Balian E.V., L?v?que C., Segers H., Martens K. (Eds.) Freshwater Animal Diversity Assessment

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ranges encompassing Eurasia, Africa, and North

America. The genus Lepthesteriella, in contrast,

seems Gondwanian, and occurs in Africa, Madagas-

car, and Asia. Lynceids are cosmopolitan, save in

Antarctica. Cyclestheria hislopi (if representing only

one species, which is uncertain) is pantropical.

Human related issues

Worldwide changes in land use, brought about

mainly by agriculture and urbanization, has led to a

global loss and deterioration of temporary pools. It

has been estimated that during the post-war period in

England, nearly 40% of ephemeral ponds have

disappeared, which is a rate loss of *1% per year.

Several studies indicate a loss of between 50% and

90% of original Californian vernal pools (Bauder,

1986). For most regions of the world, and especially

the developing countries, no such ﬁgures are avail-

able, but are likely to be as dramatic, due to

uncontrolled spraying with insecticides, mining, and

agricultural activities (Brendonck & Williams, 2000).

Large branchiopods are used in the United States

for assessment of ephemeral wetland habitat func-

tions and values (Rogers, 1998), and ﬁve US species

are currently listed as threatened or endangered in

California and Oregon. Another anostracan species

has been petitioned for federal endangered status and

is protected as a ‘‘rare, threatened or endangered

species’’ under the California Environmental Quality

Act. Another species was proposed for federal listing

in the state of Georgia, four species are proposed for

endangered species status in New Mexico, one

species is listed as vulnerable in Aus tralia, and one

in Brazil. About 32 large branchiopod species are red

listed by the IUCN (IUCN, 2000). Furthermore, the

world’s ﬁrst large branchiopod preserve has been

developed in Austria (Eder & Ho

dl, 2002) protecting

the anostracan Chirocephalus shadinii. Five of the six

endangered fairy shrimp in California are endemic to

that state, and were only discovered since 1990.

Fig. 3 Species and genus diversity of Anostraca (An), Notostraca (Not), Spinicaudata (Spin) and Laevicaudata (Laev) in each of the

seven zoogeographical regions. PA, Palaearctic; NA, Nearctic; NT, Neotropical; AT, Afrotropical; OL, Oriental; AU, Australasian;

PAC, Paciﬁc Oceanic Islands; ANT, Antarctic

Hydrobiologia (2008) 595:167–176 173

123

Some large branchiopod species are important

economically, includin g species that are: harvested

for aquaculture (Artemia spp.), for applications in

aquatic toxicology (Brendonck & Persoone, 1993),

for human consumption (Streptocephalus sirindhor-

nae), or as pets (‘‘Sea Monkeys’’ Artemia ‘‘nyos’’ )

(Sanoamuang et al., 2000).

Some Triops forms (Notostraca) are pests in rice

ﬁelds in at least seven countries on four continents

(Grigarick et al., 1961). Alternatively, these shrimp

have been used to control weeds in rice ﬁelds of Japan

(Takahashi, 1977), and also have been proposed as a

biological control agent of mosquitoes (Tietze &

Mulla, 1991). Given their rapid growth, early matu-

ration, and uniparental reproduction via resting eggs,

two Triops forms are being studied for their potential

use in aquaculture (Obrego

n-Barboza et al., 2001).

References

Bauder, E. T., 1986. San Diego vernal pools: Recent and

projected losses; their condition; and threats to their

existence, 1979–1990. Report prepared for Endangered

Plant Project, California Department of Fish and Game,

Sacramento, CA.

Belk, D., 1982. Branchiopoda. In Parker, S. P. (ed.), Synopsis

and Classiﬁcation of Living Organisms, Vol. 2. McGraw-

Hill, New York, 174–180.

Belk, D., 1989. Identifying species in the conchostracan genus

Eulimnadia by egg shell morphology. Journal of Crusta-

cean Biology 9(1): 115–125.

Belk, D., 1991. Anostracan mating behavior: A case of

scramble-competition polygyny. In Bauer, R. T. & J. W.

Martin (eds), Crustacean Sexual Biology. Columbia

University Press, New York, 111–125.

Belk, D., 1995. Uncovering the Laurasian roots of Eubran-

chipus. Hydrobiologia 298: 241–243.

Belk, D., 1998. Global status and trends in ephemeral pool

invertebrate conservation: implications for California

fairy shrimp. In Witham, C. W., & E. T. Bauder (eds),

147–150.

Belk, D. & J. Brtek, 1995. Checklist of the Anostraca. Hyd-

robiologia 298: 315–353.

Belk, D. & J. Brtek, 1997. Supplement to ‘checklist of the

Anostraca’. Hydrobiologia 359: 243–245.

Boudrias, M. A. & J. Pires, 2002. Unusual sensory setae of the

raptorial Branchinecta gigas (Branchiopoda: Anostraca).

Hydrobiologia 486: 19–27.

Braband, A. R., R. Hiesel & G. Scholtz, (2002). Phylogenetic

relationships within the phyllopoda (Crustacea, Bran-

chiopoda) based on mitochondrial and nuclear markers.

Molecular Phylogenetics and Evolution 25: 229–244.

Brendonck, L., 1989. A review of the phyllopods (Crustacea:

Anostraca, Notostraca, Conchostraca) of the Belgian

fauna. Verhandelingen van het symposium ‘‘Invertebraten

van Belgie

’’ 1989: 129–135.

Brendonck, L., 1993a. Feeding in the fairy shrimp Strepto-

cephalus proboscideus (Frauenfeld) (Branchiopoda:

Anostraca). I. Aspects of the feeding biology. Journal of

Crustacean Biology 13: 235–244.

Brendonck, L., 1993b. Feeding in the fairy shrimp Strepto-

cephalus proboscideus (Frauenfeld) (Branchiopoda:

Anostraca). II. Inﬂuence of environmental conditions on

feeding rate. Journal of Crustacean Biology 13: 245–255.

Brendonck, L., 1995. An updated diagnosis of the branchio-

podid genera (Branchiopoda: Anostraca, Branchipodidae)

with reﬂections on the genus concept by Dubois (1988)

and the importance of genital morphology in anostracan

taxonomy. Archiv fu

r Hydrobiologie 107(2): 149–186.

Brendonck, L., 1996. Diapause, quiescence, hatching require-

ments: What we can learn from large freshwater

branchiopods (Crustacea: Branchiopoda: Anostraca, No-

tostraca, Conchostraca). Hydrobiologia 320: 85–97.

Brendonck, L. & D. Belk, 1997. On potentials and relevance of

the use of copulatory structures in anostracan taxonomy.

Hydrobiologia 359: 83–92.

Brendonck, L. & A. Coomans, 1994a. Egg morphology in

African Streptocephalidae (Crustacea: Branchiopoda:

Anostraca). Part 1: South of Zambezi and Kunene rivers.

Archiv fu

r Hydrobiologie/Suppl. 99 (Monographische

Beitra

ge) 3: 313–334.

Brendonck, L. & A. Coomans, 1994b. Egg morphology in

African Streptocephalidae (Crustacea: Branchiopoda:

Anostraca). Part 2: North of Zambezi and Kunene rivers,

and Madagascar. Archiv fu

r Hydrobiologie/Suppl. 99

(Monographische Beitra

ge) 3: 335–356.

Brendonck, L. & G. Persoone, 1993. Biological/ecological

characteristics of large freshwater branchiopods from

endorheic regions and consequences for their use in cyst-

based toxicity tests. In Soares, A. M. V. M. & P. Calow

(eds), Progress in Standardization of Aquatic Toxicity

Tests, Chapt. 2. CRC Press, Boca Raton, FL, 7–35.

Brendonck, L. & W. D. Williams, 2000. Biodiversity in wet-

lands of dry regions (drylands). In Gopal, B., W. J. Junk &

J. A. Davis (eds), Biodiversity in Wetlands: Assessment,

Function and Conservation, Vol. 1. Backhuys Publishers,

Leiden, The Netherlands, 181–194.

Brendonck, L., M. Hamer & A. Thiery, 1992. The occurrence

of tetrahedral eggs in the Streptocephalidae (Crusta-

cea:Branchiopoda:Anostraca) with descriptions of a new

subgenus: Parastreptocephalus and a new species:

Streptocephalus (Parastreptocephalus)zuluensis Bren-

donck and Hamer. Journal of Crustacean Biology 12:

282–297.

Brendonck, L., B. J. Riddoch, V. Van de Weghe & T. Van

Dooren, 1998. The maintenance of egg banks in very

short-lived pools—a case study with anostracans (Bran-

chiopoda). In Brendonck, L., L. De Meester & N. G.

Hairston Jr. (eds), Evolutionary and Ecological Aspects of

Crustacean Diapause. Archiv fu

r Hydrobiologie (Special

issues) 52: 141–161.

Colburn,

A., 2004. Vernal Pools. Natural History and

Conservation. McNaughton & Gunn, Inc., Saline, MI.

Daniels, S. R., M. Hamer & D. C. Rogers, 2004. Molecular

evidence suggests an ancient radiation for the fairy shrimp

174 Hydrobiologia (2008) 595:167–176

123

genus Streptocephalus (Branchiopoda: Anostraca). Bio-

logical Journal of the Linnean Society 82: 313–327.

de Waard, J. R., V. Sacherova, M. E. A. Cristescu, E. Remigio,

T. Crease & P. D. N. Hebert, 2006. Probing the rela-

tionships of the branchiopod crustaceans. Molecular

Phylogenetics and Evolution 39: 491–502.

Duff, R. J., W. R. Hoeh, D. G. Cook & S. C. Weeks, 2004.

Isolation and characterization of thirteen polymorphic

microsatellite loci from the clam shrimp Eulimnadia tex-

ana (Crustacea: Spinicaudata). Molecular Ecology Notes

4: 397–399.

Dumont, H. J. & S. Negrea, 2002. Introduction to the Class

Branchiopoda. Guides to the Microinvertebrates of the

Continental Waters of the World. Backhuys, Leiden.

Eder, E. & W. Ho

dl, 2002. Large freshwater branchiopods in

Austria: Diversity, threats, conservational status. In

Escobar-Briones E. & F. Alvarez (eds), Modern Approa-

ches to the Study of Crustacea. Kluwer Academic/Plenum

Publishers, New York, 281–289.

Ellner, S. & N.G. Hairston Jr., 1994. Role of overlapping

generations in maintaining genetic variation in a ﬂuctu-

ating environment. The American Naturalists 143: 403–

417.

Fryer, G., 1987. A new classiﬁcation of the branchiopod

Crustacea. Zoological Journal of the Linnean Society 91:

357–383.

Grigarick, A. A., W. H. Lange & D. C. Finfrock, 1961. Control

of the tadpole shrimp, Triops longicaudatus, in California

rice ﬁelds. Journal of Economic Entomology 54: 36–40.

Hamer, M., L. Brendonck, A. Coomans & C. C. Appleton,

1994a. A review of African Streptocephalidae (Crustacea:

Branchiopoda: Anostraca) Part 1: South of Zambezi and

Kunene rivers. Archiv fu

r Hydrobiologie 3: 235–277.

Hamer, M., L. Brendonck, A. Coomans & C. C. Appleton,

1994b. A review of African Streptocephalidae (Crustacea:

Branchiopoda: Anostraca). Part 2: North of Zambezi and

Kunene rivers, and Madagascar. Archiv fu

r Hydrobiologie

3: 279–311.

Hildrew, A. G., 1985. A quantitative study of the life history of

a fairy shrimp (Branchiopoda: Anostraca) in relation to

the temporary nature of its habitat, a Kenyan rainpool.

The Journal of Animal Ecology 54: 99–110.

Hoeh, W. R., N. D. Smallwood, D. M. Senyo, E. G. Chapman

& S. C. Weeks, 2006. Evaluating the monophyly of Eu-

limnadia and the Limnadiinae (Branchiopoda:

Spinicaudata) using DNA sequences. Journal of Crusta-

cean Biology 26(2): 182–192.

International Union of Conservation of Nature and Natural

Resources, 2000, 2000. IUCN red list of threatened spe-

cies, compiled by Craig Hilton-Taylor. IUCN – The

World Conservation Union Species Survival Commission.

King, J. L. & R. Hanner, 1998. Cryptic species in a ‘‘living

fossil’’ lineage: taxonomic and phylogenetic relationships

within the genus Lepidurus (Crustacea: Notostraca) in

North America. Molecular Phylogenetics and Evolution

10: 23–36.

Linder, F., 1941. Contributions to the morphology and the

taxonomy of the Branchiopoda Anostraca. Zoologiska

Bidrag fran Uppsala, 10: 101–302.

Linder, F., 1952. Contributions to the morphology and taxon-

omy of the Branchiopoda, Notostraca, with special

reference to the North American species. Proceedings

United States National Museum 102: 1–69.

Longhurst, A. R., 1955. A review of the Notostraca. Bulletin

(British Alabama Museum of Natural History, Zoology) 3:

3–57.

Maeda-Martinez, A. M., D. Belk, H. Obregon-Barboza & H. J.

Dumont, 1995a. Diagnosis and phylogeny of the New

World Streptocephalidae (Branchiopoda: Anostraca).

Hydrobiologia 298: 15–44.

Maeda-Martinez, A. M., D. Belk, H. Obregon-Barboza & H. J.

Dumont, 1995b. A contribution to the systematics of the

Streptocephalidae (Branchiopoda: Anostraca). Hydrobio-

logia 298: 203–232.

Maeda-Martı

nez, A. M., H. Obrego

n-Barboza, H. Garcı

a-Ve-

lazco & G. Murugan, 2000. A proposal on the phylogeny

and the historical biogeography of the tadpole shrimp

Triops. Anostracan News 8: 1–4.

Martin, J. W., 1989. Eulimnadia belki, a new clam shrimp from

Cozumel, Mexico (Conchostraca: Limnadiidae), with a

review of Central and South American species of the genus

Eulimnadia. Journal of Crustacean Biology 9: 104–114.

Martin, J. W., 1992. Branchiopoda. In Harrison, F. W. & A. G.

Humes (eds), Microscopic Anatomy of Invertebrates,

Crustacea, Vol. 9. Wiley-Liss, New York, 25–224.

Martin, J. W. & D. Belk, 1989. Eulimnadia ovilunata

and

ovisimilis,

new species of clam shrimps (Crustacea,

Branchiopoda, Spinicaudata) from South America. Pro-

ceedings of the Biological Society of Washington 102(4):

894–900.

Martin, J. W. & C. Cash-Clark, 1995. The external morphology

of the onychopod ‘‘cladoceran’’ genus Bythotrephes

(Crustacea, Branchiopoda, Onychopoda, Cercopagididae),

with notes on the morphology and phylogeny of the order

Onychopoda. Zoologica Scripta 24: 61–90.

Martin, J. W. & G. E. Davis, 2001 An Updated Classiﬁcation

of the Recent Crustacea Natural History. Museum of Los

Angeles County, Los Angeles.

Mura, G., 1986. SEM morphological survey on the egg shell in

the italian Anostracans (Crustacea, Branchiopoda). Hyd-

robiologia 134: 273–286.

Murugan, G., A. M. Maeda-Martı

nez, H. Obrego

n-Barboza &

N. Y. Herna

ndez-Saavedra, 2002. Molecular character-

ization of the tadpole shrimp Triops (Branchiopoda:

Notostraca) from the Baja California Peninsula, Me

xico:

New insights on species diversity and phylogeny of the

genus. Studies on Large Branchiopod Biology, Hydrobi-

ologia 486: 101–113.

Obrego

n-Barboza, H., A. M. Maeda-Martı

nez & G. Murugan,

2001. Reproduction, molting, and growth of two Mexican

uniparental forms of the tadpole shrimp Triops (Bran-

chiopoda: Notostraca), under a recirculating culture

system. Hydrobiologia 462: 173–184.

Olesen, J., (1998). A phylogenetic analysis of the Conchostraca

and Cladocera (Crustacea, Branchiopoda, Diplostraca).

Zoological Journal of the Linnean Society 122: 491–536.

Remigio, E. A. & P. D. N. Hebert, 2000. Afﬁnities among

anostracan (Crustacea: Branchiopoda) families inferred

from phylogenetic analyses of multiple gene sequences.

Molecular Phylogenetic Evolution 17: 117–128.

Remigio, E. A., P. D. N. Hebert & A. Savage, 2001. Phylo-

genetic relationships and remarkable radiation in

Hydrobiologia (2008) 595:167–176 175

123

Parartemia (Crustacea:Anostraca), the endemic brine

shrimp of Australia: Evidence from mitochondrial DNA

sequences. Biological Journal of Linnean Society 74:

59–71.

Remigio, E. A., B. V. Timms & P. D. N. Hebert, 2003. Phy-

logenetic systematics of the Australian fairy shrimp genus

Branchinella based on mitochondrial DNA sequences.

Journal of Crustacean Biology 23: 436–442.

Rogers, D. C., 1998. Aquatic macroinvertebrate occurrences

and population trends in constructed and natural vernal

pools in Folsom, California. In Witham, C. W., E. T.

Bauder, D. Belk, W. R. Ferrin Jr. & R. Orduff (eds),

Ecology, Conservation, and Management of Vernal Pool

Ecosystems—Proceedings from a 1996 Conference. Cal-

ifornia Native Plant Society, Sacramento, CA, 224–235.

Rogers, D. C., 2001. Revision of the Nearctic Lepidurus (No-

tostraca). Journal of Crustacean Biology 21: 991–1006.

Rogers, D. C., 2002a. Amplexial morphology of selected

Anostraca. Hydrobiologia 486: 1–18.

Rogers, D. C., 2002b. A morphological re-evaluation of the

anostracan families Linderiellidae and Polyartemiidae,

with a redescription of the linderiellid Dexteria ﬂoridana

(Dexter 1956) (Crustacea: Branchiopoda). Hydrobiologia

486: 56–61.

Rogers, D. C., D. Quinney, J. Weaver & J. Olesen, 2006. A

new giant species of predatory fairy shrimp from Idaho

(Branchiopoda: Anostraca). Journal of Crustacean Biol-

ogy, 26: 1–16.

Rogers, D. C., 2006. A genus level revision of the Thamno-

cephalidae (Crustacea: Branchiopoda: Anostraca).

Zootaxa (in press).

Sanoamuang, L. A., G. Murugan, P. H. H. Weekers & H. J.

Dumont, 2000. Streptocephalus sirindhornae, new species

of freshwater fairy shrimp (Anostraca) from Thailand.

Journal of Crustacean Biology 20: 599–565.

Sassaman, C., 1995. Sex determination and evolution of uni-

sexuality in the Conchostraca. Hydrobiologia 298: 45–65.

Sassaman, C. & S. Weeks, 1993. Sex determination in the

conchostracan Eulimnadia texana. The American Natu-

ralist 141: 314–328.

Sassaman, C., M. A. Simovich & M. Fugate, 1997. Repro-

ductive isolation and genetic differentiation in North

American species of Triops (Crustacea: Branchiopoda:

Notostraca). Hydrobiologia 359: 125–147.

Scanabissi, F., M. Cesari, S. K. Reed & S. C. Weeks, 2006.

Ultrastructure of the male gonad and male gametogenesis in

Eulimnadia texana (Packard, 1871) (Crustacea, Branchio-

poda, Spinicaudata). Invertebrate Biology 125: 117–124.

Spears, T. & L. G. Abele, 2000. Branchiopod monophyly and

interordinal phyogeny inferred from 18S ribosomal RNA.

Journal of Crustacean Biology 20(1): 1–24.

Stenderup, J. T., J. Olesen & H. Glenner, 2006. Molecular

phylogeny of the Branchiopoda (Crustacea)-Multiple

approaches suggest a ‘diplostracan’ ancestry of the

Notostraca. Molecular Phylogenetics and Evolution 41:

182–194.

Suno-Uchi, N., F. Sasaki, S. Chiba & M. Kawata, 1997.

Morphological stasis and phylogenetic relationships in

tadpole shrimps, Triops (Crustacea: Notostraca). Biolog-

ical Journal of the Linnean Society 61: 439–457.

Takahashi, F., 1977. Pioneer life of the tadpole shrimps, Triops

spp. (Notostraca: Triopsidae). Applied Journal of Ento-

molgy and Zoology 12: 104–117.

Tietze, N. S. & M. S. Mulla, 1991. Biological control of Culex

mosquitoes (Diptera: Culicidae) by the tadpole shrimp,

Triops longicaudataus (Notostraca: Triopsidae). Journal

of Medical Entomology 28: 24–31.

Walossek, D., (1993). The Upper Cambrian Rehbachiella and

the phylogeny of Branchiopoda and Crustacea. Fossils and

Strata 32: 1–202.

Weekers, P. H. H., G. Murugan, J. Vanﬂeteren, D. Belk & H. J.

Dumont, 2002. Phylogenetic analysis of anostracans

(Branchiopoda: Anostraca) inferred from SSU rDNA

sequences. Molecular Phylogenetics and Evolution 25:

535–544.

Weeks, S. C., R. T. Posgai, M. Cesari & F. Scanabissi, 2005.

Androdioecy inferred in the clam shrimp Eulimnadia a-

gassizii (Spinicaudata: Limnadiidae). Journal of

Crustacean Biology 25: 323–328.

Weeks, S. C., C. Benvenuto & S. K. Reed, 2006a. When males

and hermaphrodites coexist: a review of androdioecy

in animals. Integrative and Comparative Biology 46:

449–464.

Weeks, S. C., T. F. Sanderson, S. K. Reed, M. Zofkova, B.

Knott, U. Balaraman, G. Pereira, D. M. Senyo & W. R.

Hoeh, 2006b. Ancient androdioecy in the freshwater

crustacean Eulimnadia. Proceedings of the Royal Society

of London, Series B 273: 725–734.

176 Hydrobiologia (2008) 595:167–176

123

FRESHWATER ANIMAL DIVERSITY ASSESSMENT

Global diversity of cladocerans (Cladocera; Crustacea)

in freshwater

L. Forro

Æ N. M. Korovchinsky Æ A. A. Kotov Æ

A. Petrusek

 Springer Science+Business Media B.V. 2007

Abstract Cladocera is a primarily-freshwater

monophyletic group, an important component of the

microcrustacean zooplankton. They inhabit most

types of continental fresh and saline water habitats,

occurring more abundantly in both temporary and

permanent stagnant waters. Cladocera is an ancient

group of Palaeozoic origin. About 620 species are

currently known, but we estimate that the real number

of species is 2–4 times higher. A number of currently-

recognised widespread species can be expected to

harbour extensive cryptic diversity.

Keywords Cladocera  Species richness 

Global assessment  Biogeography 

Endemicity

Introduction

Cladocerans (‘‘water ﬂeas’’) are primarily-freshwater

small-sized (0.2–6 mm, and up to 18 mm in single

case of Leptodora kindtii) branchiopod crustaceans,

inhabiting pelagic, littoral, and benthic zones. Four

cladoceran orders are recognised (Fryer, 1987):

Anomopoda, Ctenopoda, Onychopoda, and the

monotypic Haplopoda (see Fig. 1 A–D for represen-

tatives of each order). Most species occur in

continental fresh or saline waters, although two

ctenopods and several onychopods from the family

Podonidae are truly marine, and a few more cteno-

pod, anomopod and onychopod species occur in

brackish waters. Seven known species may be

regarded as true inhabitants of subterranean environ-

ment, and a few others (of the family Chydoridae)

live in semi-terrestrial conditions.

The trunk and appendages of most cladocerans

(Anomopoda and Ctenopoda) are enclosed in a

bivalved carapace. Tagmosis of the body is obscure

(except in Leptodora kindtii, the single representative

of Haplopoda), and a single eye and ocellus are

usually present. Antennules are uniramous, while

antennae are biramous (except in females of Holope-

dium), natatory, with 2–4 segments per branch. Four

to six pairs of trunk limbs are either mostly similar in

shape (Ctenopoda, Onychopoda, Haplopoda) or mod-

iﬁed individually for various functions (Anomopoda).

Water ﬂeas are important components of the

fauna of fresh waters; they are particularly

Guest editors: E. V. Balian, C. Le

que, H. Segers &

K. Martens

Freshwater Animal Diversity Assessment

L. Forro

(&)

Department of Zoology, Hungarian Natural History

Museum, Baross u. 13, 1088 Budapest, Hungary

e-mail: forro@zoo.zoo.nhmus.hu

N. M. Korovchinsky  A. A. Kotov

A.N. Severtsov Institute of Ecology and Evolution,

Leninsky prospect 33, 119071 Moscow, Russia

A. Petrusek

Department of Ecology, Faculty of Science, Charles

University in Prague, Vinic

7, 12844 Prague 2, Czechia

123

Hydrobiologia (2008) 595:177–184

DOI 10.1007/s10750-007-9013-5

signiﬁcant in the food web of stagnant waters. Most

species are ﬁlter-feeders; onychopods and haplopods

are predatory. They usually reproduce by cyclical

parthenogenesis (but asexual lineages are known as

well), and populations are mostly dominated by

females. Sexual dimorphism is normally rather

distinct. Sexual ly produced diapausing eggs are

resistant to desiccation and other unfavourable

Fig. 1 A––Sida crystallina

B––Bythotrephes

longimanus

C––Alona sp.

D––Leptodora kindti

(original drawings by G O

Sars)

178 Hydrobiologia (2008) 595:177–184

123

conditions, and may even survive passage through

the digestive track of birds (Figuerola & Green,

2002); thus, they are important propagules for

passive dispersal.

The ﬁrst information on Cladocera date from the

17th century; the histor y of research has been divided

into three to seven major phases (for a detailed

discussion see Korovchinsky, 1997; Dumont &

Negrea, 2002). An important change of paradigm,

characterised by the rejection of the prevailing

assumption of cosmopolitanism of cladoceran spe-

cies, occurred around the 1950–1980s with a new

approach to the taxonomy and phylogeny of Chydo-

ridae (Frey, 1959, 1982, 1987). Subsequently, the

concept of non-cosmopolitanism has been supported

by numerous morphological, as well as molecular,

studies. The increasing use of molecular tools in

recent years has and will continue to have a strong

impact on our understanding of cladoceran diversity,

phylogeny and biogeography (e.g., Adamowicz et al.,

2004; Cox & Hebert, 2001; Schwenk et al., 2000;

Taylor et al., 2002).

Species diversity

The currently accepted number of cladoceran species

based on existing descriptions is around 620. The

tables summarize the currently known number of

species and genera within orders and families of the

group (Tables 1, 2, Fig. 2), based on recent major

publications (Smirnov, 1992a, 1996; Korovchinsky,

1996, 2004; Orlova-Bienkowskaja, 2001; Dumont &

Negrea, 2002; Benzie, 2005; Kotov & Stifter, 2006)

and additional published or as yet unpublished

sources. The described taxonomic diversity of Clado-

cera, however, underestimates the reality, and even

higher-ranked taxa are still being discovered, e.g., a

new family, Dumontiidae (Santos-Flores & Dodson,

2003).

Only about 45–50% of the species may be

considered to be more or less well described and

valid, while the status of other species is vague,

and many of them likely represe nt cryptic com-

plexes (Korovchinsky, 1996). The families

Chydoridae, Daphniidae, Ilyocryptidae, and Sididae

have been stud ied comparatively better. The largest

number of valid species is known from Europe,

North America, Australia, and South America, and

the smallest number from Africa and Southern

Asia. This, however, at least partly reﬂects the

intensity of research rather than real patterns of

diversity.

Adamowicz & Purvis (2005) estimated three cor-

rection factors to extrapolate global branchiopod

diversity from the diversity of described species, and

Fig. 2 Global Distribution

of species and genus

diversity by zoogeographic

region (Species Number/

Genus Number).

PA––Palaearctic;

NA––Nearctic;

NT––Neotropical;

AT––Afrotropical ;

OL––Oriental;

AU––Australasian;

PAC––Paciﬁc Oceanic

Islands; ANT––Antarctic

Hydrobiologia (2008) 595:177–184 179

123

predicted that there are about 2.1 times more bran-

chiopod species in nature than currently known. The

overall cladoceran species richness is probably up to 4

times higher than currently known. This is supported

by the results of molecular studies. Detailed studies,

combining morphological analyses and molecular

tools, are especially promising for delineating species

boundaries in groups with relatively uniform mor-

phology, fewer qualitative characters, and widespread

phenotypic plasticity. Although most molecular anal-

yses have so far focused on a single model genus,

Daphnia, within a relatively short time this led to the

discovery of an unprecedented number of cryptic

lineages. According to Hebert & Taylor (1997), the

global total for the genus Daphnia (including Daph-

niopsis) is likely closer to 200 species instead of 75

included in the last monograph on the genus (Benzie,

2005). Similar patterns of widespread cryptic diversity

and high numbers of undescribed lineages can be seen

in other groups, e.g., in Moina (Petrusek et al., 2004

and unpublished data), Ilyocryptus (Kotov & S

tifter,

2006) and several genera of the Chydoridae.

Phylogeny and historical processes

Cladocerans probably derived from large bodied

branchiopod ancestors. Recent molecular analyzes

have suggested two alternative phylogenetic relation-

ships among cladoceran orders. The monophyly of

Gymnomera (Haplopoda and Onychopoda) is sup-

ported in both cases but one hypothesis suggests a

sister relationship between Anomopoda and the

remaining three orders (Swain & Taylor, 2003),

while the other clusters Anomopoda and Ctenop oda

together (De Waard et al., 2006). Other authors (see

review in Negrea et al., 1999) have recently proposed

alternative hypotheses on ordinal-level relationships

Table 1 Number of Cladocera species currently known in the

main biogeographic areas. PA: Palaearctic; NA: Nearctic; NT:

Neotropical; AT: Afrotropical; OL: Oriental; AU:

Australasian; PAC: Paciﬁc Oceanic Islands; ANT: Antarctic.

(numbers in parentheses indicate endemic species)

PA NA NT AT OL AU PAC ANT World

Order Anomopoda 195 (83) 169 (66) 170 (89) 125 (24) 89 (20) 149 (78) 29 (0) 12 (6) 537

Family Daphniidae* 58 (21) 58 (25) 32 (13) 25 (1) 17 (1) 26 (13) 6 (0) 3 (2) 121

Family Moinidae* 13 (6) 7 (2) 10 (5) 10 (1) 3 (0) 7 (3) 4 (0) 0 29

Family Dumontiidae 0 1 (1) 0 000001

Family Ilyocryptidae 11 (3) 10 (2) 9 (4) 8 (3) 5 (3) 5 (3) 1 (0) 1 (0) 28

Family Bosminidae 4 (0) 8 (3) 7 (3) 3 (0) 4 (1) 3 (0) 1 (0) 0 14

Family Acantholeberidae 1 (0) 1 (0) 0 000001

Family Ophryoxidae 3 (1) 3 (1) 0 000003

Family Macrothricidae 16 (10) 10 (5) 21 (12) 12 (2) 12 (4) 20 (9) 2 (0) 3 (1) 60

Family Neothricidae 0 0 0 0 0 3 (3) 0 0 3

Family Eurycercidae 4 (2) 5 (3) 2 (1) 1 (0) 1 (0) 0 0 0 8

Family Chydoridae 85 (40) 66 (24) 89 (51) 66 (17) 48 (11) 85 (47) 15 (0) 5 (3) 269

Order Ctenopoda 17 (5) 18 (7) 16 (9) 9 (0) 15 (4) 9 (5) 4 (1) 0 50

Family Holopediidae 1 (0) 2 (1) 1 (1) 000003

Family Sididae 16 (5) 16 (6) 15 (8) 9 (0) 15 (4) 9 (5) 4 (1) 0 47

Order Haplopoda 1 (0) 1 (0) 0 0 1 (0) 0 0 0 1

Family Leptodoridae 1 (0) 1 (0) 0 000001

Order Onychopoda 32 (31) 1 (0) 0 0 1 (0) 0 0 0 32

Family Polyphemidae 2 (1) 1 (0) 0 0 1 (0) 0 0 0 2

Family Podonidae 17 (17) 0 0 0000017

Family Cercopagidae* 13 (13) 0 0 0000013

Total 245 (119) 189 (73) 186 (98) 134 (24) 107 (24) 158 (83) 33 (1) 12 (6) 620

* Invasive species not considered

180 Hydrobiologia (2008) 595:177–184

123

for the Cladocera; these were, however, based only

on classical (Fryer, 1987) or cladistic analyses of

morphological traits.

Cladocera is an ancient group of Palaeozoic

origin (Dumont & Negrea, 2002), but their unam-

biguous fossil remains are known only from the

Mesozoic (Smirnov, 1971, 1992b; Kotov & Korov-

chinsky, 2006). Recently, Anderson et al. (2004)

described crustaceans similar to the Cladocera from

the Early Devonian. Molecular phylogenetic data

have revealed that the subfamilies of Chydoridae

(Anomopoda) were separated in the Middle Palae-

ozoic (about 400 Myr ago; Sacherova

& Hebert,

2003) and representatives of the genus Daphnia

differentiated at least 200 Myr ago (Colbourne &

Hebert, 1996). Any Mesozoic scenarios, such as

‘Gondwana-Laurasia’ (e.g., Benzie, 2005), are only

moderately applicable to cladoceran groups, espe-

cially at a generic and subgeneric level. In spite of

the general antiquity of Cladocera, radiation within

some groups is only recent or even contemporary,

e.g., in some Holarctic Daphnia and Bosmina

(Colbourne & Hebert, 1996; Taylor et al., 2002).

Present distribution and main areas of endemicity

The distribution and patterns of endemicity of higher-

level taxonomic groups are relatively well-known.

The known species diversity, as well as the number of

endemic taxa, is neverthe less bound to increase with

further faunistic research, especially from non-north-

ern temperate regions, and with the application of

detailed morphological and molecular tools to resolve

cryptic species complexes. Some endemic species

have narrow distributions, and it is therefore likely

that many remain overlooked.

The Holarctic cladoceran inland fauna is rich and

composed of all four order s. Two orders (Haplopoda

and Onychopoda), three families, 13 genera

Table 2 Number of Cladocera genera currently known in the

main biogeographic areas. PA: Palaearctic; NA: Nearctic; NT:

Neotropical; AT: Afrotropical; OL: Oriental; AU:

Australasian; PAC: Paciﬁc Oceanic Islands; ANT: Antarctic.

(numbers in parentheses indicate endemic genera)

PA NA NT AT OL AU PAC ANT World

Order Anomopoda 44 (4) 43 (3) 44 (3) 42 (1) 36 (1) 48 (11) 19 (0) 7 (0) 76

Family Daphniidae 5 (0) 5 (0) 4 (0) 5 (0) 4 (0) 4 (0) 4 (0) 2 (0) 5

Family Moinidae 1 (0) 2 (0) 2 (0) 2 (0) 2 (0) 2 (0) 1 (0) 0 2

Family Dumontiidae 0 1 (1) 0 0 0 0 0 0 1

Family Ilyocryptidae 1 (0) 1 (0) 1 (0) 1 (0) 1 (0) 1 (0) 1 (0) 1 (0) 1

Family Bosminidae 2 (0) 2 (0) 2 (0) 2 (0) 2 (0) 2 (0) 1 (0) 0 2

Family Acantholeberidae 1 (0) 1 (0) 0 0 0 0 0 0 1

Family Ophryoxidae 1 (0) 2 (1) 0 0 0 0 0 0 2

Family Macrothricidae 6 (0) 6 (0) 7 (2) 5 (0) 4 (0) 5 (0) 1 (0) 1 (0) 11

Family Neothricidae 0 00001(1)001

Family Eurycercidae 1 (0) 1 (0) 1 (0) 1 (0) 1 (0) 0 0 0 1

Family Chydoridae 26 (4) 22 (1) 27 (1) 26 (1) 22 (1) 33 (10) 11 (0) 3 (0) 49

Order Ctenopoda 7 (1) 7 (0) 6 (0) 4 (0) 6 (0) 4 (0) 2 (0) 0 8

Family Holopediidae 1 (0) 1 (0) 1 (0) 0 0 0 0 0 1

Family Sididae 6 (1) 6 (0) 5 (0) 4 (0) 6 (0) 4 (0) 2 (0) 0 7

Order Haplopoda 1 (0) 1 (0) 0 0 1 (0) 0 0 0 1

Family Leptodoridae 1 (0) 1 (0) 0 0 0 0 0 0 1

Order Onychopoda 8 (5) 1 (0) 0 0 1 (0) 0 0 0 10

Family Polyphemidae 1 (0) 1 (0) 0 0 1 (0) 0 0 0 1

Family Podonidae 5 (3) 00000 007

Family Cercopagidae 2 (2) 00000 002

Total 60 (10) 52 (3) 50 (3) 46 (1) 44 (1) 52 (11) 21 (0) 7 (0) 95

Hydrobiologia (2008) 595:177–184 181

123

(including those of the Caspian Sea and Lake Baikal),

and about 250 known species are endemic for the

region. Many taxa are presumably old and phyloge-

netically divergent, monotypic, or composed of a few

species. The Palaearctic taxa are more diverse than

those of the Nearctic due to, ﬁrst of all, the presence

of numerous Caspian and Baikalian endemics.

Among other zoogeographical regions, Australasia

is rich in endemics, represented by one family, one

subfamily, one tribe, 11 genera, and 83+ species,

while known endemics in Oriental and Neotropical

regions are of a lower rank or fewer (one tribe, one

genus and 21+ species, and three genera and 98+

species, respectively). The Afrotropical region,

though poorly studied, seems to be especially

deprived of known higher-level endemic cladoceran

taxa, being represented by a single endemic genus

and 24+ endemic species.

Cladoceran species richness does not change

evenly with latitude but concentrates in the warm

temperate to subtropical zone of both hemispheres

(*25–50, including mountain areas within the true

tropics) (Korovchinsky, 2006). In the belt from the

Mediterranean through Central Asia including the

Pontocaspian region, northern India to East Asia

(Amur region and China), ﬁve genera and over 100

known endemic species occur, while those in the

North Palaearctic do not exceed 55–60. In North

America, the area embracing the United States,

Mexican plateau, and southern Canada, is inhabited

by many endemics, including one family (Dumontii-

dae), two genera, and over 70 species. Southern

Australia, Tasmania, and New Zealand are rich in

endemics of high taxonomic rank: one subfamil y

(Sayciinae), one tribe (Australospilini), 8 genera, and

about 80 currently known species compose altogether

most of the known Australasian endemics. Only ﬁve

of them are shared between Australia and New

Zealand, which itself has seven endemic species/

subspecies. As the African cladoceran fauna has been

relatively poorly studied, there are only a few

described endemi cs (one genus and about 10 species),

but e.g., endemic Daphnia fauna of the Ethiopian

biogeographic region is certainly signiﬁcantly more

numerous (Mergeay et al., unpublished). Cladocerans

of subtropical and temperate South America include

two endemic genera and 17+ endemic species. In

total, the species richness of the sout hern temperate––

subtropical zone amounts to more than 100 endemic

species. The intermediate tropical zone, from which

altogether 163 species are known, is characterized by

fewer endemic taxa of comparatively lower taxo-

nomic rank: one tribe (Indialonini) and nine genera

(Korovchinsky, 2006).

A bipolar (antitropical) disjunct distribution of

faunal complexes and taxa (Daphnia, Pleuroxus,

Tretocephala etc.), the wide ranges of some species

(though some of these likely form species complexes)

and the narrow restriction of others, the p resence of

isolated populations, and concentration of endemics in

the warm temperate––subtropical zone of both hemi-

spheres, are typical traits of cladoceran zoogeography.

Such patterns stimulated the analysis of cladoceran

faunal formation by the modern version of the

concept of ‘ejected relicts’ instead of vicariance.

This hypothesis consider the extant Cladocera as a

relict group (Korovchinsky, 2006), whose taxa were

widely distributed in the past. Tertiary climatic

changes, primarily within the present tropical and

boreal latitudes, resulted in mass extinction of their

biotas, while the warm temperate––subtropical

regions remained comparatively unchanged. Addi-

tional factors (e.g., the radiation of freshwater

planktivorous ﬁsh) could have operated in conjunc-

tion with climate changes as well.

While this scenario might be likely for most of

cladocerans, molecular data suggest that vicariance

processes and allopatric speciation at both the intercon-

tinental level and in regional refugia within continents

plays a signiﬁcant role in shaping species diversity in at

least some genera (e.g., Daphnia). Sweepstake inter-

continental dispersals, followed by a local radiation,

seems to have been important factors in augmenting the

diversity in the different biogeographic regions. Foun-

der effects coupled with habitat shifts, such as pond-lake

transitions (Lynch, 1985) or, possibly, shifts among

substrates in littoral groups, are also regarded as

potentially important drivers of speciation. Finally,

interspeciﬁc hybridization and hybrid speciation plays

an important role in dynamic young species complexes

in Daphnia, though reports of other hybridizing cladoc-

erans are scarce (Schwenk & Spaak, 1995).

Human related issues

Cladocerans (especially Daph nia) are important model

organisms in both basic and appl ied research, due to

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