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density (Petersen & Luxton, 1982) and in species

number (Stork, 1988; Stork & Blackburn, 1993), even

if their participation to the biomass remains low

(Petersen, 1994). Though no Collembola are adapted

to life in water, many have developed successful

mechanisms to exploit water surface or water ﬁlm in

extremely humid environments. First, Collembola

have a strongly water-repellant body cuticle which

allows them to ﬂoat on the surface of water (Noble-

Nesbitt, 1963). Thus, ﬂooding acts as a major

dispersal agent for springtails. Second, claws and

mucro are often strongly modiﬁed in hydrophilous

Collembola, especially among neustonic species.

Claws tend to becom e thin and elongated compared

to thos e of non-hydrophilous relatives. This trend is

observed in many species living in habitats where

free water is abundan t (along streams, around ponds,

in caves), whatever the group (Christiansen, 1961;

Deharveng, 1988; D’Haese, 2003). Claw elongation

is considered an adaptive character as it appears

independently in phyletically remote lineages. Chris-

tiansen (1965) investigated its functional basis, and

showed that it allowed cave Collembola to walk on

water. The same is probably true for non-cave

species, which exhibit the same modiﬁcation of the

claw morphology. Another morphol ogical modiﬁ ca-

tion limited to neustonic Symphypleona and

Neelipleona, consists in a considerable enlargement

of the mucro.

Water-dependent Collembola may be allocated to

four main categories according to the conditions of

water availability, and the relative importance of non-

hydric factors of their habitats.

(1) Epigean hydrophilous species. They live on the

surface of freshwater, on soil or on low

vegetation close to water.

(2) Cryophilous species. They are active on, under

or at the edge of permanent snow ﬁelds or

glaciers. They are also present on or near small

water puddles which forms at the edge of snow

ﬁelds or on ice surface during summer. Perma-

nently cold habitats occur either at high

elevation in temperate and tropical mountains,

or at high latitudes (Antarctic and Arctic

regions). Collembola are relatively diversiﬁed

in such extreme habitats, where they tend to

represent the largest com ponent of the inverte-

brate fauna (Cass agnau, 1961).

(3) Cave hydrophilous species. They are strictly

associated with water-saturated atmosphere of

large subterranean voids (caves and ‘‘Milieu

Souterrain Superﬁciel’’, Juberthie, 2000).

Collembola represent the most diversiﬁed and

numerous invertebrates in these subterranean

habitats. Thibaud (1970) showed experimentally

that several cave Hypogastruridae cannot sur-

vive more than a few hours below 96% of

relative humidity. Collembola only found on or

very close to free water (waterfall, underground

stream, ﬁlm of water on rocks) with extremely

elongate appendages and claws are the most

sensitive (Sket, 2004).

(4) A fourth category, interstitial hydrophilous

species, is likely to exist inside humid sandy

sediments, at the level of water table. However,

ecological data are lacking to ascertain whether

some of the numerous minute interstitial species

which have been discovered these last two

decades (Thibaud & Christian, 1997) may be

considered as hydrophilous, and they will not be

dealt with here.

The ecological categories deﬁned above are more

or less relevant to anchialine or brackish water fauna

as well. The tolerance to freshwater of most euryha-

line Collembola is unknown but a few Pseudanurida

and Yuukianura are found both near marine water and

near inland streams.

All ecological categories listed above concern

Collembola which may live at the surface of water or

Fig. 1 Isotomurus alticolus (Carl) from Austria, non-cave (a)

and cave (b) specimens (after Stach, 1947)

330 Hydrobiologia (2008) 595:329–338

123

ice, but there also are records of species collected

underwater, including sea water (Jacquemart & Jac-

ques, 1980). Some records are probably pollution of

water samples by epigean species. But some species

(not necessarily hydrophilous) have been experimen-

tally shown to be able to survive submersion in sea

water for days (Coulson et al., 2002). This phenom-

enon is more documented among freshwater species.

Some Onychiuridae, Tomoceridae, Entomobryidae

(Ruppel, 1953) and Symphypleona (Falkenhan & Fox,

1932; Delamare Deboutteville, 1952) may live a few

days underwater. Thibaud (1970) tested different

Hypogastruridae species in this respect. The animals

were able to move and survive under water up to

40 days after hatchin g and up to 36 days for adults (in

Typhlogastrura balazuci, a cave-dwelling species),

but were usually unable to moult (though moulting of

Ceratophysella armata was obtained by Britt (1951)).

However, there is no evidence so far that the total life

cycle of any species may occur underwa ter. In the

ﬁeld, eggs may hatch after long period of ﬂooding

(Beck, 1976; Tamm, 1984). To which extent this

ability may contribute to explain the rapid recover of

some riparian communities after ﬂooding, as docu-

mented in Russell et al. (2004) and Russel & Griegel

(2006), remai ns to be explored.

Taxonomic diversity

With 7,761 species and 643 genera (Janssens, 2006),

including 2,000 species in Europe (Fauna Europaea,

2004), Collembola are not among the most speciose

groups of living invertebrates. However, being

among the less known, the group experiences a high

annual rate of species richness increase (Deharveng,

2004), and its relative contribution to global biodi-

versity is growing more than that of most other

invertebrates, with a total number of species possibly

as high as 50,000 according to Hopkin (1997, 1998).

Collembola are distributed in four orders (Podur-

omorpha, Entomobryomorpha, Symphypleona and

Neelipleona) and 30 families (Deharveng, 2004). The

proportion of hydrophilous species is difﬁcult to

establish, because ecological and biological data on

most species, especially in the tropics, are lacking.

An analysis based on 3,788 species (half the total

number of Collembolan species) for which a minimal

information on habitats was available or was

extrapolated from our knowledge of the concerned

genera, allows to recognize 109 marine species, 414

freshwater species in three of the four categories

deﬁned above (103 epigean hydrophilous, 26 cryoph-

ilous, 285 cave hydrophilous), and 2 marine species

which are also freshwater hydrophilous. Tables 1 and

2 reﬂect the general pattern of hydrophilous taxo-

nomic richness in the group acro ss the four orders of

Collembola and the different biogeographical regions

on Earth. The contribution of the different orders to

the biodiversity of each habitat is quite uneven, and

the same degree of unevenness is retrieved inside the

orders themselves. Among Entomobryomorpha for

instance, Isotomidae bring about half of the total

number of hydrophilous Collembola , including most

of the cryophilous ones, but it has practically no cave

species. Conversely, Entomobryidae, Oncopoduridae

and Tomoceridae, all three Entomobryomorpha fam-

ilies, contribute signiﬁcantly to cave biodiversity, but

nothing for other habitats. Among biogeographical

regions, striking patterns are emerging. Northern

temperate and boreal regions (Nearctic and Palaearc-

tic) have about 15% of hydrophilous species in their

fauna, while tropical and southern hemisphere regions

have 1–6% (4% when pooled together).

However, these values of species richness are

rapidly changing. In Europe, we begin to realize that

the classical Isotomurus palustris Mueller, 1776, the

most abundant ‘‘specie s’’ of holarctic temperate

wetlands, is probably a cluster of closely related forms

that are awaiting description (Cassagnau, 1987;

Deharveng & Lek, 1993; Carapelli et al., 2001).

Hydrophilous species of several other genera have

been overlooked in the literature (Pseudisotoma

microchaeta Cassag nau, for instance) or were recently

discovered in Europe (Isotomiella hygrophila Ster-

zynska & Kaprus, 2001). According to Murphy (1990),

the Sminthurididae (genera Sminthurides, Sphaeridia

and their allies) which are dominant in damp habitats

of the tropics may also encompass a large diversity

of undescribed species (Murphy, 1966). This points

to the necessity of a taxonomic re-appraisal of

existing species, as well as increased sampling effort.

Origin and phylogeny of Collembola

Collembola provided the ﬁrst known hexapods fossils

from the early Devonian (Hirst & Maulik, 1926;

Hydrobiologia (2008) 595:329–338 331

123

Table 1 Number of Collembolan species in the different regions of the world (total, WDpt or freshwater-dependent species, including Epig or epigean hydrophilous species, see

Introduction) and the four orders of Collembola

Number of Species PA NA AT NT OL

Total WDpt Epig Total WDpt Epig Total WDpt Epig Total WDpt Epig Total WDpt Epig

Entomobryomorpha 928 151 33 168 22 13 147 1 1 245 8 3 319 24 5

Neelipleona 6 3 1 4 0 0 3 0 0 4 0 0 2 0 0

Poduromorpha 999 145 16 190 17 12 97 2 1 247 10 4 203 1 0

Symphypleona 169 39 18 54 10 7 49 3 3 59 10 10 41 9 8

Total 2,102 338 68 416 49 32 296 6 5 555 28 17 565 34 13

Number of Species AU PAC ANT Total

Total WDpt Epig Total WDpt Epig Total WDpt Epig Total WDpt Epig

Entomobryomorpha 113 4 4 107 2 2 23 0 0 1,711 190 43

Neelipleona 2 0 0 4 0 0 1 0 0 10 3 1

Poduromorpha 91 0 0 84 0 0 26 1 1 1,677 157 19

Symphypleona 57 2 2 30 1 1 6 0 0 390 64 40

Total 263 6 6 225 3 3 56 1 1 3,788 414 103

Values derived from a subset of 3,788 species of Collembola. The main bias is the inclusion of Nepal in the palaearctic region, whereas its Collembolan fauna is relatively well

documented and mostly of tropical or tropical montane afﬁnities. PA—Palaearctic; NA—Nearctic; NT—Neotropical; AT—Afrotropical; OL—Oriental; AU—Australasian;

PAC—Paciﬁc Oceanic Islands; ANT—Antarctic

332 Hydrobiologia (2008) 595:329–338

123

Cassagnau, 1990). However, phylogenetic analyses

cannot integrate this information because the known

fossils are in too poor condition, and we have to rely

entirely on extant species to reconstruct evolutionary

history of the group. Collembola were long consid-

ered as a group of Insects. Actual ly, they represent a

class in their own, of controversial afﬁnities among

Arthropods. A detailed phylogeny of Collembola

based on morphological and molecular characters has

been produced by D’Haese (2002, 2003, Fig. 2), who

tested the hypothesis of an aquatic origin for the

group. Physiological data suggest that Collembola

might have evolved from marine ancestors (haemol-

ymph with high osmotic pressure and mainly

composed of inorganic salts), but this is still a matter

of debate in the broader context of hexapod origin

(Little, 1983, 1990). The secondary transition from

terrestrial to freshwater habitats is physiologically

more convincing. The phylo geny of D’Haese (2003)

shows that life on water was not ancestral but

derived, supporting a terrestrial (edaphic) origin for

the springtails. Among terrestrial Collembola, those

lacking molar plate on the mandible (Neanuridae,

Brachystomellidae, Odontellidae) are supposed to

ingest the water ﬁlm which surrounds soil particles,

rich in bacteria and Protozoa (‘‘suspension feeders’’,

Adams & Salmon, 1972). This aquatic type of

feeding, considered as derived phylogenet ically, has

however rarely led to adaptation to aquatic life.

Actually, only a few Collembolan lineages developed

adaptations to aquatic life, and these adaptations

probably arose independently. But they have been

surprisingly successful. Podura aqu atica Linne

1758, the only species of the family Poduridae, is

extremely frequent and abundant in northern temper-

ate countries. The genus Isotomurus has undergone a

tremendous diversiﬁcation in the Holarctic region,

colonizing all kinds of damp habitats. Sminthurides,

often very abundant near and on freshwater world-

wide, exhibits an intermediate level of species

diversiﬁcation.

Geographical distribution

Hydrophilous Collembola are present worldwide

(Thibaud & Massoud, 1986; Christiansen & Snider,

1996). However, the scarcity of information about

this category of springtails prevents any sound

Table 2 Number of Collembolan genera in the different regions of the world and the four orders of Collembola

Number of Genera PA NA AT NT OL AU PAC ANT Total

WDpt Epig WDpt Epig WDpt Epig WDpt Epig WDpt Epig WDpt Epig WDpt Epig WDpt Epig WDpt Epig

Entomobryomorpha 31 13 10 5 1 1 4 1 6 1 1 1 1 1 0 0 33 13

Neelipleona 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1

Poduromorpha 35 6 7 4 2 1 8 4 1 0 0 0 0 0 1 1 35 7

Symphypleona 4 4 5 4 2 2 3 3 3 2 2 2 1 1 0 0 9 8

Total 71 24 22 13 5 4 15 8 10 3 3 3 2 2 1 1 78 29

Hydrobiologia (2008) 595:329–338 333

123

biogeographical analysis. Data for tropical Africa are

limited to Gambia (Murphy, 1960). For tropical Asia,

almost all original information comes from Singapore

(Murphy, 1990), with a few from Indonesia (Hands-

chin, 1931). The work of Deharveng & Bedos (2004)

on aquatic Collembola of Malaysia is a simple

synthesis of existing literature data. Publications not

speciﬁcally devoted to water-dependent Collembola,

especially the excellent taxonomic works of Yosii

(1971) on tropical Asia, remain elusive about the

ecology of species. Many distributional data for the

new world are found in the works of Christiansen &

Bellinger (1998), and Mari Mutt & Bellinger (1990),

but species ecology is not documented. The compre-

hensive study of Fjellberg (1985) about Alaskan

Poduromorpha, more informative in this respect, is

geographically more limited. The informat ion avail-

able about Australian and Subantarctic Collembola

rarely include data about the dependence of species

on water (Deharveng, 1981; Greenslade, 1994). In

Europe, several important works deal with Collem-

bolan fauna of wetlands (e.g. Strenzke, 1955; Palissa,

2000), and may be used for regional biogeography

but some are taxonomically outdated. Detailed

regional fauna (Fjellberg, 1994; Jordana et al.,

1997), local ecological studies (Deharveng & Lek,

1995; Lek-Ang et al., 1999; Sterzynska & Ehrnsber-

ger, 1999; Lek-Ang & Deharveng, 2002; Russell

et al., 2004; Russel & Griegel, 2006) as well as the

classical works of Stach (for instanc e on Isotomidae

in 1947, on Sminthuridae in 1956) and Gisin (1960)

provide sound ecological information on water-

dependent species. Yosii published many articles

(1977) on Japanese Collembola, with very limi ted

information or no data on species ecology. The

synopses of the Palaearctic fauna (Zimdars & Dun-

ger, 1994; Bretfeld, 1999; Potapov, 2001; Thibaud

et al., 2004) put together a huge amount of ecological

Fig. 2 A phylogeny of Collembola based on morphological data after D’Haese (2003). Water-adapted specie s are indicated in bold

(4 marine species, and 3 freshwater-dependent species: Sminthurides sp., Podura aquatica and Anurida granaria).

334 Hydrobiologia (2008) 595:329–338

123

as well as distributional information, and represent

the richest source of information for the old world

(Fig. 3).

In spite of this uneven information, available data

point to a surprising contrast between tropical and

non-tropical regions in each ecological category. Two

of the main genera of tropical, non-interstitial, fully

marine Collembola (Pseudanurida, Oudemansia) are

widely distributed across at least two continents, but

are absent in temperate and boreal regio ns, with the

noticeable exception of the Koreo-Japanese area.

Conversely, Friesea, Anuridella, Halisotoma, well-

represented in the holarctic region, are rare or absent

in marine habitats of the tropics. Similar contrast is

retrieved among interstitial forms (Thibaud & Chris-

tian, 1997), some of which are probably hydrophilous

as well. Among epigean hydrophilous species,

Isotomurus is dominant in non-tropical regions while

Sminthurides is more diversiﬁed in the tropics and the

southern hemisp here. Among cave restricted species

the contrast is also striking between temperate

regions, where Onychiuridae and Hypo gastruridae

are well diversiﬁed, and tropical regions, where these

genera are virtually absent, but Paronellidae very

diversiﬁed in caves.

Aquatic Collembola include a few widespread

(like Podura aquatica ) and many endemic species.

Endemism patterns are extremely contrasted among

regions, habitats and taxa, in terms of proportion of

endemics in the fauna and range size of the species.

Considering the species with a known range smaller

than 500,000 sq km as endemic, and keeping in mind

that distributional information is still poor for ascer-

tain endemic status of many species in the tropics, a

few general features nevertheless emerge from

available data.

Possibly as a result of quaternary glaciations,

hydrophilous endemics seem to be absent in boreal

and northern temperate regions of Europe. In con-

trast, Mediterranean regions exhibit a high richness in

endemics, many of them undescribed (unpublished

data). At the world scale, high endemi city areas for

hydrophilous Collembola are difﬁcult to spot with

precision given the huge geographical gaps in our

Fig. 3 Global distribution of freshwater-dependent species of

Collembola identiﬁed from a subset of 3,788 species (species

number/genera number). PA—Palaearctic; NA—Nearctic;

NT—Neotropical; AT—Afrotropical; OL—Oriental; AU—

Australasian; PAC—Paciﬁc Oceanic Islands; ANT—Antarctic

Hydrobiologia (2008) 595:329–338 335

123

knowledge. However, supra-speciﬁc diversity peaks

in at least three areas: the Far-East of the Palaearctic

region (Japan, Korea, Primorye), the Pyreneo-Canta-

bric range and the Dinarides. The Far-East

Palaearctic hot- spot reﬂects the mixing of mostly-

tropical lineage s with palaear ctic ones. The two other

regions, located in western Europe, are world hot-

spots for cave biodiversity, perhaps as a result of an

history of available habitats and high productivity

levels longer than in surrounding regions (Culver

et al., 2006).

The link between habitats and endemicity is also

obvious. Endemicity among marine hydrophilous

Collembola is lower than among freshwater ones.

Marine Collembola have in particular several species

widespread in Europe or in large regions of the world.

The less fragmented structure of littoral habitats may

explain this pattern. Not surprisingly, endemism rates

vary also greatly in the recognized categories of

freshwater Collembola, in relation to the proneness of

habitats to fragmentation. Several monospeciﬁc and

phyletically isolated genera exist among cave

hydrophilous springtails (Gnatho folsomia in the

Austrian Alps, Bessoniella and Ongulogastrura in

the Pyrenees, Ongulonychiurus in the Cantabrian

range, are the most remarkable). Subterranean hab-

itats have also many endemi c species belonging to

more widespread genera. Cryophilous springtails

have only one endemic genus (Gnathisotoma,

restricted to the Pyrenees) with two species; all other

endemics of this ecol ogical category belong to non-

endemic genera. No genus is know n as endemic of

epigean wet habitats. Most are widespread, with

either one or a few species of very large distribution

(Podura and Pseudobourletiella, mostly in northern

hemisphere, for instance), or several largely distrib-

uted species (Sminthurides), or many closely related

and often endemic species ( Isotomurus). Many

endemic species are expected in this last genus, as

a result of a recent re-appraisal of its taxonomic

forms (Carapelli et al., 2001).

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123

FRESHWATER ANIMAL DIVERSITY ASSESSMENT

Global diversity of mayﬂies (Ephemeroptera, Insecta)

in freshwater

Helen M. Barber-James Æ Jean-Luc Gattolliat Æ

Michel Sartori Æ Michael D. Hubbard

 Springer Science+Business Media B.V. 2007

Abstract The extant global Ephemeroptera fauna is

represented by over 3,000 described species in 42

families and more than 400 genera. The highest generic

diversity occurs in the Neotropics, with a correspond-

ingly high species diversity, while the Palaearctic has

the lowest generic diversity, but a high species

diversity. Such distribution patterns may relate to how

long evolutionary processes have been carrying on in

isolation in a bioregion. Over an extended period, there

may be extinction of species, but evolution of more

genera. Dramatic extinction events such as the K-T

mass extinction have affected current mayﬂy diversity

and distribution. Climatic history plays an important

role in the rate of speciation in an area, with regions

which have been climatically stable over long periods

having fewer species per genus, when compared to

regions subjected to climatic stresses, such as glacia-

tion. A total of 13 families are endemic to speciﬁc

bioregions, with eight among them being monospeciﬁc.

Most of these have restricted distributions which may

be the result of them being the relict of a previously

more diverse, but presently almost completely extinct

family, or may be the consequence of vicariance events,

resulting from evolution due to long-term isolation.

Keywords Ephemeroptera  Bioregion 

Endemicity  Diversity  Distribution  Introgression

Introduction

The Ephemeroptera (mayﬂies) are an ancient lineage

of insects, dating back to the late Carboniferous or

early Permian periods, some 290 mya. It is thought

that they attained their highest diversity during the

Mesozoic (Brittain & Sartori, 2003). They are clearly

the most primitive and ancient of the extant insect

groups (Edmunds & McCaf ferty, 1988). The rela-

tionship of Ephemeroptera with other modern winged

insects is still a subject of debate. Together with the

Odonata, mayﬂies were traditionally placed in the

Paleoptera, which was considered the sister group of

all other extant primarily winged orders (Kukalova

Peck, 1991). More recently, it was suggested that

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

que, H. Segers &

K. Martens

Freshwater Animal Diversity Assessment

H. M. Barber-James (&)

Department of Freshwater Invertebrates, Albany Museum,

Grahamstown, South Africa

e-mail: H.James@ru.ac.za

H. M. Barber-James

Department of Entomology, Rhodes University,

Grahamstown, South Africa

J.-L. Gattolliat  M. Sartori

Museum of Zoology, Lausanne, Switzerland

M. D. Hubbard

Laboratory of Aquatic Entomology,

Florida A & M University,

Tallahassee, FL, USA

123

Hydrobiologia (2008) 595:339–350

DOI 10.1007/s10750-007-9028-y