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account for a large percentage of mysid species

diversity.

Tethyan subterranean relicts

All of the subterranean/groundwater mysid genera

considered here (Antromysis ––6 spp., Spelaeomysis––

7 spp., Stygiomysis––5 spp., and the monotypic

Troglomysis vjetrenicensis) are found in a

distributional pattern suggesting a Tethy an origin,

likely colo nizing groundwater habitats due to the uplift

and stranding of marine ancestors during Miocene

regressions of the Tethys and Mediterranean seas

(Boxshall & Jaume, 2000). These hypothesized Tethy-

ian dispersal events resulted in ancient mysid genera

successfully colonizing early regions of Central Amer-

ica, the Caribbean, and Indian and Mediterranean

basins, resulting in widespread genera that were later

isolated, and presently represented by species often

Table 2 Total number of inland mysid genera from the major

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

tribe () containing at least one inland genus. Only genera 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: Australasian; PAC: Paciﬁc & Oceanic

Islands; ANT: Antarctic

PA NA NT AT OL AU PAC ANT Total inland genera

**Lepidomysidae 1 1 10100 0 1

**Stygiomysidae 1 0 10000 0 1

**Mysidae 13 6 41510 0 23

*Rhopalophthalminae 0 0 00100 0 1

*Mysinae 13 6 41410 0 22

Heteromysini 0 1 0000 0 0 1

Leptomysini 0 1 10010 0 2

Mysini 13 4 31400 0 19

Total 15 7 61610 0 25

Note: Several genera occur in more than one geographical region, resulting in higher numbers of occurrences (36) than inland genera

(25)

Fig. 2 Biogeographic

regions indicating the

numbers of inland mysid

species and genera (SP/GN)

found in each: PA––

Palaearctic; NA––Nearctic;

NT––Neotropical; AT––

Afrotropical; OL––Oriental;

AU––Australasian; PAC––

Paciﬁc; ANT––Antarctica;

Grey circles indicate areas

of high biodiversity, i.e., the

Ponto-Caspian region and

the distribution of

Lepidopmysidae and

Stygiomysidae continental

subterranean species (lines

connect disjunct regions of

occurrence)

Hydrobiologia (2008) 595:213–218 215

123

endemic to a single groundwater system. Additionally,

the surface genera Parvimysis (2 spp.) and Surinamysis

(3 spp.), distributed in the Ca ribbean and South

America, are closely related to the genus Antromysis

and may also be a part of this Tethyan distribution.

Autochthonous Ponto-Caspian endemics

The Ponto-Caspian basin, consisting of the Black,

Azov, and Caspian Seas, is composed of inland seas

with complex geological histories dating back to the

Paratethys Sea (20 Mya), including periods as lacus-

trine environments (Banarescu, 1991). The Ponto-

Caspian mysid fauna (autochthonous + ‘glacial re-

licts’, see below) are generally considered the center

of inland mysid species diversity. The autochthonous

mysids that evolved in these enclosed continental

basins, occur in fresh and brackish water portions of

the basins includi ng rivers, lakes, and estuaries, and

are endemic to one basin or are found in parts of all

three (Table 3). Although this fauna consists of seven

genera and 20 species, a large portion of the endemic

mysid dive rsity in these basins is the result of a

radiation in the genus Paramysis (Table 3).

In a phylogeographic study of Limnomysis bene-

deni and six Paramysis species across the Ponto-

caspian region (Audzijonyte

et al., 2006), three main

patterns were identiﬁed: (1) no deep subdivisions

across the entire region, (2) genealogical splits

matching geographical borders among basins, and

(3) divergent lineages occurring only within the

Caspian. The discordant molecular subdivisions

among these co-distributed mysid species suggest

that the similar zoogeographic patterns were formed

at different times (Audzijonyte

et al., 2006).

Mysis ‘glacial relicts’

The genus Mys is comprises 14 species, of which

eight have inland freshwater distributions and can be

divided into two groups: (1) The M. relicta group (M.

relicta, M. diluviana, M. salemaai, M. segerstralei)

with a circumpolar distribution from boreal and

subarctic lakes of the previously glaciated areas of

Europe and North America (Audzijonyte

&Va

ino

2005), and (2) four Caspian Sea endemics (Table 3).

Phylogenetic analyses indicate that the inland Mysis

species are a monophyletic assemblage, sister to

circumarctic marine species (Audzijonyte

et al.,

2005). The separation of freshwater/continental Mysis

spp. and circumarctic marine species is estimated to

have taken place 3–7 Mya (Audzijonyte

et al., 2005),

and does not correspond to the timing in general

hypotheses of continental invasions, such as mid-

Pleistocene glaciation events or mid-Tertiary separa-

tion of the Arctic and Caspian basins (see Va

ino

1995). Furthermore, molecular divergences among

the boreal Nearctic (M. diluviana) and Palearctic (M.

relicta, M. salemaai, M. segerstralei) species and the

Caspian endemics indicate inland colonizations

Table 3 List of species endemic to the Black, Azov, and

Caspian Seas, including the occurrence in each basin (data from

Audzijonyte

, 2006). In addition to the species counted here,

there are at least four additional endemic Ponto-Caspian mysids

(Diamysis mecznikovi, Hemimysis serrata, Paramysis agigen-

sis, Paramysis pontica) whose distributions are unclear or do

not occur in freshwaters and therefore have not been included

in the species list

Black Sea Azov Sea Caspian Sea

‘Glacial-relict’

Mysis amblyops ·

Mysis caspia ·

Mysis macrolepis ·

Mysis microphthalma ·

Autochtonous Ponto-Caspian

Caspiomysis knipowitschi ·

Diamysis pengoi ·

Diamysis pusilla ·

Hemimysis anomala ···

Katamysis warpachowsky ···

Limnomysis benedeni ···

Paramysis baeri ···

Paramysis eurylepis ·

Paramysis grimmi ·

Paramysis incerta ·

Paramysis inﬂata ·

Paramysis intermedia ···

Paramysis kessleri ··

Paramysis kosswigi ·

Paramysis kroyeri ·

Paramysis lacustris ···

Paramysis loxolepis ·

Paramysis sowinskii ··

Paramysis ullskyi ···

Schistomysis elegans ·

216 Hydrobiologia (2008) 595:213–218

123

occurring at different times (A udzijonyte

&Va

ino

2006). M. relicta and M. diluviana are considered the

oldest of Mysis freshwater species that independently

colonized their respective European and American

ranges during early Pleistocene (Va

ino

et al.,

1994). In contrast, M. salemaai and M. segerstralei

are younger closely related species that have pene-

trated freshwaters more recently (Audzijonyte

ino

, 2006). As for the four endemic Caspian

Mysis species, small molecular divergences suggest a

recent, possibly late Pleistocene, sympatric radiation,

possibly driven by adaptation to a deep pelagic

habitat by M. amblyops and M. microphthalma

(Va

ino

, 1995; Audzijonyte

et al., 2005).

Euryhaline estuarine fauna

Most of the remaining mys ids (20 spp., 14 genera) are

euryhaline species with at least one population

occurring in marginal freshwaters. These species

have only very recently invaded freshwaters in

portions of their distributions.

Phylogeny

The taxonomic position and phylogenetic afﬁliations

within the Mysida are currently under debate.

Historically, the Mysida were considered members

of the crustacean superorder Peracarida, placed as a

sister taxon to the Lophogastrida in the order

Mysidacea. More recently, molecular studies have

led to raising the Mysida (and Lophogastrida) to

ordinal rank (Spears et al., 2005). With respect to

those families containing inland fauna, there is also

taxonomic and phylogenetic uncertainty. For exam-

ple, molecular and morphological data show that the

subterranean family Stygiomysidae is more closely

related to the order Mictacea than to other Mysida

families, suggesting that they be removed from the

order Mysida and placed within a separate order,

Stygiomysida, comprising the families Stygiomysi-

dae and Lepidomysidae (Meland & Willassen, 2007).

Conservation issues

At least 19 inland mysid species are associated with

groundwater habitats (caves, wells, and crab burrows)

having very limited areas of distribution that are

highly susceptible to pollution from the surface.

These species in particular are in need of assessment

for conservation ranking, as they are often found in

aquifers important to local communities as a source

of freshwater and may serve as indicators of water

quality. Many freshwater mysid species have also

served an important role in both US and European

ﬁsheries, where they have been introduced into lakes

and reservoirs to serve as food for commercially

important ﬁsh species (Mordukhai-Boltovskoi, 1979;

Northcote, 1991).

In contrast, the autochthonous Ponto-Caspian

mysids are currently invading aquatic ecosystems of

Northern Europe as a result of human activities (De

Vaate et al., 2002; Leppakoski et al., 2002). The

impact of invasive mysid species on native lacustrine

and riverine ecosystems can be large, including a

severe reduction in zooplankton abundance, with

concomitant negative effects on higher consumers

(Spencer et al., 1991; Ketelaars et al., 1999).

Acknowledgments We thank R. Va

ino

for advice

concerning the Ponto-Caspian mysid fauna and for the

helpful suggestions of one anonymous reviewer. T. bowmani

and A. almyra images by E. Peebles from ‘Common and

Scientiﬁc Names of Aquatic Invertebrates from the United

States and Canada: Crustaceans’ are courtesy of the American

Fisheries Society. This work was supported by NSF grant

DEB-0206537.

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218 Hydrobiologia (2008) 595:213–218

123

FRESHWATER ANIMAL DIVERSITY ASSESSMENT

Global diversity of spelaeogriphaceans

& thermosbaenaceans (Crustacea; Spelaeogriphacea

& Thermosbaenacea) in freshwater

D. Jaume

 Springer Science+Business Media B.V. 2007

Abstract Spelaeogriphaceans and thermosbaena-

ceans are two orders of eyeless, unpigmented perac-

arid crustaceans represented by very few species from

subterranean waters. Spelaeogriphaceans occur only

in continental waters, either running or still, in

limestone or sandstone caves, or in calcrete aquifers.

The four species known are limnic except one

occurring in slightly brackish water loosely associ-

ated with an endorheic basin. The Thermosbaenacea

are primarily marine, with only 18 species recorded

in limnic conditions or in brackish inland waters

whose salinity does not derive from dilution of

seawater. They occur in limestone caves, the inter-

stitial medium associated to alluvial deposits, or in

thermo-mineral springs. Spelaeogriphaceans are

found on the southern continents, in ancient cratons

not affected by sea transgressions at least since the

Early Cretaceous, when Gondwana started to break-

up. The former integration of these terranes into

Gondwana suggests that the penetration of spelaeo-

griphaceans in continental waters took place previous

to the fragmentation of this super continent (starting

ca. 140 Ma), and that their current distribution

pattern was driven by continental drift. The distribu-

tion of the Thermosbaenacea matches precisely the

area covered by the ancient Tethys Sea or its

coastlines. They are most probably relicts of a once

widespread shallow-water marine Tethyan fauna

stranded in interstitial or crevicular groundwater

during marine regressions.

Keywords Freshwater  Global assessment 

Species richness  Peracarida  Crustacea

Introduction

Spelaeogriphaceans and thermosbaenaceans are two

orders of eyeless, unpigmented peracarid crustaceans

represented by very few species from subterranean

waters. The Spelaeogriphacea Gordon, 1957 occur

only in continental waters, either running or still, in

limestone or sandstone caves, or in calcrete aquifers.

All species are limnic except one occurring in slightly

brackish water loosely associated to an endorheic

basin.

Thermosbaenacea Monod, 1927, in contrast, seem

to be primarily marine although only 5 out of the 34

species known are euhaline, living in lava tubes or in

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

que, H. Segers &

K. Martens

Freshwater Animal Diversity Assessment

Electronic supplementary material The online version of

this article (doi:10.1007/s10750-007-9017-1) contains

supplementary material, which is available to authorized

users.

D. Jaume (&)

IMEDEA (CSIC-UIB), Instituto Mediterra

neo de Estudios

Avanzados, c/Miquel Marque

s 21, Esporles, Illes Balears

07190, Spain

e-mail: d.jaume@uib.es

123

Hydrobiologia (2008) 595:219–224

DOI 10.1007/s10750-007-9017-1

the interstices between submersed coarse sand grains,

volcanic tephra or coral rubble (Wagner, 1994). Most

taxa occur in the anchialine environment associa ted

with marine coastal areas, where water salinity varies

sharply across the water column and derives from

dilution of sea water. Only 18 species have been

recorded in limnic conditions or in brac kish inland

waters whose salinity does not derive from dilution of

seawater. They occur in limestone caves, the inter-

stitial medium associated to alluvial deposits or in

thermo-mineral springs.

The spelaeogriphacean body is roughly cylindri-

cal, with the cephalothorax incorporating only the

ﬁrst thoracomere (which carries the maxillipeds)

and with a short, posteriorly directed carapace cover-

ing only part of the second thoracomere dorsally

(Fig. 1A). The pereion consists of seven free pereio-

meres, each with a pair of similar biramous stenop-

odial pereiopods, and the pleon comprises six free

pleomeres, each with a pair of well-developed

biramous pleopods. The telson is free, articulated to

the 6th pleomere. The most remarkable autapom or-

phy of the group is the transformation of the exopods

of the posterior pairs of pereiopods into non-setose,

respiratory paddles.

Contrary to spelaeogriphaceans, whose inclusion

in the Peracarida is undeniable since brooding

females display a thoracic ventral marsupium formed

by oo

stegites (= foliaceous medial exte nsions of the

pereiopodal coxae), the inclusion of the Thermos-

baenacea in this group is debatable since here the

embryos are carried in a dorsal brood pouch derived

from the carapace (Richter & Schol tz, 2001). Apart

of this, the rest of features of the therm osbaenacean

body plan is roughly similar to the spelaeogripha-

ceans (Fig. 1B); just notice that the pleopods are now

vestigial and present only on pleomeres 1 and 2, and

that the condition of the posterior extension of the

carapace varies from covering the second thoraco-

mere only (in males and non-brooding females of

Thermosbaenidae, Monodellidae and Halosbaenidae)

to cover all thoracomeres (Tulumellidae). Thermos-

baena mirabilis is a highly modiﬁed therm osbaena-

cean that separates from the ordinary morphology of

the group in having a pleotelson (formed by the 6th

pleomere and telson), and the display of only ﬁve

pairs of pereiopods.

The mouthparts of spelaeogriphaceans and ther-

mosbaenaceans conform to a functional series of

scrapers in an arrangement considered to ﬁt for

Fig. 1 General aspect of Spelaeogriphacea and Thermosbaen-

acea. (A) Spelaeogriphus lepidops Gordon 1957, a spelaeogri-

phacean from Table Mountain, South Africa (after Gordon,

1960). (B) male of Tethysbaena atlantomaroccana (Boutin &

Cals, 1985), a freshwater thermosbaenacean from Morocco;

notice that ornamentation of pereopodal exopods is omitted

from ﬁgure (after Cals & Boutin, 1985)

Table 1 Global diversity of Spelaeogriphacea

Distribution Habitat Salinity range

Spelaeogriphidae

Mangkurtu Poore & Humphreys, 1998

mityula Poore & Humphreys, 1998 NW Australia Borehole wells Limnic

kutjarra Poore & Humphreys, 2003 NW Australia Borehole wells Oligohaline

Potiicoara Pires, 1987

brasiliensis Pires, 1987 Brazil Caves Limnic

Spelaeogriphus Gordon, 1957

lepidops Gordon, 1957 South Africa Caves Limnic

220 Hydrobiologia (2008) 595:219–224

123

scraping small food particles from a substrate (Fryer,

1964).

Species diversity

The Spelaeogriphacea comprises a single family, the

Spelaeogriphidae Gordon, 1957 with three genera:

Speleaeogriphus and Potiicoara (both monotypic),

and Mangkurtu, with two species (Table 1). Three

fossil forms from the Carboniferous of Canada, the

Upper Jurassic of China, and the Lower Cretaceous

of Spain are treated also as spelaeogriphaceans by

some authors (Schram, 1974; Shen et al., 1998;

1999). Nevertheless, none of these fossil forms

preserves the diagnostic pereopodal exopods, and

their body tagmosis and short carapace do not

preclude their allocation to other peracaridan orders.

The Thermosbaenacea embraces four families, of

which only three include non-marine species

(Table 2). The family Thermosbaenidae is mono-

typic. The Monodellidae includes two genera:

Table 2 Global diversity of non-marine Thermosbaenacea

Distribution Habitat Salinity range

Thermosbaenidae

Thermosbaena Monod, 1924

mirabilis Monod, 1924 Tunisia Thermo-mineral springs Presumed oligohaline

Monodellidae

Tethysbaena Wagner, 1994

juriaani Wagner, 1994 Dominican Rep. Anchialine limestone wells

and spring

Limnic to oligohaline

gaweini Wagner, 1994 Dominican Rep. Wells in alluvial sediments Limnic to oligohaline

haitiensis Wagner, 1994 Haiti Spring, wells and alluvial

sediments

Limnic to oligohaline

juglandis Wagner, 1994 Haiti Wells in alluvial sediments Limnic

lazarei Wagner, 1994 Cuba Cave and river interstitial Limnic to oligohaline

tinima Wagner, 1994 Cuba Wells in limestone Limnic

calsi Wagner, 1994 Saint John; Tortola (British Virgin

Islands)

Wells in alluvial deposits Limnic to oligohaline

relicta (Po

r, 1962) Israel Thermo-mineral springs Oligohaline to hyperhaline

somala (Chelazzi & Messana,

1982)

Somalia Wells in limestone Oligohaline to polyhaline

(evaporites)

atlantomaroccana (Cals &

Boutin, 1885)

Morocco Wells in alluvial deposits Presumed limnic

tarsiensis Wagner, 1994 Spain Well in alluvial deposit Limnic

texana (Maguire, 1965) Texas (U.S.A.) Artesian wells Limnic

vinabayesi Wagner, 1994 Isla Juventud (Cuba) Cave Limnic

Halosbaenidae

Limnosbaena Stock, 1976

ﬁnki (Mestrov & Lattinger-

Penko, 1969)

Bosnia-Hercegovina; Italy Interstitial of river alluvia;

caves

Limnic

sp. Wagner, 1994 France Well Limnic

Halosbaena Stock, 1976

tulki Poore & Humphreys, 1992 NW Australia Calcrete aquifers Oligohaline (evaporites)

Theosbaena Cals & Boutin,

1985

cambodjana Cals & Boutin,

1985

Cambodia; Thailand Caves Limnic

Salinity tolerance ranges: Limnic: <0.5%; Oligohaline: 0.5–5%; Polyhaline: 18–30%; Hyperhaline: >40%

Hydrobiologia (2008) 595:219–224 221

123

Monodella (monotypic) and Tethysbaena (23 spe-

cies), of which only 13 species of Tethysbaena are

considered here as non-marine. The Halosbaenidae

consists of the truly limnic genera Limnosbaena (two

species, one of them not formally described) and

Theosbaena (monotypic), plus the genus Halosbaena,

with two marine species plus one from non-marine,

brackish inland waters. The fourth thermosbaen acean

family Tulumellidae, comprising the single genus

Tulumella (three species), is fully marine. There is

no fossil record of the Thermosbaenacea known to

date.

Many species of thermosbaenaceans have been

reported from oligohaline water in coastal aquifers or

anchialine environments only, frequently around

haloclines where water salinity changes abruptly

from limnic-oligohaline to marine euhaline. These

taxa (shown in Appendix Table 1) are not included in

the total estimate of non-marine species since none

has been reported from pure fresh waters, nor from

marine euhaline water either.

Present distribution and historical processes

Living spelaeogriphaceans appear associated with

freshwater in southern continents, in ancient cratons

not affected by sea transgressions at least since the

Early Cretaceous, when Gondwana started to break-

up (Fig. 2; Table 3; Appendix Fig. A). Spelaeogri-

phus lepidops is known only from two caves in South

Africa excavated in Ordovician quartzites (Gordon,

1957). Potiicoara brasiliensis, from two caves in

Upper Proterozoic limestone at Mato Grosso do Sul

(Brazil; Pires, 1987). And the two species of Mang-

kurtu, from borehole wells in calcrete of Middle to

Late Tertiary age on north-western Australia,

although these deposits overlie and are in direct

contact with Early Cretaceous alluvial conglomerates

(Poore & Humphreys, 1998; 2003). The former

integration of these terranes in Gondwana suggests

that the penetration of spelaeogriphaceans in conti-

nental waters took place previous to the fragmenta-

tion of this supercont inent (starting ca. 140 Ma), and

Fig. 2 Global distribution of Spelaeogriphacea and continen-

tal water Thermosbaenacea (Species number/Genus number).

PA––Palaearctic, NA––Nearctic, NT––Neotropical, AT––

Afrotropical, OL––Oriental, AU––Australasian, PAC––Paciﬁc

Oceanc Islands, ANT––Antarctic

222 Hydrobiologia (2008) 595:219–224

123

that their current distribution pattern was driven by

continental drift.

The distribution of the order Thermosbaenacea

matches precisely the area covered by the ancient

Tethys Sea or its coastlines (Fig. 2, 3; Table 3; see

also Appendix Figs A and B). They are most

probably relic ts of a once widespread shallow-water

marine Tethyan fauna, stranded in interstitial or

crevicular groundwater during episodes of marine

regression. The amphi-Atlantic distribution of Teth-

ysbaena, or the so-called ‘‘full Tethyan track’’

displayed by Halosbaena (with species in the Carib-

bean, Canary Islands and Australia) suggest the

origin of these genera dates back to at lea st the ﬁnal

opening of the Atlantic (95 Ma) and the maximum

extent of the Tethys sea (120 Ma), respectively. The

timing of colonisation of continental waters by

thermosbaenacean lineages is probably more recent

(all inland water stations fall within areas covered by

the sea in Pliocene or more recent times), and the

speciation process can have been peripatric.

The distribution of the Thermosbaenidae casts

doubts on whether they represent a separate family

from its closest relative, the Monodellidae. The

single representative of the family dwells in a

Tunisian thermo-mineral spring placed in a zone

recently covered by the sea (Late Pliocene). This

station falls within the area covered by the Monod-

ellidae, which comprises a widespread amphi-Atlan-

tic genus (indicative of the ancient origin for the

family; see above) present also in N Africa (Teth-

ysbaena), plus a second genus Monodella, mono-

typic and kn ow only from a single-anchialine cave in

Italy. The Thermosbaenidae are probably a young,

Fig. 3 Translocation of present distribution of Thermosbaenacea (including both marine and continental water taxa) to an Aptian

(Lower Cretaceous; 120 Ma) palaeo-coastline map. Shaded areas denote emerged lands

Table 3 Global and per biogeographic region diversity (species number) of Spelaeogriphacea and non-marine Thermosbaenacea

PA NA NT AT OL AU PAC World

Spelaeogriphacea

Spelaeogriphidae – – 1 (1) 1 (1) – 2 (1) – 4 (3)

Thermosbaenacea

Thermosbaenidae 1 (1) – – – – – – 1 (1)

Monodellidae 3 (1) 1 (1) 8 (1) 1 (1) – – – 13 (1)

Halosbaenidae 2 (1) – – – 1 (1) 1 (1) – 4 (3)

Total Thermosbaenacea 6 (3) 1 (1) 8 (1) 1 (1) 1 (1) 1 (1) – 18 (5)

In brackets, number of genera. No records of these groups exist from Paciﬁc oceanic islands or Antarctica. PA: Palaearctic, NA:

Nearctic, NT: Neotropical, AT: Afrotropical, OL: Oriental, AU: Australasian, PAC: Paciﬁc Oceanc Islands, ANT: Antarctic

Hydrobiologia (2008) 595:219–224 223

123

highly modiﬁed member of the family Monodellidae

adapted to an exceptional habitat (hot springs; see

Wagner, 1994: 317).

Acknowledgement This is a contribution to Spanish MEC

project CGL2005-02217/ BOS.

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