VanDamme Kotov Dumont 2010 Alona Checklist

Zootaxa 2330: 163 (2010)
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ZOOTAXA
ISSN 1175-5334 (online edition)
ZOOTAXA
2330
A checklist of names in Alona Baird 1843

(Crustacea: Cladocera: Chydoridae) and their current status:
an analysis of the taxonomy of a lump genus
KAY VAN DAMME1,3, ALEXEY A. KOTOV2 & HENRI J. DUMONT1
1
Department of Biology, Ghent University, K.L. Ledeganckstraat 35, B9000 Belgium. E-mail: kay.vandamme@ugent.be
2
A. N. Severtsov Institute of Ecology and Evolution, Leninsky Prospect 33, Moscow 119071, Russia.
E-mail: alexey-a-kotov@yandex.ru
3
Corresponding author
Magnolia Press
Auckland, New Zealand
Accepted by M. Alonso: 30 Nov. 2009; published: 7 Jan. 2010
Kay Van Damme, Alexey A. Kotov & Henri J. Dumont

A checklist of names in Alona Baird 1843 (Crustacea: Cladocera: Chydoridae) and their current status:
an analysis of the taxonomy of a lump genus
(Zootaxa 2330)
63 pp.; 30 cm.
7 January 2010
ISBN 978-1-86977-455-4 (paperback)
ISBN 978-1-86977-456-1 (Online edition)
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ISSN 1175-5326
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2 Zootaxa 2330 2010 Magnolia Press
VAN DAMME ET AL.
Table of contents
Abstract ............................................................................................................................................................................... 3
Introduction ......................................................................................................................................................................... 4
Materials and methods for analysis..................................................................................................................................... 6
Results ................................................................................................................................................................................ 14
Taxonomy.......................................................................................................................................................................... 14
Order Anomopoda Sars, 1865 ........................................................................................................................................... 14
Family Chydoridae Dybowski & Grochowski, 1894 emend. Frey, 1967 ......................................................................... 14
Subfamily Aloninae Dybowski & Grochowski, 1894 emend. Frey, 1967 ........................................................................ 14
Checklist of names in Alona Baird, 1843 sensu lato ......................................................................................................... 15
Original varieties and subspecies in Alona Baird, 1843 (trinomini) ................................................................................ 47
Valid genera separated from Alona sensu lato since 1843 ................................................................................................ 48
Existing genera to which Alona species or groups are allocated ...................................................................................... 49
Unavailable names ............................................................................................................................................................ 49
Discussion ......................................................................................................................................................................... 50
Discussion of the analysis: towards an increased stability ................................................................................................ 51
Acknowledgements .......................................................................................................................................................... 53
References ......................................................................................................................................................................... 53
Abstract
We discuss current progress with the revision of Alona Baird, 1843 (Crustacea: Cladocera: Chydoridae), the largest
cladoceran lump genus. We present the first inventory of these Aloninae since the 1970s and include an updated
checklist of names, with comments on the current status and position of each taxon. We discuss validity, affinities and
synonymy of ca. 240 names, including subspecies and varieties. Recent taxonomic shifts have lead to a better delineation
of natural groups in the Aloninae but the Alona puzzle remains incomplete. The majority of taxa are grouped into
species-complexes or separate genera. We count 14 Aloninae genera, now considered valid, that were split from Alona.
The status of a significant portion remains unclear, due to poor original descriptions and/or loss of type material. Even
with detailed morphological descriptions, the phylogenetic position of many Aloninae remains unsettled. Analysis of
Alona taxonomy based on the checklist shows historical trends and long periods of instability in the systematics of these
micro-crustaceans.
Key words: Cladocera, Chydoridae, Anomopoda, Aloninae, Alona, checklist, taxonomy, nomenclature, revision
A CHECKLIST OF NAMES IN ALONA
Zootaxa 2330 2010 Magnolia Press
Introduction
Around the turn of this millennium, the large cladoceran subfamily Aloninae Dybowski & Grochowski, 1894
(Cladocera: Anomopoda: Chydoridae) was under intense revision. Several new genera were created based on
morphology (Chiambeng & Dumont 1999, Dumont & Silva-Briano 2000, Kotov 2000a, Smirnov 2001, Sinev
2004a, b, Van Damme et al. 2005, Sinev & Shiel 2008, Van Damme & Dumont 2008a-b, 2009, Van Damme et
al. 2009). A number of existing genera were revised on a world scale, accompanied with detailed descriptions
(Hudec 2000, Sinev et al. 2005, Van Damme et al. 2003, Kotov 2009). The Aloninae were enriched with
plenty of new taxa, in contrast to the second chydorid subfamily, the Chydorinae, where only one genus and
few species were added since 2006, a single species group was partly revised, along with limb morphology of
a few common species (Smirnov 2007, Smirnov et al. 2006, Kotov 2008 and Van Damme & Dumont 2007).
Our knowledge of the Aloninae has thus improved significantly over two decades.
Despite the efforts, our increased knowledge of biodiversity of the worlds Aloninae was not accompanied
by an inventory of formal taxa. Alona sensu lato forms the bulk of the alonines, yet its size is considered
unnatural (Van Damme & Dumont 2008a). Previous inventories of Alona sensu lato were made, e.g., by
Brehm (1933a), Herrick & Turner (1895) and Smirnov (1971). The latter is no doubt the most impressive
work, published almost 40 years ago. Smirnovs contributions have had a major influence on chydorid
taxonomy. Since the 1970s, the family underwent serious changes, most particularly in the Aloninae. The
developments were followed up closely by few specialists, yet continue to escape the eye of most of the
Cladocera students.
For example, the genus Biapertura Smirnov, 1971, separated by Smirnov (1971) from Alona Baird, 1843,
as originally defined, is an artificial assemblage (Fryer 1987, Sinev 1999a, Dumont & Silva-Briano 2000).
Having a designated type species, it remains an available name if ever species related to Alona affinis (Leydig,
1860), would require a separate (sub)genus (Van Damme & Dumont 2008b). The majority of newly created
genera in the Aloninae include species, earlier regarded as members of either Alona or Biapertura. Attempts
at splitting Alona into a few large genera were unsuccessful and the taxon is now being dismantled into
smaller natural entities until its taxonomy stabilizes (Van Damme & Dumont, 2008a). As with any lump
genus, Alona species look alike at first glance and the genus had a poor original diagnosis, leading to immense
confusion from the start. Recently, two major morphological groups were recognized, the Coronatella- and
Hexalona-branches (Van Damme & Dumont 2008b), which facilitates further interpretation. The trajectory of
some taxa that went from one genus to another can be very complicated and requires thorough literature
research. Even different interpretations of the type species of the Aloninae, A. quadrangularis, have caused
confusion (Van Damme & Dumont 2008a).
The aim of this paper is to present an updated inventory of all taxa ever housed in the genus Alona since
its creation by Baird (1843). We provide comments for each taxon, including data on the types (original
localities and museum collections) and allocate it to its species group or genus. Our compilation is based on
gathering over a decade of information and forms part of a taxonomical revision of the genus. Based on the
list, we provide a formal analysis of history and tendencies of the genus, as a case study in Cladocera.
Abbreviations of the recorded museum collections
AM
AZUM
BAU
BGI
CMN
DAD
ECO-CH-ZOO
DGF
Australian Museum, Sydney, Australia.

Akdeniz University Zoology Museum, Antalya, Turkey.
Department of Ecology, Barcelona University, Spain.
Biological-Geographical Institute at the Irkutsk State University, Irkutsk, Russia.
Canadian Museum of Nature, Ottawa, Canada (See also NMNS).
Collection Dadayana, the Hungarian Natural History Museum, Budapest, Hungary.
Zoological collection of the Chetumal branch of ECOSUR, Chetumal, Q. Roo, Mexico.
Collection of D. G. Frey, Support Center of the Smithsonian Institution Museum of Natural
VAN DAMME ET AL.
GOS
HUZM
IBUL
IHW
IZK
IZUR
IZP
MGU
NHM
NMNS
NMO
MMA
MNSM
PUA
RBINS
SMNH
SUG
UNAM
USMN
USNM
UUZM
ZIN
ZMHU
ZMUC
ZSI
History in Suitland, Maryland, U.S.A.

Collection of G. O. Sars, Zoological Museum of the Oslo University, Norway.
Zoological Museum of the Hamburg University, Germany.
Institute of Biology, University of Ljubljana, Slovenia.
Institute of Hydrobiology of Academica Sinica, Wuhan, China.
Institute of Zoology, Kiev, Ukraine.
Istituto di Zoologia dellUniversita di Roma, Italy.
Institute of Zoology, Academia Sinica, Peking.
Zoological Museum of Moscow State University, Moscow, Russia.
Natural History Museum (Britisch Museum), London, United Kingdom.
National Museum of Natural Sciences, Ottawa, Canada (see also CMN).
Naturmuseum Olten, Olten, Switzeland (collection of T. Stingelin).
Museum voor Midden Afrika, Tervuren, Belgium.
Collection of noninsect invertebrates, Museum of Natural Sciences of Madrid, Spain.
Museum of Department of Zoology, Punjab Agricultural University, Ludhiana, India.
Royal Belgian Institute of Natural Sciences, Brussels, Belgium.
Swedish Museum of Natural History, Stockholm, Sweden.
Laboratory of Ecology (now Limnology Unit), Biology Dpt, State University of Ghent,
Ghent, Belgium.
Reference Collection of National Autonomus University of Mexico (UNAM), Mexico City,
Mexico.
U.S. National Museum, New York, USA.
The Smithsonian Institution Museum of Natural History, Washington, D.C., U.S.A.
Zoological Museum at Uppsala University, Sweden.
Zoological Institute of Russian Academy of Sciences, St. Petersburg, Russia.
Zoologisches Museum fr Naturkunde der Humboldt-Universitt, Berlin, Germany.
Zoological Museum, University of Copenhagen, Denmark (i.e. Collection of P. E. Mller).
Zoological Museum of Zoological Survey of India, Calcutta, India.
Note on authors names and dates used in the checklist

The use of Sars, 1861 or Sars, 1862 for names can be confusing, different ways appear in literature. We
follow Sars (1890, 1993) and Frey (1988) for these dates. For example, we use Alona rectangula Sars, 1861
(not Alona rectangula Sars, 1862), Alonopsis elongata (Sars, 1861), Rhynchotalona falcata (Sars, 1861),
Alona guttata Sars, 1862. See also comments by Frey (1982) and checklist by Frey in Sars (1993).
The date of Kurz's main publication (1874 or 1875) is also confusing; Frey and Sars used both 1874. The
title page of this paper contains information that this is "LXX. Band. I. Abtheilung. Jahrgang 1874. Heft I bis
V.", but on the bottom of the same page, the date reads "1875". We understand that the first portion of the 70th
volume of these Proceedings was planned to be published in 1874, but was de facto published in 1875, so we
use Kurz (1875).
Also different ways of transcribing names of Chinese authors may cause confusion. Smirnov (1971)
applied the phonetic transcription of unsimplified Chinese characters used at that time (e.g., Chiang Sieh-chih
or Chen Kou-hsiaao). Here, we use pinyin (e.g., Jiang Xie-zhi or Chen Guo-xiao), a system for transliterating
Chinese characters into the Roman alphabet adopted officially since 1979. First names are abbreviated in the
references, listed here in full: Chen G.X. (Chen Guo-xiao or Guoxiao) for Chen Kou-hsiaao; Chen S.Z. (Chen
Shou-zhung or Shouzhung) for Zhong Chu-Jung; Jiang X.Z. (Jiang Xie-zhi or Xiezhi) for Chiang Sieh-chih;
Shen J.R. (Shen Jia-rui or Jiarui) for Shen Chia-jui; Song D.X. (Song Da-xiang or Daxiang) for Sung Tahsiang; Yang J.Z. (Yang Jing-zhu or Jingzhu); Zhang J.F. (Zhang Jia-fu or Jiafu).
Materials and methods for analysis

Information about Alona-like species under consideration is summarized in Table 1. We separated all taxa into
several groups according to their status, namely: IS = Incertae Sedis; NN = Nomen Nudum; S =
Synonymized; SUP = Suppressed Name; S? = Likely a synonym; V = Valid; NO = Nomen oblitum; V(?) =
Good taxon but unresolved issues in nomenclature;
U = Status Unresolved; U(V) = Status unresolved, likely a valid (sub)species; U(S) = Status unresolved,
likely a synonym.
TABLE 1. Checklist of available names in Alona Baird, 1843, current status and their assignment to genus or to species
group. Trinomini included. For levels of description and current status, see under Material and Methods. For revision, nn:
revision not necessary. For types, u: type material present, but unassigned, (+): type material listed, but inaccessible. *to
be removed from Alona.
Names of species,
subspecies and varieties in
Alona
Orig. Status
Level
Allocated
to another
genus
Revised Types Current higher classification
References
abbreviata Sars, 1896
L2
Armatalona Sinev, 2004
Sinev (2004b)
acanthocercoides (Fischer,
1854)
L1
Leydigia Kurz, 1875
Kurz (1875); Kotov

(2009)
aculeata Werestschagin,
1911
L1
Coronatella Dybowski &

Grochowski, 1894
Smirnov (1971)
acuticostata Sars, 1903
L2
U(V)

Grochowski, 1894
Sars (1903), Sinev

(2004a), Van Damme
& Dumont (2008b)
affinis (Leydig, 1860)
L1
Alona affinis-group
Alonso (1996), Sinev

(1997; 2009)
africana Daday, 1908
L2
U(IS)
?gen. nov. 1 (Alona

verrucosa-group)
aliensis Jiang, 1980
L2
Unknown
Jiang (1980)
alluaudi Richard, 1898
L1
Richard (1898)
IS
Unknown
alonopsiformis Brehm, 1933 L1
U(IS)
gen. nov. 1 (Alona verrucosa- Brehm (1933b)

group)
alsafadii Dumont &

Brancelj, 1994
L2
Karualona Dumont & SilvaBriano, 2000
Dumont & SilvaBriano (2000)
ambigua (Lilljeborg, 1900)
L2
Tretocephala Frey, 1965
Smirnov (1966), Frey

(1965b), Alonso
(1996), Kotov (2000)
americana Flner & Frey,

1970
L2
Alona costata-group
Flner & Frey (1970)
anglica Hellich, 1874
L1
Alona guttata-group
Smirnov (1971)
angulata Birge, 1879
L1
IS
Unknown
Smirnov (1971)
angulata Weigold, 1911
L1
IN

Grochowski, 1894
Frey (1988)
angusticaudata Hudendorff,
1876
L1
Kurzia Dybowski &

Grochowski, 1894
Hudec (2000), Kotov

(2004)
anodonta Daday, 1905
L2
Unknown
Daday (1905)
archeri Sars, 1888
L2
Alona pulchella-group
Sinev (2002b)
archeroides Brehm, 1933
L2
Alona costata-group
Brehm (1933a)
arcuata Herbst, 1964
L1
IN
Alona elegans-group*
(Coronatella-branch)
Smirnov (2001)
continued next page
VAN DAMME ET AL.
TABLE 1. (continued)
Names of species,
Alona
Orig. Status
Level
Allocated
to another
genus
References
arcuata Sars, 1916
L2
U(S)
Sars (1916), Sinev

(2007)

Grochowski, 1894
arenaria Brehm, 1933
L1
Brehm (1933a)
azorica Frenzel & Alonso,

1988
L3
nn
Frenzel & Alonso

(1988)
bairdi King, 1853
L1
IS
?Alonopsis Sars, 1862
Smirnov (1996a, 1971)
balatonica Daday, 1895
L1
IS
Leydigia Kurz, 1875
Kotov (2009)
barbata Brehm, 1935
L1
Alona affinis-group
Brehm (1935)
barbulata Megard, 1967
L2
Alona guttata-group
Megard, 1967
barroisi Moniez, 1888
L1
IS
Unknown
Smirnov (1971)
bergi Ren, 1992
L2
Alona guttata-group
Ren (1992), Smirnov

(2001)
bessei Dumont, 1983
L2
Alona guttata-group
Dumont (1983)
beverleyae Smirnov, 1989
L2
Smirnov (1989); Sinev

(2002b)
bicolor Frey, 1965
L2
Alona costata-group
Frey (1965a), Sinev

(2009b)
boliviana Sinev & Coronel,

2006
L3
nn
Alona quadrangularis-group
Sinev & Coronel

(2006)
borealis Chengalath & Hann, L2

1981
Chengalath & Hann

(1981)
brandorffi Sinev &

Hollwedel, 2002
L3
nn
gen. nov. 1 (Alona verrucosa- Sinev & Hollwedel

group)
(2002)
brevicaudata Chen, 1991
L2
U(S?)
(+)
?Maraura Sinev & Shiel,

2008
Chen (1991)
broaensis MatsumuraTundisi & Smirnov, 1984
L2
Alona dentifera-group*
Matsumura-Tundisi &
Smirnov (1984)
braziliensis (Bergamin,
1935)
L1
Sinev et al. (2004)
bromelicola Smirnov, 1988
L2
Sinev (2002a)
bukobensis Weltner, 1897
L2
V(?)

Grochowski, 1894
Van Damme & Dumont

(2008b)
cambouei de Guerne &

Richard, 1893
L2
Sinev (2001a)
camptocercoides Schoedler,
1862
L1
Oxyurella Dybowski &

Grochowski, 1894
Kurz (1875)
cannellata Brehm, 1934
L1
U(V)
?Coronatella Dybowski &

Grochowski, 1894
Smirnov (1971)
capensis Ruhe, 1921
L1

Grochowski, 1894
Smirnov (1971)
caucasica Shiklejew, 1933
L1

Grochowski, 1894
Smirnov (1971)
cheni Sinev, 1999
L3
Alona costata-group
Sinev (1999a)
circumfimbriata Megard,
1967
L2

Grochowski, 1894
Van Damme & Dumont

(2008b), Sinev (2009b)
clathrata Sars, 1888
L2
U(V?) -
Alona guttata-group
Sars (1888)
continued next page
Names of species,
Alona
Orig. Status
Level
Allocated
to another
genus
References
coronata Kurz, 1875
L2

Grochowski, 1894
Smirnov (1971); Van

Damme & Dumont
(2008b)
costata Sars, 1862
L2
Alona costata-group
Sinev (1999a)
crassicauda Sars, 1916
L2
U(V?) -
Alona guttata-group
Sars (1916)
davidi Richard, 1895
L1
Leberis Smirnov, 1989
Sinev et al. (2005)
dentata PE Mller, 1867
L1
Rhynchotalona Norman, 1903 Frey (1962)
dentata Werestschagin, 1911 L1
IN
Alona affinis-group
Sinev (1997)
dentifera (Sars, 1901)
L2
Sinev et al. (2004)
dhilloni Battish, 1981
L1

Grochowski, 1894
Battish (1981)
diaphana King, 1853
L2
Sinev et al. (2005)
dilatata Stingelin, 1906
L1
Smirnov (1971)
duoodontha Henry, 1922
L2
Sinev (2004b)
elegans Kurz, 1875
L2
Alonso (1996)
elegantula Brehm, 1933
L1
Alona affinis-group
Brehm (1933b)
elliptica Sinev, 1997
L3
nn
Alona affinis-group
Sinev (1997)
elongata Sars, 1861
L2
Alonopsis Sars, 1862
Smirnov (1965)
esocirostris Schoedler, 1858
L1
Graptoleberis Sars, 1862
Smirnov (1971)
estonica Mamets, 1958
L1
Alona costata-group
Frey (1965a)
eucostata Sars, 1895
L2

Grochowski, 1894
Sars (1895); Van

Damme & Dumont
(2008b)
eximia Kiser, 1948
L2
Nicsmirnovius Chiambeng &

Dumont, 1999
Van Damme et al.

(2003), Kotov &
Sanoamuang (2004)
fabricii Ren, 1992
L2
Alona costata-group
Ren (1992), Smirnov

(2001)
falcata (Sars, 1862)
L2
Rhynchotalona Norman, 1903 Kurz (1875), Smirnov

(1971); Alonso (1996)
fasciculata Daday, 1905
L2
U(V)
?Alona pulchella group
Smirnov (1971)
fennicaudis Klocke, 1893
L1

Grochowski, 1894
Smirnov (1971)
freyi Idris & Fernando, 1981 L2
Unknown
Idris & Fernando

(1981)
gauthieri Brehm, 1934
L1

Grochowski, 1894
Smirnov (1971)
glabra Sars, 1901
L2
Sinev (2001c)
glacialis Birge, 1879
L1
IS
Unknown
Smirnov (1971)
globulosa Daday, 1898
L1
Indialona Rajapaksa &

Fernando, 1987
Rajapaksa & Fernando

(1987)
grisea (Fischer, 1854)
L1
Alonella Sars, 1862

(CHYDORINAE)
Herrick & Turner

(1895)
continued next page
VAN DAMME ET AL.
Names of species,
Alona
Orig. Status
Level
Allocated
to another
genus
References
gurneyi Smirnov, 2001
L2
Sinev (2004b)
guttata Sars, 1962
L2
Alona guttata-group
Alonso (1996)
hardingi Brehm, 1957
L1
U(V)

Grochowski, 1894
Harding (1957); Van

Damme & Dumont, in
prep.
harpularia Sars, 1916
L1
U(S)

Grochowski, 1894
Sinev (2007), Van

Damme & Dumont
(2008b)
herbsti Smirnov, 2001
L1
Smirnov (2001)
hercegovinae Brancelj, 1990 L3
gen. nov. 2 (Alona

hercegovinae-group)
Van Damme & Sinev

(in press)
holdeni Green, 1962
L2
+
(synt) Grochowski, 1894
Van Damme & Dumont

(2008)
hudeci Sinev, 1999
L2
Alona costata-group
Hudec (1998), Sinev

(1999)
hyalina (Chen & Gao, 1989) L2
(+)
Chen & Gao (1989)
iberica Alonso & Pretus,

1989
L3
iheringi Richard, 1895
L2
U(S)
Sinev et al. (2005)
iheringi Sars, 1901
L2
IN
Alona costata-group
Sinev (2001d), Kotov

& Sinev (2004)
iheringula Sinev & Kotov,

2004
L2-3
Alona costata-group
Kotov & Sinev (2004)
imitatoria (Smirnov, 1989)
L2
Sinev (2004b)
incredibilis Smirnov, 1984
L2
Nicsmirnovius Chiambeng &

Dumont
Kotov (2003)
inermis Pesta, 1928
L1
Ovalona Van Damme &

Dumont, 2008
Smirnov (1971), Sinev

(2004), Van Damme &
Dumont (2008b)
inkiti Shiklejew, 1929
L1
IS
Unknown
Smirnov (1971)
inornata Hudendorff, 1876
L1

Grochowski, 1894
Smirnov (1971)
inovata Smirnov, 2001
L2-3
Unknown
Rey & Vsquez (1986),

Smirnov (2001)
inreticulata Shen, Song &

Chen, 1964
L2
(+)
?Alona pulchella-group
Smirnov (1971)
intermedia Sars, 1862
L2
Alona intermedia-group
Alonso (1996)
investis Smirnov & Timms,

1983
L2
Unknown
Smirnov & Timms

(1983)
karelica Stenroos, 1897
L1
Flner (2000)
karua King, 1853
L2
kendallensis Henry, 1919
L1
V(?)
Alona affinis-group
Sinev (1997)
kolwezii Van Damme &

Dumont, 2008
L3
nn
Van Damme & Dumont

(2008a)
continued next page
Names of species,
Alona
Orig. Status
Level
Allocated
to another
genus
References
labrosa Vasiljeva &

Smirnov, 1969
L2
Phreatalona Van Damme et

al., 2009
Sinev & Kotov (2000),

Van Damme et al.
(2009)
lacustris Daday, 1888
L1
IS
?Alona costata-group
Smirnov (1971)
ladacensis Brehm, 1936
L1
IS
Unknown
Smirnov (1971)
laevis Herrick, 1884
L1
IS
Unknown
Smirnov (1971)
laevissima Sars, 1888
L2
Sars (1888); Sinev

(2001a)
lapidicola Chengalath &

Hann, 1981
L2
Chengalath & Hann

(1981)
lepida Birge, 1892
L1
U(V?) -
Birge (1892)
leydigii Schoedler, 1863
L1
Leydigia Kurz, 1875
Kurz (1875), Kotov

(2009)
leydigiodes Brehm, 1908
L1
IS
Unknown
Brehm (1933a)
lineata (Fischer, 1854)
L1
NN

Grochowski, 1894
Smirnov (1971), Frey

(1988)
lineolata (Sars, 1901)
L2
U(V)
Unknown
Sars (1901)
lineolata Chen & Li, 1991
L2
IN
(+)
gen. nov. 1 (Alona verrucosa- Chen et al. (1993)

group)
longirostris Daday, 1898
L1
Kurzia Dybowski &

Grochowski, 1894
Hudec (1999)
longirostris Henry, 1919
L1
IN
Alona affinis-group
Smirnov (1971)
longispina Chen & Yang,

1989
L2
U(V)
(+)
Cheng & Yang (1989)
macracantha Smirnov &

Timms, 1983
L2
Maraura Sinev & Shiel, 2008 Smirnov & Timms

(1983)
macrocopa Sars, 1894
L2
Sinev (2004b)
macronyx Daday, 1898
L1
Unknown*

(1985)
macrops Daday, 1898
L2
Chydorinae - Dadaya Sars,

1901 (CHYDORINAE)
Sars (1901), Rajapaksa

& Fernando (1982b)
macrops Motas and

Orghidan, 1948
L1
Smirnov (1971)
macrorhyncha Daday, 1901
L1
Kurzia Dybowski &

Grochowski, 1894
Smirnov (1971)
martensi Sinev, 2009
L3
nn
Alona affinis-group
Sinev (2009)
mascula King, 1853
L1
IS
?Alonopsis Sars, 1862
Smirnov (1966a)
mediterranea Yalim, 2005
L3
Gen. nov. 1 (Alona verrucosa- Yalim & Ciplak (2005)

group)
meridionalis Sinev, 2007
L3
nn
Ovalona Van Damme &

Dumont, 2008
Sinev (2007), Van

Damme & Dumont,
2008
microtata Henry, 1922
L1
?Armatalona Sinev, 2004
milleri Kiser, 1948
L2
Unknown
Kiser (1948)
minor Stingelin, 1905
L1
Unknown
Stingelin (1905)
minuta Poggenpol, 1874
L1
Alona guttata-group
Smirnov (1971)
continued next page
VAN DAMME ET AL.
Names of species,
Alona
Orig. Status
Level
Allocated
to another
genus
References
modesta Herrick, 1884
L1
IS
Unknown
Smirnov (1971)
monacantha Sars, 1901
L2

Grochowski, 1894
Sinev (2004a)
moniezi Richard, 1888
L1
U(S)

Grochowski, 1894
Richard (1888)
muelleri Richard, 1897
L1
Sinev & Hollwedel

(2005)
muelleri Ren, 1992
L2
IN
Alona costata-group
Ren (1992), Smirnov

(2001)
nalibokiana Dybowski &

Grochowski, 1895
L1
NN
Unknown
Smirnov (1971)
natalensis Sinev, 2008
L3
Alona costata-group
Sinev (2008)
neglecta Scott, 1895
L1
Alona intermedia-group
Brehm (1933)
nigra Smirnov, 1996
L2
?Alona pulchella group
Smirnov (1996)
novae-zealandiae Sars, 1904 L2

Grochowski, 1894
Van Damme & Dumont

(2008)
nuragica Margaritora, 1971
L2
Alonso (1996)
oblonga PE Mller, 1868
L1
Alona affinis-group
Smirnov (1971)
orellanai Alonso, 1996
L2
Alonso (1996)
ornata Shiklejew, 1930
L1
IN
gen. nov. 1 (Alona verrucosa- Shiklejew (1930),

group)
Sinev (2002c)
ornata Stingelin, 1895
L1
Alona affinis-group
Sinev (1997)
ossiani Sinev, 1998
L3
V(?)
nn
Alona affinis-group
Sinev (1998)
ovata Baird, 1850
L1
IS
Unknown
Baird (1850), Kurz

(1875)
ovata Rey & Vsquez, 1986
L2-3
IN
(+)
Unknown; Coronatellabranch?
Rey & Vsquez (1986),

Smirnov (2001)
parva (Daday, 1905)
L1
Parvalona Van Damme et al., Van Damme et al.

2005
(2005)
parvula Kurz, 1875
L1
Alona guttata-group
Smirnov (1971)
patagonica Ekman, 1900
L1
U(V?) -
Ekman (1900)
pectinata Elas-Gutterez &

Suarez-Morales, 1999
L3
Gen. nov. 1 (Alona verrucosa- Elas-Gutterez &

group)
Suarez-Morales (1999)
phreatica Dumont, 1983
L2

al., 2008
Van Damme et al.

(2008)
poppei Richard, 1897
L1

Grochowski, 1894
Rey & Vsquez (1986)
porrecta Birge, 1879
L1
IS
- (male specimen)
Smirnov (1971)
protzi Hartwig, 1900
L1

al., 2009
Van Damme et al.

(2009)
pseudanodonta Brehm, 1933 L1
U(IS)
? gen. nov. 1 (Alona

verrucosa-group)
Brehm (1933b),
Smirnov (1971)
pseudopulchra Smirnov,
1968
Unknown
Smirnov (1971)
L1
continued next page
11
Names of species,
Alona
Orig. Status
Level
Allocated
to another
genus
pseudoverrucosa (Smirnov,
1971)
L1
gen. nov. 1 (Alona verrucosa- Smirnov (1971)

group)
pulchella King, 1853
L1
Sinev (2001a)
pulchra Hellich, 1874
L1
SUP

Grochowski, 1894
Sinev (2001b)
punctata Daday, 1898
L1
Alonella (CHYDORINAE)
pygmaea Sars, 1862
L1
Chydorinae - Alonella
Kurz (1875)
quadrangularis (OF Mller,

1776)
L1
Van Damme & Dumont

(2008a)
rectangula Sars, 1861
L2

Grochowski, 1894
Van Damme & Dumont

(2008b)
reiseri Spandl, 1926
L1

Grochowski, 1894
Smirnov (1971)
reticulata Baird, 1843
L1
NO
Smirnov (1971)
reticulata Hartmann, 1917
L1
IN
Alona guttata-group
richardi Stingelin, 1895
L1
U(S)

Grochowski, 1894
Smirnov (1971); Sinev

(2001b)
rigidicaudis (Smirnov, 1971) L2
Unknown
Smirnov (1971),
Smirnov & Timms
(1983)
rustica Scott, 1895
L1
Alona costata-group
Alonso (1996)
rusticoides (Smirnov &

Timms, 1983)
L2
Unknown
Smirnov & Timms

(1983)
rusticoides Hudec, 1998
L2
IN
Alona costata-group
Hudec (1998); Sinev

(1999a)
salina Alonso, 1996
L2
Alonso (1996)
sanguinea PE Mller, 1867
L1
Smirnov (1971)
sarasinorum Stingelin, 1900 L1
Unknown
Frey (1991)
sarsi Richard, 1894
L1
IS
Unknown
Richard (1894)
sculpta (Sars, 1901)
L1
Notoalona Rajapaksa &

Fernando, 1987

(1987)
serrata Daday, 1898
L2
U(S)

Grochowski, 1894
Megard (1967)
Sinev (1999)
References
setigera Brehm, 1931
L1
Alona costata-group
setosocaudata Vasiljeva &

Smirnov, 1969
L2
?Alona quadrangularis-group Sinev & Kotov
setuloides Smirnov &

Timms, 1983
L2
Smirnov & Timms

(1983)
setulosa Megard, 1967
L2
Megard (1967), Sinev

(2009b)
shiklejewi Brehm, 1933
L1

al., 2009
Van Damme et al.

(2009)
siamensis Sinev &

Sanoamuang, 2008
L3
nn
Alona dentifera-group
Sinev & Sanoamuang

(2008)
continued next page
VAN DAMME ET AL.
Names of species,
Alona
Orig. Status
Level
Allocated
to another
genus
References
similis (Dybowski &

Grochowski, 1898)
L1
IS
Unknown
Smirnov (1971)
simonei Dumont, 1981
L2
Matralona Van Damme &

Dumont, 2009
Dumont (1981) ; Van

Damme & Dumont
(2009)
sketi Brancelj, 1992
L3
gen. nov. 2 (Alona

hercegovinae-group)
Van Damme & Sinev

(in press)
smirnovi Petkovski &

Flner, 1972
L2-3
Phreatalona Van Damme,

Brancelj & Dumont (2009)
Van Damme et al.

(2009)
socors (OF Mller, 1785)
L1
IS
Unknown
Smirnov (1971)
spinifera Schoedler, 1858
L1
Alona affinis-group
Smirnov (1971)
spinulosa Linko, 1900
L1
Alona guttata-group
Smirnov (1971)
stagnalis Daday, 1888
L1
IS
Unknown
Daday (1888)
stochi Brancelj, 1997
L3
gen. nov. 2 (Alona

hercegovinae-group)
Brancelj (1997), Van

Damme & Sinev
(2009a).
striolata Sars, 1916
L2
U(V)
Alona elegans-group
Sars (1916)
subantarctica Ekman, 1908
L2
U(V)
Ovalona Van Damme &

Dumont, 2008
Frey (1988), Van

Damme & Dumont
(2008b)
sulcata Schoedler, 1858
L1
Smirnov (1971)
switeziana Dybowski &

Grochowski, 1895
L1
NN
Unknown
Smirnov (1971)
taraporevalae Shirgur &

Naik, 1977
L2
Unknown
Frey (1991)
tenuicaudis Sars, 1862
L2

Grochowski, 1894
Dybowski &
Grochowski (1894);
Alonso (1996)
tenuirostris Hellich, 1874
L1
Kurzia Dybowski &

Grochowski, 1894
Smirnov (1971)
testudinaria Fischer, 1848
L1
Smirnov (1971)
tovelana Baldi, 1941
L1
Alona affinis-group
Smirnov (1971)
trachystriata Chen, Zhang & L2

Liu, 1993
(+)

Grochowski, 1894
Chen et al. (1993), Van

Damme & Dumont
(2008b)
transversa PE Mller, 1867
L1
Alonella Sars, 1862

(CHYDORINAE)
Smirnov (1971)
tridentata Stingelin, 1905
L1

Grochowski, 1894
Frenzel (1987)
truncata Smirnov, 1989
L2
U(V)
Unknown
Smirnov (1989)
tuberculata Herrick, 1884
L1
IN
Alona guttata-group
Smirnov (1971)
tuberculata Hudendorff,
1876
L1
SUP

Grochowski, 1894
Smirnov (1971)
tuberculata Kurz, 1875
L1
Alona guttata-group
Smirnov (1971)
tuberculata Mamets, 1958
L1
Alona costata-group
Frey (1965a)
continued next page
13
Names of species,
Alona
Orig. Status
Level
Allocated
to another
genus
References
unguiculata Smirnov, 1989
L2
Smirnov (1989)
vermiculata Smirnov &

Timms, 1983
L2
U(V?) +
Sinev et al. (2005)
verrucosa Lutz, 1879
L1
SUP
Unknown
Smirnov (1971)
verrucosa Sars, 1901
L2
gen. nov. 1 (Alona verrucosa- Sinev & Hollwedel

group)
(2004)
virago Brehm, 1935
L2
Alona costata-group
Brehm (1935),
Smirnov (1971)
wallaciana Henry, 1919
L1

Grochowski, 1894
Smirnov (1971)
weinecki Studer, 1878
L2
Ovalona Van Damme &

Dumont, 2008
Frey (1988), Van

Damme & Dumont
(2008b)
weltneri Keilhack, 1905
L1
Alona costata-group
Keilhack (1905),
Flner (2000)
werestschagini Sinev, 1999
L3
nn
Alona guttata-group
Sinev (1999b)
whiteleggei Sars, 1896
L2
U(V)
Alona affinis-group
Sars (1896)
willisi (Smirnov, 1981)
L1
Unknown
Smirnov (1981)
We used a solidly evaluated system to separate taxa into three levels according to level of description,
based on the original descriptions. Examples in Figure 1. L1. Original description may not allow decision on
closest affinities. Figures may be absent. If present, include habitus and/or postabdomen, but these are
insufficient in most details to determine the exact status, yet assignment to a higher taxon may be possible.
Example: Alona cannellata Brehm, 1934 (Fig. 1A) and Alona inkiti Shiklejew, 1929 (Fig. 1B). L2. Original
description limited to mainly classical characters, habitus and/or postabdomen, but with important
diagnostic characters. Taxon can be valid but affinities may be unclear due to lack of limb characters.
Examples (Figs 1C-G): Alona clathrata Sars, 1888, Alona whiteleggei Sars, 1896 (Fig. 1F). L3. Original
description up to modern standards, including limb characters and allowing hypotheses on phylogenetic
affinities. Example: Alona azorica Frenzel & Alonso, 1988 (Fig. 1H).
Using data from Table 1, we conducted a formal analysis of history and tendencies in the taxonomy of
Alona-like species under consideration, constructing accumulation curves for general number of taxa, number
of valid taxa, and number of taxa with level of description L3 + L2-3 (Fig. 2A). All taxa were grouped
according to time of publication in thirteen 20-year periods (from 1776 to 2009). For each period, we
calculated the general number of described taxa, number of valid [V + U(V)], possibly valid [V(?) + U(V?)],
and other taxa (Fig. 2B), as well as number of taxa with level of description L1, L2 and L3 + L2-3 (Fig. 2C).
Results
Taxonomy
Order Anomopoda Sars, 1865
Family Chydoridae Dybowski & Grochowski, 1894 emend. Frey, 1967
Subfamily Aloninae Dybowski & Grochowski, 1894 emend. Frey, 1967
VAN DAMME ET AL.
Note on the authorship of the Chydoridae. It is widely accepted that Stebbing (1902) is the author of the
family Chydoridae (Smirnov 1971, 1996a), hence in all publications, this family is referred to as Chydoridae
Stebbing, 1902. However, Dybowski & Grochowski (1894) already created "Grupa Chydorinae Nob.",
together with "Grupa Eurycercinae Kurz", "Grupa Acroperinae Nob.", "Grupa Aloninae Nob." and "Grupa
Pleuroxinae Nob." within the (original) family Lynceidae. Among these, the Eurycercidae is now regarded as
a separate family and the Aloninae as a subfamily of the Chydoridae (see Dumont & Silva-Briano 1998).
However, the Chydorinae, as a subfamily of Lynceidae, was also created by Dybowski & Grochowski (1894),
its status validated both logically and grammatically (ae) by the authors. Dybowski & Grochowski (1894) did
not assign the type genus in their text, but the type species of the Chydorinae Dybowski & Grochowski, 1894
is Chydorus (as "the name-bearing type of a nominal family-group taxon", Article 63 of ICZN 1999). So, the
Chydorinae (as a subfamily) was typified by Dybowski & Grochowski (1894). Like the Aloninae, which in
recent understanding (Frey 1967, Smirnov 1971, 1996a) includes two goups named by Dybowski &
Grochowski, "Grupa Aloninae" and "Grupa Acroperinae", the Chydorinae includes "Grupa Chydorinae" and
"Grupa Pleuroxinae" (Monospilus was transferred from the Chydorinae to Aloninae). The Family Chydoridae,
used in Stebbing (1902) must be regarded as a result of raising the Chydorinae Dybowski & Grochowski,
1894 to higher rank; this "family-group taxon" has priority over any others, should keep the original authors
and is based on Chydorus as the type genus. Hence, Chydoridae Stebbing, 1902 is a junior homonym of
Chydorinae Dybowski & Grochowski, 1894, raised to family level, this is Chydoridae Dybowski &
Grochowski, 1894.
Checklist of names in Alona Baird, 1843 sensu lato

abbreviata. Described as Alona abreviata Sars, 1896 from Sydney, Australia (Sars 1896), but amended to
Alona abbreviata Sars, 1896 (Shiel & Dickson 1995). Material, which is likely from type locality, in GOS
(Sinev 2004b). Shiel & Dickson (1995) revised the spelling of the name, The spelling of the species name
with a single b as abreviata in the original description (Sars 1896: 40) appears to be a typographical error, as
it is later spelt (p. 43 text; p. 79 fig. caption) as abbreviata. Listed as Biapertura abreviata by Smirnov &
Timms (1983). Junior synonym of Alona macrocopa Sars, 1894 (Sinev 2004b). Moved to Armatalona Sinev,
2004, now known as Armatalona macrocopa (Sars, 1894), see Sinev (2004b). See also A. macrocopa Sars,
1894.
acanthocercoides. Described as Lynceus acanthocercoides Fischer, 1854 on material from European
Russia and Madeira Island (Fischer 1854). Types lost (Kotov 2009). A valid species, now Leydigia
(Neoleydigia) acanthocercoides (Fischer, 1854), temporarily housed in Alona by P. E. Mller (1867). Kurz
(1875) removed it from Alona when creating Leydigia Kurz, 1875. Both genera, Leydigia and Alona, were
frequently confused due to different interpretations of Lynceus quadrangularis O. F. Mller, 1776. See
comments on Lynceus quadrangularis by Van Damme & Dumont (2008a) and revision of Leydigia in Kotov
(2009).
aculeata. Described as Alona aculeata Werestschagin, 1911 from a few localities in Georgia and the
North Caucasian part of Russia (Werestschagin 1911a). No information on types. Status unclear. According to
Smirnov (1971) and Frey (1988), this is a junior synonym of A. rectangula Sars, 1861, now Coronatella
rectangula (Sars, 1861) (Van Damme & Dumont 2008b). In Europe, the situation of the C. rectangula-group
is complicated and needs unraveling. See also A. rectangula Sars, 1861 and A. rectangula var. caucasica
Shiklejew, 1933.
acuticostata. Described as Alona acuticostata Sars, 1903 from Sumatra (Sars 1903). Material from type
locality in GOS, but no types assigned yet. Detailed morphology unknown. A. acuticostata is related to A.
monacantha Sars, 1901 (Sars 1903, Sinev 2004a). As an available name for South East Asian populations of
the Alona monacantha-group, the taxon needs revision based on new material from SE-Asia (Sinev 2004a).
Examination of specimens from Thailand suggests that it is a valid species (Van Damme, unp.). The A.
15
monacantha species group does not belong in Alona, but is much closer to, or part of, Coronatella (Van
Damme & Dumont 2008b). We prefer to use the new combination Coronatella acuticostata (Sars, 1903)
comb. nov. See also A. acuticostata var. tridentata Stingelin, 1905, A. hardingi Brehm, 1957, A. monacantha
Sars, 1901 and A. rectangula Sars, 1861.
VAN DAMME ET AL.
affinis. Described as Lynceus affinis Leydig, 1860 from Friedrichshafen and Langenargen, Bodensee
(Lake Constance), Germany (Leydig 1860). Types probably lost. Valid species. Smirnov (1971) selected A.
affinis as type species of Biapertura, an artificial assemblage that was later abandoned (Fryer 1987, Sinev
1997, 1999a, Van Damme & Dumont 2008a). The genus name is available should the A. affinis-group ever
require a separate, higher taxon (Van Damme & Dumont 2008a). The species can be regarded for now as
Alona affinis (Leydig, 1860). A. affinis and A. quadrangularis (O.F. Mller, 1776) were easily confused and
sometimes synonymized, but they are definitely different species. European populations are characterized in
Alonso (1996) and Sinev (1997, 2009a), the latter with descriptions of limb morphology. Distributed in
Eurasia, but the group is cosmopolitan: A. affinis has a number of siblings worldwide (e.g., Sinev 1997,
2009a). Smirnov listed five subspecies (affinis, ornata elegantula, barbata, dentata) (Smirnov 1971; as
Biapertura). These are regarded as varieties by Sinev (1997), but several may be good taxa. See under the five
subspecies of Smirnov (1971) listed above and also A. affinis vietnamica Dang Ngoc Than, 1980. For siblings
of A. affinis, see A. elliptica Sinev, 1997, A. ossiani Sinev, 1998, A. kendallensis Henry, 1919, A. longirostris
Henry, 1919, A. martensi Sinev, 2009 and A. whiteleggei Sars, 1896. See also synonym A. oblonga P. E.
Mller, 1868.
africana. Described as Alona rectangula var. africana Daday, 1908 from Lake Malawi (connection with
Mbasi River), Tanzania (Daday 1910). Types lost. Status unclear. Daday (1910) shows an animal with Sshaped postabdomen and short basal spine, both typical for the Alona verrucosa-group and atypical for the
Alona rectangula-group (Coronatella), which has a longer basal spine. Several specimens show tuberculate
valves (Daday 1910), also common in A. verrucosa-complex. So, Alona rectangula var. africana may be an
African species of the A. verrucosa-complex. Smirnov (1971) lists it as a synonym of Alona pseudanodonta
anodonta Daday, 1905, but A. anodonta Daday, 1905 is from Paraguay; synonymy of an African with a
Neotropical chydorid species is now considered as unlikely. See also A. alonopsiformis Brehm, 1933, A.
mediterranea Yalim, 2005, A. pseudanodonta Brehm, 1933 and A. verrucosa Sars, 1901.
aliensis. Described as Alona aliensis Jiang, 1980 From Ali (Ngari) District, North Xizang Province, Tibet
(Jiang 1980). Types in IHW. Status unclear. Only known from the original description. The peculiar, narrow
postabdomen almost resembles a male postabdomen. See also other Chinese Alona taxa, Alona brevicaudata
Chen, Zhang & Yang 1991, A. hyalina (Chen & Gao, 1989) A. inreticulata Shen, Sung & Chen, 1964 and A.
virago Brehm, 1935.
FIGURE 1. Examples of description levels and uncertainties in Alona species. All specimens depicted are adult
parthenogenetic females with postabdomen, from the original descriptions. A. Alona cannellata Brehm, 1934, Burkina
Faso. Basic description (L1), which does not allow a decision on affinities at the species level. Status uncertain, likely
situated in the Coronatella-branch. No types exist. B. Alona inkiti Shiklejew, 1929, Abkhazia. Unclear taxon without
types and with L1 description, listed as incertae sedis. No types exist. C. Alona milleri Kiser, 1948 from Pearl River,
China. Good drawings of habitus and postabdomen (description level L2) and types exist. Not found since its
description, yet the species is considered valid due to its peculiar morphology. D. Alona aliensis Jiang, 1980, Tibet. L2
description with a peculiar postabdomen (resembles that of a male) and unclear affinities. Types exist but inaccessible. E.
Alona brevicaudata Chen, Zhang & Yang, 1991, China. Good description of postabdomen and habitus (L2), suggest this
animal may belong to Maraura Sinev & Shiel, 2008 and is likely a synonym of A. macracantha Smirnov & Timms,
1983. Types exist but are inaccessible. F. Alona whiteleggei Sars, 1896, Australia. L2 description. Drawings and
descriptions by G.O. Sars were meticulously prepared and allow hypotheses on affinities. Likely A. affinis-group and
may have priority over A. kendallensis Henry, 1922. Types unknown. G. Alona freyi Idris & Fernando, 1981, Malaysia.
Valid species with peculiar morphology, well described (L2). Limbs unknown and further affinities unclear, maybe part
of Matralona Van Damme & Dumont, 2009. H. Alona azorica Frenzel & Alonso, 1988 from the Azores. Valid species,
described in detail, with limb morphology (L3 description). Externally similar to Coronatella, but limbs link it to the A.
pulchella group.
17
alluaudi. Described as Alona alluaudi Richard, 1898 from Tenerife, Canary Islands (Richard 1898).
Types probably lost, absent in DGF. No drawings exist. The description in Richard (1898) exists of a single
line of text, with main diagnostic character that this is a very large species (1 mm). Nomen nudum in Smirnov
(1971). In fact, A. alluaudi is better listed as incerta sedis (instead of a nomen nudum); however short, the
author did provide a description.
alonopsiformis. Described as Alona alonopsiformis Brehm, 1933 from Dagiangan (Mindanao Island,
Philippines) (Brehm 1933b). No information on types. Status unclear. From the drawings (Brehm 1933b), the
animal seems related to Neotropical Alona verrucosa Sars, 1901. Used for African populations (Dumont et al.
1984), but this Indonesian taxon probably does not occur in Africa. Needs revision on Indonesian material.
See also A. anodonta Daday, 1905, A. mediterranea Yalim, 2005, A. pseudanodonta Brehm, 1933 and A.
verrucosa Sars, 1901.
alsafadii. Described as Alona alsafadi Dumont & Brancelj, 1994 from Yemen (Dumont & Brancelj
1994), amended here to alsafadii because the taxon was named after Dr. M. M. Al-Safadi. Holotype in
RBINS, paratypes in RBINS, USNM, HNHM, IBUL, SUG. Lives in groundwater. Valid species, allocated to
Karualona by Dumont & Silva-Briano (2000), now known as Karualona alsafadii (Dumont & Brancelj,
1994). See also Alona karua King, 1853.
ambigua. Described as Alonopsis ambigua Lilljeborg, 1901 from Sweden (Lilljeborg 1901). Lectotype
and paralectotype in UUZM. Smirnov (1966a) moved the animal to Alona, but it is part of the genus
Tretocephala Frey, 1965, as one of two species, Tretocephala ambigua (Lilljeborg, 1901) and T. colletti (Sars,
1895) (Frey 1965b). A valid species, now called Tretocephala ambigua (Lilljeborg, 1901). Morphology of T.
ambigua, including limbs, in Smirnov (1966b), Alonso (1996) and Kotov (2000b).
americana. Described as Alona rustica americana Flssner & Frey, 1970 from Indiana, USA (Flssner &
Frey 1970) as one of two subspecies of A. rustica Scott, 1895 (the other is Alona rustica rustica Scott, 1895).
Holotype in NHM, paratypes in NHM, USNM, USMN and USMN. Distributed in the Nearctic. Valid
subspecies for North American populations of A. rustica Scott, 1895 (see under this name). See also A.
bicolor Frey, 1965 and A. rustica Scott, 1895.
anglica. Described as Alona anglica Hellich, 1874 from Bohemia, Czech Republic (Hellich 1874). No
information on types. Junior synonym of Alona guttata Sars, 1862 (Herrick & Turner 1895, Smirnov 1971).
See also A. guttata Sars, 1862 and A. parvula Kurz, 1875.
angulata. Described as Alona angulata Birge, 1879 from Cambridge (Massachusetts, USA) (Birge 1879).
No information on types. Status unknown. A single figure of the habitus in Birge (1879) does not allow
identification (to genus level), but it is definitely unrelated to Alona Baird, 1843; may even be a member of
the Chydorinae because of the body shape. Correctly listed as incertae sedis by Smirnov (1971).
angulata-2. Described as Alona rectangula angulata Weigold, 1911 from Saxony, Germany (Weigold
1911). No information on types. Synonym of A. rectangula Sars, 1861 according to Brehm (1933a) and Frey
(1988). Listed as nomen nudum in Smirnov (1971). But, any case, it is a primary homonym of A. angulata
Birge, 1879.
angusticaudata. Described as Alona angusticaudata Hudendorff, 1876 from Rjasan Area, European
Russia (Hudendorff 1876). No information on types. Junior synonym of Kurzia latissima (Kurz, 1875). See
Hudec (2000) and Kotov (2004).
anodonta. Described as Alona anodonta Daday, 1905 from Paraguay (Daday 1905). Types probably lost,
absent in DAD (Forr & Frey 1982). Seems similar to A. verrucosa complex or Coronatella, with tuberculate
valves but without a proximal denticle on the labral keel (Daday 1905). In the same publication and from the
same samples, Daday (1905) lists Alona verrucosa and A. rectangula, now respectively two genera. Alona
anodonta may belong to either of these genera; Daday (1910) later considered it a synonym of A. rectangula.
Details and status unknown. Smirnov (1971) lists A. anodonta as tuberculate subspecies of Biapertura
pseudoanodonta (Brehm, 1933), both with very different origins. Naming an earlier described taxon
(anodonta) as subspecies of a later described one (pseudoanodonta) was incorrect. These are both likely
different taxa, see under pseudoanodonta. Rajapaksa & Fernando (1982a) depict an Alona cf. anodonta from
VAN DAMME ET AL.
Sri Lanka, which clearly belongs to Coronatella. Alona anodonta may be incertae sedis in Coronatella, but
the status of both A. anodonta and A. pseudoanodonta remains unclear. See also A. pseudoanodonta Brehm,
1933, A. rectangula var. africana Daday, 1910 and A. verrucosa Sars, 1901.
archeri. Described as Alona archeri Sars, 1888 from Rockhampton, Queensland, Australia (Sars 1888),
also present in South East Asia. Lectotype and paralectotypes in GOS. Valid species of the Alona pulchellacomplex. Redescribed by Sinev (2002b), including limb morphology. See also A. beverleyae Smirnov, 1989,
A. laevissima Sars, 1888 and A. pulchella King, 1853.
archeroides. Described as Alona archeroides Brehm, 1933 from Sumatra (Toba) and Java (Telaga
Pengilon) (Brehm 1933a). No information on types. Status unknown. Ventral setules on labral keel and shape
of postabdomen suggests this taxon belongs to the Alona costata-group as Brehm (1933a) proposed himself.
Found at low pH (5.3 according to Brehm 1933a) and may be related to A. rustica Scott, 1895. Sinev (1999b)
listed it as a subspecies of A. setigera Brehm, 1931. See also A. cheni Sinev, 1999, A. costata Sars, 1862, A.
setigera Brehm, 1931 and A. rustica Scott, 1895.
arcuata. Described as Alona arcuata Sars, 1916 from Port Elizabeth, South Africa (Sars 1916). Types
lost. Sars (1916) as well as Brehm (1933a) indicated that this might be a form of A. harpularia Sars, 1916,
while A. arcuata and A. harpularia could be junior synonyms of Alona bukobensis Weltner, 1897, now
Coronatella, yet type material of A. harpularia is lost (Sinev, 2006). Now named Coronatella arcuata (Sars,
1916) and considered an African taxon of the C. rectangula complex with unclear identity (Van Damme &
Dumont 2008b). The status of African Coronatella species is confusing, with C. bukobensis the oldest
available name in the region (Van Damme & Dumont 2008b). The habitus of Coronatella arcuata (Sars,
1916) is different, with highly arched dorsum (Sars 1916). See also A. bukobensis Weltner, 1897, A. elegans
arcuata Herbst, 1964, A. harpularia Sars, 1916 and A. rectangula Sars, 1861.
arcuata-2. Described as Alona elegans arcuata Herbst, 1964 from Lower Rhine, Krefeld, Western
Germany (Herbst 1964). From the description, this form is likely true A. elegans Kurz, 1875. Invalid name,
homonym of South African A. arcuata Sars, 1916 (Smirnov 2001, Marrone et al. 2006). Smirnov (2001)
suggested A. elegans herbsti Smirnov, 2001 as a substitution. See A. elegans Kurz, 1875, A. elegans herbsti
Smirnov, 2001 and also A. arcuata Sars, 1916.
arenaria. Described as Alona arenaria Brehm, 1933 from Bali (Brehm 1933a). No information on types.
Related to A. pulchella King, 1853 according to Brehm (1933a), perhaps a synonym. Name refers to dots in
the valves, giving it a grainy (arenaria) appearance, a character common in A. pulchella King, 1853 and
related species. Without comparative material from Bali, we cannot decide on the validity of this taxon, which
may indeed be a junior synonym of A. pulchella (e.g., Chiang & Du 1979). See also A. pulchella King, 1853.
australis. Described as Leydigia australis Sars, 1885 from Queensland, Australia (Sars 1885). Possible
type material in GOS. Regarded as Alona by Daday (1898). A valid species, called Leydigia (Neoleydigia)
australis Sars, 1885. See Kotov (2009).
azorica. Described as Alona azorica Frenzel & Alonso, 1988 from the Iberian Peninsula, Baleares and
Azores (Frenzel & Alonso 1988). Holotype in HUZM, paratypes in HUZM, NHM, DGF. Valid species of the
Alona pulchella-complex but with the external appearance of a Coronatella (see Van Damme & Dumont
2008b). May also have tubercles on the carapace. Detailed drawings, including limbs, in Frenzel & Alonso
(1988) and Alonso (1996). Distribution in Frenzel & Alonso (1988). See also A. pulchella King, 1853.
bairdi. Described as Alona bairdii King, 1853 from New South Wales, Australia (King 1853). We revise
the spelling here, which now reads Alona bairdi King, 1853, because named after W. Baird. Types lost. Status
unclear. Smirnov (1966a) stated that this might be a synonym of Alonopsis elongata (Sars, 1861), a European
species (Kubersky 1977). Later, the author listed A. bairdii King, 1853 as incertae sedis (Smirnov 1971,
Smirnov & Timms 1983). Figures in King (1853) do not allow a decision on this taxon. Sars (1888) noted
similarities between A. bairdii King, 1853 and A. laevissima Sars, 1888, a member of the A. pulchella group.
balatonica. Described as Alona balatonica Daday, 1888 from Lake Balaton, Hungary (Daday 1888).
Types absent in DAD. Status unclear. Belongs to Leydigia Kurz, 1875 but was forgotten after the original
publication, even by Daday (Forr & Frey 1982, Kotov 2009). See also two other species in the same
19
publication, A. lacustris Daday, 1888 and A. stagnalis Daday, 1888.

barbata. Described as Alona affinis var. barbata Brehm, 1935 from Mt Elgon and Mt Kenya, East Africa
(Brehm 1935), later found in Sudan (Sinev 1997). No information on types. The original description was poor
and contained only a description of the antennule (Brehm 1935). Long setules on basis of this structure are a
valuable character, accompanied by others in which this animal differs from true A. affinis; it may be a
separate subspecies, Alona affinis barbata Brehm, 1935, or good species, closely related to A. martensi Sinev,
2009 (Van Damme, unpubl.); however, the name A. barbata has priority if both should be identical.
Populations from Eastern Africa (montane areas) are yet to be compared in detail with those from the
Palaearctic. See also A. affinis (Leydig, 1860) and A. martensi Sinev, 2009.
barbulata. Described as Alona barbulata Megard, 1967 from Wyoming, USA (Megard 1967). Types in
NHM and DGF. Valid species of A. guttata-group, distributed in the Nearctic (Megard 1967, Chengalath
1987). See also A. guttata Sars, 1862.
barroisi. Described as Alona barroisi Moniez, 1888 from the Island of Terceira, Azores (Moniez 1888).
Described as small species, with fine striae and a convex dorsum. No information on types and status
unknown. Incertae sedis in Smirnov (1971).
bergi. Described as Alona bergi Ren, 1992 from North West Territories in Canada (type locality) and
Greenland (Ren 1992). The author mentions types, but did not inform where they are. Types in ZMUC.
Related to Alona guttata Sars, 1862 but status unclear. According to Smirnov (2001), A. bergi is a junior
synonym of A. guttata and the description is insufficient for a separate species. See also A. barbulata Megard,
1967, A. clathrata Sars, 1916, A. guttata Sars, 1862 and A. werestschagini Sinev, 1999.
bessei. Described as Alona bessei Dumont, 1983 from Auvergne, France (Dumont 1983). Holotype and
paratype in SUG. Valid groundwater species of the Alona guttata-complex (Dumont 1983). Sinev (2008)
classifies A. bessei under the A. costata-complex due to the transverse lateral head pores. Examination of
types at SUG suggests that it is closer to A. guttata Sars, 1862. See also A. costata Sars, 1862, A. guttata Sars,
1862 and A. rustica Scott, 1895.
beverleyae. Described as Alona beverleyae Smirnov, 1989 from Queensland, Australia (Smirnov 1989).
Type in AM. Junior synonym of Alona archeri Sars, 1888 according to Sinev (2002b). See also A. archeri
Sars, 1888, A. archeroides Brehm, 1933 and A. pulchella King, 1853.
bicolor. Described as Alona bicolor Frey, 1965 from Massachusetts, USA (Frey 1965a). Nearctic.
Holotype in NHM, paratypes in NHM and DGF. Valid species related to Alona rustica Scott, 1895 of the A.
costata-group. In North America, the closest relative is A. rustica americana Flssner & Frey, 1970.
Morphology described in Frey (1965a) and Sinev (2009b). See also A. iheringula Kotov & Sinev, 2004, A.
rustica Scott, 1895, Alona rustica americana Flner & Frey, 1970.
boliviana. Described as Alona boliviana Sinev & Coronel, 2006 from the High Andes, Bolivia (Sinev &
Coronel 2006). Holotype and paratypes in MGU. One of three valid species of the A. quadrangulariscomplex. Detailed description in Sinev & Coronel (2006), notes on affinities in Van Damme & Dumont
(2008a). See also A. quadrangularis (O.F. Mller, 1776) and A. kolwezii Van Damme & Dumont, 2008.
borealis. Described as Alona borealis Chengalath & Hann, 1981 from Trappers Lake, Prince Albert
National Park, Saskatchewan, Canada (Chengalath & Hann 1981). Holotype in NMNS, paratypes in NHM,
USNM, DGF, NMNS. Valid species. Sibling species of A. lapidicola Chengalath & Hann, 1981, both
widespread in Canada (Chengalath 1987). Related to North American A. setulosa Megard, 1967 (Chengalath
& Hann 1981). If so, this species belongs to the A. pulchella-group, but fine morphology (limb characters) of
A. borealis unknown. See also A. lapidicola Chengalath & Hann, 1981, A. pulchella King, 1853 and A.
setulosa Megard, 1967.
brandorffi. Described as Alona brandorffi Sinev & Hollwedel, 2002 from Boa Vista, Brazil (Sinev &
Hollwedel 2002). Holotype in INPA, paratypes in INPA, MGU and ZMHU. Valid species related to A.
verrucosa, complete description in Sinev & Hollwedel (2002). The A. verrucosa-complex will be allocated to
a separate genus (Gen.nov. 1). See also A. verrucosa Sars, 1901.
braziliensis. Described as Alonella brazilienis Bergamin, 1935 from So Paulo, Brazil (Bergamin 1935).
VAN DAMME ET AL.
No information on types. Junior synonym of Alona dentifera (Sars, 1901), redescribed in Sinev et al. (2004).
The latter belongs to the Coronatella branch and needs removal from Alona (Van Damme & Dumont 2008b).
See also A. broaensis Matsumura-Tundisi & Smirnov, 1984, A. dentifera (Sars, 1901) and A. siamensis Sinev
& Sanoamuang, 2008.
brevicaudata. Described as Alona brevicaudata Chen, Zhang & Yang, 1991 from Jialingjiang River
(Hechuan, Chongqing), Sichuan Province, China (Chen et al. 1991). Types in IHW. Good drawings in Chen et
al. (1991) show a postabdomen that is short and wide, which does not fit the major Alona groups. In addition,
the animal has a remarkably large ocellus and a long basal spine. These characters remind most of Alona
macracantha Sars, 1894, now placed in Maraura Sinev & Shiel, 2008 (see Sinev & Shiel 2008). It may be a
junior synonym of the latter, or a separate (Chinese) species of the genus Maraura. Deeper affinities are
unclear. See also A. macracantha Smirnov & Timms, 1983.
broaensis. Described as Alona broaensis Matsumura-Tundisi & Smirnov, 1984 from Broa Reservoir,
Brazil (Matsumura-Tundisi & Smirnov 1984). Holotype in MGU. We list it here as a junior synonym of A.
dentifera (Sars, 1901) (see Sinev et al. 2004 for latter taxon). Shape of the rostrum and IDL setulation in the
description of A. broaensis differ and more information should be collected on its morphology (Sinev, pers.
comm.) A. dentifera belongs in the Coronatella-branch and needs removal from Alona (Van Damme &
Dumont 2008b). See also A. brazilienis Bergamin, 1935 and A. dentifera (Sars, 1901).
bromelicola. Described as Alona bromelicola Smirnov, 1988 from Nicaragua, Central America (Smirnov
1988). Holotype in MGU, paratypes in MGU, NHM and GOS. Valid species of the Alona pulchella-complex,
redescribed in detail by Sinev (2002a). Lives in bromeliads and is similar to A. karelica, with a small basal
spine on the postabdominal claw (Smirnov 1988). See also A. pulchella King, 1853, A. karelica Stenroos,
1897 and A. glabra Sars, 1901.
bukobensis. Described as Alona bukobensis Weltner, 1897 from Bukoba, Lake Victoria, Uganda (Weltner
1897). No information on types. Likely widespread in Africa (Van Damme & Dumont 2008b). Now regarded
as Coronatella bukobensis (Weltner, 1897), African member of the C. rectangula group. Likely a valid
species, but the animal needs redescription in comparison with true C. rectangula and its status is unclear.
Confused with Coronatella anemae Van Damme & Dumont, 2008 in Northern Africa, but C. bukobensis is
likely smaller, with typical C. rectangula appearance (Van Damme & Dumont 2008b). As in Europe, the
taxonomy for the C. rectangula-complex in Africa is confused. A. bukobensis has priority over two South
African taxa: A. harpularia Sars, 1916 and A. arcuata Sars, 1916. The latter may be junior synonyms (Sinev
2006) or separate (sub)species, but belong to Coronatella, not Alona. Notes on identity, confusion and some
characters in Van Damme & Dumont (2008b). See Sars (1916) and Sinev (2006). See also A. arcuata Sars,
1916, A. hardingi Brehm, 1957, A. harpularia Sars, 1916 and A. rectangula Sars, 1861.
cambouei. Described as Alona cambouei de Guerne & Richard, 1893 from Antananarivo, Madagascar (de
Guerne & Richard 1893). Material from type locality in Richards collection at DGF. Valid species, likely
widespread on the African continent (Ekman 1901); according to Sinev (2001a), its distribution extends to
tropical Asia. Sinev (2001a) described specimens from Iraq and Uzbekistan and notes that the eastern margin
is in China (Chen 1993). Often confused with the Australian A. pulchella King, 1853 and Neotropical A.
glabra Sars, 1901, but these are geographically separate species. A. cambouei Guerne & Richard, 1893 is of
the Alona pulchella-complex, with three connected head pores, disconnected in A. pulchella King, 1853.
Redescription of Northern African and Asian populations in Sinev (2001a), but morphology of the Malagasy
populations, which are true A. cambouei, still unstudied, the possibility of more than one subspecies
between Madagascar and Asia should be taken in consideration. Names of close forms are available in Asia,
such as Alona inreticulata Shen, Song & Chen, 1964. See also A. glabra Sars, 1901, Alona inreticulata Shen,
Song & Chen, 1964, A. nuragica Margaritora, 1971 and A. pulchella King, 1853.
camptocercoides. Described as Alona camptocercoides Schoedler, 1862 from Berlin area (now
apparently within the town), Germany (Schoedler 1862). Types in ZMHU (Forr 1983). Synonymy with
Alona tenuicaudis Sars, 1862 in Kurz (1875), later with Oxyurella tenuicaudis (Sars, 1862), which is accepted
now (Smirnov 1971). See also A. tenuicaudis Sars, 1862.
21
cannellata. Described as Alona cannellata Brehm, 1934 from Burkina Faso, Africa (Brehm 1934). No
information on types. Status uncertain, not recorded since first description. Figures are poor, but indicate a
member of the Coronatella-branch, e.g., strong striation and long basal spine on postabdomen, typical for
several Coronatella (e.g. rectangula-group). Brehm (1934) mentions that it is close to A. monacantha Sars,
1901 and has thickened setae in the posteroventral corner, almost merging into a denticle, which confirms a
close link to this species group. He compares it with A. acuticostata Sars, 1903. So, A. cannellata may be
another West African species of the A. monacantha-complex, besides the wider spread African A. hardingi
Brehm, 1957. The A. monacantha group does not belong to Alona sensu stricto and is close to or part of
Coronatella (Van Damme & Dumont 2008a), here listed under Coronatella. See also A. acuticostata Sars,
1904, A. bukobensis Weltner, 1897, A. hardingi Brehm, 1957, A. monacantha Sars, 1901 and A. rectangula
Sars, 1861.
capensis. Described as Alona capensis Rhe, 1914 from Southern Cape between Fishhoek and
Chapmansbai, South Africa (Rhe 1914). Types in ZMHU. A valid species, not recorded since the first
description. Rhe (1914) suggests a relationship with A. novae-zealandiae and A. poppei, which now both
belong to Coronatella. We examined the type material (Van Damme, unpubl.) and affinities with Coronatella
were not confirmed. Body of A. capensis is long (two times as long as high), size 0.59-0.62 mm, postabdomen
with moderate basal spine, strong marginal teeth and no large striations on the carapace. The animal shows a
resemblance with the A. pulchella group. But, it is peculiar and definitely a valid species. Drawings
reproduced in Smirnov (1971) but limb morphology unpublished. See also A. pulchella King, 1853.
caucasica. Described as Alona rectangula var. caucasica Shiklejew, 1933 from vicinities of Tbilisi and
Abkhazia (near Black Sea), Caucasus (Shiklejew 1933, Behning 1941). No information on types. Behning
(1941) considers it a subspecies of Alona rectangula Sars, 1861. Junior synonym of A. rectangula in Smirnov
(1971) and Frey (1988). Status unclear. See also A. aculeata Werestschagin, 1911, A. coronata Kurz, 1875 and
A. rectangula Sars, 1861.
ceylonica. Described as Alona australis var. ceylonica Daday, 1898 from Madatugama, Sri Lanka (Daday
1898). Types lost, not in DAD. Raised by Smirnov (1971) to level of subspecies. Not Alona, likely a Leydigia.
Incerta sedis, but may be a valid species, Leydigia ceylonica (Daday, 1898), see Kotov (2009).
cheni. Described as Alona cheni Sinev, 1999 from India (type locality) and other regions of South and
South East Asia (Sinev 1999a). Holotype and paratypes in MGU. Member of the A. costata-complex (Sinev
1999a, 2008). See also A. costata Sars, 1862, A. archeroides Brehm, 1933, A. setigera Brehm, 1933, and A.
virago Brehm, 1935.
ciliocaudata. Described as Alona cilio-caudata Sovinsky, 1891 from Chernigov Area, Ukraine (Sovinsky
1891). The dash should be removed from the species epitheton (ICZN, 1999). No information on types. It is
clearly a species of Dunhevedia and most probably a junior synonym of Dunhevedia crassa King, 1853,
member of a different subfamily, the Chydorinae (see Smirnov 1971).
circumfimbriata. Described as Alona circumfimbriata Megard, 1967 from Minnesota, USA (Megard
1967). Holotype in NHM, paratypes in NHM and DGF. Valid Nearctic species of the Coronatella rectangulacomplex, replaces the latter in North America. Now Coronatella circumfimbriata (Megard, 1967), see Van
Damme & Dumont (2008b). Redescribed by Sinev (2009b, as Alona). See also A. monacantha Sars, 1901, A.
poppei Richard, 1897 and A. rectangula Sars, 1861.
clathrata. Described as Alona clathrata Sars, 1888 from Crescent Lagoon, Rockhampton, Queensland,
Australia (Sars 1888). Material from the type locality is likely present in GOS, but needs confirmation.
Drawings in Sars (1888) of male and female leave no doubt that this animal belongs to the A. guttata-group.
The name may be applied to Australian populations of A. guttata Sars, 1862 if these should prove different
from true A. guttata. The A. guttata-group needs revision, containing several species worldwide (e.g., Sinev
1999b). See also A. guttata Sars, 1862 and A. werestschagini Sinev, 1999.
coronata. Described as Alona coronata Kurz, 1875 from Bohemia, Czech Republic (Kurz 1875). No
information on types. Subspecies of Coronatella rectangula (Sars, 1861) in Smirnov (1971; Alona), synonym
of C. rectangula in Brehm (1933a; Alona), Alonso (1996; Alona). Type species of the genus Coronatella
VAN DAMME ET AL.
Dybowski & Grochowski, 1894, reinstated by Van Damme & Dumont (2008b). The exact status of
Coronatella coronata (Kurz, 1875) is unknown, possibly more than one species of the C. rectangula complex
exist in Europe (Frey 1988, Van Damme & Dumont 2008b), which may be revealed with molecular methods.
In any case, A. coronata is definitely Coronatella, not Alona. Description of C. rectangula from Norway in
Frey (1988) and Belgium in Van Damme & Dumont (2008b). Original description of C. coronata contains a
depiction of male and female. Kurz (1875) compared it with the more abundant C. rectangula Sars, 1861 (as
Alona). His notes should not be forgotten and C. rectangula may need more attention in future, maybe very
similar subspecies or even species are still present in Europe! See also A. lineata (Fischer, 1854), A.
rectangula Sars, 1861 and A. richardi Stingelin, 1895.
costata. Described as Alona costata Sars, 1862 from Lake stenjvand, Norway (Sars 1862, 1993).
Material that may be from the type locality in GOS. Eurasia, populations outside this region need revision as
this name harbours several cryptic species (Sinev 1999a, 2008). The A. costata group contains a number of
species, of which several are in the related A. rustica group. A key to the A. costata- and A. rustica- species
complexes was recently published by Sinev (2008). Morphology of Iberian populations of A. costata
described in Alonso (1996), and of Russian populations in Sinev (1999a), including limb characters. See also
A. cheni Sinev, 1999, A. natalensis Sinev, 2008, A. rustica Scott, 1895, A. setigera Brehm, 1931, A. virago
Brehm, 1935 and A. weltneri Keilhack, 1905.
crassicauda. Described as Alona crassicauda Sars, 1916 from Bergvliet, South Africa (Sars 1916). Types
probably lost, absent in GOS (or not labeled under this name). Drawings of postabdomen in Sars (1916) show
this animal belongs to the A. guttata group. May be a separate species but detailed morphology unknown. See
also A. clathrata Sars, 1888, A. guttata Sars, 1862 and A. werestschagini Sinev, 1999.
davidi. Descibed as Alona davidi Richard, 1895 from Brouillard, Port-au-Prince, Haiti (Richard 1895).
Types probably lost, not in Richards collection at DGF (Sinev et al. 2005). This is no longer an Alona,
removed to Leberis Smirnov, 1989 with revision of populations from Mexico (Sinev et al. 2005). May be
widespread in Neotropics but true Haitian populations were never studied (Sinev et al. 2005). Richard (1897)
considers Haitian (true) Leberis davidi as different from the Brazilian animals, which are called L. davidi
var. iheringi Richard, 1895. If careful examination of populations from Haiti would reveal differences from
the common Neotropical form, the latter should be named L. davidi iheringi (Richard, 1897) and the Haitian
would be true L. davidi davidi (Richard, 1895). Such a decision would depend on study of topotypical
material. L. davidi is confused and synonymized in the pantropics with A. diaphana King, 1853, now L.
diaphanus (King, 1853), but these are two different species with distinctly different distributions; L. davidi
(Richard, 1895) does not occur in Africa or Asia (Sinev et al. 2005). African populations of Leberis,
mentioned in literature as Alona or Alonella davidi and diaphana, remain unstudied and contain more
than one unnamed species (Van Damme, unpubl.). The same is true for the Neotropics, where recently a
cryptic species Leberis chihuahuaensis Elias-Gutierrez & Valdez Moreno, 2008 was described (Mexico). See
also A. davidi iheringi Richard, 1895, A. davidi punctata Daday, 1898, A. davidi vermiculata Smirnov &
Timms, 1983 and A. diaphana King, 1853.
delaticiana. Recorded as Alona quadrangularis var. delaticiana Dybowski & Grochowski, 1895. Nomen
nudum (Smirnov 1971).
dentata. Described as Alona dentata P.E. Mller, 1867 from Denmark (Mller 1867). Type at ZMUC.
Original drawings suggest this is not an Alona but likely Rhynchotalona falcata (Sars, 1862), except for the
shorter rostrum (Frey 1962); the latter seems to be broken off (Frey 1989). Kurz (1875) notes that A. dentata
P. E. Mller, 1857 has one or two denticles in the posteroventral corner. See also A. dentata Werestschagin,
1911.
dentata-2. Proposed as Alona affinis var. dentata Werestschagin, 1911 from Novgorod Area, European
Russia (Werestschagin 1911b). No information on types. Werestschagin (1911b) made no description, but
refers to the description of A. affinis after Lilljeborg (1901) for his form. Raised to subspecies A. affinis
dentata, with a description by Ren (1968) and regarded by Sinev (1997) as a variety. Northern form of A.
affinis with one to three denticles in posteroventral valve corner (Sinev 1997). Details of this taxon are
23
unknown. Found regularly in subfossil remains of Scandinavian lakes and indicated under this name (e.g.,
Sarmaja-Korjonen et al. 2003). However, A. dentata Werestschagin, 1911 is a primary homonym of A.
dentata P. E. Mller, 1867. See also A. affinis (Leydig, 1860), A. dentata P.E. Mller, 1867 and A.
quadrangularis mph. dilatata Werestschagin, 1911.
dentifera. Described as Alonella dentifera Sars, 1901 from So Paulo, Brazil (Sars 1901). Lectotype and
paralectotypes in GOS. Valid Neotropical species. Smirnov (1996a) kept this species as a member of Alonella.
Redescription with detailed morphology in Sinev et al. (2004) shows that it is a peculiar species (e.g., no head
pores in adult females and two in juveniles). However, it is not an Alona, instead a member of the
Coronatella-branch that should be removed from and placed closer to (or part of) Coronatella (see Van
Damme & Dumont 2008b). Forms a small species complex, as a sibling species was recently described from
South East Asia, corresponding to A. cf. dentifera in Idris 1983 (Sinev & Sanoamuang 2007). See also A.
broaensis Matsumura-Tundisi & Smirnov, 1984, A. siamensis Sinev & Sanoamuang, 2007 and Alonella
brazilienis Bergamin, 1935.
diaphana. Described as Alona diaphana King, 1853 from Sydney, New South Wales, Australia (King
1853). No information on types, probably lost (Sinev et al. 2005). Known from Australia and South East Asia.
Allocation to Leberis Smirnov, 1989 by Sinev et al. (2005), and now called Leberis diaphanus (King, 1853).
Status of African Leberis unclear, these populations need revision (see under davidi). Also in the New World,
species are still found. Recently, L. chihuahuaensis Elas-Gutterez& Valdez Moreno, 2008 was described
from Mexico (Elas-Gutterez& Valdez Moreno 2008). See also A. davidi Richard, 1895, A. davidi iheringi
Richard, 1895, A. davidi punctata Daday, 1898, A. davidi vermiculata Smirnov & Timms, 1983 and A.
hyalina (Chen & Gao 1989).
dilatata. Proposed as Alona quadrangularis mph. dilatata Werestchagin, 1911 from Novgorod Area,
European Russia (Werestschagin, 1911b). Types unknown. No original description, Werestchagin (1911b)
refers to the description in Lilljeborg (1901) on which he based this morph. Variety of Alona quadrangularis
(O.F. Mller, 1776).
dhilloni. Described as Alona dhilloni Battish, 1981 from Patiala District, Punjab, India (Battish 1981).
Type material at PUA but will be transferred to ZSI. Status unclear. Animal with three main pores and strong
longitudinal striation, but description insufficient for a decision on its identity. Battish (1981) notes that it
resembles A. bukobensis Weltner, 1897. The latter is now an African taxon of Coronatella. Material of A.
dhilloni should be revised. See also A. bukobensis Weltner, 1897.
duoodonta. Described as Alonella duoodonta Henry, 1922 from New South Wales (Henry 1922).
Syntypes in AM. Placed in Biapertura by Smirnov & Timms (1983). Junior synonym of Armatalona
macrocopa Sars, 1894 according to Sinev (2004b). See also A. macrocopa (Sars, 1894).
elegans. Described as Alona elegans Kurz, 1875 from Bohemia, Czech Republic (Kurz 1875). No
information on types. Distributed in Western Europe (Van Damme & Dumont 2008b). Valid species, oldest
name in the Alona elegans-complex. According to Smirnov (1971), A. elegans is a subspecies of A.
rectangula, but the taxon deserves at least species rank. Dumont et al. (1979) suggested A. elegans and A.
rectangula (now C. rectangula) may form hybrids. Limbs of Iberian populations are depicted in Alonso
(1996), see remarks under A. elegans lebes. The A. elegans-complex is a member of the Coronatella branch,
does not belong in Alona and should be removed from the latter genus. Its position is close to, or even in,
Coronatella; a revision of the A. elegans group is, however, lacking. Comments on relationships and
distribution in Van Damme & Dumont (2008b). Three valid species of the A. elegans complex are present in
Spain (Alonso 1996) and two subspecies of A. elegans appear in literature (their status unclear). See A.
elegans arcuata Herbst, 1934, A. elegans herbsti Smirnov, 2001 and A. elegans lebes Dumont & Van De
Velde, 1975, A. orellanai Alonso, 1996, A. salina Alonso, 1996 and A. striolata Sars, 1916.
elegantula. Described as Alona affinis var. elegantula Brehm, 1933 from Indonesia (Brehm 1933b). No
information on types. Brehms (1933b) description does not allow a decision on the identity of this form.
Subspecies of A. affinis according to Smirnov (1971; as Biapertura), variety of the latter according to Sinev
(1997). We cannot decide on its rank here, but the name should remain available for A. affinis populations
VAN DAMME ET AL.
from Indonesia. A. affinis (Leydig, 1860) forms a species complex and has two valid species in Australia, A.
elliptica Sinev, 1997 and A. kendallensis Henry, 1919 and one name with unclear status, A. whiteleggei Sars,
1896. See under these names and A. affinis (Leydig, 1860).
elliptica. Described as Alona elliptica Sinev, 1997 from New South Wales, Australia (Sinev 1997).
Holotype in AM, paratypes in MGU. Valid species of the Alona affinis-complex, described in detail (Sinev
1997). See also A. affinis (Leydig, 1860), A. kendallensis Henry, 1919 and A. whiteleggei Sars, 1896.
elongata. Described as Alona elongata Sars, 1861 from vicinity of Christiania, Norway (Sars 1862).
Material of this taxon in GOS, which may be from type locality. Palaearctic. Allocated to a new genus
Alonopsis Sars, 1862 by Sars (1862), shortly after his description of Alona elongata Sars, 1861, see Smirnov
(1966a) and comments by Frey (in Sars 1993). Listed under Acroperus in Smirnov (1971), but Alonopsis is
definitely a separate genus. See Kubersky (1977).
esocirostris. Listed as Alona esocirostris Schoedler, 1858 (nomen nudum) and later described by
Schoedler (1862) from vicinity of Berlin, Germany. Types probably lost, absent in ZMHU. Junior synonym of
Graptoleberis testudinaria (Fischer, 1848) (Kurz 1875, Smirnov 1971). The word esocirostris means with
the snout of a pike, an interesting way in which Schoedler noted the typical rostrum of Graptoleberis. See
also A. reticulata Baird, 1843 and Lynceus testudinarius Fischer, 1848.
esteparica. Listed as Alona esteparica Chergui et al., 1999 from Morocco (Chergui et al. 1999). Nomen
nudum.
estonica. Described as Alona estonica Memets, 1958 from Estonia (Memets 1958). Paratype in NHM.
Frey (1965a) revised and depicted material from Memets and concluded that A. estonica is a junior synonym
of A. rustica Scott, 1895. See A. rustica Scott, 1895 and Alona estonica tuberculata Memets, 1958.
eucostata. Described as Alona eucostata Sars, 1894 from Dunedin, New Zealand (Sars 1894). Material
from type locality in GOS, labelled as cf. A. rectangula, New Zealand contains males and females and
likely corresponds to this taxon (Van Damme & Dumont 2008b). Valid species. Removed together with the
Alona rectangula-group to Coronatella (see Van Damme & Dumont 2008b), now regarded as Coronatella
eucostata (Sars, 1894). Detailed morphology unknown, drawings of male and female postabdomen in Van
Damme & Dumont (2008a). Smirnov (1971) lists A. eucostata Sars, 1894 as junior synonym of the European
A. richardi Stingelin, 1895, but considering the origin (far from terra typica), the name eucostata should not
be dismissed. It has priority over Coronatella novae-zealandiae (Sars, 1904), which may be a junior synonym.
See also A. novae-zealandiae Sars, 1904, A. rectangula Sars, 1861 and A. richardi Stingelin, 1895.
eximia. Described as Alona eximia Kiser, 1948 from Pearl River, Canton, China (Kiser 1948). Holotype
and paratypes in USNM. Present in South and South East Asia, as far south as Borneo (Van Damme et al.
2003, Kotov & Sanoamuang 2004, Van Damme et al. 2009). Valid species. Removed from Alona and
allocated to Nicsmirnovius Chiambeng & Dumont, 1999 by Van Damme et al. (2003), now named
Nicsmirnovius eximius (Kiser, 1948). Limbs very different from Alona sensu stricto (A. quadrangularisgroup) and Nicsmirnovius is adapted to life in riverine conditions (Van Damme et al. 2003), like Phreatalona
(see Van Damme et al. 2009). Records of A. eximia King, 1948 outside Asia represent different species, e.g.,
most African populations (e.g., Green 1962) earlier mentioned as A. eximia are now regarded as N. greeni
Van Damme et al., 2003, while in the Neotropics, they fall under N. fitzpatricki (Chien, 1970) and N.
incredibilis (Smirnov, 1984) (Van Damme et al. 2003, Kotov 2003). Morphology of N. eximius in Van Damme
et al. (2003) and in Kotov & Sanoamuang (2004), including males and limbs. See also A. incredibilis
Smirnov, 1984.
fabricii. Described as Alona fabricii Ren, 1992 from Greenland (Ren 1992). Types in ZMUC.
Drawings in Ren (1992) show an animal from the A. costata-complex that is closest to A. rustica Scott, 1895.
Status unclear, revision is necessary. According to Ren (1992) this is not A. rustica but an endemic from
Greenland with long lateral pores and tuberculate valves. Smirnov (2001) lists it as a synonym of A. rustica
Scott, 1895. Of the latter, there are two subspecies in the Holarctic, A. rustica rustica Scott, 1895 and A.
rustica americana Flner & Frey, 1970, see Flssner & Frey (1970). See also A. bergi Ren, 1992, A.
muelleri Ren, 1992 and A. rustica Scott, 1895.
25
falcata. Described as Alona falcata Sars, 1861 from Norway, later placed in Harporhynchus Sars, 1862 as
Harporhynchus falcatus (Sars, 1861) (see Sars 1862). Possible types in GOS. See introduction by Frey in Sars
(1993) for comments on Alona falcata versus Harporhynchus falcatus. Later, this animal was moved to
Rhynchotalona Norman, 1903 because older genus names were preoccupied (nomen novum pro
Leptorhynchus Herrick 1884), it is now known as Rhynchotalona falcata (Sars, 1861). Limbs in Alonso
(1996). See also A. dentata P.E. Mller, 1867.
fasciculata. Described as Alona fasciculata Daday, 1905 from Aregua-Yuguari River, Paraguay (Daday
1905). Types lost, absent in DAD (Forr & Frey 1982). Likely a valid species (Brehm 1933a, Smirnov 1971)
but not recorded since original description. Details remain unknown. Small animal (0.25mm) with
longitudinal stripes on carapace, elongate postabdomen with straight margins and a sharp dorsodistal corner
(not rounded), labral keel short and round (Daday 1905). Shape of the postabdomen may suggest this belongs
to Leberis Smirnov, 1989 or the Alona pulchella-group. Daday noted in a later publication (Daday 1910) that
A. fasciculata Daday, 1905 is a junior synonym of A. pulchella King, 1853. The latter does not occur in the
Neotropics and is replaced here by Alona glabra Sars, 1901. Unrelated to alonine Euryalona fasciculata
Daday, 1905. See also A. glabra Sars, 1901, A. pulchella King, 1853 and A. anodonta Daday, 1905.
fennicaudis. Described as Alona fennicaudis Klocke, 1893 from Switzerland (Klocke 1893). Most likely,
this is a misspelling. No information on types. Junior synonym of Oxyurella tenuicaudis (Sars, 1862), see
Smirnov (1971).
freyi. Described as Alona freyi Idris & Fernando, 1981 from Malaysia (Idris & Fernando 1983, Idris
1983). Holotype and paratypes in CMN. Valid species. The original description (Idris & Fernando 1981) did
not include limb morphology but drawings of habitus and postabdomen are clear. Further relationships
indefinite, the animal has two main head pores and long setae on valve margin, which remind of Matralona
Van Damme & Dumont, 2009 (see Van Damme & Dumont 2009). Note: the epitheton freyi also occurs in
Notoalona freyi Rajapaksa & Fernando, 1987 but these are two different taxa. See also A. intermedia Sars,
1862 and A. simonei Dumont, 1981.
fulcata. Recorded as Alona fulcata Schoedler, 1858 from the vicinity of Berlin, Germany (Schoedler
1858). Invalid, nomen nudum and a misspelling. It was later described as A. sulcata by Schoedler (1862) (see
latter).
gauthieri. Described as Alona gauthieri Brehm, 1934 from Burkina Faso (Brehm 1934). No information
on types. Drawings of the postabdomen in Brehm (1934) show an Oxyurella, not an Alona. Junior synonym of
Oxyurella singalensis (Daday, 1898), see Smirnov (1971). See also A. cannellata Brehm, 1934, same
publication.
glabra. Described as Alona glabra Sars, 1901 from Argentina (Sars 1901). Lectotype and paralectotypes
at GOS, selected by Sinev (2001c). Widsespread in Neotropics. Valid species. Redescription in Sinev (2001c),
valid member of the Alona pulchella-complex. Often confused in South America with A. cambouei de Guerne
& Richard, 1893 and A. pulchella King, 1853 but these two species do not occur in the New World (Sinev
2001a). See also A. cambouei de Guerne & Richard, 1893, A. fasciculata Daday, 1905 and A. pulchella King,
1853.
glacialis. Described as Alona glacialis Birge, 1879 from Cambridge, Massachusets, USA (Birge 1879).
No information on types, which could be present in DGF. No drawings with original description. Birge (1879)
notes that it is close to A. lineata (Fischer, 1854) (likely A. rectangula Sars, 1861), but with a broader
behind. According to Herrick & Turner (1895) this is A. guttata. The taxon and its varieties (A. glacialis
laevis and A. glacialis tuberculata) are correctly listed as incertae sedis in Smirnov (1971). See also A.
angulata Birge, 1879, A. glacialis var. laevis Herrick, 1884, A. glacialis var. tuberculata Herrick, 1884, A.
lineata (Fischer, 1854), A. porrecta Birge, 1879 and A. rectangula Sars, 1861.
globulosa. Described as Alona globulosa Daday, 1898 from Kalawewa Lake, Sri Lanka (Daday 1898).
Lectotype and paralectotypes in DAD. Distributed in South East Asia. First assigned to Indialona by Smirnov
(1971), later separated as type of new genus Notoalona Rajapaksa & Fernando, 1987 together with N. freyi
Rajapaksa & Fernando, 1987. The authors included a revision of Dadays types and detailed morphology
(Rajapaksa & Fernando 1987). Now known as Notoalona globulosa (Daday, 1898).
VAN DAMME ET AL.
grisea. Described as Lynceus griseus Fischer, 1854 from vicinities of St. Peterburg, European Russia
(Fischer 1854). Sporadically mentioned as Alona grisea (Fischer, 1854) but belongs to the Chydorinae genus
Alonella Sars, 1862 (e.g., Herrick & Turner 1895, Balvay 1984).
gurneyi. Described as Alona gurneyi Smirnov, 2001 from Queensland, Australia (Smirnov 2001).
Holotype in MGU. Junior synonym of Armatalona macrocopa (Sars, 1894) according to Sinev (2004b). See
also A. macrocopa Sars, 1894.
guttata. Described as Alona guttata Sars, 1862 from stenjvand, Norway (Sars 1862). Material from
type locality seems present in GOS (indicated as Egeberg or Ekeberg), but no types have been assigned. A
slide with the lectotype of A. rectangula Sars, 1861 (Frey 1988) contains several specimens of A. guttata and
this is likely from the type locality of A. guttata. Valid species. Widespread, may contain yet undescribed
sibling species worldwide. In the Holarctic, two well described siblings: A. barbulata Megard, 1967 from
USA (Megard 1967) and Alona werestschagini Sinev, 1999 from Eastern Russia (Sinev 1999b). Even in
Eurasia identification should be done carefully; A. guttata may be found sympatric with the post-glacial relict
A. werestschagini (Sarmaja-Korjonen & Sinev 2008). In Malaysia, records of A. guttata likely refer to a
different species (Idris 1983), but should be studied with care, as a name is available for the region (the
Australian A. clathrata Sars, 1888). Drawings of A. guttata in Alonso (1996) and Sinev (1999b). Smirnov
(1971) lists six subspecies of A. guttata: A. guttata guttata Sars, 1862, A. guttata parvula Kurz, 1875, A.
guttata setigera Brehm, 1931, A. guttata reticulata Hartmann, 1917, A. guttata spinulosa Linko, 1900 and A.
guttata tuberculata Kurz, 1875. Of these, the Australian A. setigera Brehm, 1931 was removed by Sinev
(1999a) as a species of the A. costata-group, the rest refers to European populations. A. tuberculata Kurz,
1875 is invalid as a species or subspecies, separated by large rounded pits on the carapace, a typical variable
character within A. guttata populations. The sculptured form could be indicated as a true variety, var.
tuberculata, yet the true A. guttata was described by Sars (1862) on tuberculate specimens. A detailed
taxonomical revision of the A. guttata-group worldwide is necessary, as it is among the most common
Aloninae. A. bergi Ren, 1992, A. reticulata Hartmann, 1917 and A. parvula Kurz, 1875 are likely synonyms
of A. guttata (e.g., Alonso 1996). See forms Alona guttata var. spinulosa Linko, 1900, A. tuberculata Kurz,
1875, Alona guttata var. typica rmek-Huek et al. and A. reticulata Hartmann, 1917. See also A. barbulata
Megard, 1967, A. bergi Ren, 1992, A. bessei Dumont, 1983, A. clathrata Sars, 1888, A. crassicaudata Sars,
1916, and Alona werestschagini Sinev, 1999.
hardingi. Described as Alona hardingi Brehm, 1957 from Ludjinge, Lake Tanganyika, East Africa
(Brehm 1957). No information on types. Likely a valid species, but needs redescription. Related to A.
monacantha Sars, 1901 and frequently listed under this name in African records (e.g., Konek 1984), but the
latter is a Neotropical sibling. The A. monacantha group belongs to Coronatella, and this African taxon can
now be called Coronatella hardingi (Brehm, 1957) comb.nov. See also A. cannellata Brehm, 1934, A.
monacantha Sars, 1901 and A. rectangula Sars, 1861.
harpularia. Described as Alona harpularia Sars, 1916 from Port Elisabeth, South Africa (Sars 1916).
Material from type locality in GOS. Now Coronatella harpularia (Sars, 1916), an African member of the C.
rectangula-complex. It remains uncertain whether this is a separate species, may be a junior synonym of
Coronatella bukobensis (Weltner, 1897) (Van Damme & Dumont 2008b). Alternatively, it is possible that
more than one sympatric species related to C. rectangula occurs in (South) Africa, similar to the complex
situation in Eurasia. See also A. arcuata Sars, 1916, A. bukobensis Weltner, 1897 and A. rectangula Sars,
1861.
herbsti. Originally described as A. elegans arcuata Herbst, 1964 from Western Germany and renamed as
Alona elegans herbsti Smirnov, 2001. A. elegans arcuata is an invalid trinomen, junior homonym of A.
arcuata Sars, 1916 (Smirnov 2001, Marrone et al. 2006). Status unclear, but probably true Alona elegans,
which was originally discovered nearby (Czech Republic; Kurz 1875). See A. elegans Kurz, 1875 and A.
elegans arcuata Herbst, 1964.
hercegovinae. Described as Alona hercegovinae Brancelj, 1990 from Herzegovina, Dinaric Region
(Brancelj 1990). Holotype at IBUL, paratypes at IBUL, ZMUC and SUG. Valid species. Forms a natural
27
group with two other stygobionts A. sketi Brancelj, 1992 and A. stochi Brancelj, 1992. The A. hercegovinaecomplex is being moved into a separate genus (Van Damme & Sinev, unpubl.). Species revised by Van
Damme & Sinev (unpubl.). See also A. sketi Brancelj, 1992 and A. stochi Brancelj, 1997.
holdeni. Described as Alona holdeni Green, 1962 from Fesafari, Niger Basin, Nigeria (Green 1962). West
African endemic. Syntypes from collection of J. Green in SUG. Valid species with modified spine on second
antenna. Redescribed and allocated to Coronatella by Van Damme & Dumont (2008b), now Coronatella
holdeni (Green, 1962). See also A. rectangula Sars, 1861.
hudeci. Renamed Alona hudeci Sinev, 1999 from Venezuela, originally described as A. rusticoides Hudec,
1998. Holotypes and paratypes in NHM, as Alona rusticoides Hudec, 1998. Valid species, similar to A.
rustica Scott, 1895. Nomen novum for Alona rusticoides Hudec, 1998. The latter became a junior homonym
after assigning taxa from Biapertura Smirnov, 1971 back to Alona, see Sinev (1999a). Morphology, including
limbs, in Hudec (1998). See also A. costata Sars, 1862, A. rustica Scott, 1895 and A. rusticoides Hudec, 1998.
hyalina. Described as Alonella hyalina Chen & Gao, 1989 from Dian Bai town, Guangdong Province,
China (Chen et al. 1989). Transferred to Alona by Zhang et al. (1997), based on head pore morphology. Types
in IHW. Based on the figures by Chen et al. (1989), we think that it belongs to neither Alona nor Alonella, but
to Leberis Smirnov, 1989. May be a junior synonym of Leberis diaphanus (King, 1853). See also A. davidi
Richard, 1895, A. diaphana King, 1853 and A. punctata Daday, 1898.
iberica. Described as Alona iberica Alonso & Pretus, 1989 from Iberian Peninsula (Alonso & Pretus
1989). Holotype in MNSM, paratypes in MNSM and BAU. Likely widespread in N-Africa (Van Damme,
unp.). Limb characters described (Alonso & Pretus 1989), removed from Alona by Dumont & Silva-Briano
(2000), now known as Karualona iberica (Alonso & Pretus, 1989). See also A. karua (King, 1853).
iheringi. Described as Alona davidi var. iheringi Richard, 1895 from Rio Grande do Sul, Brazil (Richard
1895). Types probably lost, not in DGF. Frey (1991) classified this taxon as junior synonym of A. davidi
Richard, 1895. Not an Alona; moved to Leberis since the allocation of the Alona diaphana-group (see Sinev
et al. 2005); now Leberis davidi iheringi (Richard, 1897). True Leberis davidi is originally from Haiti,
whereas var. iheringi is South American. Richard (1897) notes differences in postabdomen shape and
armature, suggesting that iheringi and davidi may be two different taxa. Detailed morphology of Mexican
populations of davidi in Sinev et al. (2005). A revision of the Haitian populations (= true davidi) is still
lacking. See also A. davidi Richard, 1895, A. diaphana King, 1853, A. iheringi Sars, 1901 and A. iheringula
Kotov & Sinev, 2004.
iheringi-2. Described as Alona iheringi Sars, 1901 from So Paulo, Brazil (Sars 1901). Lectotype and
paralectotypes in GOS. Neotropical member of A. costata-complex. Originally synonymized with A. rustica
by Frey (1965a), but later proven to be a separate species (Sinev 2001d). Alona iheringi is a junior homonym
of Alona davidi iheringi Richard, 1897 (Leberis davidi (Richard, 1897) in our current understanding).
Corrected by Kotov & Sinev (2004), now Alona iheringula Kotov & Sinev, 2004. See also A. davidi iheringi
Richard, 1897, A. iheringula Kotov & Sinev, 2004 and A. rustica Scott, 1895.
iheringula. Created as Alona iheringula Kotov & Sinev, 2004, a nomen novum for Alona iheringi Sars,
1901. See under iheringi-2for types. Valid Neotropical species of the A. costata-complex and closest to A.
rustica Scott, 1895 (see Kotov & Sinev 2004), full description with limb morphology in Sinev (2001d) as A.
iheringi Sars, 1901. See also A. bicolor Frey, 1965, A. iheringi Sars, 1901 and A. rustica Scott, 1895.
imitatoria. Described as Biapertura imitatoria (Smirnov, 1989) from Northcliffe, West Australia
(Smirnov 1989). Holotype in AM, paratypes in AM and MGU. Allocated to Armatalona by Sinev (2004b) and
redescribed, now Armatalona imitatoria (Smirnov, 1989). See also A. macrocopa Sars, 1894.
incredibilis. Described as Alona incredibilis Smirnov, 1984 from Rio Negro, Brazil (Smirnov 1984).
Holotype and paratypes in MGU. Allocated to Nicsmirnovius Chiambeng & Dumont, 1999 by Kotov (2003),
and now called Nicsmirnovius incredibilis (Smirnov, 1984). Redescribed, including limbs (Kotov 2003).
Other Nicsmirnovius species in Van Damme et al. (2003). See also A. eximia Kiser, 1853.
inermis. Described as Alona inermis Pesta, 1928 from South Georgia, Sub-Antarctic (Pesta 1928). No
information on types. Status unknown. Likely invalid. According to Sinev (2004b), it may be a species of the
VAN DAMME ET AL.
Australian genus Armatalona Sinev, 2004. More likely, A. inermis belongs to the only Alona-like genus
common on South Georgia and other islands of the Sub-Antarctic, Ovalona Van Damme & Dumont, 2008
(Frey 1988, Van Damme & Dumont 2008b). A. inermis Pesta, 1928 corresponds to this genus in postabdomen
shape. If so, it is likely a junior synonym of subantarctica, which occurs in South Georgia (see Van Damme &
Dumont 2008b). Detailed morphology of A. inermis Pesta, 1928 unknown, drawings in Pesta (1928) poor
(reproduced in Smirnov 1971). See also A. investis Smirnov & Timms, 1983, A. macrocopa Sars, 1894, A.
subantarctica Ekman, 1905 and A. weinecki Studer, 1878.
inkiti. Described as Alona inkiti Shiklejew, 1929 from Lake Inkit, Abkhazia (Shiklejew 1929). No
information on types. Obscure taxon (see Smirnov 1971), impossible to derive from original description and
drawings, maybe with an aberrant postabdomen. A search by Dr. N.N. Smirnov in the type locality revealed
no specimens (Sinev, pers. comm..). May be best listed as incertae sedis.
inornata. Described as Alona inornata Hudendorff, 1876 from Ryazan Area, European Russia
(Hudendorff 1876). No information on types. Status unclear. Figure shows a postabdomen with long basal
spine, reaching halfway strongly curved terminal claw (Hudendorff 1876). Junior synonym of Alona
rectangula Sars, 1861 in Lilljeborg (1901; as Lynceus), Brehm (1933a), Frey (1988) and Smirnov (1971). We
regard A. inornata Hudendorff, 1876 as a member of Coronatella and a synonym of Coronatella rectangula
(Sars, 1861). See also A. rectangula Sars, 1861.
inovata. Created as Alona inovata Smirnov, 2001 from Venezuela as nomen novum for Alona ovata Rey
& Vsquez, 1986, which is a primary homonym (Smirnov 2001). Valid species but position unclear, likely not
an Alona; drawings of Rey & Vsquez (1986) include some limb features. See A. ovata Rey & Vsquez, 1986.
inreticulata. Described as Alona inreticulata Shen, Song & Chen, 1964 from Chowkoutien, vicinity of
Peking, China (Shen et al. 1964). Types in IZP. Recorded later from Australia (Smirnov & Timms 1983).
Status and affinities unclear. Figures of male and female postabdomen and divided marginal denticles
(Smirnov 1971) suggest a member of the A. pulchella-complex. Needs status revised in comparison with A.
pulchella, may be a synonym or form of A. cambouei. Drawings in Chiang & Du (1979) and Smirnov &
Timms (1983). See also A. pulchella King, 1853.
insulcata. Described as Alonella sculpta var. insulcata Stingelin, 1905 from Vietnam (Stingelin 1905).
Material may be in NMO, but needs to be checked. Occurs sporadically as Alona globulosa insulcata, e.g. in
Monakov (1969). Junior synonym of Notoalona globulosa (Daday, 1898) in Rajapaksa & Fernando (1987).
intermedia. Described as Alona intermedia Sars, 1862 from Lake Maridalsvand, Norway (Sars 1862).
Sars did not assign types, but material from type locality may be present in GOS for the assignment of
lectotypes. Occurs worldwide (e.g., Flssner 2000, Idris 1983). A complex with different species, not yet
unraveled or revised (Smirnov 1971, Chengalath 1987, Van Damme & Dumont 2009). Drawings of Iberian
populations in Alonso (1996) (partim limb morphology). Listed as Biapertura intermedia in Smirnov (1971)
with two subspecies, Biapertura intermedia intermedia and Biapertura intermedia minor (see under latter
name). See also A. intermedia var. minor Stingelin, 1905 and A. simonei Dumont, 1981.
investis. Described as Alona investis Smirnov & Timms, 1983 from Victoria, Australia (Smirnov &
Timms 1983). Holotype in AM. Valid species. Peculiar, definitely not related to Alona sensu stricto, but
further position unknown as with several Australian taxa. Postabdomen short with sparse lateral armature, like
A. inermis Pesta, 1928 (Smirnov & Timms 1983). May be related to (or part of?) Armatalona Sinev, 2004. See
also A. brevicaudata Chen & Zhang, 1991, A. inermis Pesta, 1928, A. macrocopa Sars, 1894.
jabalpurensis. Described as Indialona jabalpurensis Rane, 1983 from Madhya Pradesh, India (Rane
1983). The author mentions types and catalogue numbers, but did not specify where they are deposited. Junior
synonym of A. macronyx Daday, 1898, See Rajapaksa & Fernando (1987) and Sarma & Sarma (1990).
karelica. Described as Alona karelica Stenroos, 1897 from Finnish Karelia, Finland (Stenroos 1897). No
information on types. Weigold (1911) lists karelica as a form of A. rectangula, but these are unrelated. A.
karelica Stenroos, 1897 is from the A. pulchella-group and A. rectangula belongs to Coronatella, two distinct
groups (see A. rectangula Sars, 1861). This is a valid species of the Alona pulchella-complex from Europe,
but extremely rare. Drawings in Flssner (2000), but limb characters unknown (no one ever found enough
29
specimens). Characterized by a long basal claw with short basal spine, this species seems close to
Australasian A. archeri Sars, 1888. Records from SE Asia as A. karelica may belong to the latter species (e.g.,
in Idris 1983). Reports from Mexico as A. cf. karelica, show animals similar to A. karelica Stenroos, 1897 or
to A. bromelicola Smirnov, 1988 (Elas-Gutterez et al. 2008; Van Damme et al., in press). Until further notice
we consider A. karelica a European species (Van Damme et al., in press). See also A. bromelicola Smirnov,
1988 and A. pulchella King, 1853.
karua. Described as Alona karua King, 1853 from River Karua, Stroud, New South Wales, Australia
(King 1853). No information on types. Widespread in South East Asia. Temporarily placed in Paralonella
Birge, 1910 by Birge (1910) and in Biapertura by Smirnov (1971) and Smirnov & Timms (1983). Removed
from Alona to Karualona by Dumont & Silva-Briano (2000), now it is Karualona karua (King, 1853).
Redescribed by Dumont & Silva-Briano (2000) based on specimens from Western Australia, as opposed to
the type locality, in NSW. More than one species may be present in Australia under this name (Shiel &
Dickson 1995). See also A. alsafadii Dumont & Brancelj, 1994, A. iberica Alonso & Pretus, 1989 and A.
muelleri Richard, 1897.
kendallensis. Described as Alona kendallensis Henry, 1919 from New South Wales, Australia (Henry
1919). Widespread in Australia (Shiel & Dickson 1995). Holotype in AM. Moved to Biapertura by Smirnov
(1971) and listed as such in later publications (Smirnov & Timms 1983, Shiel & Dickson 1995). Valid species
of the A. affinis-complex, revised by Sinev (1997) including limbs. May be a synonym of A. whiteleggei Sars,
1896, but status of the latter remains unclear. See also A. affinis (Leydig, 1960), A. affinis var. elegantula
Brehm, 1933, A. elliptica Sinev, 1997, A. longirostris Henry, 1919 and A. whiteleggei Sars, 1896.
kolwezii. Described as Alona kolwezii Van Damme & Dumont, 2008 from DR Congo (Van Damme &
Dumont 2008a). Holotype and paratypes in RBINS. Valid species of A. quadrangularis-complex, described in
detail, including limbs (Van Damme & Dumont 2008a). See also A. boliviana Sinev & Coronel, 2006 and A.
quadrangularis (O.F. Mller, 1776).
kwangsiensis. Described as Alona kwangsiensis Jiang, 1963 from Nanning (type locality) and Kweihsien,
Kwangsi, China (Jiang 1963). Holotype and paratypes in IHW. Form with four to five denticles in the
posteroventral corner. Smirnov (1971) placed this taxon in the genus Biapertura and notes it as a junior
synonym of B. karua King, 1853, now Karualona karua (King, 1853), see Chiambeng & Dumont (2000).
Latter authors confirmed this synonymy, after the species was temporarily reinstated by Ventakaram (1999).
See also A. karua King, 1853.
labrosa. Described as Alona labrosa Vasiljeva & Smirnov, 1969 from Lake Baikal (Vasiljeva & Smirnov
1969). Holotype and allotype in MGU, paratypes in ZIN and BGI. Redescription in Sinev & Kotov (2000),
including limb morphology. Valid species, moved to Phreatalona by Van Damme et al. (2009), now
Phreatalona labrosa (Vasiljeva & Smirnov, 1969). See also A. protzi Hartwig, 1900, A. phreatica Dumont,
1983, A. smirnovi Petkovski & Flner, 1972 and another Baikal endemic, A. setosocaudata Vasiljeva &
Smirnov, 1969.
lacustris. Described as Alona lacustris Daday, 1888 from Hungary (Daday 1888). Types probably lost,
not in DAD. According to Frey (1965a), this may be an older name for A. rustica Scott, 1895 but its identity
remains unclear. Smirnov (1971) listed it correctly as incertae sedis.
ladacensis. Described as Alona ladacensis Brehm, 1936 from North India (Brehm 1936). No information
on types. Incertae sedis in Smirnov (1971).
laevis. Described as Alona glacialis var. laevis Herrick, 1884 from Minnesota, USA (Herrick 1884).
Types lost. Incertae sedis (Smirnov 1971). See A. glacialis Birge, 1879.
laevissima. Described as Alona laevissima Sars, 1888 from Malchi Lagoon (Gracemere), Queensland,
Australia (Sars 1888). No types assigned, but material, likely from type locality, in GOS. Sars (1888)
recognized this animal as close to A. archeri (Sars, 1888). Member of the A. pulchella-complex (Brehm
1933a). Types revised by Sinev (2001a) and listed as junior synonym of A. pulchella. See also A. archeri Sars,
1888 and A. pulchella King, 1853.
VAN DAMME ET AL.
lapidicola. Described as Alona lapidicola Chengalath & Hann, 1981 from Upper Waterton Lake,
Waterton Lakes National Park, Alberta, Canada (Chengalath & Hann 1981, Chengalath 1987). Holotype and
allotype in NMNC, paratypes in NHM, NMNS and USNM. Valid species, related to Alona setulosa Megard,
1967, both of the Alona pulchella complex, but finer morphology remains unknown. Good drawings of
general features (e.g., postabdomen) in Chengalath & Hann (1981). See also A. borealis Chengalath & Hann,
1981, A. pulchella King, 1853 and A. setulosa Megard, 1967.
latissima. Described as Alonopsis latissima Kurz, 1875 from Zbraslavic, Bohemia (Kurz 1875), now
Czech Republic. No information on types. Now Kurzia latissima (Kurz, 1875). Daday (1884) lists it as Alona.
See Hudec (2000) and Kotov (2004) for Kurzia revision.
lebes. Described as Alona elegans lebes Dumont & Van De Velde, 1975 from NW Sahara (Dumont & Van
de Velde 1975). Holotype in RBINS, paratypes in SUG. Details (limb characters) unknown. According to
Alonso (1996), who examined material of A. elegans lebes (Dumont, op.cit.), this is A. elegans sensu stricto.
The possibility that the Mediterranean (below the Alps and Pyrenees) and Northern African populations differ
from European, true A. elegans, should not be excluded, in which case drawings in Alonso (1996) may even
refer to A. elegans lebes. This may be a valid subspecies. See also A. elegans Kurz, 1875 and A. striolata Sars,
1916.
lepida. Described as Alona lepida Birge, 1892 from USA (Birge 1892). No information on types, but
these may be present in DGF. Junior synonym of Alona quadrangularis according to Smirnov (1971), but
revision of populations from the Nearctic in comparison with Palearctic Alona quadrangularis lacking. A.
lepida Birge, 1892 may be a valid name if North American populations of A. quadrangularis would be
different from the European. It is not unusual to find cryptic siblings of Eurasian chydorids in North America
(e.g., Michael & Frey 1984, Megard 1967). See also A. quadrangularis (O.F. Mller, 1776).
leydigi. Described as Alona leydigii Schoedler, 1863 from vicinity of Berlin, Germany (Schoedler 1863,
Kotov 2003, 2009). Types lost, not in ZMHU. Kurz (1875) moved this species to Leydigia when creating the
genus. Later, rmek-Huek et al. (1962) marked L. leydigi as type species of the genus. Now Leydigia
(Leydigia) leydigi (Schoedler, 1863). Correct spelling is leydigi, not leydigii (Kotov 2003, 2009).
leydigioides. Described as as Lynceus rectangulus var. leydigioides Brehm, 1908 from Dalmatia (now
Croatia) (Brehm 1908). No information on types. Insufficiently described, status unclear even according to
Brehm himself (Brehm 1933a). Incertae sedis.
lineata. Described as Lynceus lineatus Fischer, 1854 from vicinity of St. Peterburg (Sergiefskoje near
Peterhof), Europian Russia (Fischer 1854). No information on types. Even though this taxon is likely identical
to Alona rectangula Sars, 1861 and has priority, A. lineata was kept as nomen nudum and A. rectangula Sars,
1861 as the correct name for stability (Frey 1988). Sars (1862) became only aware of Fischers taxon after
naming A. rectangula Sars, 1861, and later used A. lineata (Fischer, 1854) (Sars 1993). The name appears in
several early publications (e.g., Sars 1862, Kurz 1875) but was subsequently replaced by A. rectangula (Frey
1988). Nomenclature in the Coronatella rectangula group is most confusing, particularly within Europe. See
also A. rectangula Sars, 1861.
lineolata. Described as Alonella lineolata Sars, 1901 from So Paulo, Brazil (Sars 1901). No information
on types at GOS. Valid species. Elmoor-Loureiro et al. (2008) rediscovered this animal in Brazil and showed
that it is a member of the Aloninae, closer to Alona or a derived genus. The animal is under description,
including limb characters.
lineolata-2. Described as Alona verrucosa lineolata Chen & Li, 1991 from reservoir in Dongshan Village,
Huarong County, Hunan Province (Chen et al. 1991). Types in IHW. Drawings show this taxon is related to
Alona verrucosa but the status is unclear. It is likely a separate species, with strong similarities to A.
mediterranea Yalim, 2005 (and with priority). But Alona lineolata Chen & Li, 1991 is a secondary homonym
of A. lineolata (Sars, 1901). In case of conspecify of Alona lineolata Chen & Li, 1991 with A. mediterranea
Yalim, 2005, the former name can be replaced by the latter which is the "next oldest available name from
among its synonyms" (article 23.3.5 of ICZN 1999). In case of two different taxa, a replacement name for A.
lineolata Chen & Li, 1991 is needed. See also A. verrucosa Sars, 1901.
31
longinqua. Described as Biapertura longinqua Smirnov, 1971 from New South Wales, Australia
(Smirnov 1971). Types in AM, MGU and ZIN. Likely a valid species but affinities unclear, this animal needs
revision. Cannot be assigned to an existing group at present.
longirostris. Described as Alona longirostris Daday, 1898 from Colombo, Sri Lanka (Daday 1898).
Lectotype and paralectotypes in DAD. Valid species that belongs to Kurzia Dybowski & Grochowski, 1894
under the name Kurzia longirostris (Daday, 1898). See Hudec (2000) and Kotov (2004) for Kurzia revision.
longirostris-2. Described as Alona longirostris Henry, 1919 from New South Wales, Australia (Henry
1919). Holotype in AM. Affinities unclear (Brehm 1933a), synonym of Alona affinis (Leydig, 1860)
according to Smirnov (1971). True A. affinis does not occur in Australia (Sinev 1997) so synonymy is
unlikely. Two other species of the A. affinis group are considered valid for this continent, A. elliptica Sinev,
1997 and A. kendallensis Henry, 1919 and a third taxon, of unclear status, A. whiteleggei Sars, 1896. In any
case, the name A. longirostris Henry, 1919 is a primary homonym of Alona longirostris Daday, 1898. So, in
case of conspecify of A. longirostris Henry, 1919 with A. elliptica Sinev, 1997, A. kendallensis Henry, 1919 or
A. whiteleggei Sars, 1896 (the chance is very high!), the former name can be replaced by the "next oldest
available name from among its synonyms" (case 23.3.5 of ICZN, 1999). See also A. affinis (Leydig, 1860), A.
elliptica Sinev, 1997, A. kendallensis Henry, 1919 and A. whiteleggei Sars, 1896.
longispina. Described as Alona longispina Chen & Yang, 1989 from Guizhou Province, China (Chen et
al. 1989). Types at IHW. Status unclear, may be valid but description insufficient. Postabdomen with general
shape of A. pulchella-group, but with relatively long basal spine (longispina) and ventral indentation of the
labral keel. Johnson (1956a) reported a similar animal from Sumatra, under A. cf. glabra, with the same
type of postabdomen (long basal spine), so this form may have a wider distribution in Asia. The A. pulchellagroup in this region is complex. See also A. archeri Sars, 1888, A. arenaria Brehm, 1937 and A. pulchella
King, 1853.
macracantha. Described as Alona macracantha Smirnov & Timms, 1983 from New South Wales,
Australia (Smirnov & Timms 1983). Holotype in MGU, paratypes in MGU and ZIN. Australian endemic with
large ocellus, long basal spine, and peculiar postabdomen (Smirnov & Timms 1983). Valid species, not an
Alona. Moved to a new genus by Sinev & Shiel (2008), now Maraura macracantha (Smirnov & Timms,
1983). See also A. brevicaudata Chen, Zhang & Yang, 1991 and A. macrocopa Sars, 1894.
macrocopa. Described as Alona macrocopa Sars, 1894 from Kaitaia, North Island, New Zealand (Sars
1894). Lectotypes and paralectotypes selected by Frey in GOS (Sinev 2004b). Valid species. Placed in
Biapertura by Smirnov & Timms (1983), later removed from Alona or Biapertura and allocated to the genus
Armatalona Sinev, 2004 by Sinev (2004b), now known as Armatalona macrocopa (Sars, 1894). See also A.
abbreviata Sars, 1895 and A. imitatoria (Smirnov, 1989).
macronyx. Described as Alona macronyx Daday, 1898 from Colombo Lake, Sri Lanka (Daday 1898).
Lectotype and paralectotypes in DAD (Rajapaska & Fernando 1985). Redescription, including limbs, in
Rajapaksa & Fernando (1985). Widespread in SE Asia (e.g., Idris 1983, Sarma & Sarma 1990). Valid species
but definitely not a member of Alona sensu stricto. Needs revision. This species was assigned to Alona, then
moved to Indialona Petkovski, 1966 (Smirnov 1971) but belongs to neither (Rajapaksa & Fernando 1985).
Rajapaksa & Fernando (1985) suggest Alona macronyx Daday, 1898 may need assignment to a new,
monotypic genus, but until revision, its position remains unclear. Indialona ganapati Petkovski, 1966, which
has a peculiar morphology, had a similar fate (Kotov 2000). See also A. globulosa Daday, 1898.
macrops. Described as Alona macrops Daday, 1898 from Madatugama and Kalawewa Lake, Sri Lanka
(Daday 1898). Lectotype and paralectotypes in DAD. Removed from Alona by Sars (1901) when creating the
genus Dadaya Sars, 1901. This is a valid species but far from Alona. It is now called Dadaya macrops (Daday,
1898) and is a member of the Chydorinae, not of Aloninae (Smirnov 1996a). Sars (1901) based his definition
of Dadaya macrops on Neotropical specimens, but more than one species may be present in the tropics. A
second name is available from Brazil, Dadaya ocellata Bergamin, 1940. See Rajapaksa & Fernando (1982b)
for details on Dadaya macrops (Daday, 1898).
VAN DAMME ET AL.
macrops-2. Described as Alona macrops Mota & Orghidan, 1948 from Rumania (Mota & Orghidan
1948). No information on types. Description based on single specimen with large ocellus, which may be an
artefact. This animal is in reality Alona quadrangularis (O.F. Mller, 1776). A primary homonym of Alona
macrops Daday, 1898. See also A. quadrangularis (O.F. Mller, 1776).
macrorhyncha. Described as Alona macrorhyncha Daday, 1900 from New Guinea (Daday 1900, 1901).
Lectotype in DAD. This is a junior synonym of Kurzia longirostris (Daday, 1898) (e.g., in Harding 1957). See
Hudec (2000) and Kotov (2004) for Kurzia revision.
maduensis. Proposed as Alona Madensis Keilhack, 1905 from Madusee (Jezioro Miedwie), Pomerania,
Poland (Keilhack 1905). No information on specimens, but part of Weltners collection (on which Keilhack
based his study) is present at ZMHU in the collection of D. Flssner. Status unclear. Keilhack (1905) proposes
a separate name for specimens of the Alona rectangula-group from Poland, which he describes as Alona sp.
(in German): I propose the name A. Madensis in case a revision of Alona would indicate the herein
described specimens as an independent species. The note comes after description of the specimens. The
name is also mentioned in Keilhack (1908), but was not picked up by other authors. We regard A. maduensis a
form of Coronatella rectangula (Sars, 1861), a species group for which the situation in Europe is complex.
See also A. rectangula Sars, 1861.
martensi. Described as Alona martensi Sinev, 2009 from Drakensberg Region, Republic of South Africa
(Sinev 2009a). Holotype and paratypes in MGU. A valid species, member of Alona affinis group, described in
detail. Endemic to the Drakensberg Mountains (Sinev 2009a). See also A. affinis (Leydig, 1860) and A. affinis
barbata Brehm, 1935.
mascula. Described as Alona mascula King, 1853 New South Wales, Australia (King 1853). No
information on types. According to Sars (1888), this is a male of A. bairdi King, 1853 (see under that name);
Smirnov (1966a) considers A. mascula as the male of Alonopsis elongata (Sars, 1861). The latter is unlikely:
true A. elongata is a Palaearctic species that does not occur in Australia (Kubersky 1977). Later listed as
incertae sedis (Smirnov 1971, Smirnov & Timms 1983).
mediterranea. Described as Alona mediterranea Yalim, 2005 from Antalya, Turkey (Yalim & Ciplak
2005). The authors noted that the description of the species was the responsibility of E.B. Yalim only (Yalim
& Ciplak 2005). Holotype and paratypes in AZUM. Described from Turkey, but with wide distribution in
Africa and Arabia (Van Damme, unpubl.). Member of the Alona verrucosa-complex, description contains
limb characters (Yalim & Ciplak 2005). A. mediterranea may be identical to chinese A. lineolata Chen et al.,
1991. Another, earlier name available for African A. verrucosa-like animals, is Alona rectangula var. africana
Daday, 1908 from Malawi (Daday 1908). See also A. lineolata Chen & Li, 1991, A. rectangula var. africana
Daday, 1908 and A. verrucosa Sars, 1901.
meridionalis. Described as Alona meridionalis Sinev, 2007 from Drakensberg region, Republic of South
Africa (Sinev 2007), likely endemic. Holotype and paratypes in MGU. Valid species. Removed from Alona
and allocated by Van Damme & Dumont (2008b) to Ovalona. Peculiar because of its transverse lateral head
pores. Now known as Ovalona meridionalis (Sinev, 2007). See also A. subantarctica Ekman, 1905 and A.
weinecki Studer, 1878.
microtata. Described as Alona microtata Henry, 1922 from New South Wales, Australia (Henry 1922).
Syntypes in AM. Status unknown. Drawings show an aberrant morphology, with angular postabdomen. Status
unclear. Shiel & Dickson (1995) list it as a synonym of A. guttata Sars, 1862, but the poor description makes
it impossible to determine affinities or validity of this small (0.28 mm) animal. Drawings reproduced in
Smirnov (1971).
milleri. Described as Alona milleri Kiser, 1948 from Pearl River, Canton, China (Kiser 1948, Chiang &
Du 1979). Paratypes in DGF and USNM. Holotype at USNM. One of the rarest Aloninae. Good drawings by
Kiser (1948) show an animal with unmistakable postabdomen, short basal spine and long lateral fascicles.
Affinities unknown, Kiser (1948) notes that it is related with A. rectangula. We have no doubt Kisers species
is valid, but a redescription is necessary to clarify its position; we are not sure it is related to Coronatella. No
figures of this animal exist besides Kisers (1948). See also A. rectangula Sars, 1861.
33
minor. Described as Alona intermedia var. minor Stingelin, 1905 from Saigon, Vietnam (Stingelin 1905).
Type material in NMO. Only known from the original description by Stingelin (1905), which does not allow a
decision on its status. Raised to subspecies rank by Smirnov (1971), as Biapertura. Likely more than one
cryptic species is hiding under the cosmopolitan A. intermedia Sars, 1862, but Stingelins form has likely
nothing to do with the latter (Brehm 1933a). So, status unclear, likely unrelated to A. intermedia. See also A.
intermedia Sars, 1862.
minuta. Described as Alona minuta Poggenpol, 1874 from Moscow town, Russia (Poggenpol 1874). No
information on types. Synonym of Alona guttata Sars, 1862 in Smirnov (1971). See also A. guttata Sars,
1862.
modesta. Described as Alona modesta Herrick, 1884 from Minnesota, USA (Herrick 1884). Types lost.
Status unclear. Incertae sedis in Smirnov (1971).
monacantha. Described as Alona monacantha Sars, 1901 from So Paulo and Ipiranga, Brazil (Sars
1901). Lectotype and paralectotypes in GOS selected by Sinev (2004a), other samples of Sars in the same
collection. Distributed in Neotropics. Records outside Neotropics need revision (e.g., A. monacantha in Idris
1983) or refer to other species (e.g., A. monacantha in Korinek 1984 is the African sibling A. hardingi). Valid
species related to A. rectangula. Morphology redescribed by Sinev (2004a). Like A. rectangula, the A.
monacantha group does not belong to Alona. A. monacantha is much closer to Coronatella and differs only in
minor characters (e.g., denticles on labrum or carapace), see Van Damme & Dumont (2008b); specimens
without denticles on the valves may be present in the Neotropics (Sinev, pers. comm.). We list it here as
Coronatella monacantha (Sars, 1901) comb. nov.. Smirnov (1971) discerns two subspecies (A. monacantha
monacantha Sars, 1901 and A. monacantha tridentata Stingelin, 1905), but the situation is complex (see under
tridentata). See also A. acuticostata Sars, 1903, A. acuticostata var. tridentata Stingelin, 1905, A. cannellata
Brehm, 1934, A. circumfimbriata Megard, 1967, A. hardingi Brehm, 1957, A. rectangula Sars, 1861 and A.
reiseri Spandl, 1926.
moniezi. Described as Alona moniezi Richard, 1888 from Auvergne, France (Richard 1888) without
drawings. Never reported since. Type material in DGF, includes males. Examination of the types shows that A.
moniezi is nearly identical to A. elegans, but females show features of C. rectangula. In comparison to C.
rectangula, it lacks strong denticles on IDL setae of P1 and has different features in the postabdomen. Status
and redescription will be discussed in a separate paper, but it is likely an invalid taxon. Revision of the types
and comments on this taxon in Van Damme et al. (in press). See also A. rectangula Sars, 1861.
muelleri. Described as Alona Mlleri Richard, 1897 from Lunache, Chile (Richard 1897). Neotropics.
Valid species. No information on types. Allocated to Karualona Dumont & Silva-Briano, 2000 by Sinev &
Hollwedel (2005), now as Karualona muelleri (Richard, 1897). One of two well described Neotropical
Karualona species, the other is K. penuelasi Dumont & Silva-Briano, 2000. Redescribed in detail by Sinev &
Hollwedel (2005). See also A. karua King, 1853.
muelleri-2. Alona muelleri Ren, 1992 from North West Territories, Canada (Ren 1992). Types in
ZMUC. Ren (1992) notes that this taxon differs from A. costata in the shape of the labral plate, weak
striation on the carapace, and rounded preanal corner. According to Smirnov (2001), it is A. rustica Scott,
1895, like A. fabricii Ren, 1992. To us, A. muelleri seems more similar to Alona costata Sars, 1862. In any
case, this is a junior homonym of Alona muelleri Richard, 1897. See also A. costata Sars, 1862, A. fabricii
Ren, 1992, A. rustica Scott, 1895.
nalibokiana. Described as Coronatella nalibokiana Dybowski & Grochowski, 1895 from Poland
(Dybowski & Grochowski 1895). No information on types. Status unclear and no figures; nomen nudum
(Smirnov 1971).
natalensis. Described as Alona natalensis Sinev, 2008 from KwaZulu-Natal, Republic of South Africa
(Sinev 2008). Holotype and paratypes in MGU. Valid species of the A. costata-complex, described in detail
(Sinev 2008). See also A. costata Sars, 1862.
neglecta. Described as Alona neglecta Scott, 1895 from Shetland Islands, Scotland, United Kingdom
(Scott 1895). No information on types. Junior synonym of Alona intermedia Sars, 1862 (e.g., in Brehm 1933a,
VAN DAMME ET AL.
Alonso 1996). In Smirnov (1971) listed under Biapertura intermedia (Sars, 1862). See A. intermedia Sars,
1862.
nigra. Described as Alona nigra Smirnov, 1996 from Bolivia (Smirnov 1996b). Holotype and paratypes in
MGU. Valid species, but detailed morphology unknown. Peculiar in having three separate head pores, a rare
character, known in the A. pulchella group (e.g., A. nuragica Margaritora, 1971). See also A. pulchella King,
1853.
novaezealandiae. Described as Alona novae-zealandiae Sars, 1904 from DUrville Island, New Zealand
(Sars 1904). Types probably lost, not in GOS. Drawings of limbs in Smirnov (1971) from Australian
populations. Allocated to Coronatella by Van Damme & Dumont (2008b). Now named Coronatella
novaezealandiae (Sars, 1904). Name also applied to Africa, for example by Jenkin (1934) and Rey & St-Jean
(1969), but these populations belong to C. anemae Van Damme & Dumont, 2008. Needs redescription;
perhaps there are two separate species of the C. rectangula-complex in New Zealand. If not, C.
novaezealandiae may be a junior synonym of C. eucostata (Sars, 1894) (Van Damme & Dumont 2008b), both
clearly belong to the C. rectangula-complex. See also A. eucostata Sars, 1894 and A. rectangula Sars, 1861.
nuragica. Described as Alona nuragica Margaritora, 1971 from Sardinia, Italy (Margaritora 1971). Types
in IZUR. Circum-Mediterranean species (Spain, Italy, Sicily). Drawings in Alonso (1996). Valid species.
Unusual member of the Alona pulchella-complex with three separate main head pores, like A. cambouei.
Limb details unpublished but clearly related to A. pulchella (Van Damme, unp.). See also A. pulchella King,
1853.
oblonga. Described as Alona oblonga P. E. Mller, 1867 from Denmark (Mller 1867). Types in ZMUC
and some of P. E. Mllers material is also present in Lilljeborgs collection at UUZM. Junior synonym of A.
affinis (Leydig, 1860) (Lilljeborg 1901, Brehm 1933a, Smirnov 1971, Sinev 1997). See also A. affinis
(Leydig, 1860).
orellanai. Described as Alona orellanai Alonso, 1996 from Spain (Alonso 1996). Holotype and paratypes
in MNSM. Iberian Peninsula. Valid species. Member of the A. elegans-complex, description in Alonso
(1996). See also A. elegans Kurz, 1875 and A. salina Alonso, 1996.
ornata. Described as Alona affinis var. ornata Stingelin, 1895 from Lake Titicaca, Peru (Stingelin 1895).
Holotype in NMO (Frenzel 1987). A confusing taxon, status unclear. Stingelin created A. affinis var. ornata
based on a peculiarity of this form from Titicaca Lake, namely carapaces having a fine striation. Later,
Stingelin (1895) found the same fine sculpturing in A. affinis from Switzerland (Basel) and noted Alona affinis
var. ornata Stingelin, 1895 for the first time, with distribution Peru and Switzerland. So, the original,
available type of Alona affinis ornata is from Lake Titicaca, but the name appears first in a publication on
Europe. From two different continents, these are likely two different taxa, a typical transfer of names. Frenzel
(1987) notes that type material of A. affinis ornata Stingelin, 1895 has different head pores from the European
A. affinis (Leydig, 1860) (anterior pore is much smaller, posterior large and with chitineous ring). These types
should be checked, as A. affinis ornata Stingelin, 1895 has priority on another South American relative, A.
ossiani Sinev, 1998, should these two be identical. Alternatively, A. affinis ornata may be an endemic of Lake
Titicaca. Its status remains unclear until the types and new material of the A. affinis group from Titicaca can
be examined. In Europe, mainly in palaeolimnological remains, the name A. affinis ornata Stingelin, 1895 is
used as a variety of A. affinis with fine striation; Parenzan (1932) even considered it a full species, A. ornata.
The ornamentation is considered a variable character between populations in A. affinis, and in Europe it is
considered a morph (Sinev 1997). Listed by Smirnov (1971) as one of the subspecies of A. affinis. See also A.
affinis (Leydig, 1860) and A. ossiani Sinev, 1998.
ornata-2. Described as Alona rectangula var. ornata Shiklejew, 1929 from Abkhazia, Caucasus
(Shiklejew 1929, 1930). No information on types. Behning (1941) lists this taxon as synonym of Alona
rectangula caucasica Shiklejew, 1933, which is an illogical step keeping priority of the former in mind. May
be a member of the A. verrucosa-group shape of male and female postabdomen are similar and A. cf.
verrucosa occurs in Abkhazia (Sinev 2002c). Smirnov (1971) as synonym of the Australian A. archeri Sars,
1888, but the latter is highly unlikely. Status remains unclear. In any case, it is a junior homonym of Alona
35
affinis var. ornata Stingelin, 1895. See also Alona rectangula caucasica Shiklejew, 1933 and Alona affinis var.
ornata Stingelin, 1895.
ossiani. Described as Alona ossiani Sinev, 1998 from So Paulo, Brazil (Sinev 1998). Holotype and
paratypes in GOS. Widely distributed in Neotropics. Description in Sinev (1998) including limbs, see also
drawings in Elas-Gutirrez et al. (2006). Valid species and member of the A. affinis-complex, but priority and
validity of A. affinis ornata Stingelin, 1895 of which types are from Lake Titicaca, should be checked. See
also A. affinis (Leydig, 1860) and A. affinis ornata Stingelin, 1895.
ovata. Described as Alona ovata Baird, 1850 from Blackbeath, UK (Baird 1850). This material is present
at NHM, according to White (1847), and although deposited before the description, may be the original
specimens Baird (1850) based his description on. Status unknown. Description insufficient, original material
if retrieved should be studied. Incertae sedis in Kurz (1875), Smirnov (1971).
ovata-2. Described as Alona ovata Rey & Vsquez, 1986 from Venezuela (Rey & Vsquez 1986). Types
in the collection of Rey and Vsquez, but no information on location of this collection. Drawings in Rey &
Vsquez (1986) suggest a member of the Coronatella-branch, but further affinities unknown. Invalid name,
primary homonym of A. ovata Baird, 1850. Smirnov (2001) suggested a nomen novum, A. inovata Smirnov,
2001 for this taxon. See A. inovata Smirnov, 2001.
parva. Described as Alona parva Daday, 1905 from Paraguay-Paran River Basin, Paraguay (Daday
1905). Lectotype and paralectotypes in DAD. Initially described as Leydigia and moved to Alona by Smirnov
(1971). Limbs show that this taxon belongs to neither, removed and allocated to monotypic genus Parvalona
by Van Damme et al. (2005). Now known as Parvalona parva (Daday, 1905).
parvula. Described as Alona parvula Kurz, 1875 from Deutschbrod and Rokycan, Bohemia, Czech
Republic (Kurz 1875). No information on types, status unclear. Form of Alona guttata Sars, 1862, with faint
striation. Regarded as synonym (Lilljeborg 1901, Brehm 1933a, Alonso 1996) or subspecies (Smirnov 1971)
of A. guttata Sars, 1862. We regard it as a form, but the A. guttata-group needs revision. See also A. guttata
Sars, 1862.
patagonica. Described as Alona cambouei var. patagonica Ekman, 1900 from South Patagonia, Chile
(Ekman 1900). Lectotype and paralectotypes in SMNH (Frey 1988, Kotov & Gololobova 2005); material
labeled as A. cambouei from this region in DAD likely corresponds to the same species. Listed as synonym
of A. pulchella King, 1853 in Smirnov (1971). Ekman (1900) did not include drawings with the original
description, but notes that patagonica differs in postabdomen armature, e.g., longer distal spines in lateral
fascicles and a clear spinule adjacent to the basal spine. The author confirmed differences of this South
American form again after comparison with A. cambouei from the White Nile, Africa (Ekman 1901).
Synonymy with A. pulchella King, 1853 is unlikely, as latter is replaced in South America by A. glabra Sars,
1901. Status of A. patagonica unclear; it may be a valid species. Ekman (1900) notes it to be common and
abundant in Southern Patagonia, also recorded here by Daday (1902). In DAD there are several tubes with
rich material from Patagonia labelled as A. cambouei de Guerne & Richard, 1893 (Forr & Frey 1982). We
can confirm this material belongs to the A. pulchella-group and contains animals close to A. glabra Sars,
1901; the rich material from Dadays collection could be useful for a redescription. See also A. cambouei de
Guerne & Richard, 1893, A. glabra Sars, 1901 and A. pulchella King, 1853.
pectinata. Described as Alona pectinata Elas-Gutterez & Suarez-Morales, 1999 from South Mexico
(Elas-Gutterez & Suarez-Morales 1999). Holotype in USNM, paratypes in USNM, UNAM, ECO-CH-ZOO.
Material in USNM listed as Biapertura pectinata. Member of A. verrucosa-complex, limb morphology
described by Elas-Gutterez & Suarez-Morales (1999). Closely related to Neotropical Alona verrucosa Sars,
1901 and will be part of a new genus. See also A. verrucosa Sars, 1901.
phreatica. Described as Alona phreatica Dumont, 1983 from Auvergne, France (Dumont 1983).
Holotype and paratypes in SUG. Distributed in Western Europe, an obligate interstitial. Valid species.
Allocated to Phreatalona by Van Damme et al. (2009), now named Phreatalona phreatica (Dumont, 1983).
Males depicted in Sabater (1987), Alonso (1996) and Van Damme et al. (2009), description of limbs in the
latter publication. See also A. labrosa Petkovski & Flssner, 1972, A. protzi Hartwig, 1900, A. smirnovi
Vasiljeva & Smirnov, 1969.
VAN DAMME ET AL.
poppei. Described as Alona poppei Richard, 1897 from Lunache, Chile (Richard 1897). Types most
probably lost, not in DGF. Valid species. Not Alona, likely a valid Coronatella (Van Damme & Dumont
2008b), now Coronatella poppei (Richard, 1897). The name was transferred to Africa and used for taxa
related to C. rectangula (Sars, 1861) (e.g., Delachaux 1917). These records are now considered to belong to
Coronatella anemae Van Damme & Dumont, 2008. C. poppei (Richard, 1897) is unlikely to occur in Africa
(Van Damme & Dumont 2008b) and also records from Australia (Smirnov & Timms 1983) most likely belong
to another species. Drawings of morphology are shown by Rey & Vsquez (1986) from Venezuela and
confirm it as a separate species, but C. poppei needs redescription based on material from Chile. In DAD,
abundant material labelled as A. poppei Richard, 1897 is present from Patagonia, but should be rechecked
(may belong to A. cambouei patagonica). See also A. circumfimbriata Megard, 1967, A. eucostata Sars, 1894,
A. novae-zealandiae Sars, 1904 and A. rectangula Sars, 1861.
porrecta. Described as Alona porrecta Birge, 1879 from Massachusets and Wisconsin, USA (Birge
1879). No information on types. Birges (1879) single figure does not allow identification. Drawing refer
likely to a male alonine the name refers to the pores in the postabdomen, opening of the spermiducts (Sinev,
pers. comm.). Incertae sedis in Smirnov (1971).
protzi. Described as Alona protzi Hartwig, 1900 from Knigsberg (now Kaliningrad, Russia), bank of
Hellsee by Biesenthai, Brandenburg, Germany (Hartwig 1900). No types assigned; one male from from the
type locality used for original description (to be designated as allotype) and several specimens from
Brandenburg in the Wiener and Flner Collections at ZMHU (Van Damme et al. 2009). Valid species.
Distributed in Danubian Europe. Easily recognized as the only European Alona-like species with three
denticles on posteroventral valve corner. Peculiar ecology, prefers interstitial; removed from Alona to
Phreatalona Van Damme, Brancelj & Dumont, 2009. A. protzi is the type species of the genus (Van Damme et
al. 2009), now named Phreatalona protzi (Hartwig, 1900). See A. protzi typica Behning, 1941 and A. protzi
shiklejewi Brehm, 1933 (forms from the Caucasus), A. labrosa Vasiljeva & Smirnov, 1969, A. phreatica
Dumont, 1983 and A. smirnovi Petkovski & Flssner, 1972.
pseudanodonta. Described as Alona pseudanodonta Brehm, 1933 from Mindanao Island, Philippines
(Brehm 1933b), locality in Brehm (1938). No information on types. May be close to A. verrucosa, but true
affinities unclear. In Smirnov (1971), this is a valid species of Biapertura, with two subspecies, B.
pseudoanodonta pseudoanodonta Brehm, 1933 and B. pseudanodonta anodonta Daday, 1905. This was
taxonomically incorrect, see anodonta. In addition, the assignment of two subspecies under pseudoanodonta
by Smirnov (1971) is based on presence or absence of tubercles, a character that describes varieties (e.g., A.
tuberculata Kurz, 1875). For now, we prefer to list A. pseudanodonta Brehm, 1933 and A. anodonta Daday,
1905 as two separate taxa of which the details are unknown. Both may belong to either Coronatella or Gen.
Nov. 1. Alona cf. anodonta, depicted in Rajapaksa & Fernando (1982a) may correspond to this taxon, but due
to the absence of types and unclear description by Brehm (1933b), it is impossible to decide. The latter (A. cf.
anodonta in Rajapaksa & Fernando 1982a) is clearly a Coronatella. See also A. anodonta Daday, 1905.
pseudopulchra. Proposed as Alona rectangula pseudopulchra Smirnov, 1968 in his Dr. Sci. Dissertation
(Smirnov 1968), so this is a manuscript name. No information on types. The taxon was proposed to
distinguish between morphs with smooth and tuberculate valves, both described with the same name (A.
pulchra) by Hellich (1874). Valve ornamentation in Coronatella rectangula (Sars, 1861) is a variable trait
(Sinev 2001b). Later synonimized with A. rectangula rectangula by Smirnov (1971). See also A. pulchra
Hellich, 1874 and A. rectangula Sars, 1861.
pseudoverrucosa. Described as Biapertura pseudoverrucosa Smirnov, 1971 from Paraguay, based on a
record of A. rectangula in Daday (1905). Samples containing A. rectangula from Paraguay are absent in DAD.
Dadays figures show a clear member of the A. verrucosa-group, although the author notes a few differences.
Smirnov (1971) moved A. verrucosa Sars, 1901 to Biapertura and created a new name Biapertura
pseudoverrucosa Smirnov, 1971, with two subspecies: B. pseudoverrucosa pseudoverrucosa Smirnov, 1971
and B. pseudoverrucosa verrucosa (Sars, 1901). As for A. pseudoanodonta (see above), proposing an earlier
described taxon as subspecies of one described later, is incorrect. So, A. pseudoverrucosa is a junior synonym
of Alona verrucosa (Sars, 1901). See also A. anodonta Daday, 1905 and A. verrucosa Sars, 1901.
37
pulchella. Described as Alona pulchella King, 1853 from Varroville, Sydney, New South Wales, Australia
(King 1853). Australian taxon, no reliable records exist as yet from outside latter continent (Sinev, 2001a). No
information on types. Frequently confused with Neotropical A. glabra Sars, 1901 and with Alona cambouei de
Guerne & Richard, 1893. The latter is a sibling from Madagascar that is likely widespread on the African
continent and extends into Asia (Sinev 2001a). True A. pulchella does not occur in South America. A.
cambouei is close to A. pulchella, but has three separate main head pores (Sinev 2001a). Oldest name in the A.
pulchella-complex, a group widespread in the southern hemisphere, with several species. Redescription of A.
pulchella, including limbs, in Sinev (2001a). In South East Asia and Australia, several species exist of the A.
pulchella-complex. See also A. archeri Sars, 1888, A. arenaria Brehm, 1933, A. azorica Frenzel & Alonso, A.
beverleyae Smirnov, 1989, A. borealis Chengalath & Hann, 1981, A. bromelicola Smirnov, 1988, A. cambouei
de Guerne & Richard, 1893, A. cambouei var. patagonica Ekman, 1900, A. fasciculata Daday, 1905, A. glabra
Sars, 1901, A. inreticulata Shen, Song & Chen, 1964, A. laevissima Sars, 1888, A. lapidicola Chengalath &
Hann, 1981, A. longispina Chen & Yang, 1989, A. nigra Smirnov, 1966, A. nuragica Margaritora, 1971, A.
setuloides Smirnov & Timms, 1983, A. setulosa Megard, 1967 and A. unguiculata Smirnov, 1989.
pulchra (suppressed). Described as Alona pulchra Hellich, 1874 from Bohemia (Hellich 1874), Czech
Republic. No information on types. Brehm (1933a) lists it as separate species; Smirnov (1971, 1996b) notes it
as a subspecies of A. rectangula; Sars labeled several specimens of A. rectangula of his collection as pulchra.
The form is likely a tuberculate variety of A. rectangula, there is no clear morphological difference between
both (Frey 1988, Sinev 2001b). A similar case exists with the tuberculate forms (tuberculata) of A. rustica and
A. guttata. In any case, this name is suppressed by ICZN (1958) and placed in the "Official Index of Rejected
and Invalid Species Names in Zoology". Being an ecological morph of Coronatella rectangula (Sars, 1861),
tuberculate rectangula can be called the "pulchra" form without taxonomic meaning. See also A. rectangula
Sars, 1861, A. rectangula pseudopulchra Smirnov, 1968 and A. tuberculata Kurz, 1875.
punctata. Described as Alona punctata Daday, 1898 from Madatugama, Kalawewa and Mahaveliganga,
Sri Lanka (Daday 1898). Types in DAD. Present also in India. Sometimes listed as Alona davidi punctata.
Figures in Daday (1898) and an examination of type material indicate this is Leberis, not Alona (Sinev et al.
2005). This Indian form may be a junior synonym of Alona diaphana King, 1853 according to Frey (1991),
now Leberis diaphanus (King, 1853) (Sinev et al. 2005). See also A. davidi Richard, 1895, A. diaphana King,
1853 and A. hyalina (Chen & Gao 1989).
pygmaea. Described as Alona pygmaea Sars, 1862 from Norway (Sars 1862). Types not in GOS. Junior
synonym of Alonella nana (Baird, 1850), see Kurz (1875) and Lilljeborg (1901).
quadrangularis. Described as Lynceus quadrangularis O.F. Mller, 1776 from Denmark (Mller 1776).
Types lost, but most Cladocera described by O.F. Mller (1776, 1785) were likely collected near the castle in
Frederiksdal where he resided in summer. Neotypes of A. quadrangularis should be designated from the
direct vicinity, as for other cladocerans (e.g., from Lake Huul S, see Korovchinsky 2008 or Lake Sjaels, see
Frey 1980). Nominal species of Alona sensu stricto or the A. quadrangularis-complex and type species of the
subfamily Aloninae. Designation to Alona Baird, 1843 by Johnson (1956b), who marked quadrangularis as
type species of the genus (confirmed by ICZN 1958). Considered a Palaearctic species. Populations outside
the region need revision and may contain cryptic species, in addition to two recently described siblings from
South America and Africa (Sinev & Coronel 2006, Van Damme & Dumont 2008a). General morphology in
Alonso (1996), Sinev & Coronel (2006), limb morphology in Smirnov (1966b) and Van Damme & Dumont
(2008a). Overview of the history of this species and a key to species in Van Damme & Dumont (2008a). See
also A. boliviana Sinev & Coronel, 2006, A. kolwezii Van Damme & Dumont, 2008, A. lepida Birge, 1892 and
A. whiteleggei Sars, 1896.
rectangula. Described as Alona (Lynceus) rectangula Sars, 1861 from vicinity of Oslo, Norway (Sars
1862). Lectotype and paralectotypes in GOS selected from Dam paa Egeberg, see Frey (1988); the latter
locality is likely Lake stenjvand, Ekeberg, Oslo, Norway, see remarks under A. guttata Sars, 1862. True
rectangula is considered restricted to Eurasia (Sinev 2001b). Forms a complex in Europe and species look
very similar worldwide. May be a junior synonym of A. lineata (Fischer, 1854) but A. rectangula was retained
for stability (Frey 1988); Sars was unaware of Fischers taxon and used the name A. lineata as well (e.g., Sars
VAN DAMME ET AL.
1862). See notes by Frey in Sars (1993). Both names appear to indicate the same taxon (e.g., Kurz 1875).
Coronatella rectangula Sars, 1861 has a confusing synonymy and contains most available names of all Alona
species (Sinev 2001b). Smirnov (1971) lists seven subspecies of Coronatella rectangula (as A. rectangula):
rectangula, coronata, richardi, elegans, novae-zealandiae, pulchra and serrata; status as subspecies is not
certain for all, some are good species (e.g. A. elegans). Even more names exist in literature for the C.
rectangula group (e.g. caucasica, pseudopulchra, madensis, trachystriata). See under these names. The
tuberculate form (not subspecies), can be called the pulchra-form after C. rectangula var. pulchra (Hellich,
1874). Examples of names with an unclear status are A. inornata Hudendorff, 1876, A. madensis Keilhack,
1905 or A. richardi Stingelin, 1895. C. rectangula is most likely to still harbour cryptic species in Europe
(Frey 1988). The complex in Eurasia needs detailed unraveling and assignment of names (e.g., aculeata,
richardi, coronata, trachystriata). Differences may be beyond classical morphological studies and the
widespread C. rectangula-complex would benefit from molecular work. Outside Europe, the situation for
relatives of C. rectangula is even more confusing (e.g., bukobensis, arcuata, harpularia, novae-zealandiae,
eucostata, poppei). Populations are highly variable but differences between taxa are sometimes small (see for
example C. anemae in Van Damme & Dumont 2008b). Notes on morphology of populations from Norway by
Frey (1988), from Russia by Sinev (2001b), description of C. rectangula limbs from Belgium in Van Damme
& Dumont (2008b). All A. rectangula forms are far from Alona. Van Damme & Dumont (2008b) assigned this
group to a separate genus, for which thorough literature research revealed an available name. Housed in the
genus Coronatella Dybowski & Grochowski, 1894, it is now known as Coronatella rectangula (Sars, 1861).
The genus, with Leberis and others, forms a typical group of the Coronatella-branch (Van Damme & Dumont
2008b). See also A. aculeata Werestschagin, 1911, A. acuticostata Sars, 1903, A. angulata Weigold, 1911, A.
arcuata Sars, 1916, A. cannellata Brehm, 1934, A. circumfimbriata Megard, 1967, A. coronata Kurz, 1875, A.
elegans Kurz, 1875, A. eucostata Sars, 1894, A. hardingi Brehm, 1957, A. harpularia Sars, 1916, A. holdeni
Green, 1962, A. inornata Hudendorff, 1876, A. lineata (Fischer, 1854), A. madensis Keilhack, 1905, A.
monacantha Sars, 1901, A. moniezi Richard, 1888, A. novae-zealandiae Sars, 1904, A. poppei Richard, 1897,
A. pulchra Hellich, 1874, A. rectangula var. caucasica Shiklejew, 1933, Alona rectangula var. ornata
Shiklejew, 1929, A. rectangula pseudopulchra Smirnov, 1968, Alona rectangula serrata Daday, 1908, Alona
rectangula trachystriata Chen, Zhang & Liu, 1993 and A. richardi Stingelin, 1895.
rectirostris. Described as Phryxura rectirostris P. E. Mller, 1867 from Denmark (Mller 1867).
Description based on "a single specimen with a much distorted postabdomen" (Fryer 1997). The same
conclusion was earlier made by Smirnov (1971), who decided that P.E. Mller (1867) described an atypical A.
costata, so Phryxura rectirostris was regarded as a junior synonym of the former. Later, a slide labelled by P.
E. Mller was found in ZMUC by Frey (1989). It contains Disparalona rostrata (Koch, 1841), so, P.
rectirostris is a junior synonym of the former. The generic name is Phryxura P. E. Mller, 1867 is suppressed,
while Disparalona Fryer, 1968 was placed in the Official List of Generic Names in Zoology (Fryer 1997).
reiseri. Described as Alona reiseri Spandl, 1926 from Paranagua, Brazil (Pesta 1926). No information on
types. Junior synonym of A. monacantha Sars, 1901 according to Smirnov (1971).
reticulata (nomen oblitum). Described as Lynceus (Alona) reticulata Baird, 1843 from Southall,
Middlesex, United Kingdom (Baird 1843). Types at NHM (White 1847). Drawings clearly show that this is
Graptoleberis Sars, 1862, currently known as Graptoleberis testudinaria (Fischer, 1848). Graptoleberis
reticulata (Baird, 1843) had priority over the latter and reticulata was selected as type species of the genus (by
monotypy, see Sars 1862). But Bairds name was later forgotten, while G. testudinaria became widely used
during the XXth century. According to cases 23.9.1.1 and 23.9.1.2 of the ICZN (1999), authors did not require
to replace a name which has been used as valid by at least ten authors in 25 publications during the past 50
years, by a major synonym not in use after 1899. So, the type species of the genus Graptoleberis is Lynceus
reticulata Baird (nomen oblitum), a synonym of Lynceus testudinarius Fischer (nomen protectum).
Graptoleberis testudinaria (Sars, 1862) is cosmopolitan, with several subspecies indicated by Smirnov
(1971), but an updated revision is lacking. Limbs in Alonso (1996), as Graptoleberis testudinaria. See also A.
esocirostris Schoedler, 1858 and A. testudinaria (Fischer, 1848).
39
reticulata-2. Described as Alona guttata var. reticulata Hartmann, 1917 from Austria (Hartmann 1917).
No information on types. Status unknown. Smirnov (1971) kept it as a subspecies of A. guttata Sars, 1862.
Name invalid, primary junior homonym of Alona reticulata Baird, 1843. See also A. guttata Sars, 1862.
richardi. Described as Alona richardi Stingelin, 1895 from Basel, Switzerland (Stingelin 1895).
Lectotype in NMO. Brehm (1933a) lists it as a separate species, Smirnov (1971) as subspecies Alona
rectangula richardi Stingelin, 1895. Junior synonym of C. rectangula according to Frenzel (1987, as Alona)
and Sinev (2001b, as Alona), but status remains unclear. Drawings in Frenzel (1987) based on type material,
show strong differences in postabdomen and armature of IDL setae with true C. rectangula from Norway.
Flner (1962) remarks richardi from Germany. In any case, this taxon does not belong in Alona but
Coronatella, as all A. rectangula forms (Van Damme & Dumont 2008b), but its subspecies or species status is
unclear. Until a thorough redescription or molecular data on the C. rectangula-group, we keep it as a form of
Coronatella rectangula (Sars, 1861). See also remarks under A. rectangula Sars, 1861.
rigidicaudis. Described as Biapertura rigicaudis Smirnov (1971) from New South Wales, Australia
(Smirnov 1971, Smirnov & Timms 1983). Holotype in AM, paratypes in AM, MGU, ZIN. Valid species but
affinities unknown. Not Alona or Biapertura but may be closer to either Coronatella or Karualona, with
reduced limbs (Van Damme & Dumont 2009, unp.). With two main head pores and a peculiar postabdomen, it
is a relatively peculiar animal. See also Biapertura willisi Smirnov, 1981 or Alona willisi (Smirnov, 1981).
rostrata. Described as Lynceus rostratus Koch, 1841 from Germany (Koch 1841). No information on
types. The taxon was housed in Alonella and Alona, see Daday (1884), Lilljeborg (1901). A valid species,
now named Disparalona rostrata (Koch, 1841). For Disparalona, see Smirnov (1992) and Michael & Frey
(1984).
rustica. Described as Alona rustica Scott, 1895 from the Shetland Islands, Scotland, UK (Scott 1895). No
information on types. True A. rustica is a valid Palaearctic species. Morphology in Frey (1965a), Alonso
(1996) and Sinev (1999a). Two sibling species in the New World, A. bicolor Frey, 1965 and A. iheringula
Kotov & Sinev, 2004. Two subspecies (Flssner & Frey 1970), the Palaearctic one is called Alona rustica
rustica Scott, 1895, Nearctic is A. rustica americana Flssner & Frey, 1970. The tuberculate variety in Europe
is known as tuberculata (see below). See also A. archeroides Brehm, 1933, A. bicolor Frey, 1965, A. costata
Sars, 1862, A. iheringula Kotov & Sinev, 2004, A. neglecta Scott, 1895, A. rustica americana Flssner &
Frey, 1970 and A. rustica tuberculata Mamets, 1958.
rusticoides. Described as Biapertura rusticoides Smirnov & Timms, 1983 from Tasmania, Australia
(Smirnov & Timms 1983). Holotype in AM, paratypes in NHM and ZIN. Belongs to Alona after collapse of
Biapertura by Sinev (1999). From original figures, this is clearly a valid species but affinities remain
unknown and limb characters are unstudied. Priority over A. rusticoides Hudec, 1998.
rusticoides-2. Described as Alona rusticoides Hudec, 1998 from Venezuela (Hudec 1998). Holotype and
paratypes in NHM, and in authors collection. Name invalid, junior homonym according to Sinev (1999) after
the collapse of Biapertura Smirnov, 1971. Sinev (1999) suggested a replacement name, A. hudeci Sinev, 1999,
in his revision of the A. costata-group. A key to A. costata species can be found in Sinev (2008). See also A.
rusticoides (Smirnov & Timms, 1983).
salina. Described as Alona salina Alonso, 1996 from Iberian Peninsula, Spain (Alonso 1996). Holotype
and paratypes in MNSM. Valid species of the A. elegans-complex; peculiar in ecology, living in saline waters
(Boronat et al. 2001). Morphology in Alonso (1996) but no limb details. Forms a small species group with
two new species in saline waters, from South-East Russia and Mongolia (Sinev et al. in press). See also A.
elegans Kurz, 1875 and A. orellanai Alonso, 1996.
sanguinea. Described as Alona sanguinea P.E. Mller, 1867 from Denmark (Mller 1867). Types in
ZMUC (as A. anguinea). Junior synonym of Alona quadrangularis in Lilljeborg (1901), Smirnov (1971).
See A. quadrangularis (O. F. Mller, 1776).
sarasinorum. Described as Alona sarasinorum Stingelin, 1900 from Celebes, Sulawesi, Indonesia
(Stingelin 1900, Brehm 1933b). Type in NMO (Frenzel 1987). Reported from SE Asia (Frey 1991, Idris
1983). Valid species with unmistakable characters but insufficiently described. Peculiar in its preference for
VAN DAMME ET AL.
saline waters (Frey 1991). Seems closest to Leberis Smirnov, 1989 due to dorsal keel, postabdomen and
habitus but limbs are unpublished (Supiyanit 2005) and may reveal surprises. Drawings in Frey (1991) under
its junior synonym A. taraporevalae Shirgur & Naik, 1977 and in Idris (1983) under A. cf. sarasinorum. See
also A. taraporevalae Shirgur & Naik, 1977.
sarsi. Described as Alona sarsi Richard, 1891 from Luwu, South Sulawesi, Indonesia (Richard 1891).
Types in DGF. According to Stingelin (1900), it is close to A. sarasinorum Stingelin, 1900 in postabdomen
morphology, but status unclear. See also A. sarasinorum Stingelin, 1900.
sculpta. Described as Alonella sculpta Sars, 1901 from Ipiranga, Brazil (Sars 1901). Lectotype and
paralectotypes in GOS. Mentioned as Alona sculpta in Rajapaksa & Fernando (1987: 132), but it is likely a
type error, further as Alonella sculpta. Synonymized by the same authors and now falls under Notoalona
globulosa (Daday, 1898). See also A. globulosa Daday, 1898.
serrata. Described as Alona rectangula serrata Daday, 1908 from Kotakota, Zambia, Africa (Daday
1908). Types lost, not in DAD (Forr & Frey 1982). Variety listed in Smirnov (1971) as one of seven
rectangula subspecies. From the original description, it is unclear what this taxon is. Five to six denticles on
the posteroventral corner and shape of postabdomen suggests that it may belong to Coronatella or even
Karualona, therefore not Alona. May be related to A. monacantha Sars, 1901 (Megard 1967). See also A.
hardingi Brehm, 1957, A. monacantha Sars, 1901 and A. rectangula Sars, 1861.
setigera. Described as Alona guttata setigera Brehm, 1931 from New Zealand (Brehm 1931). No
information on types. Distributed in Australia, New Zealand (Sinev 1999b) and Oceania (Schabetsberger et
al. 2009). Subspecies of A. guttata in Smirnov (1971) and placed in Biapertura by Smirnov & Timms (1983),
but does not belong to either. Revised by Sinev (1999b). Valid, the only species of the Alona costata-complex
with two main head pores. See also A. costata Sars, 1862.
setosocaudata. Described as Alona setosocaudata Vasiljeva & Smirnov, 1969 from Lake Baikal, Asian
Russia (Vasiljeva & Smirnov 1969). Holotype and allotype in MGU, paratypes in ZIN and BGI. Valid species
that seems closest to Alona quadrangularis-group, details, including limbs, in Sinev & Kotov (2001). See also
A. labrosa Vasiljeva & Smirnov, 1969 and A. quadrangularis (O. F. Mller, 1776).
setuloides. Described as Alona setuloides Smirnov & Timms, 1983 from Western Australia (Smirnov &
Timms 1983). Holotype in AM, paratypes in MGU, ZIN and NHM. Valid species, close to the American A.
setulosa Megard, 1967 (Smirnov & Timms 1983) but fine morphology unknown. Three separated head pores
and postabdomen refer to the A. pulchella-group. See also A. pulchella King, 1853 and A. setulosa Megard,
1967.
setulosa. Described as Alona setulosa Megard, 1967 from New Mexico, USA (Megard 1967). Holotype
in NHM. Valid species of the A. pulchella-complex, redescribed by Sinev (2009b). See also A. borealis
Chengalath & Hann, 1981, A. lapidicola Chengalath & Hann, 1981 and A. pulchella King, 1853.
shiklejewi Proposed as Alona protzi Shiklejewi Brehm, 1933 from Caucasus (Brehm 1933a), without
description. No information on types. Brehm (1933a: 718) assigned a name to the A. protzi described by
Shiklejew (1930) from the Caucasus. This form should be Phreatalona protzi (Hartwig, 1900) (Van Damme et
al. 2009). The main diagnostic character, presence of lateral fascicles on postabdomen, is present in true P.
protzi as well. See also A. protzi Hartwig, 1900 and A. protzi typica Behning, 1941.
siamensis. Described as Alona siamensis Sinev & Sanoamuang, 2007 from Thailand (Sinev &
Sanoamuang 2007). Holotype and paratypes in MGU. Distributed in South East Asia, related to Neotropical
Alona dentifera Sars, 1901. Corresponds with A. cf. dentifera from Malaysia reported in Idris (1983) (Sinev &
Sanoamuang 2007). A. dentifera and its sibling do not belong to Alona but are part of the Coronatella-branch
and still need to be removed (Van Damme & Dumont 2008b). See also A. dentifera (Sars, 1901).
similis. Described as Coronatella similis Dybowski & Grochowski, 1898 from the area of Lwow and
Lake Svityaz, Ukraine (Dybowski & Grochowski 1895). No information on types. Originally a nomen nudum
(Dybowski & Grochowski 1895) but described later (Dybowski & Grochowski 1898). Incertae sedis.
simoneae. Described as Alona simonei Dumont, 1981 from Guinea, W. Africa (Dumont 1981). Holotype
in MMA, paratypes in SUG. Examination of paratypes showed that this is a valid species, with six limbs.
41
Peculiar morphology suggests a separate genus Matralona Van Damme & Dumont 2009. The name was
amended to simoneae and is now Matralona simoneae (Dumont, 1981). Shows close affinities with
Armatalona Sinev, 2004 (Van Damme & Dumont 2009). See also A. freyi Idris & Fernando, 1981, A.
intermedia Sars, 1862 and A. macrocopa Sars, 1894.
sketi. Described as Alona sketi Brancelj, 1992 from Slovenia, Dinaric region (Brancelj 1992). Types in
NHM and author's collection (IBUL), syntypes at SUG. Stygobiont. Valid species, now revised and removed
from Alona in a new genus (Van Damme & Sinev, unpubl.). See also A. hercegovinae Brancelj, 1990 and A.
stochi Brancelj, 1992.
smirnovi. Described as Alona smirnovi Petkovski & Flssner, 1972 from Lake Ohrid, Macedonia
(Petkovski & Flssner 1972). Holotype and paratypes in ZMHU. Moved to the (hypo)rheic genus
Phreatalona Van Damme, Brancelj & Dumont, 2009, now Phreatalona smirnovi (Petkovski & Flssner,
1972). See Van Damme et al. (2009). See also A. labrosa Vasiljeva & Smirnov, 1969, A. phreatica Dumont,
1983 and A. protzi Hartwig, 1900.
socors. Described as Lynceus socors O.F. Mller, 1776 from Denmark (Mller 1776). Appears as Alona
socors (O.F. Mller, 1776), for example in Schoedler (1863). Types lost. Incertae sedis (Kurz 1875, Smirnov
1971).
spinifera. Recorded as nomen nudum (Schoedler 1858) and later described as Alona spinifera Schoedler,
1862 from Spree (Treptow) and Berlin-Spandauer Channel, Berlin, Germany (Schoedler 1862). Types from
Berlin-Spandauer Channel (or Charit-Graben as in the original publication) still present in ZMHU (Forr
1983). Junior synonym of Alona affinis according to Smirnov (1971; Biapertura). Schoedler (1863) and Kurz
(1875) remark two points in which this form differs from true A. affinis, 1. smaller body (0.4 mm) and a 2.
lower head. Status unclear, type material should be examined. See A. affinis (Leydig, 1860).
spinulosa. Described as Alona guttata var. spinulosa Linko, 1900 from Solovietski Islands in the White
Sea, European Russia (Linko 1900). No information on types. Status unknown, originally listed as a variety
and raised to subspecies by Smirnov (1971). The A. guttata group needs revision and contains several species
worldwide. See also A. bergi Ren, 1992, A. guttata Sars, 1862 and A. werestschagini Sinev, 1999.
stagnalis. Described as Alona stagnalis Daday, 1888 from Hungary (Daday 1888). Types lost (Frey
1965a). May be an older and valid name for A. rustica Scott, 1895 but impossible to confirm due to lack of
type material and poor description (Forr & Frey 1982, Frey 1965a). Alonso (1996) lists it as synonym of
Alona rectangula Sars, 1861. Incertae sedis in Smirnov (1971). See also two other uncertain Hungarian
species by Daday (1888), A. lacustris Daday, 1888 and A. balatonica Daday, 1888.
stochi. Described as Alona stochi Brancelj, 1997 from Slovenia, Dinaric Region (Brancelj 1997).
Holotype and paratypes in authors collection at IBUL, other paratypes in NHM. Additional material from
type locality in SUG. Valid species, revised and moved from Alona into a new genus (Van Damme & Sinev,
unpubl.). See also A. hercegovinae Brancelj, 1990 and A. sketi Brancelj, 1992.
striolata. Described as Alona striolata Sars, 1916 from Green Point Common, Cape Town, Republic of
South Africa (Sars 1916). Material from type locality in GOS. Status unclear, the taxon was never revised.
May be valid a member of the A. elegans-complex (Sars 1916), but far outside the Western Palaearctic
distribution of the latter (Van Damme & Dumont 2008b). Only reported once in South Africa (Smirnov 2008);
also a record from Australia should be checked (Shiel & Dickson 1995). The A. elegans-group remains
otherwise unrecorded from both regions. See also A. arcuata Sars, 1916, A. elegans Kurz, 1875 and A.
harpularia Sars, 1916.
subantarctica. Described as Alona bukobensis var. subantarctica Ekman, 1905 from South Georgia and
Falklands, Subantarctic (Ekman 1905). Material in Ekmans collection, SMNH (Frey 1988, Kotov &
Gololobova 2005). Raised to species level by Sars (1909), who called it A. subantarctica Ekman, 1905. This
taxon has no direct relation with A. bukobensis (now Coronatella bukobensis of the C. rectangula-species
group). A. subantarctica was synonymized with A. weinecki Studer, 1878 by Rhe (1914) and Frey (1988)
who studied Ekmans (1905) material. Finally, both these taxa were moved from Alona to Ovalona by Van
Damme & Dumont (2008b) with redescription of limb morphology and notes on distribution. Ovalona
VAN DAMME ET AL.
subantarctica (Ekman, 1905), as it is now called, may be a valid (sub)species and not just a synonym of O.
weinecki, which could be limited to the eastern subantarctic islands (See Frey 1988, Van Damme & Dumont
2008b). See also A. weinecki Studer, 1878.
sulcata. Recorded as Alona fulcata Schoedler, 1858 (nomen nudum and apparent misspelling) and later
described as Alona sulcata Schoedler, 1862 from Spree, Pltzensee and Berlin-Spandauer Channel, Berlin,
Germany (Schoedler 1862). Tube containing A. sulcata in ZMHU can be regarded as type material (Forr
1983). Junior synonym of Alona quadrangularis (O. F. Mller, 1776) in Kurz (1875), Smirnov (1971).
switeziana. Described as Alona witeziana Dybowski & Grochowski, 1895 from Lake Svityaz, Ukraine
(Dybowski & Grochowski 1895). No information on types. No drawings in original description. Nomen
nudum (Smirnov 1971).
taraporevalae. Described as Alona taraporevalae Shirgur & Naik, 1977 from Back Bay, Bombay, India
(at salinities of 26) (Shirgur & Naik 1977). Holotype and paratypes in ZSI, one paratype in DGF. Revision
of material of A. taraporevalae Shirgur & Naik, 1977 from DGF (Frey 1991) and specimens of A.
sarasinorum Stingelin, 1900 from Thailand (Supiyanit 2005) confirm synonymy with A. sarasinorum and the
latter as a valid species. Affinities of A. sarasinorum unclear, but definitely not Alona. Drawings, no limbs, in
Frey (1991) who compared this animal with Leberis (former A. diaphana-group), but with better knowledge
of the latter, such affinities are uncertain. See also A. sarasinorum Stingelin, 1900 and A. diaphana King,
1853.
tenuicaudis. Described as Alona tenuicaudis Sars, 1862 from Norway (Sars 1862). Types in GOS.
Removed from Alona by Dybowski & Grochowski (1894) to Oxyurella Dybowski & Grochowski, 1894 and
type species of the genus. Valid species, now Oxyurella tenuicaudis (Sars, 1862).
tenuirostris. Described as Alona tenuirostris Hellich, 1874 from Czech Republic (Hellich 1874). No
information on types. Junior synonym of Kurzia latissima (Kurz, 1875). See Hudec (2000) and Kotov (2004)
for revision of the genus Kurzia.
testudinaria (nomen protectum). See reticulata for discussion of priority. Described as Lynceus
testudinarius Fischer, 1848 from the vicinity of St. Petersburg, European Russia (Fischer 1848). Neotype in
IKZ. Hellich (1877) regarded the species as an Alona. It belongs to Graptoleberis Sars, 1862 (see Sars 1862,
1993), now regarded as Graptoleberis testudinaria (Fischer, 1848), valid protected name. Graptoleberis has
several subspecies worldwide, listed by Smirnov (1971). A contemporary revision is lacking. See Lilljeborg
(1900) and Smirnov (1971). See also A. esocirostris Schoedler, 1858 and A. reticulata Baird, 1843.
titi. Described as Alonopsis titi Harding, 1955 from Lake Titicaca, Peru (Harding 1955). Holotype in
NHM. Smirnov (pers, comm.) suggests that this taxon may be closer to Alona than to Alonopsis. Listed as
Acroperus titi (Harding, 1955) in Smirnov (1971). Status unclear, needs to be redescribed.
tovelana. Described as Alona affinis f. tovelana Baldi, 1941 from Lago di Tovel, Italy (Baldi 1941). No
information on types. Identical to true A. affinis (Leydig, 1860) according to Smirnov (1971). See also A.
affinis (Leydig, 1860).
trachystriata. Described as Alona rectangula trachystriata Chen, Zhang & Liu, 1993 from Guixi County,
Jiangxi Province, China (Chen et al., 1993). Types in IHW. Status unclear, species needs revision. Not Alona,
but Coronatella. Related to rectangula and may be a separate species or subspecies of this complex,
Coronatella rectangula trachystriata (Chen, Zhang & Liu 1993). See A. rectangula Sars, 1861.
transversa. Described as Pleuroxus transversus Schoedler, 1862 from vicinities of Berlin (Schoedler
1862). Mentioned as Alona transversa (Schoedler, 1862) by P.E. Mller (1897), see Lilljeborg (1901). Types
unknown. Belongs to a different subfamily, most likely a junior synonym of Alonella nana (Baird, 1850)
(Lilljeborg 1901). See also A. pygmaea Sars, 1862.
tridentata. Described as Alona acuticostata var. tridentata Stingelin, 1905 from Thailand (Stingelin
1905). Types in NMO (Frenzel 1987). Status unclear. Regarded by Smirnov (1971) as a subspecies of A.
monacantha Sars, 1901. Indeed, A. acuticostata Sars, 1903 and A. monacantha Sars, 1901 are related, but the
latter is restricted to the Neotropics (Sinev 2004a) whereas A. acuticostata Sars, 1903 was described from
Sumatra (and needs revision). A. acuticostata var. tridentata Stingelin, 1905 is likely a form of A. acuticostata
43
Sars, 1903 with three denticles in the posteroventral corner of the valves, a variable trait. Three denticles may
occur, though rarely, in A. monacantha specimens (Van Damme, unpubl.) Authors synonymized tridentata
with monacantha (Harding 1957, Megard 1967). The A. monacantha group is not Alona but most likely
Coronatella (see Van Damme & Dumont 2008b), combined with the A. rectangula group. See also A.
acuticostata Sars, 1903, A. hardingi Brehm, 1957, A. monacantha Sars, 1901 and A. rectangula serrata
Daday, 1898.
truncata. Described as Alona truncata Smirnov, 1989 from Queensland, Australia (Smirnov 1989).
Holotype and paratype in AM. Likely a valid species, with short postabdomen and long basal claw, but
affinities of this Australian taxon unclear.
tuberculata. Described as Alona tuberculata Kurz, 1875 from Seitenthal-Deutschbrod, Ssava Stream,
Bohemia, Czech Republic (Kurz 1875). No information on types. Kurz (1875) designated several tuberculated
ecomorphs as separate taxa. Valve tubercles as main character invalid for (sub)species, this is variable within
populations. What induces the carapace ornamentation is unknown (may have an underlying ecological
trigger). The form tuberculata should be used as variety, as done in palaeolimnological works (e.g.,
Szeroczynska & Sarmaja-Korjonen 2007). It was regarded as a synonym of Alona guttata Sars, 1862 (Brehm
1933a, Alonso 1996) or subspecies in (Smirnov 1971). Kurzs (1875) form must be regarded as no more than
a tuberculate variety of A. guttata. See A. guttata Sars, 1862.
tuberculata-2 (suppressed). Described as Alona tuberculata Hudendorff, 1876 from Ryazan Area,
European Russia (Hudendorff 1876). No information on types. Description reveals little except valve
tubercles in longitudinal rows and a female postabdomen as in A. inornata Hudendorff, 1876; figure shows
only male postabdomen. Like A. inornata, A. tuberculata is thought identical to A. rectangula Sars, 1861
(Lilljeborg 1901, as Lynceus). It may be another name for tuberculate forms of C. rectangula, like A. pulchra
Hellich, 1874. In any case, A. tuberculata Hudendorff, 1876 is a junior homonym of A. tuberculata Kurz,
1875. The former is a suppressed name (ICZN 1958, 2009) placed in the "Official Index of Rejected and
Invalid Species Names in Zoology". See also A. inornata Hudendorff, 1876 and A. rectangula Sars, 1861.
tuberculata-3. Described as Alona glacialis var. tuberculata Herrick, 1884 from Minnesota, USA
(Herrick 1884). No information on types. Incertae sedis (Smirnov 1971). Junior honomym of A. tuberculata
Kurz, 1875. See also A. glacialis Birge, 1879
tuberculata-4. Described as Alona estonica tuberculata Memets, 1958 from Estonia (Memets 1958).
Tuberculate form of Alona rustica Scott, 1895 (Frey, 1965a). Junior homonym of A. tuberculata Kurz, 1875.
See A. rustica Scott, 1895.
tuberculata-5. Described as Alona rustica tuberculata Mamets, 1958 from Estonia (Memets 1958).
Tuberculate form of A. rustica (Frey 1965a). Junior homonym of A. tuberculata Kurz, 1875. See A. rustica
Scott, 1895.
typica. Described as Alona protzi typica in Behning, 1941 from the Caucasus (Behning 1941). No
information on types. Most probably a junior synonym of Phreatalona protzi (Hartwig, 1900). Although
Behning (1941) may not have meant to create a special taxon using the epithethon typica, it is a trinomen. In
any case, it is a junior synonym, as Brehm (1933a) already assigned a name to A. protzi form from the
Caucasus, Alona protzi shiklejewi Brehm, 1933. The unfortunate tradition of using of typica (similarly with
tuberculata) was continued by rmek-Huek et al. (1962) for Alona guttata var. typica, listed hereunder.
See also A. protzi Hartwig, 1900 and A. protzi shiklejewi Brehm, 1933.
typica-2. Described as Alona guttata var. typica rmek-Huek, Stracraba & Brtek, 1962 from former
Czechoslovakia (rmek-Huek et al. 1962). Junior homonym of Alona protzi typica in Behning (1941). See
also A. guttata Sars, 1862.
unguiculata. Described as Alona unguiculata Smirnov, 1989 from Queensland, Australia (Smirnov
1989). Holotype in AM, paratype in MGU. Affinities unknown, postabdominal shape and armature suggest
this may be related to (or a member of) the A. pulchella-complex. May be a valid species, but presently
unresolved. Two revised Australian species of this group are A. pulchella King, 1853 and A. archeri Sars,
1888. See also A. beverleyae Smirnov, 1989 and A. laevissima Sars, 1888
VAN DAMME ET AL.
vermiculata. Described as Alona davidi vermiculata Smirnov & Timms, 1983 from New South Wales,
Australia (Smirnov & Timms 1983). Types in AM, MGU, ZIN. Form with fine striation. Detailed morphology
unknown. Not Alona, but a member of Leberis Smirnov, 1989, maybe a species related to Leberis diaphanus
(King, 1853), or a variety or subspecies of the latter. See A. davidi Richard, 1895 and A. diaphana King, 1853.
verrucosa-2 (suppressed). Recorded as Alona verrucosa Lutz, 1879 from Switzerland as nomen nudum
(Smirnov 1971). Suppressed name (Johnson 1956b, ICZN 1958, 2009) placed in the "Official Index of
Rejected and Invalid Species Names in Zoology". See also A. verrucosa Sars, 1901.
verrucosa. Described as Alona verrucosa Sars, 1901 from So Paulo, Brazil (Sars 1901). Lectotype and
paralectotypes assigned by Valdivia-Villar, in GOS. A. verrucosa Sars, 1901 is in fact a primary junior
homonym of A. verrucosa Lutz, 1878, but the former was retained for stability (Johnson 1956b). Records
worldwide, but true verrucosa is likely limited to Neotropics. Redescription of A. verrucosa Sars, 1901 from
Brazil, including limbs, in Sinev & Hollwedel (2002). The verrucosa-complex does not belong in Alona and
sibling species are described in an upcoming revision of this group; the group will be moved into a new genus.
Smirnov (1971) moved this group to Biapertura under the name B. pseudoverrucosa Smirnov, 1971. See also
A. alonopsiformis Brehm, 1933, A. anodonta Daday, 1905, A. brandorffi Sinev & Hollwedel, 2002, A.
mediterranea Yalim, 2005, A. pectinata Elias-Gutierrez & Suarez-Morales, 1999, Alona verrucosa lineolata
Chen & Li, 1991 and Biapertura pseudoverrucosa Smirnov, 1971.
vietnamica. Described as Alona affinis vietnamica Dang Ngoc Thang, 1980 from Vietnam (Dang Ngoc
Thang et al. 1980). No information on types. The taxon is described poorly, but this is apparently a member of
affinis-group and could be a junior synonym of A. affinis (Leydig, 1860). See Alona affinis (Leydig, 1860).
virago. Described as Alona virago Brehm, 1935 from China (Brehm 1935). No information on types.
Drawings in Brehm (1935), Chiang & Du (1979) and Smirnov (1971) suggest this taxon belongs to the Alona
costata-group, closest to Alona rustica Scott, 1895. Status and details of this Brehmian species are unknown
and no material exists. See also A. cheni Sinev, 1999, A. costata Sars, 1862 and A. rustica Scott, 1895.
wallaciana. Described as Alona wallaciana Henry, 1919 from New South Wales, Australia (Henry 1919).
Holotype in AM. Not an Alona, allocated to Oxyurella by Smirnov (1971), now named Oxyurella wallaciana
(Henry, 1919). See also A. tenuicaudis Sars, 1862.
weinecki. Described as Alona weinecki Studer, 1878 from Kerguelen Island, Subantarctic (Studer 1878).
Valid species from Subantarctic Islands. Types lost, but material of A. weinecki from the Kerguelen from the
collection of Rhe (1914) at ZMHU might be useful to assign neotypes. Revised by Frey (1988) who
compared it with C. rectangula (Sars, 1861). Van Damme & Dumont (2008b) showed that this is not an Alona
and allocated it, together with A. meridionalis Sinev, 2007, to Ovalona (Coronatella-branch), including
description of the limbs. Now Ovalona weinecki (Studer, 1878). Morphological description of populations
from Easter Island (Dumont & Martens 1996) corresponds to member of the A. pulchella-complex, not to O.
weinecki (see Van Damme & Dumont 2008b). Populations on the Maritime Antarctic and Sub-Antarctic
islands may comprise two (sub)species, weinecki in the East and subantarctica in the West (Van Damme &
Dumont 2008b). See also A. bukobensis Weltner, 1897, A. meridionalis Sinev, 2007, A. novae-zealandiae
Sars, 1904 and A. subantarctica Ekman, 1908.
weltneri. Described as Alona weltneri Keilhack, 1905 from Madusee (Jezioro Miedwie), Pomerania,
Poland (Keilhack 1905). Holotype in ZMHU. Distributed in Western Palaearctic (Keilhack 1905, 1908,
Scourfield 1907, Flssner 1962, 2000, Van Damme et al., in press). Valid species of the Alona costata-group
but rarely recorded and never in high numbers. We examined the holotype and confirm this as a peculiar taxon
(Van Damme et al., in press). Together with A. karelica Stenroos, 1897, A. weltneri is one of the rarest
European Alona. Very close to European A. costata Sars, 1862, but postabdomen of A. weltneri has a round
dorsodistal end, not acute as in A. costata. Drawings in Scourfield (1907) and Flner (1962, 2000), among
the few authors ever to have seen this species, but no limbs were described. See also A. costata Sars, 1862.
werestschagini. Described as Alona werestschagini Sinev, 1999 from Kirghizia (type locality) and Russia
(Sinev 1999b). Holotype and paratypes in MGU. Valid member of the A. guttata-complex, limb characters
described in Sinev (1999b). Wide but patchy distribution in Eurasia. Found in Scandinavia, where it is
45
considered a postglacial relict; may be sympatric with A. guttata Sars, 1862 (Sarmaja-Korjonen & Sinev
2008). See also A. clathrata Sars, 1888, A. crassicauda Sars, 1916 and A. guttata Sars, 1862.
whiteleggei. Described as Alona whiteleggii Sars, 1896 from Centennial Park, Sydney, New South Wales,
Australia (Sars 1896). Sars (1896) used two spellings in the same paper (whiteleggii and whiteleggei); we
amend it here to A. whiteleggei Sars, 1896 because the taxon was named after Mr. Whitelegge and because
different spellings occur in the original description. No types in GOS. Status unclear. According to Sars
(1896), intermediate between A. quadrangularis (O. F. Mller, 1776) and A. oblonga P. E. Mller, 1868 (= A.
affinis), but with a postabdomen most like in the latter. Suggested as a synonym of A. affinis (Leydig, 1860)
(Smirnov 1971, Shiel & Dickson 1995; Biapertura), but, considering that true Alona affinis does not occur in
Australia (Sinev 1997), synonymy is unlikely. Drawings of Sars (1896) do not allow a decision on its status at
the species level. Most likely, it is indeed an Australian species of the A. affinis group, to which we think the
postabdomen resembles strongly. The more common name for Australian populations of the A. affinis-group,
A. kendallensis Henry, 1922, might even be a junior synonym of A. whiteleggei Sars, 1896. See also A. affinis
(Leydig, 1860), A. longirostris Henry, 1922, A. kendallensis Henry, 1922, and A. quadrangularis (O. F.
Mller, 1776).
willisi. Described as Biapertura willisi Smirnov, 1989 from Queensland, Australia (Smirnov 1989).
Holotype in MGU. Species with two head pores and peculiar postabdomen with long lateral spines. This is
undoubtedly a valid species with special morphology, but its affinities remain unstudied. May be closely
related to the Australian Alona rigidicaudis (Smirnov 1971; as Biapertura), which also has two main pores
and a short postabdomen. See also A. rigidicaudis (Smirnov, 1971).
Original varieties and subspecies in Alona Baird, 1843 (trinomini)

africana Daday, 1908. See A. rectangula Sars, 1861.
americana Flner & Frey, 1970. See A. rustica Scott, 1895.
angulata Weigold, 1911. Homonym of A. angulata Birge, 1879. See also A. rectangula Sars, 1861.
arcuata Herbst, 1964. See A. elegans Kurz, 1875.
barbata Brehm, 1935. See A. affinis barbata Brehm, 1935.
bukobensis Weltner, 1897. See A. bukobensis Weltner, 1897.
caucasica Shiklejew, 1933. See A. coronata Kurz, 1875 and A. rectangula Sars, 1861.
ceylonica Daday, 1898. See A. australis var. ceylonica Daday, 1898.
elegantula Brehm, 1933. See A. affinis (Leydig, 1860).
herbsti Smirnov, 2001. See A. elegans Kurz, 1875.
iheringi Richard, 1897. See A. davidi Richard, 1895.
laevis Herrick, 1884. See A. glacialis Birge, 1879.
lineolata Chen, Zhang, Yi & Guo, 1991. See under A. verrucosa Sars, 1901.
minor Stingelin, 1905. See A. intermedia Sars, 1862.
nalibokiana Dybowski & Grochowski, 1894. See A. inornata Hudendorff, 1876.
novae-zealandiae Sars, 1904. See A. rectangula Sars, 1861.
ornata Shiklejew, 1929. See A. rectangula Sars, 1861.
ornata Stingelin, 1895. See A. affinis (Leydig, 1860).
parvula Kurz, 1875. See A. guttata Sars, 1862.
patagonica Ekman, 1900. See A. cambouei de Guerne & Richard, 1893.
pseudopulchra Smirnov, 1968. See A. rectangula Sars, 1861.
pulchra Hellich, 1874. See A. rectangula Sars, 1861.
punctata Daday, 1898. See A. diaphana King, 1853.
reticulata Hartmann, 1917. See A. guttata Sars, 1862.
richardi Stingelin, 1895. See A. rectangula Sars, 1861.
VAN DAMME ET AL.
serrata Daday, 1898. See A. rectangula Sars, 1861.

setigera Brehm, 1931. See A. setigera Brehm, 1931.
shiklejewi Brehm, 1933. See A. protzi Hartwig, 1900.
spinulosa Linko, 1900. See A. guttata Sars, 1862.
subantarctica Ekman, 1908. See A. bukobensis Weltner, 1897, A. subantarctica Ekman, 1908.
trachystriata Chen, Zhang & Liu, 1993. See A. rectangula Sars, 1861.
tovelana Baldi, 1941. See A. affinis (Leydig, 1860).
tridentata Stingelin, 1905. See A. acuticostata Sars, 1903
tuberculata Herrick, 1884. See A. glacialis Birge, 1879.
tuberculata Kurz, 1875. See A. guttata Sars, 1862.
tuberculata Memets, 1958. See A. estonica Memets, 1958 and A. rustica Scott, 1895.
vermiculata Smirnov & Timms, 1983. See A. davidi Richard, 1895.
weltneri Keilhack, 1905. See A. rectangula Sars, 1861 and A. weltneri Keilhack, 1905.
Valid genera separated from Alona sensu lato since 1843

Alonopsis Sars, 1862. Type species: Alona elongata Sars, 1861. Separated by Sars (1862). Holarctic.
Armatalona Sinev, 2004. Type species: Alona macrocopa Sars, 1894. Defined by Sinev (2004b). Australian
genus with two valid species (one record from Thailand).
Coronatella Dybowski & Grochowski, 1894. Type species: Alona coronata Kurz, 1875. Contains A.
rectangula Sars, 1861, A. monacantha Sars, 1901 and related species. Not used since 1894, reinstated by
Van Damme & Dumont (2008b) for allocation of the A. rectangula-group. Cosmopolitan.
*Dadaya Sars, 1901. Type species: Alona macrops Daday, 1898. Created by Sars (1901). Pantropical. *Genus
of the Chydorinae, not Aloninae.
Graptoleberis Sars, 1862. Type species: Alona reticulata Baird, 1843 (nomen oblitum), a synonym of Lynceus
testudinarius Fischer, 1848 (nomen protectum). Well-established genus by Sars (1862), but unrevised
according to recent standards. Cosmopolitan.
Karualona Dumont & Silva-Briano, 2000. Type species: Alona karua King, 1853. Pantropical (Mediterranean).
Leydigia Kurz, 1875. Type species Alona leydigii Schoedler, 1863 (originally L. quadrangularis O. F. Mller,
1776). Large genus, revised and subdivided by Kotov (2009). Cosmopolitan.
Maraura Sinev & Shiel, 2008. Type species: Alona macracantha Smirnov & Timms, 1983. Monotypic (Sinev
& Shiel 2008). Australian.
Matralona Van Damme & Dumont, 2009. Type species: Alona simonei Dumont, 1981. Monotypic, peculiar
genus (Van Damme & Dumont 2009). West Africa (Guinea).
Notoalona Rajapaksa & Fernando, 1987. Type species: Alona globulosa Daday, 1898. Few species, described
in detail (Rajapaksa & Fernando 1987). Pantropics.
Oxyurella Dybowski & Grochowski, 1894. Type species: Alona tenuicaudis Sars, 1862. Well delineated
genus, but unrevised. Cosmopolitan.
Ovalona Van Damme & Dumont, 2008. Type species: Alona weinecki Studer, 1878. Two valid species,
discussion on a third (Van Damme & Dumont 2008b). Subantarctic and the Cape.
Parvalona Van Damme, Kotov & Dumont, 2005. Type species: Alona parva Daday, 1905. Monotypic.
Peculiar genus from South America (Van Damme et al. 2005).
Phreatalona Van Damme, Brancelj & Dumont, 2009. Type species: Alona protzi Hartwig, 1900. Four valid
species with preference for river interstitial (Van Damme et al. 2009). Western Europe and Baikal.
Rhynchotalona Norman, 1903. Type species: Alona falcata Sars, 1861. Palaearctic genus with currently two
valid species.
47
Under study:
Biapertura Smirnov, 1971. Type species: Alona affinis (Leydig, 1860). As defined in Smirnov (1971), the
genus was dismantled. Its core, the A. affinis-group, may be a valid (sub)genus and is under study.
Gen. Nov. 1 Van Damme, Sinev & Dumont. Type species: Alona verrucosa Sars, 1901. Contains A. verrucosa
and related species, manuscript yet unpublished by Van Damme, Sinev and Dumont. Pantropical.
Gen. Nov. 2 Van Damme & Sinev. Type species: Alona hercegovinae Brancelj, 1997. Contains three species,
all stygobionts, separated in yet unpublished manuscript by Van Damme & Sinev. Dinaric Region
(Europe).
Existing genera to which Alona species or groups are allocated

Kurzia Dybowski & Grochowski, 1894. Type species: Alonopsis latissima Kurz, 1875. Well-established
genus by Dybowski & Grochowski (1894). Revisions and subgeneric division in Hudec (2000) and Kotov
(2004). Cosmopolitan.
Leberis Smirnov, 1989. Type species: Leberis aenigmatosus Smirnov, 1989. Contains the "Alona" diaphanagroup (Sinev et al. 2005). Pantropical (-Mediterranean).
Nicsmirnovius Chiambeng & Dumont, 1999. Type species: Niscmirnovius camerounensis Chiambeng &
Dumont, 1999. Contains the A. eximia-group (Van Damme et al. 2003). Pantropical.
Tretocephala Frey, 1965. Type species: Alonopsis ambigua Lilljeborg, 1901. Two species with disjunct
distribution, Palaearctic-South Africa.
Unavailable names
Listed in collections, unpublished names are mentioned here in a negative sense (Case 11.5.2. of ICZN, 1999),
to avoid confusion, e.g., in case of future descriptions with similar epitheta.
Alona anastata Unpublished provisory name by unknown author (P.E. Mller?). Types in ZMUC.
Alona indica Daday Unpublished provisory name. Types in DAD.
Alona monodonta Sars. Unpublished provisory name. Types in GOS.
Alona sinostata Sars Unpublished provisory name. Types in GOS.
Alonella karua var. dentifera Daday. Unpublished provisory name. Types in DAD.
Alona tessellata Sars Unpublished provisory name for A. guttata Sars, 1862 (see Sars, 1993, Plate 99. No
types.)
Discussion
Splitting of Alona Baird, 1843
Alona Baird, 1843 underwent a serious reshuffling of taxa. Such shifts may be confusing for workers
outside Chydoridae taxonomy, but splitting of the genus is not a new phenomenon. Several genera were
separated early from Alona and remained stable since, such as Oxyurella and Graptoleberis. In total, 14
genera of the Aloninae were split from Alona (see above), about half of the total now recognized in the
subfamily. Four other genera have a type species that is not Alona, but now contain taxa that resided in it (e.g.,
Leberis, Nicsmirnovius). Three additional assemblages are under study (A. affinis-, A. verrucosa- and A.
hercegovinae-groups). Half of the genera derived from Alona were described since 2000 and result from
increased efforts in morphological revisions, which sped up the splitting. A large number of characters are
VAN DAMME ET AL.
now studied in detail. We are reaching a level of understanding in systematics that may be more realistic in
terms of diversity yet taxonomic impediments are many and time-consuming to overcome. Many taxa are still
waiting for description, the number of total species known in Cladocera now only half to a fourth of the
expected number (Forr et al., 2008). This is definitely the case for Alona-like species, and the above list
shows that many names are available for species outside Europe. More revisions are in progress, but future
disentanglement of the genus should progress with care. Genera need to be well defined and their separation
from Alona and position relative to others well argumented. For example, the position of the A. dentifera- and
A. elegans-groups still needs to be figured out, both are definitely not Alona and belong with or stand at least
close to Coronatella (see Van Damme & Dumont 2008b). In ideal cases, groups can be recognized that have a
particular morphology as well as ecology and where evolution took place within (e.g., Phreatalona Van
Damme et al. 2009).
Avoiding confusion by tuberculate morphs
Tubercles on the valves and head shield are an adaptation likely correlated with benthic life, with degree
of expression varying among individuals within single populations (Sinev 2001b, Kotov 2006). The ability to
express tubercles seems to be most common in the A. verrucosa- and A. guttata groups and in Coronatella.
The tradition of considering such tubercles as important characters has lead to confusion. They occur in both
Chydorinae and Aloninae. Morphs with dimples or tubercles show no additional characters that separate them
from smooth forms (e.g., caelatus form of Chydorus sphaericus; see Frey 1980), yet emphasis on this easily
visible character has lead to splitting. From the list above, we see that Alona rectangula, now Coronatella
rectangula, even has two names for tuberculate forms, A. pulchra Hellich, 1874 and A. tuberculata
Hudendorff, 1876. The trait is variable (Sinev 2001b). Smirnov (1971) created two subspecies for A.
verrucosa, Biapertura pseudoverrucosa pseudoverrucosa and B. pseudoverrucosa verrucosa, based on the
absence or presence of dimples. Kurz (1875) suggested a new taxon, A. tuberculata Kurz, 1875, considered a
name for tuberculate A. guttata Sars, 1862. The latter is even more confusing, as the original specimens
described by Sars (1862) were tuberculated. The name tuberculata occurs not less than five times in our list.
The tradition has also caused erroneous synonymisations. For example, Johnson (1956b) requested the ICZN
to suppress A. pulchra Hellich, 1874 (= Coronatella rectangula), because he believed it to be synonymous to
A. verrucosa Sars, 1901 based on the presence of tubercles. The request was accepted (ICZN 1958, 2009) but
the argumentation was incorrect: both taxa are not identical and even belong to two different genera now.
The exact factors that cause valve ornamention are unknown, and until we do, it may remain useful to
record tuberculate versus smooth morphs. For Coronatella rectangula, we suggest to use "forma pulchra",
and for A. guttata and A. rustica, "forma tuberculata". Tubercles are not limited to the former taxa; we found
them also in specimens of Leberis sp. from Sicily (coll. by F. Marrone) and in A. azorica from Spain (coll.
M.R. Miracle).
Prospects in Alona
Desiderata for future studies in the Aloninae and in Chydoridae in general, are:
(1) A revision of existing collections and type material of early authors (e.g., GOS, DAD) and
investigating unexplored type material, found in the collections of J. Richard and E.A. Birge (DGF), W. Baird
(NHM) and P.E. Mller (ZMUC). Specimens that belonged to G.Y. Werestchagin, A. Linko and A.L. Behning
are present in ZIN, but uncatalogued. A small part of the collection of V. Brehm was recovered and will be
deposited at NHM (V. Ko nek, pers. comm.). A few collections need thorough evaluation of the type
specimens or even a check if material is still present (e.g., IHW).
(2) If type material is lost, collection and revision of material from the terra typica are mandatory.
Lectotypes or neotypes have not been assigned for a number of species.
(3) Regional studies remain important towards a better alonine taxonomy and diversity. There is a strong
49
discrepancy between regions. Mexico moved in only a decade from one of the poorest to one of the bestknown countries in its assessment of freshwater zooplankton (e.g., Elas-Gutirrez et al. 2006, 2008).
Smirnov (2008) prepared an updated checklist for the Republic of South Africa. Recent checklists for
Australia exist (Smirnov & Timms 1983, Shiel & Dickson 1995), but many Australian taxa still have
uncertain affinities and need revision (e.g., A. willisi). Also, many checklists still contain taxa bearing
European names while they probably belong to local species for which names exist, e.g., A. clathrata for the
Australian A. guttata. Taxonomical updates should be prepared diligently, as for Mexico and South Africa.
This is not always the case. Sarma & Sarma (1990) revised types of 13 obscure Indian Cladoceran taxa
described in the 1980s and synonymized all of them. New chydorid taxa based on poor descriptions in
obscure journals should be avoided.
(4) The system of Alona is not complete yet. Several names are still to be removed. Our list includes only
three suppressed names, yet many more are invalid (Table 1). Researchers continue the Aloninae revisions of
unresolved taxa. Diversity is also higher than presently assumed, the addition of new species within existing
species complexes should not come as a surprise (e.g., Sinev et al., in press). Of several Alona-like species,
we are yet to establish the position relative to Alona sensu strictu, pending on documentation of the limbs.
Groups may still need to be separated from core Alona, but for understanding their phylogenetic boundaries,
approaches complementary to just morphology are necessary. Our checklist only presents a snapshot of the
current situation.
As for regional assessments mentioned earlier, one area that lags behind is North America. The region has
no recent comprehensive checklists or taxonomical revisions in Cladocera, yet harbours surprises such as the
recently discovered Dumontiidae (see Santos-Flores & Dodson 2003). American chydorid taxonomy slowed
to a hold after Frey, yet his legacy shows that several Palaearctic species have different Nearctic subspecies
and siblings (e.g., Frey 1965a, Flssner & Frey 1970, Michael & Frey 1984). We are convinced that there is
larger chydorid diversity in North America than currently assumed. Regional names may be available (e.g.,
Alona lepida Birge, 1892).
VAN DAMME ET AL.
FIGURE 2. A. Accumulation curve per annum for the Alona-like species under consideration. Upper curve - total
number of taxa; striped - valid and most probably valid taxa [V + U(V) in Table 1], dotted - other taxa; solid black - taxa
of levels L3 and L2-3. B. Number of taxa described during successive 20-year periods and their current status. Upper
curve - total number of taxa; solid black - valid species [V]; striped - most probably valid [U(V)]; dotted - other taxa. C.
Number of taxa during successive 20-year periods and their original description levels. Upper curve - total number of
taxa; solid black - descriptions of levels L3 or L2-3; striped - L2; dotted - L1.
51
Discussion of the analysis: towards an increased stability

Analysis of the checklist above reveals trends in Alona systematics and the importance of efforts in
descriptions in taxonomical stability of the group. Eight examples of Alona taxa differing in description level
are shown in Figure 1 (Figs 1AH), the analysis is presented in Figure 2.
O. F. Mller (1776, 1785) created the first two species and new Alona-like taxa were not added until 1843.
A period of intensive taxonomic activity started soon after and lasted until 1900. Particularly important were
contributions by Sars (1861, 1862, 1885, 1888, 1890, 1894 and 1896). From ca. 1900 to ca. 1935 the number
of described species grew at a slower pace (Fig. 2A). Pulses of increased activity, subject to individuals and
depending on key revisions, alternate long periods of stagnation in the number of valid taxa (plateaus in Fig.
2A). 19351960 is such a period of stagnation, described for Cladocera by Korovchinsky (1997). Activities
increased again in 19601980, continuing to date (Fig. 2A). Remarkably, the increase of the number of total
taxa only correlated with the number of valid taxa around 18601865 and 18901900 (in both cases, largely
due to contributions by G.O. Sars) and from 1980 to date (Fig. 2A). In contrast, periods 18651890 and 1900
1940, led to description of almost exclusively invalid or unavailable taxa (Fig. 2A)! Stagnations of numbers of
valid taxa yet increase of total numbers, versus a parallel increase of both valid and total number of taxa,
qualify as periods of respectively taxonomical instability versus stability. Instable periods may be very long.
More than half of the taxa described over a century (during six 20-year periods; 18411960 in Fig. 2B) are
currently regarded as invalid, most of them described at the "naive" level L1 (Fig. 2C). Surprisingly, in the
period 19211960, most descriptions of new taxa (Fig. 2B) were predominantly at this low description level
(Fig. 2C; examples in Figs 1AB). The latter interval cannot be considered a period of stagnation, but instead
witnessed a significant reduction in taxonomic standards, a regression after the levels reached by Sars,
Lilljeborg and few others.
In contrast, the recent increase in total number of taxa parallels an increase in number of valid taxa (Fig.
2A), signalling a period of relative taxonomical stability. The number of valid species described since the
1960s, makes up more than half of the total number of species described (Fig. 2B). Another positive trend is
the dominance of taxa with level of description L2 since the 1960s and an increase in the number of taxa with
a modern level of description (L3) since 1970s (Fig. 2C) (see Figs 1GH). Both trends coincide (Figs 2BC).
Older, basic descriptions were compiled by Smirnov (1971), who added thoracic limb characters to diagnoses,
helping to clear the field for a wide use of limb traits in taxonomy. The appearance of Smirnovs monograph is
pivotal for the increased portion of valid versus dubious taxa after the 1970s (Fig. 2B). Descriptions
including limbs started relatively soon after this and were before only practiced by careful workers like W.
Lilljeborg and G.O. Sars. The appearance of L3 descriptions was also influenced by Frey's (1980, 1982, 1983,
1991) ideas on the general level of a cladoceran description and redescription of old taxa according to new
standards (Frey 1988). Another work is that on functional morphology and adaptive radiation in the
Chydoridae by Fryer (1968), providing the first interpretation of limbs in an evolutionary context. As well, the
detailed revision of Eurycercus by Frey (1973) opened a new page in cladoceran taxonomy (Korovchinsky
1996, 1997) and continuation of revisions adds significantly to taxonomical stability in the Aloninae.
Of late (20012009 in Figs. 2BC), the number of newly described Alona-like taxa decreased in
comparison to 19812000. The latter is connected with a splitting of the genus Alona into a series of smaller
natural entities (Van Damme & Dumont), not of loss of interest in this group. Forr et al. (2008) counted about
620 valid species of the Cladocera in the world and our total of 104 valid Alona-like species makes up a
significant portion (about 17%). The positive tendencies in taxonomy revealed above for Aloninae (increase
of number of valid taxa with L3 level of description, Figs 2BC) become characteristic of other cladoceran
groups as well. Unfortunately, the Daphniidae, with the genus Daphnia O. F. Mller, 1785 (Anomopoda:
Daphniidae), are an exception (see Korovchinsky 1996, Ishida et al. 2006). Here, descriptions with levels L2
and even L1 keep being published. The use of apparently unavailable names, violating the principles of ICZN
(1999), are still common in Daphnia (e.g., Colbourne et al. 2006) and recently proposed changes remain
unsupported by thorough step-by-step revisions.
VAN DAMME ET AL.
Acknowledgements
We wish to thank curators of the different collections which hold Alona species, for allowing us to investigate
type material, in particular Dr Charles Oliver Coleman (ZMHU), Dr Marit Christiaensen (GOS), Dr Valorie
Barnes (DGF), Dr F. Ferrari & Dr Chad Walter (DGF) and Dr Lazlo Forr (DAD).
Many thanks to Steven Tracey (ICZN Secretariat) for supplying us with some literature sources and
comments on ICZN; to Mr. Xu Lei (Jinan University, Guangzhou, China) for translating papers from Chinese;
to Prof. Dr N. N. Smirnov, Dr A.Y. Sinev (M. V. Lomonosov Moscow State University, Moscow) for
comments on the text; Dr L. Nevalainen (University of Helsinki, Finland) and Dr A. Brancelj (University of
Ljubljana, Slovenia) for various help.
This work is partly supported by the Russian Foundation for Basic Research (grant 09-04-00201-a),
"Biodiversity" Program of the Russian Academy of Sciences, and Russian Science Support Foundation (for
AAK).
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VanDamme Kotov Dumont 2010 Alona Checklist

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Zootaxa 2330: 163 (2010)

ISSN 1175-5326 (print edition)

Copyright 2010 Magnolia Press

A checklist of names in Alona Baird 1843

Accepted by M. Alonso: 30 Nov. 2009; published: 7 Jan. 2010

Kay Van Damme, Alexey A. Kotov & Henri J. Dumont

FIRST PUBLISHED IN 2010 BY

2010 Magnolia Press

2 Zootaxa 2330 2010 Magnolia Press

VAN DAMME ET AL.

A CHECKLIST OF NAMES IN ALONA

Zootaxa 2330 2010 Magnolia Press

Australian Museum, Sydney, Australia.

4 Zootaxa 2330 2010 Magnolia Press

VAN DAMME ET AL.

History in Suitland, Maryland, U.S.A.

Note on authors names and dates used in the checklist

Zootaxa 2330 2010 Magnolia Press

Materials and methods for analysis

Revised Types Current higher classification

abbreviata Sars, 1896

Armatalona Sinev, 2004

Leydigia Kurz, 1875

Kurz (1875); Kotov

Coronatella Dybowski &

acuticostata Sars, 1903

Coronatella Dybowski &

Sars (1903), Sinev

affinis (Leydig, 1860)

Alonso (1996), Sinev

africana Daday, 1908

?gen. nov. 1 (Alona

aliensis Jiang, 1980

alluaudi Richard, 1898

alonopsiformis Brehm, 1933 L1

gen. nov. 1 (Alona verrucosa- Brehm (1933b)

alsafadii Dumont &

Karualona Dumont & SilvaBriano, 2000

Dumont & SilvaBriano (2000)

ambigua (Lilljeborg, 1900)

Tretocephala Frey, 1965

Smirnov (1966), Frey

americana Flner & Frey,

Flner & Frey (1970)

anglica Hellich, 1874

angulata Birge, 1879

angulata Weigold, 1911

Coronatella Dybowski &

Kurzia Dybowski &

Hudec (2000), Kotov

anodonta Daday, 1905

archeri Sars, 1888

archeroides Brehm, 1933

arcuata Herbst, 1964

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VAN DAMME ET AL.

Revised Types Current higher classification

arcuata Sars, 1916

Sars (1916), Sinev

Coronatella Dybowski &

arenaria Brehm, 1933

azorica Frenzel & Alonso,

Frenzel & Alonso