellar seta

I
I
I
I
I
I
I
I
I
I Fig. 36. A poronoticine Cryptostigmata (Oribatei Supe iares) showing pores and porose areas (see detail).

Grandjean's system of immature classification illustrates a remarkable similarity to the

I superfamilial groupings established on the b sis of adult characteristics. The plicate
condition, for example, is found in the related "a ult superfamilies" AMERONOTHROIDEA,

I PASSALOZETOIDEA and PELOPOIDEA. Th Excentrosclerosae are virtually confined to

the superfamilies GALUMNOIDEA and ORIBATULOIDEA, while Grandjean's Macro-
sclerosae are contained in the superfamilies qERATOZETOIDEA and GALUMNOIDEA.

I There is little doubt that the study of additi~ral immature forms will reveal relationships
between families and superfamilies which eRisting adult-oriented classifications cannot
illustrate. Indications are, however, that the uperfamilial groupings currently recognized

I on the strength of adult features may be well funded in this cohort.

Like the HYDROZETOIDEA of the Pyc onoticina, the poronoticine family AMERO-
I NOTHRIDAE (AMERONOTHROIDEA) is ad pted to wet habitats. Hygroribates marinus
Banks is found on rocks in tidal zones and on stuarine strands where it apparently feeds on

I
I
298
I
algae (Grandjean 1947). Fortuynia yunkeri van der Hammen is found intertidally (van der
Hammen 1963) as is F. marina van der Hammen. Species of Selenoribates also are found at
I
the edge of the sea (Grandjean 1966b). Another ameronothroid, Cymbaeremaeus, is
coliected in dry sandy pine duff at high altitudes (Woolley 1960) or on the surface of scaly I
pine or spruce twigs. Alaskozetes antarctica (Michael) is found in algae and other substrates
in the Antarctica (Covarrubias 1968a), as are other members of the superfamily (Wallwork
1967).
I
Scutovertex minutus (Koch) is a member of the superfamily PASSALOZETOIDEA
which has been experimentally infected with eggs of the tapeworm Bertiella studeri
I
(Stunkard 1939). Passalozetoids usually are moss and litter inhabitants (Sellnick 1960), as
are the closely related PELOPOIDEA. Pelops tardus Koch, P. planicornis (Schrank) and
P. acromius (Hermann) are known vectors of the rabbit tapeworm Cittotaenia ctenoides.
I
Members of the litter and moss-inhabiting ORIBATELLOIDEA also may be naturally
infected by tapeworms. Wardle and McLeod (1952) found a species of the oribatellid I
genus Achipteria to be infected by an anoplocephaline cestode. Another Achipteria species
was found to feed on leaf epidermis (Wallwork 1958). The immatures of the latter species
may be coprophagous on the feces of the adults.
I
The MICROZETOIDEA includes a single family, the MICROZETIDAE, the members
of which inhabit humus or litter (Grandjean 1936).
I
Fuscozetes fuscipes (Koch), a member of the large superfamily CERATOZETOIDEA,
feeds on vascular tissue of hemlock needles, fungal hyphae, and moist decaying birch
I
leaves. Nymphs feed on dead mites or other arthropods (Wallwork 1958). Many ceratoze-
toids are common in moss and forest litter (Schweizer 1956, Sellnick 1960), although some
may be found in the littoral zone (Punctoribates quadrivertex Halbert, for example) or in
I
subarctic or arctic situations (Melanozetes meridianus Sellnick, Edwardzetes elongatus
Wallwork, E. dentifer Hammer) (Wallwork 1967). Species of the ceratozetoid genera I
Fuscozetes and Trichoribates are vectors of tapeworms (Allred 1954).

Probably no single genus of poronoticine Oribatei is as important in the transmission

I
of cestode parasites as the galumnoid genus Galumna. G. virginiensis Jacot is an important
vector of the sheep tapeworm, Moniezia expansa. G. emarginata (Banks) and G. nigra
(Ewing) also serve as vectors for this parasite (Allred 1954). Other species of Galumna
I
transmit several kinds of tapeworms to vertebrate hosts in Europe, Asia and North
America. Members of the GALUMNOIDEA are frequently collected from moss (Sell nick
1960) or in pasture sod. G. confusa Woodring and G. parva Woodring both were found to
I
feed on fungi in culture (Woodring 1965). Orthogalumna terebrantis Wallwork was found
in hyacinth leaf burrows (Wallwork 1965), and Pergalumna omnjphagous Rockett and
Woodring was observed to prey on live nematodes in laboratory cultures (Rockett and
I
Woodring 1966).
I
The ORIBATULOIDEA, like the CERATOZETOIDEA, are found in a wide range of
habitats. Scheloribates, Oribatula and Zygoribatula are common in pasture sad (Woolley
1960), but are not restricted to this substrate. Scheloribates spp., for example, are found
I
in stored grain (Hughes 1961), in leaf litter in the soil (Wallwork 1958, Woodring 1965) or
in moss (Schweizer 1956). Other genera such as Haplozetes and Peloribates (family I
I
I
II 299
I HAPLOZETIDAE) are primarily sailor moss for s. Along with the GALUMNOIDEA and
the CERATOZETOIDEA, several species of ORI ATULOIDEA are important vectors of
I tapeworms. Sche/oribates /aevigatus (Koch) aCls as the vector for no fewer than eight
tapeworm species (Allred 1954), while cysticerc, ids of two tapeworm species have been
recovered from each of three other Sche/oribate species. Uebstadia simi/is Michael is a
I vector of Cittotaenia ctenoides, and Oribatu/a ~inuta is the intermediate host of the
cestodes Monoecocestus sigmodontis and MonieZia expansa.

I Useful References
!
!
l

I Allred, D.M. (1954).

31.44-51.
Mites as Intermediate hosts of lapeworms. Proc. Utah Acad. Arts and Letters

I Block, W.C_ (1965), The life hIStories of PlatynothrfS pettifer (Koch 1839) and Damaeus clavipes
(Hermann 1804) (Acarina Cryptostlgmata) In soils of Pennine moorland. Acarologia 7(4)'735-743.
Covarrubias, R. (1968a). Some observations on Antarc Ie Onbatel (Acarina) Liochthonius australIs sp.

I n., and two Oppla ssp. n. Acarologla 10(2).313-356

Covarrubias, R. (1968b). Observations sur Ie genre Phe~/lOdes.
Oribatei). Acarologia 10(4):657-695_ [PLATERE
[OPPIOIDEAJ

AEOIDEA]
I-Pherollodes rob/ens/s n. sp. (Acarina,

I Grand]ean, F. (1933).
Grandjean, F. (1936a).
Etude sur Ie developpement deslOribates.
Les Microzetidae n. fam. (Oebates).
Bull. Soc. Zool. France 58(1):30-61.
Bull. Soc. Zoo I. France 61(2):60·93.
Grandjean, F. (1936b). Observations sur les Oribates (rOe serie). Bull. Mus. d'Hist. Nat. Paris Ser. 2,

I 8(3):246-249. [ORIBATULOIDEA] I
Grandjean, F. (1936c). Les Oribates de Jean Frederic H1ermann et de son Pere. Ent. France 105:27·110.
[GALUMNOIDEA]

I Grandjean, F. (1947). Observations sur les Oribates 17e serie).

19(2):165-172. [AMERONOTHROIDEA]
Bull. Mus. d'Hist. Nat. Paris Ser. 2,

Grandiean, F. (1954). Essai de classification des Oribate (Acariens). Bull. Soc_ Zool. France 78:421-446.

I Grandiean, F. (1955). Sur un Acarien des lies Kerguel n, Padacarus auberti (Oribate). Mem. Mus. Nat.
Hist. Nat. (n.s.) A, Zoo I. 8(3):109-150. [AMERON~OTHROIDEA]
Grandjean, F. (1956a). Observations sur les Galumnidae. Ire serie. Rev. Fr. Ent. 23:137-146.
I Grandjean, F. (1956b). Sur deux especes nouvelles d'O ibates (Acari ens) apparentees a Oripoda elangata
Banks 1904. Arch. ZoaL Exp_ Gen. 93(2):185-218 [ORIBATULOIDEA]
r (Oribate).
Grandiean, F. (1959)_ Palypterazetescherubin Berl.191 Acarologia 1(1):147·180.
I Grandjean, F. (1960a).
ZETOIDEA]
Les Mochlozetidae n. lam ( ribates). Acarologia 2(1):101·148. [CERATO·

I Grandiean, F. (1960b).
Grandjean, F. (1963).
[OPPIOIDEA]
Les Autognetidae n. lam. (Ori ates). Acarologia 2(4) :575-609. [OPPIOIDEA]
Les Autognetidae (Oribates) Deuxieme partie. Acarologia 5(4):653-689.

I Grandiean, F. (1966a). Les Staurobatidae n. lam. (Oriba es). Acarologia 8(4) :696-727. [ER EMAEOI DEA]
Grandjean, F. (1966b).
8(1):129·154.
Selenoribates mediterraneus n. sp. et les Selenoribatidae (Oribatesl. Acarologia
[AMERONOTHROIDEA] I
I Grandjean, F_ (1967). Nouvelles observations sur lejS Oribates (5e serie).
[EREMAEOIDEA]
Hammen. L. van der (1952). The Oribatei (Acari) of the Netherlands_
Acarologia 9( 1) :242-272.

Zool. Verhand. 17:1-139.

I Hammen, L. van der (1959). Berlese's primitive cribatid mites.

Hammen, L. van der (1963). Description of
Zoo I. Verhand. Leiden 40: 1-93.
Fortuvnia! yunker; nov. spec., and notes on the Fortuynidae
nov. fam. (Acarida, Oribatei). Acarologia 5(1):15r167. [AMERONOTHROIDEA]

I Hammer. M. (1944). Studies on the oribatids and collem ales of Greenland_ Medd. Gronland 141 (3): 1·21 O.

I
II
300
I
Hammer, M. (19581. Investigations on the oribatid fauna of the Andes Mountains. I. The Argentine &
Bolivia. BioI. Skrift. Danske Vid. Selsk. 10:1-162.
I
Hughes, A.M. (1961). The Mites of Stored Food. Ministry Agr. Fish. and Food Tech. Bull. 9:287 pp. + vi.
Kates, K.C. and C.E. Runkel (1948). Observations on oribatid mite vectors of Moniezia expansa on
pastures, with a report of several new vectors from the United States. Proc. Helminth. Soc. Wash.
I
15(1):18-33.
Michael, A.D. (1884). British Oribatidae. Ray Society, London 1:336 + xi + plates. I
Michael, A.D. (1888). British Oribatidae. Ray Society, London 2:337-657 + xi + plates.
Oudemans, A.C. (1917). Notizen liber Acari. 26 Reihe (Oribatoidea). Arch. f. Naturg. 83:1-84.
Riha, G. (1951). Okologie der Oribatiden im Kalksteinboden. Zool. Jahrb. 80:407-450.
I
Rockett, C.L. and J.P. Woodring (1966). Biological investigations on a new species of Ceratozetes and of
Pergalumna (Acarina: Cryptostigmata). Acarologia 8(3):511-520.
Schweizer, J. (1956). Die Landmilben des Schweizerischen Nationalparkes. 3. Sarcoptiformes Reuter
I
1909. Soc. Helvet. Sci. Nat. Pare. Nat. 5(N.F.) (34):215-377.
Schweizer, J. (1957). Die Landmilben des Schweizerischen Nationalparkes. 4. Ihr Lebensraum, ihre
Vergesellschaftung unter sich und ihre Lebensweise. Soc. Helvet. Sci. Nat. Pare. Nat. 6(N.F.) (37):
I
11-107.
Sellnick, M. (1928). Formenkreis: Hornmilben, Oribatei. Tierw. Mitteleur. 3:1-42.
Sell nick, M. (1960). Formenkreis: Hornmilben, Oribatei. Tierw. Mitteleur. 3:45-134 + plates.
I
Stunkard, H.W. (1939). The role of oribatld mites as transmitting agents and intermediate hosts of ovine
cestodes. Int. Kong. Ent. Berlin vii. 3:1671-1674.
Stunkard, H.W. (1941). Studies on the life history of the anoplocephaline cestodes of hares and rabbits.
I
Jour. Parasit. 27:299-325.
Stunkard, H.W. (1944). Studies on the life history of the oribatid mite, Galumna sp., intermediate host of
Moniezia expansa. Anat. Rec. 89(4) :550.
I
Trave, J. (1959). Sur Ie genre Niphocepheus Balogh 19431es Niphocepheidae, famille nouvelle. (Acariens,
Oribates). Acarologia 1(4):475·498. [CARABODOIDEA]
Trave, J. (1963). Oribates des Pyrenees-Orientales, 2e serie, Zetorchestidae (1re partie): Saxicolestes
I
pollinivorus n. sp. Vie et Milieu 14(2) :449-455. [ZETORCHESTOI DEAl
Wallwork, J.A. (1958). Notes on the feeding behaviour of some forest soil Acarina. Oikos 9(2):260-271.
Wallwork, J.A. (1959). The distribution and dynamics of some forest soil mites. Ecol. 40(4):557-563.
I
Wallwork, J.A. (1965). A leaf-boring galumnoid mite (Acari: Cryptostigmata) from Uruguay. Acarologia
7(4) :758-764.
Wallwork, J.A. (1967). Cryptostigmata (orbatid mites). Antarctic Res. Ser. 10:105-122.
I
Wardle, R.A. and J.A. McLeod (1952). Zoology of the Tapeworms. Unlv. Minnesota Press, Minneapolis:
780 pp. I
Woodring, J.P. (1965). The biology of five new species of oribatids from Louisiana. Acarologia 7(3):564-
576. [GALUMNOIDEA,ORIBATULOIDEA]
Woodring, J.P. and E.F. Cook (1962). The biology of Ceratozetes eisalpinus Berlese, Scheloribates
laevigatus Koch, and Oppia neer/andica Oudemans (Oribatei) with a description of all stages.
I
Acarologia 4(1 ):101-137.
Woolley, T.A. (1960). Some interesting aspects of oribatid ecology (Acarina). Ann. Ent. Soc. Amer.
53(1) :251-253.
I
Woolley, T.A. (1965). Eutegaeidae, a new family of oribatid mites, with a description of a new species
from New Zealand (Acarina: Oribatei). Acarologia 7(2):382-388. [CARABODOIDEA] I
I
I
I
I
I 301

I Suborder Crypt stigmata

I (Plates 128 to 144, pp. 304-320)

KEY TO THE SUP!RFAMILIES

I 1. Femora of legs divided. Gnathosoma visible from above (astegasimy). Tarsi may not
I be monodactylous in all immature stages. S all weakly sclerotized species with four

~~I.e~~~i~.o.n ;~~se~~~~;r~~~~A'EACA'Ri ipI1~e' ; 3D) S~~~~f~~ii; PA'LAEACAROI'DEA

I Femora of legs not divided. Gnathos9ma generally not visible from above
(stegasimy). Tarsi always monodactylous I'n larval (Plate 8-3) and nymphal stages.

I ~e~I~~~ .~r. ~t~~~~I:. ~cler.o.t~z.e~. ~~e.ci~s. ~~thl ~~~er. ~~~~ ~~~~ sOI.e.n.i~,a. ~~ ta~s~~ ~ of 2

I 2. With at least one of the following charac~eristics

well discernible: propodosoma
ptychoid (capable of being "jack-knifed" tto hysterosoma (Plate 131-1); tibia and
genu of about uniform length and shaPje; genital and anal shields contiguous,

I occupying entire length of ventral shield wrich may be divided into two parts by a
horizontal, semicircular or parabolical transverse suture; anal and adana I shields
often separated (Plate 129-1) , upercohort OR/BATE//NFER/ORES 4
I Without the above characteristics; that is: p opodosoma aptychoid, (not constructed
so that it may be "jack-knifed" into hy terosoma); tibia longer than genu, and

I shaped differently; genital and anal shields rounded, usually well separated and not
covering entire length of ventral side; ventr I shield without transverse suture; adana I
shields absent ,upercohort OR/BATE/ SUPER/ORES 3

I 3. With short horizontal or well developed downward bending pteromorphae pro-

truding laterally from idiosoma, and/or VI ith area porosae (Fig. 36), sacculi, and
I brachytrachea on dorsum of idiosoma (not gaster) .... Cohort PORONOTICINA ... 9
Without pteromorphae (certain L1ACAR IDEA are exceptions (Plate 142-1)); no

I 4.
area porosae, sacculi, or brachytrachea on notogaster. Cohort PYCNONOTICI NA ... 15

Propodosoma ptychoid; body generally co pressed laterally. . . . . . . . . . . . . . . . . .. 5

I Propodosoma aptychoid (certain HYPO HTHONOIDEA are exceptions); body

cylindrical or flattened dorsoventrally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 6

I 5. Genital and anal shields generally rounded, separated from each other
............................. (Plat 130) Superfamily MESOPLOPHOROIDEA
.

I Genital and anal shields extending over wh, Ie length of ventral side, contiguous ....
.............................. (PJI te 131) Superfamily PHTHIRACAROIDEA

I 6. Notogaster with 1-3 transverse sutures

Notogaster without transverse sutures
"
"
7
8

I 7. With one indistinct transverse suture 01 the notogaster; lateroabdominal glands

present (Plate 1 1) Superfamily PARHYPOCHTHONOIDEA

I
I
 I
302

With 1-3 distinct transverse sutures on the notogaster; lateroabdominal glands

I
absent (Plate 132) Superfamily HYPOCHTHONOIDEA

8. Propodosoma and hysterosoma movably connected .

I
· . .. .. .. . . . . .. .. .. . .. . .. (Plates 132, 133) Superfamily PERLOHMANNOIDEA
Propodosoma and hysterosoma immovably fused . I
· (Plates 133, 134 [see also 10-8J) Superfamily NOTHROIDEA

9. Pteromorphae greatly expanded, extending both forward and behind .

I
.. . . . . . . . .. .. . . . .. . . . . . . . .. . .. .. .. (Plate 134) Superfamily GALUMNOIDEA
Pteromorphae generally distinct but without anterior and posterior extensions ... 10 I
10. Chelicerae narrow, elongate; interlamellar hairs usually very large, leaf-shaped; body
frequently covered with thick cerotegument and some fusiform dorsal setae . I
· , (Plates 134, 135) Superfamily PELOPOIDEA
Chelicerae not as above, interlamellar hairs usually not leaf-shaped; body generally
without cerotegument and fusiform dorsal setae 11
I
11. Minute species (ca. 250 II), with relatively large lamellae; chelicerae with bacil-
liform lateral appendages (Plate 135) Superfamily MICROZETOIDEA
I
Species usually larger than 250 II; chelicerae without bacilliform lateral appendages.. 12
I
12. Lamellae extremely wide, meeting or fusing medially, covering major portion of
prodorsum; without translamellar bridge
·
.
(Plate 136) Superfamily ORIBATELLOIDEA
I
Lamellae variable, often absent; if broad, then with translamellar bridge 13
I
13. Notogaster with rough sculpturing; with coarse wrinkles, tubercles, reticulation, or
with thick, dark chitinization
· .... (Plates 136,137) Superfamilies AMERONOTHROIDEA + PASSALOZETOIDEA
.
I
Notogaster generally smooth, without sculpturing as above

14. With at least one of the following characteristics well discernible; 1-5 pairs of
14
I
genital setae; lamellae attenuated anteriorly, almost always without translamella
and cusps; pteromorphae protruding from outline of body
· (Plate 137) Superfamily ORIBATULOIDEA
. I
With 6 pairs of genital setae; lamellae usually not conspicuously attenuated
anteriorlY, translamella (Plate 138-1) and cusps generally present; pteromorphae
I
15.
curved ventrad " (Plate 138) Superfamily CERATOZETOIDEA

Hysterosoma with protruding lateral opisthonotal glands .

I
· (Plate 138) Superfamily HERMANNIELLOIDEA
Hysterosoma without lateral opisthonotal glands 16
I
16. Notogaster often with eccentric layers of nymphal exuviae. Genital shields each with
a transverse suture " " ., (Plate 139) Superfamily L10DOIDEA
I
I
I
I
303
I
~~~~;:~t~~I.~~~~i~~,.~~~~.~r~~~~t: .n.o.t. ~s. ~re... ~.e~~t~~ ~~i.e~~s. ~~t~~.u~ .t~~~s~~:s~ 17
I 17. With at least one of the following charapteristics well discernible: legs long,

I filiform, with three minute claws; genu, tibia and tarsus with socket articulations;
pseudostigmatic organ flabellate or lichen if 'rm; only 2-6 pairs of posteromarginal
notogastral hairs present (Plate 140) Superfamily PLATEREMAEOIDEA

I Legs not filiform or, if so, then monoda<fylous; genu, tibia and tarsus without
socket articulations; pseudostigmatic organs Igenerally not flabellate or licheniform;
with 7-14 pairs of notogastral hairs, some Of~hiCh are not posteromarginal 18
I 18. Legs long, each shaped like a string of earls, composed of spherical swollen
segments; apoteles monodactylous. Hyster soma usually spherical, with 8 pairs of
I ,
notogastral hairs in two longitudinal row and three pairs of posteromarginal
hairs (Plate 140) Superfamily DAMAEOIDEA

I f
Legs neither long nor with swollen joints if leg joints appear swollen, then with
less than 8 pairs of notogastral hairs). Hystersoma not as above 19

I 19. Body covered with cerotegument and ad1herent dirt.

rounded, protruding considerably beyond roftrum .,
Lamellae wide, apically
.
· (Plate 1F9) Superfamily POLYPTEROZETOIDEA
I Body not covered with cerotegument and adherent dirt. Lamellae various 20

I 20. With at least one of the following Charajters well discernible: chelicerae long,
attenuate, without movable chelae, serrate distally; leg IV modified for jumping,
with a thick spine; rostral setae near each other, plumose, flabellate or bifurcate.
I .......................... (Plates 14 ,141) Superfamily ZETORCHESTOIDEA
Without the above characteristics 21

I 21. Prodorsum with true lamellae each usually ~earing lamellar hair on cusp 22
Prodorsum without true lamellae (rarer narrow costulae present, without
I :r:t:~:::etrc:~:~.;~U'9'h'~~~I~~~;i~~;'~i~~ 'clo~;~e'~'r;n'~I~~,':U~~;~I~~ 'r~~i~~;~t;~~,23
'o'r
22.
I or at least with thick dark chitinization . .. .
·
.
(Plate 141) Superf milies CEPHEOIDEA + CARABODOIDEA

I Notogaster smooth, rarely with fine retic lation, granUlation or foveolation, often
dark but not thickened (Plate 142) Superfamily lIACAROIDEA

I 23. Pseudostigmata and pseudostigmatic org ns extremely small, sometimes absent.

Found in fresh water (I late 143) Superfamily HYDROZETOIDEA
Pseudostigmata and pseudostigmatic organ always distinct. Terrestrial forms ..... 24
I 24. Ventral shield with four pairs of setae.
(Plates 142, 143) Superfamily OPPIOIDEA
I Ventral shield with more than four pairs of setae .
· " (Plate 144) Superfamily EREMAEOIDEA
I
I
 PLATE 128
I
304
I
notogastral shields
I
pygidium (notogasterl

I
I
supraPle"u~ra~I
shield-
-I::=-_A~::J1:0 I
pleural
shield
I
adanal shield

anal shield
I
genital shield
aggenital shield
128-1
I
I
I
notagaster
I
I
I
I
I
I
128-2 I
I
128-1 and 128-2; hypothetical Cryptostigmata illustrating primitive (128-1 J and advanced {128-21 shield states. Primitive famls
such as the BRACHYTHONllDAE may have a full complement of shields as shown in 128-1 (after KnUile 1957)

I
I PLATE 1129
305
I M
a,
I ...
N

I
I
I
I
I
I

~
'~-:l:r~.
I i..) V.
'..... / .....j

~
( .
I (""; ("\

I
I
I
I
I
I
I
I
I 129-1 to 129-6; posteroventral shield states in the Cryptostigmata. 129-1; hypothetical Cryptostigmata illustrating the primitive
posteroventral shield state: 129·2; Acaronychus sp. (PA~EACAROIDEA)' with weak or indistinct shield development:
129·3; Nanhermannia 'p. (NOTHROIDEAI. illustrating di gastry: 129-4; Eulohmannia ,p. (PER LOHMANNOIDEAI.

I
illustrating pseudodiagastry: 129-5; Epilohmannia sp. (P R LOHMANNOIDEA), illustrating schizogastry: 129-6; Hermannia
sp. (NOTHROIDEA), illustrating simple genital and anal s ield condition with aggenital and adanal shields fused laterally
 PLATE 130
I
; 306
I
I
I
I
I
I
I
pygidium ---I-.,q..
I
I
I
I
I
I
I
I
I
I
130-4
I
130-1 and 130-2; superfamily PALAEACAROI DEA_ 130-1; Palaeacarus hystricinus Triigardh, dorsum (after Balogh 1961): 130-2;
Acaronychus sp. (Oregon, USA). leg I with detail of tarsus
I
130-3 and 130-4; superfamily MESOPLOPHOROIDEA, Mesoplophora sp. 130-3; lateral aspect of female: 130-4; posteroventral
region (after Baker et al_ 1958) I
I
307
I
I
I
I
131·1
I
I
I
I chelicera.--I-~

I
I -,l~ ~~
I '1.5M~~~"~·' 131-2

I 7f Ilr~ll ~
:
I X\
1-1-\~ T"·~"
I X
genital and
anal
contiguous

/~r. ·'\
:
I
131_3~m~~~~'~'
I 131-1 and 131-2; superfamily PHTHIRACAROI DEA, Euphthiracar ssp .. lateral aspect illustrating "jack-knifing" or ptychoidy:

I 131-3;
131-2; genital-anal region
superfamily PARHYPOCHTHONOIDEA, parhYPOchthoniur sp_ (British Columbia), dorsum
 PLATE 132 I
308
I
I
I
I
I
I
~~~~ _ _ genitalshield

I"x-'G-ld---"N-t'~--aggenital shieid
I
~~=--_pleural shield
I
I
I
I
I
I
I
I
I
I
I
132-1 and 132-2; superfamily HYPOCHTHONOIDEA, EobrachVthonius sp. (Oregon, USA). 132·1; venter of female: 132·2;
I
dorsum of female
132-3; superfamily PERLOHMANNOIDEA,Perlohmannia sp. (Oregon, USA), anterodorsal region
I
 II
309
I
I
I
~ empodium claw-like

I
I
I
I
I
I
I
I
I
I
I
I adoral sclerite-~-\-4-

palpal trochanter

I
I mentum_+-----...A

':I 133-1;
133-2;
r
superfamily PERLOHMANNOIDEA, Perlohmannia sp., vejter of female
superfamily NOTHROIDEA. Hermannia sp_ (Oregon, USA, venter of gnathosoma
 PLATE 134
I
310
I
I
I
I
I
I
I
I
I
I
I
I
I
pteromorp hae expanded,
extending anten
posterior
.or and I
I
134·2 I
I
I
I
NOTH~NOIDEA, . p dorsum US A)B'keretal.
OIDEA Hermann/asp"(Oregon, venter 1958 )
134·1;
134·2;
134·3;
family
superfamily
super
superfaml
'Iy GALU Galumna
PE LOP OIDEA , Eupelops sp.,S c'helicera (after a
I
I PLATE 135
311
I interlamellar hair

I pterom rphae without

anterior and posterior
I ;:--_~e.xtenSi0i"S
fixed digit with lateral
appendage

I \
I
f
I I f \ \ 135-2
I
V I \' / dollsal
co setae
manly
fus form

I
\, / "~,/\
I
I 135·'
1~t~Jrr
\ pseudostigmatic

I organ_

I
I
I
••
I
••
,. I
135-1; superfamily PELOPOIDEA, Eupelops acromios (Hermannl.rdorsum (after Balogh 19651
135-2 to 135-4; superfamily MICROZETOIDEA, Microzetes sp_ 13 -2; chelicera (after Baker et al.19581: 135-3; venter:
135-4; dorsum (after Baker et al_ 195B)
 PLATE 136 I
312
I
136-1
I
pteromorphae
with narrow
I
anterior
cusps
I
I
I
136-3
I
I
I
I
I
I
I
I
I
I
I
I
136-1 and 136-2; superfamily ORIBATELLOIDEA. 136-1; ?Achipteria sp. (Oregon, USAl. anterodorsal region: 136-1; Oribatella
sp. (Oregon, USAl. dorsum
I
136-3 and 136·4; superfamily AMERONOTHROIDEA. 136-3; Ameronothrus lineatus (Thorelll. anterodorsal region: 136-4;
Scapheremaeus palustris Sellnick (Oregon, USA), dorsum
I
I PLATE 1)37
313
I
137-1
I 1 pair of
~1l.-'l----genitalsetae

II
I
I
! '\
I 137·2

••
I
••
••
I
• 137-4

I
• 5 pairs of ---+--
genital setae

•• 137-1;
13('3
superfamily PASSALOZETOIDEA, Licneremaeus prOdig;1SUS Schuster, anterodorsal region

•
137-2 to 137-4; superfamily ORIBATULOIDEA. 137-2; Pirnodus\detectidens Grandjean, genital shields (after Balogh 19611:
137-3; Euscheloribates sp. (Colorado, USA), dorsum with ,ateral detail of (from left) Cantharozetes lucens Hammer
(after Balogh 1965) and Xv/obates sp. 137-4; Pi/oba,es prsellus (Balogh), genital shields (adapted from Balogh 19651
 PLATE 138 I
314
I
138-1

'-=~I-+---::---
lamellar seta

cusp
translamella
pseudostigmatic organ
••
I
I
I
I
I
I
I
I
lateralopisthonotal-
gland I
6 pairs of genital
I
\ /
setae

I
I
r~~ I
138-3
I
138-2 I
138·1;
138·3;
superfaml'1 Y CERAT
superfamrly
OZE TOI DEA, Trichoobate.
HERMANNIELLOID . ssp
EA , Hermann,. dorsum.
1/;'sP.
. 138-2; ?genus, genital shield
(Oregon, USA) , dorsum I
I
 •• PLATE 139
315

•• ~ \I-'!+-i-- apoteles tridactylous

I 139-2

I
I
I
I
•• lamellae wide,'
rounded
•

I
••
I
•
I 139-3

I
316
PLATE 140 I
I
pseudostigmatic organ
flabellate
I
I
legs long, composed of
L.)!70---spherical swollen segments
••
I
140-3 I
I
•
I
I
I

140-2
i __ rostral seta
••
I
socket----\'iO'-.(
articulation genu III
I
rIi!/ \
~
3'-*
I
140-4

140-1 and 140-2; superfamily PLATEREMAEO IDEA. 140-1; Licnodamaeus undulatus (Paoli). dorsum (after Balogh 1965): 140-2;
Plateremaeus sp., femur and genu III
•
I
•
140·3; superfamily DAMAEOIDEA, ?Damaeus sp. (Oregon, USA), venter and dorsum
140·4; superfamily ZETORCHESTOI DEA, Zetorchestes f1abrarius Grandjean, anterodorsal region (after Balogh 1965)
I
••
PLATE J'41
317

chelicerae attenuate, serrate

distally

I
I
•
I
I
I
I
I
I
I
I ~- notogaster pitted

I
I
•
I
141-3
I
I 141·1;
141·2;
superfamily ZETORCHESTOIOEA. Gustavia sp. 10regon, ~SA)' anteroventral aspect
superfamily CARABODOIDEA, Carabodes marginatus IMiphael), dorsum lafter Balogh 1965)
141-3; superfamily CEPHEOIDEA, Cepheus sp. 10regon, USA), drrsum
I
 PLATE 142 I
318
I

142-2
•
I
I
I
I
I
I
I
142-4
•
I
I
I
142-3 I
I
adanals

',1(0.\
1
••
notogaster
smooth
~----~~
\ 7-\\ 142-5 I
142·1 to 142-3; 142.'1 .' Liacarus
,uperfamll L1ACAROIDE:';'ing 142-1;
latus 142.4; Suctobe
(Oregon, ~:a~;'tusa
Hatenreffri:n'i.;odorsal
(Oregon,
Jacot (Conn
S
regi~~ticut,
' 42·2; Cultoribula ,p., (Oregon,
U A) , venter,
USA),1 leg I: 142.5', Multtoppt
. 'a ,p. I
U~42:5'
A) palp: OPPIOIDEA.
142-4 an10rego~, USA), poster
,uperfamlly oventral region
I
I PLATE 43 ~
•
I
319

I
I
I
I
I
I
I
I
••
I
••
•
I
I
I
•
143-1; superfamily OPPIOI DEA, Multioppia sp., dorsum
143-2; superfamily HYDROZETOIDEA, Hydrozetesconfervae (S hrank). dorsum (after Balogh 1965)
320
PLATE 144 I
I
144-1 I
without distinct lamellae
I
~+---_ interlamellar seta I
~---
I
I
I
I
I
I
144-2
I
I
I
ventral shield w i t h - -
many setae
I
I
anal s h i e l d - - - - ¥ - - - - - I I -

•
I
I
144-1 and 144-2; superfamily EREMAEOIDEA, Eremobelba flaggelaris Jacot (Connecticut, USA). 144-1; dorsum:
144-2; posteroventral region (diagrammatic)
I
I
I 321

I
I
I
I
I
I
I
I
•
I
I
I
I
I
I
••
I
I
I
 I
322

NOTES I
I
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I
•
I A
INDEX

(Numbers in bold face refer to illust ations in the text and plates)
323

I Abacarus; 149
Aberoptinae; 148
j
MERONOTHROIDEA; 288, 297, 302, 312
merono thrus; 312
Acarapis; 43 MEROSEIIDAE; 71, 88, 107
I Acari; 1, 6, 9
acariasis; 42, 232 i meroseius; 107
nactinochaeta; 25
ACARIDAE; 27, 28, 34, 38, 39, 40, 42, 229, Anactinochitinosi; 25
I 230,232,250,251,255,260,261
Acaridia; 249
i\1 nactinotrichida; 25
anal pedicel; 39, 40, 75

~
Acaridiae; 59 nal shield; see ventral shields
I Acariformes; 9, 58, 141, 143, 228, 288, 290-291
acarinarium; 39, 71
nal sucker; 260
nal valves; 67, 68
Acaroidea; 232-238, 249 na,ges; 244, 274, 275
I
I
ACARONYCHIDAE; 292
Acaronychus; 292, 305, 306
Acarophenax; 146, 187
Acaropsis; 163, 204
Acarus; 1, 38, 232
accessory gland; 26, 27, 28
i NALGIDAE; 41,243-244,253,274,275
nalgoidea; 243-245, 252
ndre's fluid; 49
f:ndrolaelaps; 119, 120
fngelothrombium; 168
1"NOETI DAE; 228, 231,249, 255, 256

I Aceria; 149
Aceropsis; 224
fnoetoidea; 31,231-232, 249
fnoetus; 231
acetabula, genital; 225 fntennomegistus; 79
I Achipteria; 298
Acomatacarus, 168
tNTENNOPHORIDAE; 78, 94,131
~ntennophorina;92
Actacarus; 155 CntennoPhoroidea; 78, 79, 93
I Actinochaeta; 25
actinochitin; 25
fntennophorus; 79,131
rnterior polar cap; 31
Actinochitinosi; 25 ~nus; 25, 26

I Actinotrichida; 25
adoral sclerite; 289, 309
tNYSTIDAE; 27, 37, 161, 179, 201
I4nystis; 161, 201
aedeagus; 21, 22, 228, 260 IAnystoidea; 156, 161, 162, 179

I Afrophioptes; 205
Agaue; 14, 189
I'i\phelacarus; 292
'Ap;onoseius; 75

~
Agistemus; 159 Plonobia; 157
I air sac pore; 64, 66
Alabidocarpus; 283
apodeme; 228, 230
Apolonia; 168, 169
Alaskozetes; 298 Aponomma; 136

I Aleuroglyphus; 232
Alicorhagia; 196
apotele, palpal; 14,62,64,65,66,67,68
apotele, tarsal; 17, 18
AlICORHAGIIDAE; 144, 178, 196 Arachnida; 1, 5

I AI/ochaetophora; 158, 211, 212

ALLOCHAETOPHORIDAE; 158, 180, 211, 212
Araneae; 5, 6, 7
Archemyobia; 163
AI/othrombium; 167, 221 ARCHEONOTHRIDAE; 292

I Amblyomma; 140
Amblypygi; 5
ARCTACARIDAE; 88, 95, 96, 106
Arctacarus; 95, 96, 106
Amblyseius; 108, 109 Argas; 137, 140

I ambulacrum; see apotele, tarsal

AMERONOTHRIDAE; 297
ARGASIDAE; 35, 136, 137,138,140
Arhodeoporus; 155

I
I
324
I
Arrenurus; 224
arrhenotokv; 31
CALIGONELLIDAE; 160,181,214,215
Caloglyphus; 34, 39, 232, 260, 261
I
Arthropoda; 5
Asca; 111
CalvoNa; 236, 263
Calyptostoma; 24,166,218 I
ASCIDAE; 37,39,40,70,71,89,111 CALYPTOSTOMIDAE; 166,181,218
Aspidilaelaps; 71
aspis (see also prodorsal shield); 304
Astacopsiphagus; 155
Camerobia; 160, 216
CAMEROBIIDAE; 181
camerostome; 89,112
I
astegasimv; 301,306
Asternolaelaps; 86
Astigmata; 12, 16, 19,21, 24, 25, 28, 31, 35, 41,
Caminella; 26,75,123
Camisia; 294
Campylochirus; 283
I
59, 228-231, 249, 292
ATOPOMELIDAE; 239, 254
canal, gland; 262
CANESTRINIIDAE; 236,238,249,258
Canestrinioidea; 238-239, 249
I
Audycoptes; 242, 269
AUDYCOPTIDAE; 230, 242, 252, 269
Austrochirus; 239
AUTOGNETHIDAE; 289
Cantharozetes; 313
Caparinia; 270
Carabodes; 317
I
AXONOPSIDAE; 172

B
CARABODIDAE; 296
CARABODOIDEA; 289, 296, 303, 317
CARPOGLYPHIDAE; 232, 233, 251, 255, 264
I
Bakericheyla; 163
Bakerocoptes; 246
Carpoglyphus; 233, 264
caruncle; 229
Cas; 114, 115
I
Balaustium; 166,217
Bdella; 153, 193
BDELLIDAE; 152, 153, 177, 193
CELAENOPSIDAE; 80, 81,94,131
Celaenopsoidea; 80-81, 93
cement laver; 11, 11
I
Bdelloidea; 152, 177
Bechsteinia; 201
Benoinyssus; 150
cephalothorax; 6, 7
CEPHEOIDEA; 289, 296, 303, 317
Cepheus; 296, 317
I
Berlese funnel; 45
Biscirus; 153
blastoderm; 31
Ceratoppia; 296
CERATOZETOIDEA; 297,298,299,302,314
CERCOMEGISTIDAE; 78, 92, 95, 128
I
Blattisocius; 71,111
blister mite (see also ERIOPHYIDAE); 148, 149
Bontiella; 242
Cercomegistina; 92
Cercomegistoidea; 78, 92
Cercomegistus; 95,128
I
Bonzia; 153
Boophilus; 136
Boydaia; 151, 200
CHAETODACTYLIDAE; 234, 251, 255, 266, 267
Chaetodactvlus; 234, 266, 267
chaetotaxy, dorsal; 69
I
BRACHYTHONIIDAE; 304
Brachythonius; 289
brain; 12, 25, 26, 28
chaetotaxv, leg; 18, 18, 119
Charletonia; 166
Chelacaropsis; 162, 163
I
Brevipalpus; 14, 22, 158, 192
Bryobia; 157
bud mite (see also ERIOPHYIDAE); 148, 149
Cheletomorpha; 14,163
Cheletosoma; 163
chelicerae; 5, 6,12,13,14,15,21,26
I
bursa copulatrix; 21, 23, 27, 28, 228, 268

C
Chelicerata; 5
CHEYLETIDAE; 14, 33, 37,162-163,179,204
Cheyletiella; 162, 163,204
I
caecae, gastric; 25, 26
CAECULIDAE; 154, 179,200,201
Caeculisoma; 166
Chevletoidea; 162-165, 174, 177, 179
Cheyletus; 33,163,204
chicken mite;41
I
Caeculoidea; 154-155, 179
Caeculus; 200, 201
Calholaspis; 102
chiggers (see also TROMBICULIDAE); 41, 168
Chinodex; 164 I
I
I
I
325
I Chirnyssus; 246 Cl[ticulin; 11, 11
Chirobia; 246, 284 C c1othorax; 80

I Chirodiscoides; 239
Chirodiscus; 239
C mbaeremaeus; 298
C~RTOLAELAPIDAE; 87
Chirorhynchobia; 240, 281 C todites; 42,247,279

I CHIRORHYNCHOBIIDAE; 228, 239, 240, 254,

281
CI TODITIDAE; 42,247,254,279
Clytoditoidea; 247-248, 251
Chorioptes; 241, 271 Crenspinskia; 234, 264

I CHORTOGLYPHIDAE; 232, 233, 250, 259

Chortoglyphus; 259
OZENSPI NSKII DAE; 251

Chrysocoptes; 239 o
I CIRCOCYLLIBANIDAE; 76,91,125
Circocyl/iba; 125
AMAEiDAE; 296, 303
AMAEOIDEA; 296, 316
circumcapitular suture; 6,12
amaeus; 316
I Claparede organ; 23, 24
classification, higher; 55-57
claw, palpal thumb; 142
ASYPONYSSIDAE; 89,115
asyponyssus; 115
emodex; 41,164,184
I claws; 15, 17
CLOACARIDAE; 164, 174, 184, 185
EMODICIDAE; 41,164,174,184
entidinychus; 76
Cloacarus; 184, 185
epiuming mite; 246
I Coccotydeus; 199
collecting techniques; 45-49
'ermacarus; 234, 263
ermacentor; 136
condylophore; 229, 250
"ERMANYSSIDAE;41, 71, 72, 90,117
I Copidognathus; 155
coprophagy; 39
Coptocheles; 214, 215
'ermanyssus; 41, 71, 117
1 ermatophagoides; 242, 270
ermatophagoidinae; 242
I copulation; 20
corniculus; 13, 67
cornu sacculus; 23, 28
ERMOGLYPHIDAE; 41,244,253,277,278
ermoglyphus; 244
eutogyne; 149
I costula; 319
coxa; 17, 18, 34
coxal field; 228, 256, 283
,eutonymph Isee also hypopus); 34, 35, 75
i eutosternum; 13, 68
eutovarial membrane; 31
I Coxequesoma; 76, 125, 126
COXEQUESOMIDAE; 76, 91,125, 126
crista metopica; 181,217,218
I eutovum; 31
iagastry; 305
IARTHROPHALLIDAE; 90,121
I Crotalomorpha (ms-l; 188
CROTALOMORPHIDAE Ims.); 146, 175, 188
CRYPTOGNATHIDAE; 159, 175, 188
iarthrophallina; 90
iarthrophalloidea; 77-78, 90
iarthrophal/us; 77, 121
I Cryptognathus; 188
Cryptostigmata; 12,16,19,20,21,24,25,31,
33,35,37,39,41, 50, 51, 59,144,288-292,
icrocheles; 33
IGAMASELLIDAE; 71, 88,108
pigamasel/us; 108
I 301, 304, 305
pinogamasus; 39

•
CTENACARIDAE; 292
pinothrombium; 168
Ctenacarus; 289
DINYCHIDAE; 76, 91,124
CTENOGL YPHIDAE; 232,233,250,265
b!PLOGYNIIDAE; 15, 80, 93, 95, 96,130
Ctenoglyphus; 232, 265
eiP,othrombium; 219
Cultoribula; 318
fipolyaspis; 83
I Cunaxa; 153, 193
CUNAXIDAE; 152, 153, 177, 193
disc, genital; 22, 24, 27,141
!discidium; 304
Cunaxoides; 153, 193
~iSCOUrel/a; 123
I custodium; 304
cuticle; 11
eISCOURELLIDAE; 75, 91,123
disjugal furrow; 17

I
I
326
I
dissection techni;ues; 50-51
Dolichotetranychus; 158, 191
Erythroides; 166
esophagus; 25, 26, 28
I
Doloisia; 169
dorsal chaetotaxy; 69
dorsal pores; 295, 297
Eulohmannia; 305
EUPALOPSELLIDAE; 22,159,175,216
Eupalopsellus; 216
I
dorsal shields; 16, 140,221,267,304
Dorsipes; 146
duct, ejaculatory; 22, 27, 28
Eupalopsis; 152
eupathidium; 13, 24, 25
Eupelops; 310, 311
I
duplex seta; 210
Dyscitaspis; 83, 91,122

E
Euphthiracarus; 294, 307
Eupodes; 150, 190, 191
EUPODI DAE; 39, 150, 176, 190, 191
I
Echinomegistus; 79, 129
Edentalges; 241
Eupodoidea; 144, 150, 176
Eupodostigmatina; 174,176,178
Euroglyphus; 242, 270
I
Edwardzetes; 298
egg; 33
ejaculatory duct; 22, 27, 28
Euscheloribates; 313
Euschiingastia; 168, 222
Eutarsopolipus; 146
I
Eleutherogonina; 20, 174
embryogenesis; 31
empodium (see also featherelaw); 15, 17,229
Eutrachytes; 34,125
EUTRACHYTIDAE; 34, 76, 91,125
Eutrombieula; 168
I
Enarthronota; 293
Endeostigmata; 141, 177
Endotrombicula; 169
Eutrombidium; 168
Euzercon; 81, 95, 132
EUZERCONIDAE; 80,81,94,95,132
I
Eniochthonius; 294
Ensliniella; 236
ENSLINIELLIDAE; 251
EVANSACARIDAE; 246, 254, 283
Evansacarus; 246, 283
EVIPHIDIDAE; 71, 90, 95,118,119
I
Enterohalacarus; 43
ENTONYSSIDAE; 42, 48, 71, 89, 90,114
Eobrachythonius; 308
Eviphis; 95,118,119
Ewingia; 42,241,280
EWINGIDAE; 42,241,254,280
I
Eotetranychus; 22, 157
EPICRIIDAE; 70, 86, 99
Epicrius; 99
Ewingoidea; 240-241, 254
Excentrosclerosae; 297
exocuticle; 11, 11
I
epicutiele; 11, 11
Epidermoptas;244,277
EPIDERMOPTIDAE; 244,253,277
exoskeleton; 5, 11
Exothorhis; 159
eyes (see also ocelli); 24
I
epigynial flap; 230
epigynial shield; 83, 84
epipharynx; 26
EYLAIDAE; 171, 172
Eylais; 171, 223 I
epistome; 6,12,13,26,67,95
EREMAEOIDEA; 23,296,303,320
Eremaeus; 23, 296
F
Falculifer; 234, 244, 277 I
Eremobelba; 320 FALCULIFERJDAE; 253
ereynetal organ; 178,200
Ereynetes; 199, 200
famulus; 24, 25, 260
feather mite; see Analgoidea I
EREYNETIDAE; 14, 15,42,151,178,199,200 featherelaw; 183
erineum; 149
Eriophyes; 149
ERIOPHYIDAE; 27, 38, 148, 174, 183
Fedrizzia; 129
FEDRIZZIIDAE; 92,129
Fedrizzioidea; 82, 92
I
Eriophyinae; 148
Eriophyoidea; 141, 146, 148, 158, 174
ERYTHRAEIDAE; 27, 41,166,181,217
femur; 17, 18
Fessonia; 218
folliele mite; see DEMODICIDAE
I
Erythraeoidea; 165-167, 181
Erythraeus; 166
Fortuynia; 298
fovae pedales; 90, 91, 121 I
I
I
I
327
I Frevana; 273 H lotvdeus; 150, 153
FREYANIDAE; 244, 252, 272, 273 H nnemania; 169

I fungivorous mites; 38-39 H plozetes; 298

•
FUSACARIDAE; 234, 250, 255, 261 HtPLOZETIDAE; 299
Fusaearus; 234, 261 H RPYRHYNCHIDAE; 22,163,180,205
Fuseazetes; 298 H, rpvrhvnehus; 22, 205
Fuseurapoda; 76,127
hiart; 25
H mipteroseius; 42,109,110

I G
gall mite (see also ERIOPHYIDAE); 148, 149
Galumna; 198, 310
H misareoptes; 238, 257
H MISARCOPTIDAE; 238, 249, 255, 257
H mitarsonemus; 146

I GALUMNOIDEA; 297, 298, 299, 302, 310

Gamasel/us; 105
Gamasiphis; 105
H ricia; 267
HfRICIIDAE; 251
Hfrmannia; 39, 294, 305, 308, 310

I Gamasina; 86
gastric caecae; 25, 26
GASTRONYSSIDAE; 248, 253, 279
Hrrmanniel/a; 314
HjERMANNIELLOIDEA; 301,314
Gastranvssus; 248, 279 H,ETEROCHEYLIDAE; 15, 164, 177,206

I Geekobia; 161,202
Geekabiel/a; 161, 202
Heterochevlus; 15, 206
Heterostigmata; 59, 174
genital acetabula; 225 terozercon; 104

I genital disc; 22, 24, 27, 141, 228, 257, 289

genital shields; see ventral shields
HETEROZERCONIDAE; 71,87,104
h gher classification; 55-57
genital tracheae; 21 h ndgut; 25, 26

I genu; 17, 18
germinal band; 31
irstiel/a; 161
irstionvssus; 120
gland, accessory; 26, 27, 28 istiasama; 231 , 256

I gland, opisthonotal; 260, 307 alaeelaena; 71,102

~
gland, salivary; 12, 26 Olostaspel/a; 13, 102
gland canal; 262 OLOTHYRIDAE; 64,65,66

I GLYCYPHAGIDAE; 13, 22, 23, 28, 38, 42, 232,

233,233,234,251,255,267,268
"olothyrina; 58
"I olothyroidea; 58
Glvevphagus; 13, 22, 23, 233, 233, 268 Pia,othvrus; 65, 66

I gnathosoma; 6,12,13,34,135
Gohieria; 233, 262
floplachevlus; 159, 212
HOPLOMEGISTIDAE; 80, 93, 129

I
grain mite; 38
Grammoliehus; 232
Grandjean's organ; 229 1 aplophara; 39, 294
aplothrambium; 168
arstia; 232, 267

~t
Gustavia; 296, 317 ouse dust mite; 242
I GUSTAVIIDAE; 296
Gvmnoglvphus; 242, 270
oyer's medium; 51-53
I ughesiel/a; 242
ullia; 242

I haemocoel; 25
H vadesia; 236, 262
II.Y ADESIIDAE; 236, 250, 262
HValamma; 136

I Haemogamasus; 95, 119

Hafenrefferia; 318
Vbalicus; 144
vdraehna; 172,223,224

~
HALACARIDAE; 14, 27, 38, 48, 50, 153, 155,
YdraChnellae;38, 42, 47, 59, 141, 169, 174
I 174, 176, 189
Halacaroidea; 141, 153, 155-156, 176
Ha/araehne; 113
YDRACHNIDAE; 172
YDRACHNOIDEA; 171-172, 182,223,224
Hvdrovolzia; 170,222

I HALARACHNIDAE; 42,43,71,89,90,113
Haller's organ; 24,136,139
HALOLAELAPIDAE; 87
b,YDROVOLZIIDAE; 170
rYDROVOLZIOIDEA; 170-171, 182, 222

I
I
328
I
Hydrozetes; 319
HYDROZETOIDEA; 296, 297, 303, 319
L I
LABIDOCARPIDAE; 239,254
Hydryphantes; 171
HYDRYPHANTIDAE; 171
HYDRYPHANTOIDEA; 171, 182,223
LABIDOPHORIDAE; 232,233,234,235,250,
255,262,263
I
Labidophorus; 234, 262
Hygroba tes; 225
HYGROBATOIDEA; 14, 172-173, 182,224,225
Hygroribates; 297
Labidostomma; 154, 194
LABIDOSTOMMIDAE; 37,154,177,194
I
Labidostommoidea; 154, 177
Hypoaspis; 120
HYPOCHTHONOIDEA; 293, 301, 302, 308
Hypodectes; 234
labrum; 13, 62, 67, 289
lacinia; 13, 123, 128
I
Lackerbaueria; 232
HYPODERIDAE; 35, 234, 235, 249, 255
hypopus; 34, 35, 231,232,233,233,234,235,
236, 255, 256, 263, 266, 267
lactophenol; 49
LAELAPI DAE; 39, 41, 71, 72, 80, 90, 95, 119, 120 I
LAELAPONYSSIDAE; 87, 103
hypostome; 12, 13, 135, 135
hysterosoma; 6, 17
hysterosomal shield; 16
Laelaponyssus; 103
Laelaps; 71, 120
I
lamella; 297, 304
HYSTRICHONYSSIDAE; 14, 90, 118
Hystrichonyssus; 14, 118, 157
Laminosioptes; 42, 247, 279
LAMINOSIOPTIDAE; 42,247,253,279 I
LARDOGLYPHIDAE; 250
I
ICHTHYOSTOMATOGASTERIDAE; 86
Lardoglyphus; 232, 260
Larinyssus; 115 I
idiosoma; 6, 12, 17 larva; 33, 34
internal mala; 13, 68
Intestine; 25, 26
lolina; 155,192
Larvacarus; 158
Lasioseius; 111
latigynial shields; 83, 95
I
IOLINIDAE; 156, 176, 192
lolinoldea; 156,176
Lawrencarinae; 151
Ledermuelleria; 159, 215
LEEUWENHOEKIIDAE; 181
I
Iphiseius; 108
Iponemus; 39, 146
Isle of Wight disease; 43
leg chaetotaxy; 18
legs; 17, 18, 24, 28 I
IXODIDAE; 136, 138, 139, 140 Leiodinychus; 76
Ixodes; 136, 139
Ixodides; 58
LEMURNYSSIDAE; 242, 251,252,268
Lemurnyssus; 242, 268 I
Ixodoidea; 136-138 Lemuroecius; 239
IXODORHYNCHIDAE; 71, 90,118
Ixodorhynchus; 118
Lemuroptes; 239
Leonardiella; 126 I
Leptotrombidium; 169
J
JOHNSTONIANIDAE; 42,169,181,219,220
Leptus; 166, 217
L1ACAROIDEA; 296, 303, 318
Liacarus; 296, 318
I
K
Kennethiella; 236
Licneremaeus; 313
Licnodamaeus; 316
Liebstadia; 299
I
Kleemannia; 107
Klinckowstroemia; 128, 129
KLiNCKOWSTROEMIiDAE; 82, 92, 95, 128, 129
Limnochares; 171
L1MNOCHARIDAE; 171
Limnozetes; 296
I
Knemidocoptes; 246, 283, 284
KNEMIDOCOPTIDAE; 246, 254, 283, 284
Kramerella; 272
Linobia; 238, 258
L1NOBIIDAE; 238, 249, 258
Linopodes; 39, 150, 190
I
L1NOTETRANIDAE; 158, 180, 211
I
I
I
I
329
I Linotetranus; 158, 211 ICROGYNIIDAE; 74, 83,96,97
Liodes; 315 icrogynium; 96, 97

I L10DOIDEA; 296, 302, 315

L1ROASPIDAE; 74
icromegistus; 79,132
icrosejus; 97
Liroaspis; 96, 98 P1icrosmaris; 166
I L1STROPHORIDAE; 239, 254, 282, 283
Listrophoroidea; 239·240, 254
Nficrotrombidium; 167
~tiCrotubercle; 183
Listrophoroides; 239 ":(crotydeus; 199

I Listrophorus; 239, 282

Loba/ges; 241, 272
fjicrozetes; 311
~ICROZETIDAE; 298
LOBALGIDAE; 241, 252, 272 ~ICROZETOIDEA; 298, 302, 311

I Lobogyniella; 96
Locustacarus; 43,146,147
~ixonomata; 293, 294
~codilied T u IIgren apparatus; 45, 46

~
Lohmannella, 189 %throngnathUS; 160

I Lohmannellinae; 155
LOHMANNIIDAE; 294
onobiacarus; 236
onoceronychus; 157
LORDALYCHIDAE; 144,178,197 onogvnaspida; 83, 84

I
I
Lorda/ychus; 197
Lordoche/es; 102
Ivrilissure; 11, 62, 65, 68 t orte/mansia; 242
r ounting techniques; 51·54
u/tioppia; 318, 319
ruridectinae; 234

~
M USCUlature;
12, 25, 26, 28
Vcetophagv; 38, 39

I Macroche/es; 6,23,71,95, 102

MACROCHELIDAE; 13,23,37,39,71,87,95, ycteronyssus; 248, 253

•
102 ydopho/eus; 236
MACRONYSSIDAE; 41, 71, 72, 89, 90,116,117 f'/yobia; 163
~YOBIIDAE; 163,177,207
Macrosclerosae; 297
Ma/ayog/yphus; 242 r-tyocoptes; 239, 281, 282
jMYOCOPTIDAE; 239, 254, 281, 282

I Malpighian tubules; 25, 26

mange (see also scabiesl; 41, 241, 244, 245, 246
Manitherionyssus; 115
yrmicotrombium; 166

I MEGISTHANIDAE; 80, 93, 130

Megisthanoidea; 79·80,93
Megisthanus; 80, 130
Nabiseius; 109
Nanhermannia; 27, 294, 305

I
Megninia; 244 Nanorchestes; 23, 27, 144, 196
Me/anozetes; 298 NANORCHESTIDAE; 23, 27, 144, 178, 196
Me/ichares; 71,111 NEMATALYCIDAE; 151, 174, 184

I
membrane, deutovarial; 31 Nemata/ycus; 184
mentum; 309 Neochey/etiella; 163
Mesa/ges; 244 Neocnemidocoptes; 246

I
mesogvnial shield; 83, 85 Neocypho/ae/aps; 71
mesonotal scutellum; 16 Neodiscopoma; 95
Mesop/ophora; 294, 306 Neofedrizzia; 81
MESOPLOPHOROIDEA; 293, 301, 306 Neomegistus; 79
I Mesostigmata; 12, 16, 19, 19,21,25,27,28,31,
33,35,39,42,50,58,67,68,69,83,84,
Neonyssus; 114, 115
Neoparholaspulus; 102
85 NEOPHYLLOBIIDAE; 160, 181, 216
I Metagynella; 126
METAGYNELLIDAE; 76, 92,126
Neophyllobius; 160, 216
Neopodocinum; 71
Neotrombieu/a; 168
I
Metarhombognathus; 155
Metastigmata; 19, 20, 21, 24, 25, 28, 31, 35, 41, Neottia/ges; 235
51,58,135-137 nervous system; 28, 29

I
•
 I
330

Nesbitt's fluid; 49 Ornithonyssus; 71,116,117

I
NOTHROIDEA; 27, 293, 294, 302, 305, 309, 310 Orthoga/umna; 298
Notoedres; 246
notogaster; 301, 304
Orthoha/arachne; 113
Otobius; 137
I
Notostigmata; 19, 19,24, 25, 58, 62, 63 Otodectes; 241
NUTTALLlELLIDAE; 136, 137, 138
Nycteridocoptes; 246
OTOPHEIDOMENIDAE;42, 43, 71, 88,109,110
Otopheidomenis; 43,109,110 I
Nycterig/yphus; 236, 259 Ou/enzia; 236
nymph; 33, 35
nymphochrysalis (see also nymphophan); 47, 47
OULENZIIDAE; 251
ovary; 26, 27, 28
I
nymphophan; 172 oviduct; 26, 27, 28

o
oviposition; 31
ovipositor; 27, 33, 288
I
ovoviviparity; 33
ocelli (see also eyes); 5, 62, 65
Odontoscirus; 153
Oxya/ges; 244
I
O/igonychus; 15,157 P
OMENTOLAELAPIDAE; 71, 87,103
Omento/ae/aps; 103
PACHYGNATHIDAE; 144, 178, 195
Pachygnathina; 177
I
Onycha/ges; 242 Pachygnathoidea; 144,177
Onychopalpida; 58
Ophioce/aeno; 80
PACHYLAELAPIDAE; 39,71,87,103
Pachy/ae/aps; 103
I
Ophiomegistus; 79 Palaeacari; 292-293, 301
Ophionyssus; 71
Ophiopneumico/a; 114
PALAEACARIDAE; 292
Palaeacariformes; 292
I
Ophioptes; 163, 205, 206 PALAEACAROIDEA; 292, 301, 305, 306
OPHIOPTIDAE; 163, 177, 205, 206
OPILIOACARIDAE; 13,63,65
Pa/aeacarus; 293, 306
palpal apotele; 14,62,64,65,66,67,68
I
Opilioacariformes; 5,58,60,61,61,62,63 palpal thumb-claw; 142, 175, 203
Opilioacaroidea; 58
Opi/ioacarus; 13, 65
Palpigradi; 5
palps (see also pedipalps); 5, 6,13,14,68
I
Opiliones; 5, 7 Panonychus; 157
opisthonotal gland; 260, 307
opisthosoma; 6, 17
Paracoroptes; 241
paragynial flap; 230
I
OPLITIDAE; 92 Para/gopsis; 242
Oppia; 296
OPPII DAE; 289
Para/orryia; 199
Para/ychus; 145
I
OPPIOIDEA; 296, 303, 318, 319 Paramealia; 242
Opsereynetes; 14
Opsonyssus; 248
PARAMEGISTIDAE; 79, 93, 94,129
Paramegistus; 79
I
Oribatei; 59 PARANTENNULIDAE; 79, 94, 132
Oribatei Inferiores; 144, 293-295, 301
Oribatei Superiores; 288, 295-300, 301
Parantennulus; 132
parasite transmission; 41, 232, 296, 298, 299
I
Oribatella; 312 Parasiten90na; 20, 25, 28, 35, 141, 179
ORIBATELLOIDEA; 298, 302, 312
Oribatu/a; 298, 299
Parasitenini; 59
PARASITIDAE; 27, 37, 39, 70, 86,100
I
ORIBATULOIDEA; 289, 297, 298, 299, 302, Parasitiformes; 9,19,58,61,63,64,67,135
313
Oribotritia; 294
parasitism; 40-43
Parasitoidea; 70, 86
I
Ornithochey/a; 163 Paratrombidium; 167
Ornithodoros; 137 PARATYDEIDAE; 151, 178, 198
I
I
I
I
331
I
•
Paratvdeus; 198 PI~RSIGIIDAE; 171
paraxial seta; 228, 229
Pi~bates; 313
Parho/aspella; 15 Pit eliaphi/us; 161
PARHOLASPIDAE; 15, 70, 87, 102 PI NIDAE; 172
Parho/aspu/us; 102 Pi nopsis; 14, 225
I Parhypoehthonius; 294, 307
PARHYPOCHTHONOIDEA; 293, 301, 307
Pi nodus; 313
P~I~NODISCIDAE;76, 91, 95,126
parthenogenesis; 31 P/fnodiscus; 95,126
I PASSALOZETOI DEA; 288,297,298,302,313
pathogen transmission; 40, 41, 72, 137, 149, 168,
p~te, stigmal; 19, 135, 139
pi te, sucker; 34
169 P ATEREMAEOIDEA; 303, 316

I pedicel; 6, 7
pedicel, anal; 39, 40, 75
PIFteremaeus; 316
P/~tv/iodes; 296, 315
Pedieu/oehe/esi 197, 198 P!('tvnothrus; 294

I PEDICULOCHELIDAE; 145, 178, 197, 198

pedipalps (see also palps); 5, 12
pl'eural shields; 304
pllieate forms; 296
pedoteetum; 304 pbeumoeoptes; 42,247,280

I Pelecanectes; 235
PELOPOIDEA; 297, 298, 302, 310, 311
P'NEUMOCOPTIDAE; 42,247,253,280
Pfeumonyssoides,- 113
Pe/ops; 298 PODAPOLIPIDAE; 42, 43,146,147,175

I Pe/oribates; 298
PENTHALEIDAE; 150, 176, 191
PEdapO/iPUS; 43,146,147
podaeephalie canal; 228, 229
Penthaleus; 150, 191 P:ODOCINIDAE; 88, 95, 104

I Pentha/odes; 190
PENTHALODIDAE; 150, 176, 190
PEdaeinum; 95,104
adam ere; 228, 258
Percanestrinia; 238 odosoma; 6

I Pergalumna; 298
Perig/isehrus; 95, 116
oecilochirus; 100
OLYASPIDAE; 30, 40, 75, 91, 122
peritreme; 19, 22, 67, 68, 135, 209, 214, 217 o/yaspinus; 123

I Perlohmannia; 308, 309

PERLOHMANNIIDAE; 38
olyaspis; 75, 91,122
o/ypterozetes; 315
PERLOHMANNOIDEA; 293, 294, 302, 305, 308, FOLYPTEROZETOIDEA; 303, 315

I 309
Petrobia; 157, 209
'omerantzia; 159,212
I OMERANTZIIDAE; 159, 180, 212
pharynx; 12, 25, 26 I , ONTARACHNIDAE; 174

I Phau/odiaspis; 127
phoresy; 39-40, 40
Ibare canal; 11, 11
ores, dorsal; 295, 297
Photia; 238 oronotieina; 288,295,296,301

I PHTHIRACAROIDEA; 23, 293, 301, 307

Phthiraearus; 23,294
orose area; 140,296,297
osterodorsal gut lobes; 25
Phylloeoptinae; 148, 149 astpedal furrow; 17

I Phyllodinyehus; 124
phylogeny; 9
Physalozercon; 79
rredation; 37-38
pregenital sclerite; 230
!preparation techniques; 49-51

I phY50gastry; 33,187
phytophagy; 38
PHYTOPTIDAE; 174
lr:roetoPhYIIOdeS; 244
PROCTOPHYLLODIDAE; 41,244,253,275,276
Proetotydeus,' 156

I Phytoptinae; 148
Phytoptipalpus; 158
Iprodinyehusi 76
prodorsal shield; 16
prodorsum (see al5a pradorsal shield); 304
Phytoptus; 149, 183

I PHYTOSEIIDAE; 37, 88,108,109

Piersigia; 171
Promata; 174
propodasoma; 6, 17,23
propad050mal shield; 16

I
I
332
I
Prostigmata; 12, 16, 19,20,21,24,25,27,28,31, Raphignathoidea; 152, 156, 159-160, 175, 180 I
33,35,42,59,141-144,174,292 Raphignathus; 160, 213
Protacarus; 144
PROTODINYCHIDAE; 77, 91, 122
receptacie, seminal; 23, 27, 28
rectal cavity; 25, 26
reproductive organs; 26,27,28
I
Protodinychoidea; 77, 91
Protodinychus; 77, 83, 122
Protomyobia; 207
Resinacarus; 187
respiration; 19
rhagidial organs; 189
I
protonymph; 34, 35
PSEUDOCHEYLIDAE; 161, 179,202
Pseudocheylus; 202
RHAGIDIIDAE; 37,150,176,189
Rhinoecius; 115
RHINONYSSIDAE; 48,71,89, gO, 114, 115
I
pseudodiagastry; 305
pseudorutellum; 258, 259
Pseudoscorpiones; 5, 8, 21
pseudostlgma; 23, 308
Rhinoseius; 71
Rhipicephalus; 136
Rhizoglyphus; 39, 232, 260
I
pseudostigmatic organ (see also trichobothrium,
sensillum); 23,24,196,297,304
Pseudotritia; 39
RHODACARIDAE; 39,71,87,105
Rhodacarus; 105
Rhombognathinae; 155
I
Psoralges; 241
PSORALGIDAE; 252
Psoralglnae; 241
RHYNCAPHYTOPTIDAE; 174
Rhyncaphytoptus; 183
Rhyncoptes; 240, 281
I
Psorergates; 163, 164, 208
PSORERGATIDAE; 163, 177,208
Psorobia; 208
RHYNCOPTIDAE; 239, 240, 254, 281
Ricardoella; 15,42,151
Ricinulei; 5
I
Psoroptes; 241
PSOROPTI DAE; 41, 229, 241, 252, 270, 271
Psoroptidia; 249
Rivoltasia; 244
Rodentopinae; 234, 255
Rodentopus; 234,235
I
Psoroptoidea; 241-243, 251
PTEROLICHIDAE; 253
Pterolichus; 278
Rodhainyssus; 248, 279
ROSENSTEINIIDAE; 236,249,259
rust mite (see also ERIOPHYIDAE); 148
I
pteromorphae; 295, 296, 304
Pterophagus; 244
Pterygosoma; 161
rutellum; 13, 62, 197,288,289,309

S
I
PTERYGOSOMIDAE; 42,161,179,202
Ptilonyssoides; 115
ptychoidy; 301, 307
sacculus foemineus; 23, 28
sacculus vestibulus; 23, 28
I
Saimiroptes; 242, 269
Punctoribates; 298
Pycnonoticina; 288, 295, 296, 297, 301
Pyemotes; 146, 187
salivary gland; 12, 26
salivary styli; 12, 13
I
SAPROGL YPHIDAE; 234, 236, 251,255,263,
PYEMOTIDAE; 5, 15,31,33,42,145, 146, 175,
187
pygidial shield (see also pygidium); 16
264
saprophagy; 39
I
Sarcoptes; 245-246, 285
pygidium; 304
Pygmephorus; 15, 187
PYROGLYPHIDAE; 241, 242, 251, 252, 270
SARCOPTIDAE; 41,228,230,245-246,254,285
Sarcoptiformes; 59
I
Sarcoptoidea; 245-247, 251
Pyroglyphinae; 242
Pyroglyphus; 242
Sasacarus; 220
scabies (see also mange); 245 I
scab mite; 41
R
Radfordia; 163, 207
scaly-leg mite; 246
ScarabaS{Jis; 119 I
Rainbowia; 166 Scheloribates; 39,298,299
ramus; 23
RAPHIGNATHIDAE; 37,160,180,213
schizeckenosy; 25, 28
Schizocoptes; 239 I
I
I
I 333
I schizogastry; 305
L";"""M '"
l
SCHIZOGYNIIDAE; 80, 81, 93,130 peleognathinae; 151

I Schizogynium; 130
Schizomida; 5
pe/eognathus; 200
pe/eorehestes; 144, 196
Schmidt layer; 11, 11 perm packet; 21,23

I
~
Sehongastia; 168 perm transfer organ; 15, 21
Sehoutedenoeoptes; 276 permatophore; 21,23

I Sciuracarus; 242, 269 PHAEROLICHIDAE; 144, 178, 195

I
Se/ero/iehus; 232 <phaero/iehus; 195
Scorpione,; 5, 8, 21 phaerotarsus; 166

I scrub typhus (see also tsutsugamushi); 40,169

SCUTACAR IDAE; 43, 175, 185
Scutacarus; 185
pider mite; see TETRANYCH IDAE
PINTURNICIDAE; 71, 89, 95,116
"pinturnix; 116

I Scutovertex; 298
scutum; 138, 140
~potted fever; 40
teatonyssus; 117
segmentation, body; 5 " teganacarus; 294
I SEJI DAE; 74, 86, 96, 98
Sejina; 83
ftegasimy; 301
fteneotarsonemus; 146, 186
Sejoidea; 74, 83 rternal shield; see ventral shields

I sejugal furrow; 17
Se/enoribates; 298
tternogynial shield; 92, 128
STIGMAEIDAE; 159, 180,214,215
~tigmal plate; 19, 135, 139
I seminal receptacle; 23, 27, 28
seminal vesicle; 22, 28
Sennertia; 234, 266
Istigmata; 19, 19, 20, 67,68,217
~tomatodex; 164
I sensillum; 193
seta, duplex; 210
seta, paraxial; 228, 229
stomatostigmatina; 177
IStrelkoviacarus; 277
styli, salivary; 12, 13, 26

I seta, supracoxal; 208, 229

setae; 18, 24
stylophore; 14, 176, 209
stylostome; 168, 168
setae, vertical; 229, 258 subcapitulum; 12, 13,26,261

I shield, epigynial; 83,84,95

shield, mesogynial; 83, 85
sucker, anal; 260
sucker, tarsal; 260

I shield, sternogynial; 92, 128 sucker plate; 34, 255, 266

shields, dorsal; 16, 140,221,267,304 Suetobe/ba; 318
shields, latigynial; 83, 95 Suidasia; 232

I shields, pleural; 304

shields, ventral; 34, 68, 84, 85, 95, 96, 139,289,
304,305
Summersiella; 159
supracoxal gland; 229
supracoxal seta; 13, 208, 229

I Sierraphytoptinae; 148
Simognathinae; 155
Syringobia; 244,278
SYRINGOBIIDAE; 253
Siteroptes; 33,145,187 SYRINGOPHI L1DAE; 41, 48,163, 177,207

I slime mite; 231

SMARIDIIDAE; 41,166,181,218
Syringophi/us; 163,207

Smaris; 166, 218 T

I So/danel/onyx; 155
solenidion; 13, 24, 228, 229
tarsal sucker; 260
TARSOCHEYLIDAE; 159,180,212

I Solifugae; 5
SPELAEORHYNCHIDAE; 71, 89, 112
TARSONEMIDAE; 5, 38, 39, 48,146,175,186
Tarsonemina; 20, 35,174

I
I
334
I
Tarsonemoidea; 145, 174 trichobothrium; 23, 24, 25 I
Tarsonemoides; 186 Trichoecius; 281
Tarsonemus; 146, 186
Tarsotomus; 161
Trichometridium; 33
Trichoribates; 298, 314 I
tarsus; 17, 18 Trigynaspida; 39, 83, 85
techniques, collecting; 45·49
techniques, dissection; 50·51
techniques, mounting; 51·54
tritonymph; 35
tritostemum; 13, 67, 68, 95 I
trochanter; 17,18
techniques, preparation; 49·51
Tectocepheus; 296
Trombicula; 168, 169, 220
TROMBICULIDAE; 23, 35, 41,168,181,220, I
tectostracum; 11, 11 222
Teinocoptes; 246, 248
TEINOCOPTIDAE; 246, 254, 284
teieiochrysaiis; 47
Trombidiformes; '59
TROMBIDIIDAE; 41,167,169,182,219,220, I
221
tenent haris; 176, 209
TENERiFFIIDAE; 161, 179,203
TENUIPALPIDAE; 14, 22, 38,156; 157,158,
Trombidioidea; 167·170, 171, 181
trombidiosis; 168
I
Trombidium; 142, 219
176,191,192
Tenuipalpus; 158, 191
TERPNACARIDAE; 145, 178, 197
Trouessartia; 275, 276
tsutsugamushi (see also scrub typhus); 169
I
Terpnacarus; 197
testes; 27, 28
tetartosternum; 92, 128
tubules, Malpighian; 25, 26
tubulus annulatus; 23, 28
Tuckerella; 15, 158,211
I
TETRANYCHIDAE; 15, 22, 37, 38,156,157,
180, 209, 210
Tetranychinae; 157
TUCKERELLIDAE; 15, 158, 180,211
Tullgren apparatus, modified; 45,46
Turbinoptes; 42, 276
I
Tetranychoidea; 148,156·159,176,180
Tetranychus; 157, 209, 210
Tetranycopsis; 209
TURBINOPTIDAE; 42,243,244,251,253,276
tutorium; 304
TYDEIDAE;37, 151, 176, 179, 198, 199
I
Tetrapodilina; 174
Tetrapolipus; 146
Tetrastigmata; 19, 19, 24, 25, 58, 64
Tydeoidea; 142, 151, 174, 178
Tydeus; 152
Typhlodromus; 109
I
thelytoky; 31
Thinozercon; 124
THINOZERCONIDAE; 76, 91, 124
Tyrophagus; 39, 232, 260

U
I
Thyas; 223
Thyreophagus; 232
tibia; 17, 18
UNIONICOLIDAE; 172
UROACTINIIDAE; 92
I
Urobovella; 76
tick fever; 1
ticks (see also Metastigmata); 1, 5, 31, 33, 35, 41,
135
Urodiaspis; 127
Urodiscella; 76
I
Urojanetia; 75
Tortonia; 232
Trachytes; 123
TRACHYTI DAE; 26, 75, 91, 123
Uropoda; 126
Uropodella; 96, 97
I
UROPODELLIDAE; 74, 86, 96, 97
Trachytoidea; 74-75, 91,121
Trachyuropoda; 75
TRACHYUROPODIDAE; 92
UROPODIDAE; 21, 27, 28, 39, 40, 75, 76,92,95,
126,127
I
Uropodina; 39, 86, 90
transmission, parasite; 41, 232, 296, 298, 299
transmission, pathogen; 40, 41, 72, 137, 149, 168,
169
Uropodoidea; 75, 90, 121
uropore; 25
I
Uropygi; 5
Treatia; 42
Trematura; 76
urstigmata; 24, 24,181,220,221,222
uterus; 26, 27, 28 I
I
I
II 335
I V W NTERSCHMIDTIIDAE; 251
W th's organs; 13, 62
vagina; 27, 28
I valves, anal; 67, 68
vas deferens; 27, 28
x
Vasates; 148 Xfnillus; 296
I Vatacarus; 169
Veigaia; 14,96,101
Xfnocastor; 233
~enorvctes; 262
VEIGAIIDAE; 14, 70, 86, 96,101 IObates; 3~3
I
I
velvet mite; see Angelothrombium or
Dinothrombium
ventral shields; 34, 65, 68, 84, 85, 95, 96,139, y ke; 31
I 289,304, 305
ventriculus; 25, 26
UNKERACARIDAE; 242,252,269
unkeracarus; 242
vertica I setae; 229, 258
I vesicle, seminal; 22, 28
Vidia; 236, 255
z
ercon; 96,110
vitellophage cell; 31

I w
ERCONIDAE; 70, 88, 96,110
etorchestes; 316
- ETORCHESTOIDEA; 296, 303, 316, 317
. etzellia; 159
I water mite; see Hydrachnellae
Winterschmidtia; 236
uluacarus; 74
umptiella; 89
vgoribatula; 298
I
I
I
I
I
I
I
I
I
I
I
I
111111111111111111111111111111111111 I111
058450
 J. Med. EDt. VoL 7, no. 6: 748
l :. BOOK REVIEW
I.../!..,'--'
,..,,'
By G. \\.. Krantz
Publish('d uy Oregr)Jl Stare L;I1iveIsily l.3ook Swre~, Tilc. Corvallis, OregoIl. 1970. 333 p.,
includin,s :H:; figwes anti 1++ piares, Spiral bound paperback. SI 1.45
This is a \\clI organized mallual prim31'ilv emphasi:-.:ing
sy,>n'malic') and providing only ba:.ic in;c.mruu ion on Inol-
og:~·. 1l100pholo~)'. collcctlon and preparaIion. Such
ballie in!orrnalion is pr('~cnl('d deally in 5 ~ pagc~ OJ 7
Ullroductory chapters. The lO-page discu~sion of col-
lection, preparation and mountillg of mite specimen~ is
particularly g-ood. The remainder or the manual is
devoted to ruiH' c1allsifimtion \0 til(> familv levc>l \\ilh Ihe
exception of tiw HydracilllE:'1lac and Crypwstigmat<l,
\\hich alc k(,).~d oniy lo ')upcrl~l.Imii('s.
T!le :nanual retl('ct') th(~ ::.late 01" IIUX of mile cbssifica-
don (odav. .\lmost:ln 2 spC'ciai::::l" a~rt>C' on dt'!ails or
hi:-r!wr d'is"ific:ation. Tht' <llllr:ul has ch(),,~'n a lo(~ic.:al
syslcm "filch ~n('orp()J',H(,S IllO:-.t or tll\~ 1(':1.tun>s ~el~(>:01!ly
aCct>Diabh> to .~caJ"OjO\!i$I'> wda\. TIc inclu<.!t,,, m!l(';- and
lIcks· l:1 the :-ltnC~a~,; .\~'3ri. cOIll'po,('d OJ'lhe order:-, Opliio-
<..\caniorme'>. P.1.l·asltl!()lmp.:i. "'ld .-\('ari,c.")trIlt·':i. Hi,; ill·
dmion 01' the Hoi()[i~~Ti(~,:,' \\ith ParasiI ifomw, rUth('l'
l!lan :,5 ,~ st>panH(~ ')I'ncr t'C'pIT"'('ri l~ :-:onH~\\'hat 01 a depar-
!lin' IrOlll i)<:~<'i ~)l·a('[ic('. btl! ;:!H~ :LlTi.lJ1'.;l'l!leI11 ' ... lov,lcai.
It i,; I"('r:'(>:::hi"!! '0 :iud the .\It>rr~..,tll<;mata. or lic:.:<,. alig:wd
bes!(i", tIle _\k..,O<,il!!:tlUla (II '!I ..' P:tmsitiiol"llH ,,>, \\ h('l'e
:!1(.'" ~", ..' ll\ to bl.'io:1'~.
SI.'\·<.'r<l.i ~b(ILll ci('nL:lo~rarn,> al(" :)n's~'~~lc-d lill:S<:r<uiut!
the pos:.!llit· rcla(!ol:~hi!)s \\ithi:1 tlh' AC3ri,
Eac!1 rn'(l(':' anti ,)ul.lOrdf'l· is Crl:1.t,tt'lC'rizl'd. toiio\\(·d h\
a blH'! <.~j:-.nl...<;ioll 01 f'ach of l:.t' 111( ludrd slllwr:amiiies.
illcl.HlJn! dia!!l1osis. mode') of iii;','. <l''bOCialion \\itJl OIiWl'
Olganisrm .:l.n~i injun.- produced. £11 sue!l iimited di.:>CllS-
"i.)IlS lJ)(.' alllhor pe!fol'ce ha::. b('en \'(>!v ~ei(>C1ive. and
occa<,ion~li1) ~()O rnllcl.l so, a:. ;:01' exampie lbe 6ijurc 10
metlW)J1 on p. :!,H· il) rH.'rp;).ra~!~i':im. or' :Jara':iitic ins('crs
b'l' C,~:'tal:1 Eplderrnoptidal". III :rc:'lc>ra!. hC)\\"('\'0l', ,he
.hoiC'(' uf examnll's i::. ~ood, anci ns l'e;)r('~('lll<1I,V(' r!':i
!lcce~~an' m all inrrodllcton- :nanll~li.
T}1(' cXI('mi\"~ keu an:' ~l("ar rmd '..!('lwl'alh- uractical
fc)r ll~C by bl'ginning studf'nrs as wcll ·as by ~e;rologi~ts
custolUariiy working with a limiled Iar.ge or laxa. The
text fi?uf/''), and particularl~ the many pl:1.tes ilIustratiH'
of kpy characr(>f"$. J('pf~',)f'nt <l major parr of thc manual.
For the mO~t pan lhe lint> drawings :1,'(' d('m' and wC'll
labeled. In a Ie" instances tht' Ch<l.l acreri"rics are
indistinctly shown or il is difliculL w determine thc srruc-
30 J)ecember 1970
rUTe labeled.
The aUI hoI' dis I ributed a li')l of I:i COlTl'Ctiono;: to hf'
made in the mallual, most or which 'were of minor impor-
tance. L·nf(JnuTlately there arc additional errors. somc
typO~T:lphical. but Others reprc:::enling errols in fact 01"
of omission. On p. IB the leg chactOtaxy sequence is
rpver~cd; thc number sub~cripls on the chaetolOtaxic
diagrams of genu and tibia should b(' rE'\"er~ed, with :ld l
at the distal Imd of thc :>egment~ Ctc. In discu.:;~ine; rc-
product ion on p. 28 thE:' author has neglected (he !mpor-
ranI work of.J. H. Yonnl:( '.196B:: d(>srribmg ill df'lal1 (he
lertiii?ation pl'Ocess in a iaelapici .. _. haemogar::w.sid\. III
r!w n'ry bri('f discussion of ti.k~ the erroneou::: slateHH'nl
is madt' on p. 136 thar RI.i/)hejJ/:,7itLJ sangwr'dlS eic'laches
!mm Ihe ori:!ti~al hO,)l onh b('lOr~ Lh(' linal moit. This
\\ould be ('o;·J't'ct :01' R. !~.':r:J/. On ill(> following page
one ~'.'('''> r:l<H Olo!,ius /l1egr.iw !(,(,(ls (llliy in rhe !<u\'at sta~t>.
~\t i('<lst rh(' 2nd nYn1!)!l n'qUIn'5 tood and must engorge
10 n'pJetiOll. In discmsmg Ihe chig-~er yenor') or' scrub
l\phll~ 0:1 p. !6~) rhe ,:l.Ulh'lr \!;ves no ':n<.:ication 01 tile'
eXlcn...,i\·(' \\0' I( impiic,~tin~ ~c\'eral :mecies or c:ti~!'~elS as
\TCTOI'~ ::1 :t<iciition 10 Ul(' 3 clas<;lcal ')pe('ies lXg'!1liot1l'c!.
III lht' .....'ction on .\~li(!Ulata on p. ~~B :, the state:ne:lt
I:Wt cht'lIc(~!a", arc cheiate-ciel,tart': in ':!cneral .his is
IfUt'. bUI :l is :nisicadinl? Il1 that som~~ Pso;'oproic!pa hu\e
::::yit'i-Jikc Chl'!'lC. ')l'veral nal~l(':'; used in lhc m::.nuaI
are oUI-dated. {'.!Z., Pr:.f:/T1l'"n's5U.I Ja/,,;ri tor Z:mljJli(;:!'l
u:;k,:rz. Pl'cilmm~rssw ((;"m/Um for P.'1.wmor!..~'s50i(if!J canillum.
dndro!aeiap\ .:.:i::~1%.i for ,J'wroiaci//!,f /a;:r':1",iIJid. Some
:'t'ler<'nCt~ll i:O figun.'s and plalC') are t'J"!ont'ou:-:. c",~ .. p. 35,
FIG. 29 ')houid read Fl(;..10: n..')8. ('ollpleL ..;. :::hOlild rC'i(~r
to Pk~t(' I';'-~ insl('ad 0;:' Ph.n' :~<3: p. 27. Plate 10- J LO
10-,i slwuid :"cad n'prOCiUni\'C' sysr('Tns rat!wr than oii-
m('!l(ar y camlls.
Pelhap':i it is surpri~ul\! jll a manual co\'('rin~ "uch a
helCro.(~('IlCOUS ~roup ,)f o! l;anlsms. tht' "uqjcc: of a {Io(l([
of rC'ccnt publications. thal tlw author ha~ illc1u(icd :lS
If>\\ errors as he ha~, .\s a whole thc publication t'<~prescnls
a \"t'ry us(,iul lOoJ, both Jor rhe !wginninl{ srllOent and a~ a
l'e!('reTlce sourc(' for practicing acaI"OIo~i'>ls.
-DC'ane P. Furman. Department of EnIOnlolo::?:y &.
P:lraSil0104"Y. Clliwrsit)' of California, nf'rkl'le~,
11111/111/1/ 1/11//1/1/1111/1/1/111/1/111
058450
 J. Med. Ent. Vol. 7, no. 6: 748
/. .... BOOK REVIEW
Iv!..;'''''''' 1
,.i.·
.\ MX'iL'AL Of' .\C.\ROLOGY
By G. \\'. Krant/.
PllbJi-;lwd by O[('gr:m SUile' CIlivCI\ily Book Sm[('s. Inc. C()rvalJi~, Oregon. 1970. 335 p ..
including :36 figures and 14+ plates. Spiral bound paperback. Sll.+.>
Tbi:: i:. a \\(,11 organized manual primarily elUpha~izing
<;ysrt'lTIati,s and prO\'idin,~ ollly basic inlOrination on biol-
og'y. ITlNpho!ogy. C'ollcnion and pn'pa,ation. Such
basic illl()rmaliorJ is prt'~cllH'd cicarl~ in .)'1 jMg(;:. or 7
introductory chapters. The IO-page dj~cuss:on of col-
If'clion, preparation and mOnnlll1r; of mite spf>cirncr:s j-;
panicuiarly good. The remainder of the manual is
dC\,(Hl::'d to milE' da:>'iification to the f~mih' lE:'vd "ith the
exception of rhe Hydr:'lchIlf>!l<lC and ("l-YPIOSti~m'H<l,
whicil are keyNi onI....- to <;up(':fi.i.mdies.
T!H.' :nanu<ll :'('i!eets tht~ stare of flux of mire d"s':>ifica-
tion 'oday. \!r::lOSl no ~ ,~p(.'c:iali'),:. a~p('\' ot~ <!('l:ds OJ'
hi;hcr c~,~:>sifl,ali()n, '1 h,-' i~lll(H):' ha~ ('!lO'if>n a logical
<,\'st(.':1'::' '.\ilich il~corpo:'~If'''' mO:"1 0;' rht' i'catllrv<; :~e!I(~raJly
accemablC' to aCrll'oJCH!!SI') wda\", He mdw:.h ~ ll1ir('~ and
tid:,,'!n the ~ll:)c!a::<, . \~"ll I, CU!l!DO~~'ti 01 dw 01 r~(':-s Opilio-
a'rlnlOrnH"S. P~~fa':>ll:I()I'm'~:.. ilnd .·\caril(lr::.:H"'. His in-
clusioll m' t!w Holothyridal' \\ii!J Parasiui()J'Jnf's ra~hf'1'
than !l!o; a "'E'p~n.lt(· Of(!(': r('p:'c,~enr'i so;::(>~\ rw.t OJ' a l:(':)ar-
1;J:e ::m:!~ pasr pl'l(,tic<'. t>m ltV' :11T<U':.;en::.('~! i, Im!:('aL
lr i~ !·'~lrc'ii1iw.: to n:ld !Iw ~1,'!a::;:I\!rr::-.ta, or ~iC:K"', :~h!.!i~('(l
bl'~ide lb~ ;"[f>.~o,r:'!!l~r!.la. of '!l\" P.:l.nl~itil()r:nc·:', \\I1('le
rhe': S('(>!~1 LO !)('kH1'!,
S(>\f'rai :hP!'ul ,i(~n(:lo'!ram') ar,' I)l('s('!l:t>d li!I:311,lliJU!
I1ll' poss!bit, rda[!(lI:.:.hirs '\\i{hi~l dl(' ·.\can.
Eac!l o:'(!('!' ami <;ubordcJ' i<; c~:,~r::.c:t('ri/ed. ~()iio\\"ed h\
a Ill'if'!" d:snl,)'5ioll of f>::.ch of :!H: incJurh-d sup('rl:-,,.uili('<;,
incluc!II1'.! dia!!nosi,~. rnod(~<; of :iit>, a::soc:ation \\ ith cn!Jcr
on~a:lisil;S anfl iniun !)rocluced. In :'llch li:1Iit<,d Ji~cas­
sions rhe amhor' perr()I"c(· ha:> he('ll \'eIT sf'lf>ctin', :1nd
occ<:!.:>ionn.i!v tOO :nuch ..,0, as for example' the fi.tilur<~ ro
:n~!lli{)r. 0:1 p. 2+.1 hype!"para~lt::-m o( p.lla,,,ir:c i:1S(',IS
by (,f'rtair! Ep!derr:::loplidac. In ';enerai. hOWCH'I'. th~
c:lOic(' uf exnrnpie~ is ~~ooci. and a:: : (~pl"(·<;("~ltall'. C' as
nCCE-!o;.-nry i!l an intl"oductor\' :nannaL
The cxt<'nsi\ (' keys are df>rJ.r ami ~cn('rail\" pr:lcri'rJ.1
fell" lISC h~ b('ginnln~ students as wdl as by ;::l.(.:arolo~~i~ts
custolUalliv \\()r~iug wirh a limit(·J rar~ge or taxa, The
text ligur<><;. and p'\I·ticulady the many pIau's illustratiH'
of kf>y cbaract('I,>. n'pn"wnl a major part (,f the m:lrlllai.
For the mo<;t pari tilt' lillL' drawings are df'm' and W('1l
laheled. Jn a few insTnnces th~' charnct('ristics are
indistinctiy shown 01 it i,~ difliculr to detcr:nille thf> struc-
30 Dec_ber 1970
ture labeled.
The author distribuled a list of 15 corn>clion-: to be
madE-' ill the rnanual~ IllOSt of which were of minor impor-
tance. t"n!()rtunately there (lre addilional errors, some
l~po~raphical. but olhers reprc~enting E:'ITOrS in fact or
or omission. On p. 13 the leg chactotaxy scq\lf>nce is
I"p\"('n:ed; the numbt>r subscripts on the chactototaxic
diagram~ of genu auel tibia should be re\"f'r~ed~ \... idl ad!
at the distal lmd or the s('~mcnr, etc, In disc\ls<;in~ re-
produclion on p. 28 the amhor has :Wtdf>Cled the irnpor-
r~Hlt \\ork ot' I, H. Youn!! I goff: dps<:riblJl(,! in df>tail the
l('nilization !;fOCC~-; in a r;wlapic! '= hacIllogrlnla:.id':. 111
til(' \ ('1'\' !n'i('!' discussion oC lick-: the ('ITOneous slatemf>nl
is made Oil p, 136 that RJ,ifnre!)I!([~llJ sangwllli/'s clct<:l.ches
frolll r1H:' oritdnai hOM onh b('IOf(' lll(! linai moiL 'fhi<;
\\ouiri ~)e CUl'n~cr 10r N, ,,',rLli, On Ihe iollowing p~g('
olle :;t>E-'S that ().'oinus 11/1?C;nini [e('ci<; onl....- ;:1 the ian'al sragf'.
,-\t It.asl the 2nd ll\"mph J'('(!uins IUoci and muSl l'ngOl:g~
,0 !c~'letion, In UiSc:.bsin!!, lhe chi1!gcr \€ctors of ::,cnlb
t'\"phu::. Oll i). 'G9 the r.ul!lor trin~s no il~tiicaliou nt' ,he
t>",(e:lsi\<~ \\01'1( implic:uiIH! sC~'eral ,,!'eci('<; or cr::.it!~?e!s as
~('(IO!S ::1 n.U<Lltion ;0 lhc .l ,bs$ica! ipecit><; !~~('ntioner!.
In d~t' :'t'ction on :hr:trlllnr<l on p. :!2B :~ thf' star('!:'W:H
l:wr c!wlice!:l~ art" citeiaH'-Ut'TtWtf>: :u z('rteral ,his is
trut>. but ir ;.~ I1llsL..'ading in that SOllle P~o'~'oni()idea h~\ ('
:-ty!<>r-:ih.c ch('];w. S('\"(·rai :laIrW<; ll~ed i:l . th~' :nanlla!
ale otll.da~eti, f>.g., P"l1lm l )f'.. :'W:,\ :)akr:ri i()!' Zum/Hif;'(t
i)tl,\'.fZ. Pr1/.I[1f!();I)',H:.LS C'il,j,;um !i:,r Pnl/lfmon..i'~Jouks ttif/l1!Um.
~,~~~~~:::;,~~)';0 ~:~;'I~~;:~ '1a~~rpi~l!,r~:~, ~;(:it;~\;.(;:::~:~:~II'(~ ~~~ . r;.o ~;~
PH;. 29 shoul(i read 11(;. '30: p, 58. coupiN + should rdi.·r
to Plate 1 !-:I- Ulswad of Plar(> 3-J: p. 27. P!atc 10-1. i:O
10-,) ')ilouid rl"ad reproducti\(> "~st('m'i l'<ltiw!' !hall ali-
rrl('ntaT~ <.:ana!<;,
P('rhap.~ it i:> surprislll'! i:l a ~~:anual cO\'erin-:r ~uch :l.
hf>tcmgclwous ~roup of 0: ganisllls. the SUbjPl't of n. flood
of recr'nt publicarion:.. that tlw author has inclltd~'d .as
t(~W PI'I'OI'S as he has. As a whole rh(' publication r<'pre<;f>nts
a \'(~ry useful tool. both J(n' the beginnin~~ studf>nt and as a
l'f'lcrcnn' SOUl.. P lor pmcricing a,arolo~-\,i;';I<;.
-Dean<, P. furman. Depal tmCIll of Entomology &
Parasiw!m;:y. l:lli\"crsity of Calili:Hllia~ Berk(>lf>y.