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

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of relatively low diversity has been widely adopted,

as a method of identifying priority areas for

conservation. Most analyses have identiﬁed ancient

lakes as hotspots for freshwater biodiversity. For

copepods, Lake Baikal with over 120 species, the

majority of which are endemic (Boxshall & Evst-

igneeva, 1994), and Lake Tanganyika with 69

species, of which 34 (49%) are endemic (Coulter,

1991; Boxshall & Strong, 2006), constitute prime

hotspots. In both the copepod fauna comprises

several species ﬂocks, deﬁned as monophyletic

clades endemic to the lake and presumed to have

originated via intralacustrine speciation. The age of

the lake (i.e. the time available for speciation to

have taken place), the heterogeneity of habitats at

all scales within the lake, and their biotic complex-

ity are all potential factors affecting the generation

and maintenance of high levels of biodiversity in

ancient lakes. Boxshall & Evstigneeva (1994)

suggested that trophic specialisation was a key

driver in the adaptive radiation of the cyclopids in

Lake Baikal in contrast to the canthocamptids

where they considered differentiation in mating

behaviour to be a key driver.

In recent decades, awareness of the diversity of

groundwater faunas has increased. Sket (1999) for

example, noted that 70% of the 245 harpacticoid

copepods and 60% of the 145 cyclopoids recorded

in the Limnofauna Europaea compendium were

stygobites. Culver & Sket (2000) recommended that

cave systems inhabited by at least 20 stygobitic

species should be designated biodiversity hotspots.

Worldwide only 20 karstic systems and two sites in

porous aquifers meet this criterion (Culver & Sket,

2000; Danielopol & Pospisil, 2002). The species

richness of the Lobau wetland in the Danube ﬂood

plain led Danielopol & Pospisil (2002) to refer to it

as a biodiversity hotspot. In a sampled area of

0.8 km

they discovered 30 copepod species of

which 10 were hypogean. The situation is not

unique: Karanovic (2004) discovered a subterranean

fauna of 31 species (24 new) in the arid Murc hison

region of western Australia, and Stoch (1995)

reported 70 species from the Venezia Giulia (N.E.

Italy and southeastern Slovenia) of which 33 (47%)

were stygobionts. Galassi (2001) reviewed the

patterns of diversity of groundwater copepods,

noting that stygobiont copepods inhabit all kinds

of aquifers (karstic, ﬁssured and porous). She

identiﬁed local heterogeneity, patchiness in geomor-

phological and hydrological characteristics, as well

as biotic interactions as causal factors affecting

diversity patterns over a range of spatial and

temporal scales.

Endemism

The level of endemism in freshwater copepods seems

remarkable. The great majority (2,558 out of 2,814

species, or 90.9%) of species occur in a single region.

The Neotropical, Afrotropical, Palaearctic and Aus-

tralasian Regions each display in excess of 80%

endemism (Table 3). Nearctic and Oriental Regions

display intermediate levels (65–75%) and the Paciﬁc

islands and Antarctica both display between 45% and

50%. Species that are not endemic to a single region

include highly vagile and cosmopolitan species that

occur in four or more regions.

Endemic genera occur in all regions except Antarc-

tica (Table 2), in all orders and across all life styles. The

Neotropical is home to the greatest number of endemic

genera (Table 2). There is evidence that in the

Ergasilidae most of the endemic genera represe nt a

single monophyletic lineage (El-Rashidy, 1999). Sim-

ilarly, Boxshall & Halsey (2004) recognized the

possibility that the cluster of ﬁve lernaeid genera

endemic to the Neotropics may be closely related (i.e.

form a monophyletic group). No phylogeny exists for

the Diaptomidae but, building on the speculation

(Boxshall & Jaume, 2000) that diaptomines only

invaded So uth America after the closure of the Panama

gap, it is possible that the cluster of 14 endemic genera

might also form a monophyletic lineage.

Faunal links

Pair-wise analysis of the occurrence of species that

inhabit more than one zoogeograph ic region (Table 4)

provides a relative measure of the faunal connectivity

between regions. The dominant links as estimated by

Sørenson’s index (SI) are those between the Palae-

arctic and Nearctic, between the Nearctic and Neo-

tropical, and between the Oriental and Australasian

regions. The Palaearctic to Nearctic link identiﬁes

Holarctic taxa and these typically belong to lineages,

the distributions of which have been profoundly

affected by the Pleistocene glaciations. The Nearctic

to Neotropical link in part testiﬁes to the strength of

204 Hydrobiologia (2008) 595:195–207

123

the faunal exchange that took place between North

and South America after the closure of the Panama

gap. The Oriental to Australasian link may indicate

the main dispersal route of the freshwater colonists of

Australia. There is also a relatively strong link

between Palaearctic and Oriental regions, which

may reﬂect the weakness of the barr ier separating

these regions, particularly in central and eastern

China. The link between Antarctic and Neotropical

regions is strong—10 out of the 17 species present in

the Antarctic are shared with the Neotropics. These

are mainly centropagids and cyclopids that are

distributed from the Falklands to the circum-Antarc-

tic islands. This faunal link is not identiﬁed by

Table 3 Number of endemic species recorded from each zoogeographic region

Taxon PA

region

PAC ANT

region

Total endemic

species number

Calanoida

Diaptomidae 121 65 77 71 81 2 – – 417

Centropagidae 3 1 14 – 1 45 2 1 67

Temoridae 11 8 – – – – – – 19

Aetideidae (Senecella)– – – – – – – –

Harpacticoida

Canthocamptidae 285 46 114 37 40 51 1 1 575

Parastenocarididae 117 9 69 36 16 1 1 – 249

Ameiridae 94 6 11 9 4 9 3 3 139

Miraciidae

(Diosaccidae)

25 – 5 11 1 6 1 – 49

Ectinosomatidae 11 – – – 1 – – – 12

Phyllognathopodidae 1 – 2 2 1 1 – 1 8

Harpacticidae 3 – – – – – – – 3

Chappuisiidae 2 – – – – – – – 2

Huntemanniidae 1 1 – 1 1 – – – 4

Laophontidae 2 – – – – 1 – – 3

Cletodidae – – – – 2 – – – 2

Darcythompsoniidae 1 – – – – – – – 1

Cyclopoida

Cyclopidae 254 56 130 123 68 43 4 2 680

Oithonidae – – 5 – – – – – 5

Ozmanidae – – 2 – – – – – 2

Lernaeidae 11 7 10 37 42 1 – – 108

Ergasilidae 50 21 63 17 20 7 1 – 179

Gelyelloida

Gelyellidae 2 – – – – – – – 2

Siphonostomatoida

Lernaeopodidae 25 5 – – 1 – – – 31

Caligidae 1 – – – – – – – 1

Dichelesthiidae – – – – – – – –

Total endemic species 1,020 225 502 344 279 167 13 8 2,558

Total species (Table 1) 1,204 347 561 405 381 205 29 17 2,814

Proportion of endemics 84.7% 64.8% 89.5% 84.9% 73.2% 81.5% 44.8% 47.1% 90.9

PA: Palaearctic, NA: Nearctic, NT: Neotropical, AT: Afrotropical, OL: Oriental, AU: Australasian, PAC: Paciﬁc Oceanic Islands,

ANT: Antarctic

Hydrobiologia (2008) 595:195–207 205

123

Sørenson’s Index, given the paucity of the Antarctic

fauna relative to the Neotropical.

Introduced species

There are numerous examples of introduced species

from all life styles, planktonic to parasitic. In free-

living copepods, at least 18 species have been

involved in intercontinental introduction (Reid &

Pinto-Coelho, 1994). The translocation of ﬁsh stocks

for aquaculture has been implicated in the introduc-

tion of parasites such as Paraergasilus longidigitus,

Neoergasilus japonicus and Lernaea cyprinacea

(Fig. 1H) Introductions have also been associated

with the transportation of tropical plants for horticul-

ture and agriculture. For example, the presence of the

Asiatic centropagid Boeckella triarticulata in Italy

was attributed to movement of Chinese carp, the

introduction of Mesocyclops ogunnus to the Cayman

Islands and Brazil may have resulted from transpor-

tation of organisms for aquaculture, and it has been

suggested that M. pepheiensis (as M. ruttneri), an

East-Asian species, was introduced to the southern

USA by rice culture. The uncontrolled exchange of

ballast water has been implicated in the introduction

of estuarine taxa, including Limnoithona, Sinoc al-

anus, Pseudodiaptomus and Acartia.

Human related issues

Copepods act as intermediate hosts for a variety of

parasites. Species of Mesocyclops and Thermocyclops

are intermediate hosts for guineaworm (Dracunculus

medinensis), a debilitating nematode parasite.

Guineaworm larvae are consumed by copepods and

remain in the body cavity until ingested by drinking

unﬁltered water. Guineaworm is less prevalent than

in the past but remains a human health problem,

particularly in West Africa and India. Other parasites

of man that utilize copepods as intermediate hosts

include the ﬁsh tapeworm, Diphyllobothrium latum.

Parasitic copepods such as Lernaea cyprinacea

and Ergasilus sieboldi can be serious pests, causing

mass mortality and signiﬁcant comme rcial losses in

freshwater aquaculture. They are often found in

inconspicuous microhabitats on the host and may be

overlooked in ﬁsh quarantine checks. Translocation

of ﬁsh stocks, without sufﬁcient checks, is responsi-

ble for the widespread introduction of these pests

outside of their natural ranges.

Free-living copepods can be voracious predators

and this functional role has been exploited in the use

of Mesocyclops species as biol ogical control agents

against mosquitoes. The spread of diseases such as

malaria and dengue fever involves mosquito vectors

and Mesocyclops have been experimentally trialled as

a biocontrol agent for mosquito larvae. The success

of thes e trials has been variable, but in Viet Nam for

example, Mesocyclops has been used effectively in

controlling mosquito larvae at local levels.

Desertiﬁcation and the overexploitation of ground

water are major conservation issues for copepods.

Groundwater copepods, especially those with narrow

geographical distributions, are at risk from intrusion

of saline waters as the groundwater table is lowered.

Acknowledgements We would like to thank Estelle Balian

and Koen Martens, for the invitation to contribute to this

volume, and Diana Galassi and an anonymous reviewer for

their suggested improvements.

Table 4 Faunal links between zoogeographic regions estimated using Sørensen’s index, based on species-level data: values

exceeding 0.1000 shown in bold

PA NA AT NT OL AU PAC

NA 0.1251

AT 0.0646 0.0771

NT 0.0453 0.1101 0.0621

OL 0.0984 0.0769 0.0941 0.0679

AU 0.0397 0.0507 0.0623 0.0574 0.1058

PAC 0.0162 0.0319 0.0461 0.0407 0.0585 0.0684

ANT 0.0066 0.0275 0.0190 0.0346 0.0101 0.0270 0.0435

206 Hydrobiologia (2008) 595:195–207

123

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123

FRESHWATER ANIMAL DIVERSITY ASSESSMENT

Global diversity of ﬁshlice (Crustacea: Branchiura:

Argulidae) in freshwater

William J. Poly

 Springer Science+Business Media B.V. 2007

Abstract The Branchiura of freshwater habitats,

consisting of the valid genera Argulus, Chonopeltis,

Dipteropeltis, and Dolops, presently contains 113

valid species and 12 undescribed species. The entire

group is composed of ectoparasitic species that

usually live on ﬁshes. The highest diversity of genera

and species occurs in the Afrotropical and Neotrop-

ical regions. The diversity of the freshwater species

surpasses that of species in marine and brackish

waters, but this could be due to inadequate study of

the fauna of the latter habitats. One species, Argulus

japonicus, has been introduced from east/southeast

Asia to all other continents, except Antarctica.

Studies of higher level relationships place the

Branchiura with either Pentastomida or Ostracoda.

Hypotheses about phylogenetic relationships of either

the genera or species in this group have not been

proposed.

Keywords Fishlice  Fishlouse  Parasites 

Nonindigenous species  Biogeography

Introduction

The subclass Branchiura contains a single family, the

Argulidae, and four valid genera: Argulus Mu

ller,

1785, Chonopeltis Thiele, 1900, Dipteropeltis Calman,

1912, and Dolops Audouin, 1837. Yamaguti (1963)

erected subfamilies within the Argulidae as well as the

family Dipteropeltidae; however, neither subfamily

designations nor the family group name Dipteropelti-

dae are used herein. Branchiurans are ectoparasites of

ﬁshes primarily but occasionally live on amphibians or

invertebrates, and they can move about freely on their

hosts (Stuhlmann, 1891; Yamaguti, 1963; Cressey,

1978; Jackson & Marcogliese, 1995; Poly, 2003).

Species of Branchiura are known by the common

name, ﬁshlouse (plural: ﬁshlice).

Branchiurans are compressed dorsoventrally with a

circular to oval shield-like carapace, a pair of com-

pound eyes anteriorly, four pairs of swimming legs on

the thorax, and a short, unsegmented abdomen poste-

riorly (Fig. 1). Total length of adults ranges from a few

millimeters to just over 30 mm, and adult females tend

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

que, H. Segers &

K. Martens

Freshwater Animal Diversity Assessment

The present paper was intended as a very brief overview of the

known diversity of the Branchiura in freshwaters only, and

regrettably is deﬁcient in mention of much information and

many relevant citations pertaining to the group.

W. J. Poly (&)

Aquatic Biodiversity Research Institute, 11205 Schroeder

Road, Saint Marys, OH 45885, USA

e-mail: wpoly@calacademy.org

W. J. Poly

Department of Ichthyology, California Academy of

Sciences, Golden Gate Park, San Francisco, CA 94118,

USA

123

Hydrobiologia (2008) 595:209–212

DOI 10.1007/s10750-007-9015-3

to be larger than males. Argulus, Chonopeltis, and

Dipteropeltis possess a pair of suction cups (modiﬁed

ﬁrst maxillae) in late juvenile and adult stages, whereas

Dolops retains the larval, claw-like appendages as

adults (Fig. 1). On legs 2–4, male Argulus spp. and

Chonopeltis spp. may have secondary sexual modiﬁ-

cations that are absent on females. Other differences

between the sexes include the presence of a pair of

testes in the male’s abdomen and a pair of spermath-

ecae in the female’s abdomen, a greater number of

support rods in suction cups of females, and also

usually a higher number of sclerites in the support rods

of females (Argulus). Eggs often can be seen in the

thorax of an adult female, and in some species eggs

occur in the thor ax and in the lobes of the carapace as

well. Shapes of respiratory areas, shapes of carapace

and abdomen, number of sclerites in suction cup

support rods and number of support rods per suction

cup (Argulus), features of the second maxillae (espe-

cially basal plate), pigmentation, and secondary sexual

characters of males are some of the most useful

taxonomic characters for distinguishing species (Wil-

son, 1902, 1944; Sikama, 1938; Meehean, 1940; Poly,

2005).

Mating takes place on the host ﬁsh, and later, the

female leaves the host to lay eggs, which are attached

with an adhesive substance to objects , such as rocks,

plants, or sticks. Sperm are stored in the spermathecae

of females in Argulus and presumably in Chonopeltis

and Dipteropeltis , whereas males of the genus Dolops

deposit a spermatophore on the females (Carvalho,

1941; Fryer, 1958, 1960). Further information on the

ecology and morphology of argulids can be found in

Jurine (1806), Clark (1902), Wilson (1902), Calman

(1912), Tokioka (1936), Loro (1964), van Niekerk &

Kok (1989), Rushton-Mellor & Boxshall (1994), and

Van As & Van As (1999).

Species diversity

Dipteropeltis contains one valid species and occurs

only in South America, Chonopeltis has 14 valid

species and is found only in Africa, and Dolops holds

13 valid species with all but two species in South

America. Species within the three aforementioned

genera live in freshwater only. Argulus contains

about 129 valid species and occurs on or around all

continents, except Antarctica, in marin e and estuarine

(n = 44) and freshwater habitats (n = 85). In total, the

diversity of freshwater Branchiura stands at 113

species. In addition, the author is describing at least

12 new species from freshwaters of North America

and Australia. The highest diversity of genera and

species occurs in the Afrotropical and Neotropical

regions (Fig. 2, Table 1). The Nearctic and Oriental

regions contain nearly equal numbers of species,

whereas the Palearc tic and Australasian regions have

much less diversity.

Phylogeny and historical processes

Phylogenetic studies concerning the Branchiura per-

tained to higher level relationships only. Results of

several studies supported a sister group relationship

between the Branchiura and the Pentastomida or the

Ostracoda (discussed in Martin & Davis, 2001;

Wingstrand, 1972; Lavrov et al., 2004; Regier et al.,

2005 and references therein). Phylogenetic studies of

the generic and speciﬁc relationships have not been

attempted yet. Further detailed taxonomic study of the

group will provide much of the data needed for more

meaningful and complete phylogenetic analyses.

Present distribution and main areas of endemicity

Biogeographic analyses of the Branchiura were made

by Fryer (1969) with regard to Dolops, which

displays a Gondwanan distribution (South America,

Fig. 1 Typical body form of a branchiuran. Dorsal (A) and

ventral (B) views of the female holotype of Dolops tasman-

ianus Fryer, 1969. Copyright  CSIRO; Reproduced from the

Australian Journal of Zoology 17: 49–64 (Fryer, 1969) with

permission of CSIRO PUBLISHING, Melbourne, Australia

(http:// www.publish.csiro.au/journals/ajz)

210 Hydrobiologia (2008) 595:209–212

123

Africa, and Tasmania), and the biogeography of

African species of Argulus, Chonopeltis, and Dolops

was discussed by Fryer (1968). Chonopeltis and

Dipteropeltis are endemic to Africa and South

America, respectively. Little else has ever been

published on biogeography of this group. Argulus

japonicus Thiele, 1900 has been introduced from

east/southeast Asia to all other continents, except

Antarctica (Table 1). Native species do not occur

across great distances in most cases, except for some

species in the Afrotropical region, such as Dolops

ranarum (Stuhlmann, 1891) and Argulus africanus

Thiele, 1900 (Fryer, 1968), and for Argulus foliaceus

(Linne

, 1758) in the Pale arctic region. Deﬁnitive

discussions about species’ distributions cannot be

made until additional species are described and

further collecting and taxonomic study reveal true

diversity more accur ately.

Fig. 2 Distribution of species of Branchiura occurring in

freshwater habitats in each of the biogeographic regions of the

world. Argulus japonicus was included in the number of

species for all regions where it is either native (east/southeast

Asia) or introduced. PA––Palearctic, NA––Nearctic, NT––

Neotropical, AT––Afrotropical, OL––Oriental, AU––Austral-

asian, PAC––Paciﬁc Oceanic Islands, ANT––Antarctic

Table 1 Number of species per genus of Branchiura occurring in freshwater habitats in each of the biogeographic regions of the

world

PA NA NT AT OL AU PAC ANT World

Argulus 81821251621 85

Chonopeltis 14 14

Dipteropeltis 11

Dolops 11 1 1 13

Total Argulidae 8 (1) 18 (1) 33 (3) 40 (3) 16 (1) 3 (2) 1 (1) 0 113

The Japanese ﬁshlouse, Argulus japonicus, was included in all regions where it is either native or introduced, but in the overall total,

this species was counted only once. Hence, the total from adding all separate regions is 119, minus 6 occurrences of A. japonicus

equals 113. Number of genera per region in parentheses. PA: Palearctic, NA: Nearctic, NT: Neotropical, AT: Afrotropical, OL:

Oriental, AU: Australasian, PAC: Paciﬁc Oceanic Islands, ANT: Antarctic

Hydrobiologia (2008) 595:209–212 211

123

Human related issues

Branchiurans occasionally attain high densities on

their hosts, resulting in ﬁsh kills in natural bodies of

water (rarely; usually lentic waters) or ﬁsh hatchery/

aquaculture operations (more commonly). Fishlice

also appea r at times in large public aquaria, home

aquaria, or small outdoor ponds and can contribute to

mortality of ornamental ﬁshes.

Acknowledgments The author expresses much appreciation

to the organizers of the meeting on Freshwater Animal

Diversity Assessment held in Mechelen, Belgium for their

careful planning, logistical support, and ﬁnancial assistance,

resulting in an immensely productive and enjoyable session,

and for all their efforts to assemble this volume. The author

also thanks Geoff Boxshall and an anonymous reviewer for

their comments on the manuscript.

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123

FRESHWATER ANIMAL DIVERSITY ASSESSMENT

Global diversity of mysids (Crustacea-Mysida) in freshwater

Megan L. Porter Æ Kenneth Meland Æ

Wayne Price

 Springer Science+Business Media B.V. 2007

Abstract In this article we present a biogeographical

assessment of species diversity within the Mysida

(Crustacea: Malacostraca: Peracarida) from inland

waters. Inland species represent 6.7% (72 species ) of

mysid diversity. These species represent three of the

four families within the Mysida (Lepidomysidae,

Stygiomysidae, and Mysidae ) and are concentrated in

the Palaearctic and Neotropical regions. The inland

mysid species distributional patterns can be explained

by four main groups representing different freshwater

invasion routes: (1) Subterranean Tethyan relicts (24

spp.); (2) Autochthonous Ponto-Caspian endemics (20

spp.); (3) Mysis spp. ‘Glacial Relicts’ (8 spp.); and (4)

Euryhaline estuarine species (20 spp.). The center of

inland mysid species diversity is the Ponto-Caspian

region, containing 24 species, a large portion of which

are the results of a radiation in the genus Paramysis.

Keywords Inland fauna  Freshwater biology 

Mysid  Diversity

Introduction

The order Mysida (Crustacea: Malacostraca: Peraca-

rida), ﬁrst described in 1776 by Mu

ller, contains over

1,000 described species distributed throughout the

waters of the world (Wittmann, 1999). Although

>90% of mysid species are exclusively marine, the

remaining species represe nt either species from

coastal habitats with direct marine connections

(e.g., estuaries, coastal rivers, marine caves) or from

true invasions of inland freshwaters (Audzijonyte

2006; Mauchline, 1980). This review deals with the

zoogeography and historical processes leading to the

current diversity of freshwater mysid species. While

most of the true freshwater mysids occur in ‘conti-

nental’ lacustrine and riverine habitats, many of the

subterranean mysids occur only from island habitats;

for our purposes we have, therefore, chosen to deﬁne

‘inland’ species, as any species with documented

populations occurring in freshwaters (salinity <3 g/l).

Inland mysids range in size from 3 to 22 mm, and due

to the brood pouch present in mature females are

Electronic Supplementary Material The online version of

this article (doi:10.1007/s10750-007-9016-2) contains supple-

mentary material, which is available to authorized users.

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

que, H. Segers and

K. Martens

Freshwater Animal Diversity Assessment

M. L. Porter (&)

Department of Biological Science, University of

Maryland, Baltimore County, 1000 Hilltop Circle,

Baltimore, MD 21250, USA

e-mail: porter@umbc.edu

K. Meland

Department of Biology, University of Bergen, Bergen,

Norway

W. Price

Department of Biology, University of Tampa, Tampa, FL

33606, USA

123

Hydrobiologia (2008) 595:213–218

DOI 10.1007/s10750-007-9016-2

often called ‘opossum shrimp’ (Fig. 1). Most inland

species are nektobenthic (although a few species are

pelagic), omnivorous suspension-feeders, or scaveng-

ers. Some species can be found in large numbers,

serving as an important food source for many other

organisms.

Species diversity

The Mysida contain *1,073 species, only 72 of

which are documented from inland waters (See

supplemental material). Out of the four families

within the Mysida sensu Martin & Davis (2001),

three contain species from inland systems (Tables 1,

2). Two of these families, the Lepidomysidae and the

Stygiomysidae, are monogeneric taxa (Spelaeomysis

and Stygiomysis, respectively) contain ing species

adapted to subterranean habitats (e.g., caves, ground-

water, wells, and crab burrows). While all of the

species in these two families (9 and 7, respectively)

are from inland habitats, many are from island or

coastal systems with direct connections to marine

environments. Nonetheless, 7 Lepidomysidae and 5

Stygiomysidae have made the transition to what can

be considered freshwater environments. In compari-

son, the family Mysidae contains the largest diversity

of inland taxa (60 species, 23 genera), with most of

this diversity found within the subfamily Mysinae,

tribe Mysini (Tables 1, 2).

Zoogeography

The inland mysid fauna are located mainly in the

Palaearctic (39 species, 15 genera) and Neotropical

(20 species, 6 genera) biogeographical regions

(Tables 1, 2; supplementary tables; Fig. 2). Within

freshwater habitats, primarily from these two broad

regions, four historical diversiﬁcation patterns can

Fig. 1 A––Taphromysis bowmani; B––Spelaeomysis sp. (Pho-

tos by E. Peebles, courtesy of the American Fisheries Society)

Table 1 Total number of inland mysid species found in the

major geographical regions from each family (**), subfamily

(*), and tribe () containing at least one inland species. Only

species numbers from each family are used to calculate the

ﬁnal totals for each region. See supplementary tables for

detailed information. PA: Palaearctic; NA: Nearctic; NT:

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

asian; PAC: Paciﬁc & Oceanic Islands; ANT: Antarctic

PA NA NT AT OL AU PAC ANT Total inland species

**Lepidomysidae 1 1 4 0 2 0 0 0 7

**Stygiomysidae 1 0 4 0 0 0 0 0 5

**Mysidae 37 10 12 1 5 1 0 0 60

*Rhopalophthalminae 0 0 0 0 1 0 0 0 1

*Mysinae 37 10 12 1 4 1 0 0 59

Heteromysini 0 1 0 0 0 0 0 0 1

Leptomysini 0 1 1 0 0 1 0 0 2

Mysini 37 8 11 1 4 0 0 0 56

Total 39 11 20 1 7 1 0 0 72

Note: Several species occur in more than one geographical region, resulting in higher numbers of occurrences (79) than species (72)

214 Hydrobiologia (2008) 595:213–218

123