-1

Obligate groundwater fauna of France: diversity

patterns and conservation implications

DAVID FERREIRA*, FLORIAN MALARD, MARIE-JOSEDOLE-OLIVIER and JANINE GIBERTUMR CNRS 5023, Ecologie des Hydrosystemes Fluviaux, Equipe Hydrobiologie et Ecologie

Souterraines, Universite Claude Bernard Lyon 1, Bat. Forel, 43 Boulevard du 11 Novembre 1918,

F-69622 Cedex, France; *Author for correspondence (e-mail: davferreira@free.fr; phone: +33-4-

72432945; fax:+33-4-72431523)

Received 31 January 2005; accepted in revised form 6 June 2005

Key words: Biodiversity, Conservation, Endemism, France, Ground water, Stygobiotic fauna

Abstract. We examined taxonomic and geographic patterns of the obligate groundwater fauna (i.e.

stygobiotic fauna) by assembling in a distributional data base all species occurrences reported from

France since 1805. A simulated annealing algorithm was used to identify conservation targets. Until

the 60s, biological surveys were restricted to caves but the proportion of sampling sites in

unconsolidated sediments increased from 1 to 16% over the last 40 years. A total of 380 species and

subspecies in 40 families were collected, 70% of which being restricted to France. As observed in

other temperate regions, the stygobiotic fauna was dominated by crustaceans (65% of species) and

molluscs (22%). The cumulative number of species did not level off over time, clearly showing that

biodiversity was underestimated. Temporal trends in the cumulative number of obligate ground-

water and surface water species suggested that groundwater comprised more crustaceans than

surface freshwater. Endemism was high although the geographic range size of species increased as

distributional data accumulated. Of 380 species, 156 were known from a single 400-km2 cell, among

which 73% were located in the southern third of France. The distribution map of species richness

changed dramatically over time, indicating that the location of richness hotspots was sensitive to

sampling effort. Less than 2% of the French landscape was needed to capture 60% of known

species. Thus, a large proportion of species could be protected by focusing habitat conservation

efforts on a few complementary species-rich aquifers located in distinct regions.

Introduction

In the last few decades, groundwater ecology has developed rapidly forming afertile discipline of aquatic ecology (Gibert et al. 1994). Whereas the subter-ranean domain has long been considered as a species-poor environment,worldwide syntheses revealed an unexpectedly high diversity of living forms ingroundwater (Botosaneanu 1986; Juberthie and Decu 1994, 1998, 2001).Botosaneanu (1986) listed about 7000 obligate groundwater species worldwide.Groundwater ecosystems harbor different kinds of animal organisms fromtypically accidental obligate-surface water species (i.e. stygoxenes) to a highlyspecialized obligate groundwater fauna (i.e. stygobionts), the members ofwhich developed adaptive strategies for life in a dark and energy-limited

Biodiversity and Conservation (2007) 16:567–596 � Springer 2007

DOI 10.1007/s10531-005-0305-7

environment (Marmonier et al. 1993; Langecker 2000). Because moststygobionts have a narrow distribution range, the risk of species extinction isexpectedly high in face of the increase in multiple anthropogenic pressures(Malard et al. 1996; Gibert and Deharveng 2002; Danielopol et al. 2003). Thehigh level of endemicity in groundwater systems requires specific protectionmeasures for maintaining their ecological integrity and biological diversity(Notenboom et al. 1994). However, the current incomplete state of knowledgeon groundwater biodiversity and the lack of sound conservation strategiesseverely constrain the implementation of protection policies (Bouchet 1990;Holsinger 1993; Gibert 2001).In order to reveal the hidden biodiversity of groundwater, ecologists have

recently begun to synthesize and map existing data on the diversity anddistribution of stygobiotic fauna. Culver et al. (2000) mapped the distribu-tion of 973 aquatic and terrestrial obligate cave species in U.S.A. andshowed that 61% of the species were found only in a single county. TheItalian Ministry of Environment maintains a distributional database ofstygobionts as a background for developing adequate conservation strategies(Stoch 2000, 2001). In France, groundwater biodiversity patterns are stillpoorly known despite the existence of syntheses on the distribution of sev-eral taxonomic groups (Lescher-Moutoue 1967; Rouch 1968; Coineau 1971;Henry 1976; Magniez 1976). Works by Ginet and Juberthie (1987) andJuberthie and Ginet (1994) were the first attempts to document globaldiversity patterns based on distribution data for a restricted number ofspecies.The objectives of this paper were (1) to summarize our present-day knowl-

edge of groundwater biodiversity in France based on distributional data col-lected since 1805; (2) to examine the taxonomic and geographic patterns ofspecies diversity; and (3) to discuss the conservation implications of thesepatterns.

Material and methods

Data sources and validation procedure

Over the last three years, we assembled in a database the occurrences of allstygobionts reported from France since 1805. With the exception of Nematoda,Tardigrada and Hydrachnidia, all invertebrate groups known to have repre-sentatives within the stygobiotic fauna were included in the database. Infor-mation available in the literature, existing databases, and personal collectionswere used to gather as many records as possible and to produce lists anddistribution maps of species for each taxonomic group. In order to minimizethe risk to include spurious records, lists of stygobiotic species, records, anddistribution maps were submitted to taxonomists for correction and validation(Ferreira et al. 2003).

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Database and mapping

In order to be exported in the European database on groundwater biodiversitydeveloped under the 4D software (Ferreira et al. 2003), all data were entered in4 Excel spreadsheets containing for each record (i.e. line) a total of 28 fields (i.e.columns). The following fields were used in the present study: (1) class, order,family, genus and species names; (2) species authorities and description year;(3) spatial coordinates of the site; (4) data sources (e.g. bibliographic reference,personal collection); (5) collection year; and (6) habitat type (e.g. karst orporous aquifers). The spatial coordinates of the centroid of the civil parish(gazetteer GEOFLA� Commune, National Geographic Institute, Paris) wasused for most records because we lacked the precise coordinates of the sam-pling site. France is divided into 36,582 civil parishes, the average area of whichis 15±15.2 km2. Data were exported into a geographical information system(ArcView 3.2 software) to map the distribution of species and species richness.

Data analysis

Because the collection data were presence-only, they could not be used toevaluate spatial heterogeneity in sampling effort. Following the proceduredeveloped by Culver et al. (2004) for the analysis of species richness in Dinariccaves, Slovenia, we utilized time snapshots of the groundwater fauna toexamine stability of diversity patterns. Based on an intensive literature search,we used the number of sampling sites at which groundwater invertebratecommunity studies were carried out to assess spatial variation in samplingeffort during three overlapping periods: prior 1907, 1960 and 2003. These datescorresponded to stepping-stones in the historical development of research ongroundwater biodiversity: the creation of the association named ‘Biospeologica’(Racovitza 1907), the emergence of several groundwater research groups in theearly 60s (Juberthie and Ginet 1994), and the launching of the present databasewithin the framework of the European program PASCALIS (i.e. Protocols forthe Assessment and Conservation of Aquatic Life In the Subsurface). Siteswhich were sampled within the framework of taxon-oriented studies (e.g. re-gional inventory of selected species) were discarded in order to restrict taxo-nomic biases in our estimate of sampling effort. The point coverage of samplingsites was intersected with a geological polygon coverage (i.e. geological map ofFrance, scale 1/1,000,000, BRGM 1997) to determine differences in the numberof sampling sites between areas comprising karst aquifers and porous aquifers.According to Geze (1973), karst areas were further divided into highly kars-tified (thick and compact limestone), well karstified (thin limestone or alter-nating with shales), and poorly karstified (chalk) areas.The year at which a species was first described or reported from French

groundwater was used to examine changes in the cumulative number of speciessince 1805. We also compared temporal changes in the cumulative numbers of

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stygoxen and stygobiotic crustacean species. Although only the descriptionyear of species was available for stygoxen crustaceans (i.e. a species could bereported from France several years after its description), the comparisonproved to be valuable for assessing the phase difference between knowledge ofbiodiversity in groundwater and surface water.The frequency distribution of range size was examined at three different

periods to determine how increasing sampling of groundwater could modifyour perception of rarity among stygobiotic species. According to Gaston(1991), we distinguished between two measures of range size: the area ofoccupancy of a species and its extent of occurrence. The number of 400-km2

cells (see below for the definition of the grid system) in which a species wascollected was used as a measure of its area of occupancy. The latitudinal extentof a species, i.e., the straight-line distance (km) between latitudinally mostwidely separated occupied sites, was used as a measure of the extent ofoccurrence.A grid-based distribution map of species richness was produced for each

period by counting the number of species present in the 400-km2 cells of thegrid coverage. The choice of the cell area was dictated by the spatial resolutionof the data; one cell contained in average 15 civil parishes. We examinedtemporal changes in the frequency distribution of species richness as well as therelationships between species richness of cells and the number of sampling sites.Marxan 1.8.6, an optimization package designed for marine protected area

site selection (Possingham et al. 2000; Ball and Possingham 2001), was used todetermine the minimum number and potential location of cells for representingall species at least once. We performed adaptive simulated annealing followedby the summed irreplaceability heuristic algorithm to select cells. Then, theswap iterative improvement algorithm was used to ensure that no selected cellswere superfluous. We ran the simulated annealing 1000 times and measured theselection frequency of each cell by recording the number of times it was in-cluded in the ‘reserve’ network. Selection frequency is a measure of theimportance of a cell for meeting the conservation goal (i.e. representing allspecies at least once).

Results

Sampling effort

The number of sampling sites at which groundwater invertebrate communitystudies were carried out increased from 53 in 1907 to 1567 in 2003. Themajority of the sampling sites was concentrated in southern and eastern Francein areas comprising highly karstified limestone (Figure 1). Almost no inverte-brate community studies were conducted in alluvial aquifers of several majorrivers (e.g. the Seine and the Loire Rivers) and aquifers in fissured rocks andunconsolidated sediments of the Paris Basin, Armoricain Massif, and Central

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Figure 1. Distribution map of sampling sites at which groundwater invertebrate community

studies were conducted prior to 1907 (upper panel), 1960 (middle panel), and 2003 (lower panel).

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Massif. Until the 60s, sampling was almost totally restricted to karst areas(99%) and more precisely to caves (95% of sampled sites). Over the last40 years, the proportion of sampling sites in areas comprising extensivedeposits of unconsolidated sediments increased from 1.2 to 15.9%.

Taxonomic patterns

The database contained 380 species and subspecies corresponding to more than5700 records of which 83.5%, 12% and 2.5% corresponded to occurrences ofcrustaceans, molluscs and annelids, respectively. The list of species is providedin Appendix 1. Species were distributed among 40 families and 100 genera(Table 1). Crustaceans and molluscs accounted for 65 and 22% of stygobioticspecies richness, respectively, whereas they represented only 6 and 15%,respectively, of all stygoxen species collected in France. Crustaceans accountedfor 60% of the generic and family richness. About 43% of crustaceans werecopepods, among which 33% belonged to the Cyclopidae family (Cyclopoida)and 63% to the Ameiridae, Canthocamptidae and Parastenocarididae families

Table 1. Number of families, genera, and species within the stygobiotic fauna of France.

Groundwater Surface water

Family Genus Species Species

Taxa

Nemertina 1 1 1 1

Planaria 2 6 24 11

Annelida

Aphanoneura 2 2 2 10

Oligochaeta 4 12 21 144

Polychaeta 1 1 1 na

Hirudinea 1 1 1 22

Molluscs 3 17 82 150

Crustacea

Branchiopoda 1 1 2 88

Decapoda 1 1 1 10

Ostracoda 3 10 23 68

Syncarida 2 11 21 0

Isopoda 5 7 49 1

Amphipoda 6 10 45 12

Copepoda Cyclopoida 1 7 35 31

Copepods Harpacticoida 4 10 68 35

Copepods Gelyelloida 1 1 1 0

Copepods Calanoida 1 1 1 24

Insects 1 1 2 3200

All taxa 40 100 380 4305

The last column provides estimates of the number of surface water obligate species in France

(na: not available). Data from different sources.

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(Harpacticoida). Out of ten crustacean groups, six had more species withinstygobiotic fauna than in stygoxen fauna. Whereas insects represented about74% of all invertebrate species known from surface water, the stygobiotic faunacomprised only two insect species belonging to the genus Siettitia (ColeopteraDytiscidae).The cumulative number of species increased exponentially with time

(Figure 2). During the last 50 years, the number of species increased three-fold(i.e. from 124 in 1951 to 380 in 2003). However, the rate of reporting of newspecies and records has decreased since the 80s. From 1978 to 2003, only 82species were reported whereas 136 species had been reported during the sameduration from 1948 to 1973. The rate of reporting of species belonging to themeiofauna (i.e. body size <1 mm) suddenly increased in the 30s, and sincethen, has remained markedly higher than the reporting rate of speciesbelonging to the macrofauna (body size >1 mm) (Figure 2). Consequently, theproportion of meiofaunal species (i.e. almost all Copepoda, Ostracoda,Syncarida and Branchiopoda) increased from 4% in 1909 to 40% in 2003.There was almost a 100-year phase difference between the curves of the

cumulative number of stygoxen and stygobiotic species of crustaceans(Figure 3). About 80% of stygoxen species were described before 1900, whereasonly 4% of stygobiotic species had been reported. Since the 70s, only fivestygoxen species have been described against 70 species for the stygobioticfauna. Despite this pronounced time lag between the curves, the present-daynumber of stygobiotic crustaceans (246 species among which 18 undescribedspecies) almost equals that of stygoxenes (266 species).

Geographic patterns

The distributions of range size were strongly skewed towards small areas ofoccupancy and latitudinal extents (Figure 4). In 2003, 41% of species werecollected from a single cell and 38% had a latitudinal extent less than 3 km.Only 5 and 2.5% of species had an area of occupancy higher than 25 cells and alatitudinal extent higher than 600 km, respectively.However, the average and maximum range sizes increased over time. From

1960 to 2003, the average area of occupancy and average latitudinal extentincreased from 2.2±3.9 to 5.6±10.4 cells and from 58±159 to102±184 km, respectively. The maximum area of occupancy increased fromthree cells in 1907 to 54 and 82 cells in 1960 and 2003, respectively. Themaximum latitudinal extent was only 82 km in 1907 but 900 and 940 km,respectively, in 1960 and 2003.Over the last 100 years, the distribution patterns of species richness changed

dramatically (Figure 5). From 1907 to 2003, the proportion of cells containingno species decreased by 35%. Meantime, the number of species in the richestcell increased from 4 to 52. However, in 2003, only 2.7% of the cells containedmore than 11 species, whereas 33% had between 1 and 10 species. Species

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richness increased linearly with the number of sampling sites (Y = 0.49x;p<0.01). Difference in the number of sampling sites between cells explained45% of the variation in species richness.Although 103 cells were needed to capture all species at least once, a high

proportion of species rapidly accumulated in relatively few cells (Figure 6). Of

Figure 2. Upper panel: cumulative numbers of new species and records of stygobiotic organisms

since 1805. Lower panel: differences in the cumulative numbers of new species and records among 3

body size groups of stygobiotic organisms. Continuous, thick broken and thin broken lines cor-

respond to body sizes <1 mm, 1–6 mm, and >6 mm, respectively.

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a total of 380 species, 227 (60%) were represented in only 17 cells and 75% ofspecies were represented in 34 cells. Beyond 34 cells, adding an additional cellresulted in an increase of only two or one species. Figure 7 indicates theselection frequency of cells during 1000 runs of the optimization algorithm.Only 145 cells out of 445 appeared in at least one solution, among which 90appeared in all solutions, representing 362 species. Of the 90 cells appearing inall solutions, 86 contained the entire distribution of one or more species andwere therefore irreplaceable. The location in southern France of two thirds ofever-selected cells reflected the concentration of endemic species. Of the 156species restricted to a single cell (i.e. single-cell endemics), 114 occurred in thesouthern third of France (Figure 8).

Discussion

Sampling effort

Groundwater fauna has certainly been more investigated in France than inmost other countries (except Slovenia). A total of 1800 caves were sampled inEurope within the framework of the Biospeologica program, among which60% were located in France (Juberthie and Ginet 1994). The French databaseincludes about the same number of records for the aquatic cave fauna (i.e. 3000records) as the U.S. database (i.e. 2774 records, but see Culver et al. 1999,2000). However, sampling is still highly incomplete and spatially concentratedin limestone regions. The disparity in sampling effort between karsts and other

Figure 3. Cumulative numbers of new species and records of stygobiotic (open squares) and

stygoxen (full diamonds) crustaceans since 1758.

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geological formations reflects the predominance of the biospeological thoughtfor more than one century (Rouch 1986; Belles 1992). Until the 60s, the bio-logical exploration of groundwater was restricted to caves as they provided aneasy access to the subterranean realm. This led to a spatially limited andanthropocentric view of groundwater domain (i.e. the voids accessible to

Figure 4. Frequency distribution of the area of occupancy (expressed as the number of 400-km2

cells in which a species occurred) and latitudinal extent of stygobiotic species during three over-

lapping periods: prior to 1907 (black pattern), 1960 (grey pattern) and 2003 (white pattern).

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Figure 5. Grid-based distribution map of species richness during three overlapping periods: prior

to 1907 (upper panel), 1960 (middle panel) and 2003 (lower panel).

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humans), which retarded the comprehension of their structure and functioning.However, research conducted during the last 40 years in a variety of uncon-solidated sediments (e.g. marine, alluvial, colluvial and morainic deposits)revealed that stygobiotic animals were likely to be found wherever there wasgroundwater. The disproportionate sampling effort allocated to karst aquifersin many regions probably biases our estimate of groundwater biodiversitybecause nearby karst and alluvial aquifers usually harbor dissimilar inverte-brate assemblages. Rouch (1988) showed that the Baget karst system, Pyrenees,and its ensuing alluvial aquifer contained 22 and 21 species, respectively, withonly 12 species in common. Similarly, the Dorvan karst system, Jura, and thecontiguous alluvial aquifer of the Albarine River harbored 22 and 21 species,respectively, but shared only 8 species Gibert et al. 2000). About 47 and 29% ofspecies collected in France are exclusive to karst and unconsolidated habitats,respectively, and 20% occur in both habitats,

Taxonomic patterns

Groundwater fauna of France is more diversified (i.e. 380 species) than thatof other countries including U.S.A. (300 species; Culver et al. 2000), Italy(265 species; Stoch 2001), Slovenia (210 species; Sket 1999a) and Croatia(about 200 species; Gottstein-Matocec et al. 2002). This elevated number ofspecies is most likely the result of a higher sampling effort in various aquifers.Although all invertebrate groups are represented in groundwater, the stygo-biotic fauna is dominated by crustaceans at different systematic levels. Thisoverrepresentation of crustaceans (i.e. 65% of species) has been observed ingroundwater of all temperate regions (Sket 1999a, b). Danielopol et al. (2000)estimated that the stygobiotic fauna accounted for 40% of the total numberof aquatic crustacean species in Europe. The lack of competitors such as

Figure 6. Species accumulation curve for 103 cells representing all species at least one.

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aquatic insects has been advocated by several authors for explaining the highdiversity of crustaceans in groundwater (Stoch 1995; Sket 1999b). Beingdependent upon air for breathing or reproducing, aquatic insects areextremely rare in groundwater whereas they represent 50% of all animalspecies living in surface water of Europe (Danielopol et al. 2000). The genus

Figure 7. Grid-based distribution map of species richness showing the location and selection

frequency of cells for representing all species at least one (out of 1000 runs of the selection algo-

rithms). Cells numbered from 1 to 17 correspond to the 17 first most-complementary cells con-

taining 60% of known species.

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Siettitia (Coleoptera Dytiscidae), which comprises two species living in thewater-table region of alluvial aquifers of the Rhone River, is one of the rareinsect lineages known from Europe (Spangler and Decu 1998).The exponential increase in the cumulative number of species since the

beginning of the 19th century clearly demonstrates that groundwater biodi-versity in France is still largely underestimated. This is particularly true for thecrustacean meiofauna, for which new species are continuously described(Galassi et al. 1999; Apostolov 2002). Temporal trends in the cumulative

Figure 8. Grid-based distribution map of endemic richness. An endemic was defined as a species

restricted to a single 400-km2 cell.

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number of obligate groundwater and surface water species suggest thatgroundwater comprises more crustaceans than surface freshwater. This lendssupport to the idea that the contribution of stygobiotic fauna to continentalfreshwater biodiversity is probably far more important than previously sus-pected (Gibert and Deharveng 2002). Decrease in the report rate of species forthe last 20 years essentially reflects the deteriorating capacity of an everdecreasing number of active taxonomists to identify biological material andpublish updated lists of species (Reid 1992; Galassi 2001; Valdecasas andCamacho 2003). As observed for two decades in all branches of biology (Oliver1988; Gaston and May 1992), this crisis in biosystematics impoverishes con-siderably the quality of taxonomic information which is, however, the mostfundamental requirement for any assessment and study of groundwater bio-diversity. A high proportion of records (i.e. from 12% for Copepoda up to 52%for Syncarida) that are not identified to species do not figure on distributionmaps and there is an increasing number of potentially new species (i.e. about 19species) still awaiting description. On the other hand, new information forunderrepresented taxonomic groups such as Oligochaeta progressively becomeavailable (Martınez-Ansemil et al. 1997; Giani et al. 2001).

Geographic patterns

Although the average latitudinal extent and area of occupancy of groundwaterspecies increased over time as distributional data accumulated, the geographicrange size of most stygobionts remained extremely small compared to those ofepigean species. Nearly 70% of stygobiotic species collected in France are re-stricted to the French landscape (including Corsica). For comparison, theFrench fauna of Ephemeroptera and Odonata (i.e. 252 species) have 80 and100% species in common with the Italian and Swiss fauna, respectively. Sket(1999a) reported that the Dinaric Mountains, probably the richest ground-water region in the world in relation to its area, also shared very fewstygobiotic species with neighboring regions (i.e. 22% of species). Recentgenetic studies reinforced the significance of endemism within groundwaterfauna by demonstrating that a number of species identified on the basis ofmorphological criteria corresponded in fact to clusters of distinct species(Cobolli-Sbordoni et al. 1990; Ketmaier et al. 2000; Lefebure et al. 2006).The exceptional richness of endemics is directly linked to the highly fragmentednature of the groundwater domain that fosters evolution processes (i.e. speci-ation) leading to the isolation of populations (Holsinger 2000; Humphreys2000). Because endemism is typically higher in groundwater than in surfacewater fauna, the contribution of groundwater fauna to the richness of conti-nental water increases with increasing spatial scale (Culver and Sket 2000;Gibert and Deharveng 2002).The high proportion of single-cell endemics in the southern third of France

(i.e. 73%) was unlikely the result of incomplete sampling. The Pyrenees

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contained five time more endemic species (52 species) than the Jura (10 species),although these two mountain areas received comparable sampling efforts.Distribution maps of stygobiotic species recently produced within the frame-work of the European project PASCALIS also revealed that more than 80% ofsingle-cell endemic species (0.2 · 0.2 decimal degree cell) occurred in SouthernEurope (J. Gibert, unpublished data). The southern concentration of endemic-rich areas supports the hypothesis that the distribution of stygobiotic species,most of which colonized groundwater several millions years ago, was stronglymodified by cold Pleistocene events (Ginet 1971; Holsinger et al. 1983; Strayeret al. 1995; Magniez 1997). Many endemic species that were particularly vul-nerable due to their small population size might have gone extinct in northernareas during the last ice age.The distribution map of species richness changed dramatically over time as

more extensive surveys provided additional records, clearly indicating that thelocation of biodiversity hotspots varied as a function of sampling effort. Sincethe 60s, sampling in alluvial aquifers of the Rhone River resulted in several newhotspots. In contrast, Culver et al. (2000) argued that the concentration ofbiodiversity within a few cave regions of the U.S.A. was not sensitive to anincrease in sampling effort. In France, most species-rich cells (>20 species)corresponded to areas that received a disproportionate amount of samplingeffort. These included areas that were located within a short distance to re-search facilities such as the subterranean laboratory in Moulis, Pyrenees (52species), the groundwater ecology laboratory in Lyon (43 species), and thelaboratory of animal biology in Dijon, Cote-d’Or (24 species). Several studiesdemonstrated that sampling was positively biased towards areas designated ashotspots of biodiversity (Nelson et al. 1990; Reddy and Davalos 2003). Con-versely, this concentration of species in highly sampled areas strongly suggestedthat species richness was underestimated in most groundwater areas of France.For example, we would expect all cells located along the southern part of theRhone River to contain at least 20 species because most interstitial stygobiontscollected in the vicinity of the Lyon city are also known to occur in alluvialgroundwater in southern France.

Conservation implications

Although several studies documented the adverse effects of groundwater con-tamination and extraction on invertebrate assemblages (Rouch et al. 1993;Malard et al. 1996; Mosslacher and Notenboom 2000), the diverse and vul-nerable groundwater fauna of France has received almost no conservationattention to date (Bouchet 1990; Juberthie 1995). Despite the extremely narrowdistribution range of most obligate groundwater species known from France,only 20 species (18 molluscs, 1 copepod and 1 coleopteran) were included in thethreatened species list of the International Union for Conservation of Natureand Natural Resources (IUCN 2004). However, listing of threatened species

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under the IUCN act or European Habitat Directive is only a minor step forconserving groundwater biodiversity. As stated by Culver et al. (2000, p. 395),‘‘there are simply too many subterranean species at risk to deal with them oneat time’’. Indeed, almost all groundwater species of France could potentially belisted in the European habitat directive based on legislative criteria used fordefining priority species (i.e. endemism, rarity, species characteristic of a singlebiogeographic region). A more efficient conservation solution is to protect aminimum network of complementary aquifers that maximizes the representa-tion of as many species as possible while minimizing the risk of biodiversity lossdue to human activities (Gaston et al. 2002; Margules et al. 2002). This strategywhich pays explicit attention to patterns of between-aquifer complementarity at anational scale enables the maintenance of regionally distinctive species-richassemblages while minimizing the duplication of conservation efforts acrossregional agencies (Mace 2000). Whereas heuristic algorithms are commonlyused for designing effective reserve networks in terrestrial and marine systems(Csuti et al. 1997; Stewart et al. 2003), the designation of priority groundwatersites is still essentially based on traditional selection methods (selection ofhotspots of species richness and of endemism). Culver et al. (2000) applied forthe first time a greedy algorithm to delineate potential subterranean prioritysites in the U.S.A and showed that 50% of obligate cave fauna occurred in lessthan 1% of the landscape. We also demonstrated using an adaptive simulatedannealing that only 1.2% (i.e. 17 cells) of the French territory was needed tocapture 60% of known stygobiotic species. Thus, it should be possible toconserve a large proportion of species by focusing habitat conservation effortsin a few complementary species-rich aquifers (i.e. conservation targets) locatedin distinct regions including the Pyrenean Mountains (cells 1, 5, 11, and 12 inFigure 7), the Roussillon Plain (9, 15), the North-Montpellieran karsts (3, 7,17), the Ardeche calcareous plateau (4, 10), the lower Rhone valley (2, 16), theOccidental Jura (13, 14), the Burgundy karsts (8), and the Rhine valley (6).Although many of these species-rich aquifers were previously identified byother scientists (Juberthie and Juberthie-Jupeau 1975; Malard et al. 1997;Gibert et al. 2000), their present-day location within each region reflects tosome extent the distribution of data-rich areas.We identified two major research avenues for implementing an effective

groundwater reserve network in France. A first important step is to shift from agrid cell selection approach to an aquifer selection approach that incorporatesnot only representation targets of species and minimum space requirement butalso socio-economic costs related to the vulnerability of aquifers and strengthof human activity in the catchment. Since the objective function of Marxanallows assigning different costs for each aquifer (Ball and Possingham 2001),the resultant reserve network would not only depend on the ecological valuesof aquifers but also on their conservation costs linked to socio-economicconstrains. The use of aquifers as planning units is all the more importantbecause aquifer classifications based on relevant ecological criteria (i.e. per-meability, pore size, hydrological connection with surface environment) can be

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used as abiotic surrogates of groundwater biodiversity in data-poor regions. Asecond critical step consists in developing appropriate sampling strategies inresponse to well-defined conservation goals. Because beta diversity amongregions (i.e. dissimilarity in species composition between regions) makes by farthe highest contribution to total richness of stygobionts in groundwater ofFrance (Ferreira et al. unpublished), the inclusion of new conservation targetsnecessitates sampling aquifers in supposedly rich regions that have not yet beeninvestigated. In contrast, increasing the flexibility of the network of conser-vation targets requires sampling additional aquifers in regions that have al-ready been sampled. Although the high level of endemism within the obligategroundwater fauna would necessarily reduce the latitude in placement of pri-ority areas for conservation, the present lack of flexibility of the reserve net-work is primarily attributable to the weakness of sampling effort in mostregions. The use of aquifers as planning units, the integration of conservationcosts, and the urgent need for updating biological data are interdisciplinarygoals that require hydrogeologists, socio-economists, and biologists to worktogether for implementing an effective groundwater reserve network.

Acknowledgements

This work was supported by the European program PASCALIS (Protocols forthe ASsessment and Conservation of Aquatic Life In the Subsurface) (EVK2-CT-2001-00121). We thank the numerous scientists who provided data andcorrections: C. Bou, N. Coineau, A. Brancelj, L. Deharveng, M. andG. Falkner, D. Galassi, N. Giani, R. Ginet, C. Jouin-Toulmond, J. Juget,F. Lescher-Moutoue, G. Magniez, J.-P. Henry, P. Marmonier, J. Mathieu,P. Richoux, and M.-J. Turquin.

Appendix 1. List of groundwater obligate species of France.

NEMERTINA

Tetrastemmatidae

Prostoma puteale de Beauchamp 1932

PLANARIA

Dendrocoelidae

Dendrocoelopsis beauchampi (Gourbault 1969)

Dendrocoelopsis bessoni Gourbault, Benazzi & Helleouet 1976

Dendrocoelopsis brementi (de Beauchamp 1919)

Dendrocoelopsis chattoni (de Beauchamp 1949)

Dendrocoelopsis garmieri (de Beauchamp 1950)

Dendrocoelopsis vandeli (de Beauchamp 1931)

Dendrocoelum (Bolbodendrocoelum) agile de Beauchamp 1932

Dendrocoelum (Dendrocoelides) barbei de Beauchamp 1956

Dendrocoelum (Dendrocoelides) coiffaiti de Beauchamp 1956

Dendrocoelum (Dendrocoelides) col1ini (de Beauchamp 1919)

584

Appendix 1. (continued)

Dendrocoelum (Dendrocoelides) lescherae Gourbault 1971

Dendrocoelum (Dendrocoelides) regnardi (de Beauchamp 1919)

Dendrocoelum (Dendrocoelides) tuzetae Gourbault 1965

Dendrocoelum (Dendrocoelum) infernale (Steinmann 1907)

Dendrocoelum (Eudendrocoelum) gineti de Beauchamp 1954

Dendrocoelum (Eudendrocoelum) remyi de Beauchamp 1926

Dendrocoelum (Polycladodes) album (Steinmann 1910)

Dendrocoelum sollaudi de Beauchamp 1931

Miodendrocoelum parisi de Beauchamp 1929

Planariidae

Atrioplanaria delamarei Gourbault 1969

Atrioplanaria notadena de Beauchamp 1937

Phagocata albissima (Vejdovsky 1883)

Phagocata vitta (Duges 1830)

Plagnolia vandeli de Beauchamp & Gourbault 1964

ANNELIDA, HIRUDINEA

Erpobdellidae

Trocheta bykowskii Gedroyc 1913

ANNELIDA, APHANONEURA

Aeolosomatidae

Aeolosoma gineti Juget 1959

Potamodrilidae

Potamodrilus fluviatilis Lastochkin 1935

ANNELIDA, POLYCHAETA

Nerillidae

Troglochaetus beranecki Delachaux 1921

ANNELIDA, OLIGOCHAETA

Enchytraeidae

Enchytraeus flavus Moszynski 1938

Pachydrilus fossor Vejdovsky 1877

Haplotaxidae

Delaya corbarensis (Delay 1972)

Haplotaxis leruethi (Hrabe 1958)

Lumbriculidae

Cookidrilus speluncaeus Rodriguez & Giani 1987

Trichodrilus capilliformis Rodriguez & Giani 1994

Trichodrilus cernosvitovi Hrabe 1937

Trichodrilus intermedius (Fauvel 1903)

Trichodrilus 1eruthi Hrabe 1937

Trichodrilus pragensis (Vejdovsky 1875)

Trichodrilus tenuis Hrabe 1960

Tubificidae

Abyssidrilus cuspis (Erseus & Dumnicka 1988)

Gianius labouichensis (Rodriguez & Giani 1989)

Gianius cavealis Juget & Des Chatelliers 2001

Haber turquini (Juget & Lafont 1979)

Krenedrilus sergei Giani, Erseus & Martinez-Ansemil 1990

Rhyacodrilus amphigenus Juget 1987

Rhyacodrilus balmensis Juget 1959

Rhyacodrilus lindbergi Hrabe 1963

Rhyacodrilus subterraneus Hrabe 1963

Spiridion phreaticola (Juget 1987)

585

Appendix 1. (continued)

MOLLUSCA

Amnicolidae

Bythinella bouloti Girardi, Bichain & Wienin 2002

Bythinella cylindracea (Paladilhe 1869)

Bythinella eutrepha (Paladilhe 1867)

Bythinella galerae Girardi, Bichain & Wienin 2002

Bythinella geisserti Boeters & Falkner 2003

Bythinella padiraci Locard 1903

Bythinella pupoides phreaticola Bernasconi 1989

Hydrobiidae

Alzoniella (Alzoniella) haicabia Boeters 2000

Alzoniella (Alzoniella) navarrensis Boeters 1999

Alzoniella (Alzoniella) pyrenaica (Boeters 1983)

Alzoniella (Alzoniella) ,junqua Boeters 2000

Alzoniella (Navariella) elliptica (Paladilhe 1874)

Avenionia bourguignati (Locard 1883)

Avenionia brevis (Draparnaud 1805)

Avenionia berenguieri (Bourguignat 1882)

Belgrandiella saxatilis (Reynies 1844)

Belgrandiella ? dunalina (Moquin-Tandon 1856)

Fissuria boui Boeters 1981

Graziana provincialis (Boeters 2000)

Graziana rayensis (Caziot 1910)

Graziana ? cezairensis Boeters 2000

Heraultiella exilis (Paladilhe 1867)

Islamia bomangiana Boeters & Falkner 2003

Islamia bourguignati (T. Letourneux 1869)

Islamia consolationis (Bernasconi 1985)

Islamia germaini Boeters et Falkner 2003

Islamia minuta (Draparnaud 1805)

Islamia moquiniana (Dupuy 1851)

Islamia spirata (Bernasconi 1985)

Istriana falkneri Boeters 2000

Moitessieriidae

Bythiospeum anglesianum (Westerlund 1890)

Bythiospeum articense Bernasconi 1985

Bythiospeum bourguignati (Paladilhe 1866)

Bythiospeum bressanum Bernasconi 1985

Bythiospeum charpyi charpyi (Paladilhe 1867)

Bythiospeum charpyi giganteum Bernasconi 1969

Bythiospeum diaphanoides Bernasconi 1985

Bythiospeum diaphanum diaphanum (Michaud 1831)

Bythiospeum diaphanum michaellense Girardi 2002

Bythiospeum dorvani Bernasconi 1985

Bythiospeum drouetianum (Cless in 1882)

Bythiospeum francomontanum Bernasconi 1973

Bythiospeum garnieri (Sayn 1889)

Bythiospeum klemmi (Boeters 1969)

Bythiospeum michaudi (Locard 1882)

Bythiospeum moussonianum (Paladilhe 1869)

Bythiospeum racovitzai (Germain 1911)

586

Appendix 1. (continued)

Bythiospeum rhenanum rhenanum (Lais 1935)

Bythiospeum terveri (Locard 1882)

Henrigirardia Wienini Girardi 2001

Moitessieria bourguignati Coutagne 1883

Moitessieria cocheti Boeters & Falkner 2003

Moitessieria heideae Boeters & Falkner 2003

Moitessieria fontsaintei Bertrand 2001

Moitessieria,juvenisanguis Boeters & E. Gittenberger 1980

Moitessieria lescherae Boeters 1981

Moitessieria locardi Coutagne 1883

Moitessieria massoti Bourguignat 1863

Moitessieria nezi Boeters & Bertrand 2001

Moitessieria olleri Altimira 1960

Moitessieria rhodani Coutagne 1883

Moitessieria rolandiana Bourguignat 1863

Moitessieria simoniana (Saint-Simon 1848)

Palacanthilhiopsis vervierii Bernasconi 1988

Palacanthilhiopsis ? margritae Boeters & Falkner 2003

Paladilhia conica Paladilhe 1867

Paladilhia gloeeri Boeters & Falkner 2003

Paladilhia pleurotoma Bourguignat 1865

Paladilhia pontmartiniana (Nicolas 1891)

Paladilhia sp.

Paladilhia umbilicata (Locard 1902)

Palaospeum bessoni bessoni (Bernasconi 1999)

Palaospeum bessoni rebenacqense Boeters & Bertrand 2001

Palaospeum? nanum Boeters & Bertrand 2001

Plagigeyeria deformata (Nicolas 1891)

Spiralix burgundina (Locard 1882)

Spiralix collieri (Nicolas 1891)

Spiralix corsica (Bernasconi 1994)

Spiralix hofmanni Boeters & Falkner 2003

Spiralix puteana (Coutagne 1883)

Spiralix rayi (Locard 1882)

Spiralix vitrea (Draparnaud 1801)

CRUSTACEA

CLADOCERA

Chydoridae

Alona bessei Dumont 1983

Alona phreatica Dumont 1983

DECAPODA

Atyidae

Troglocaris inernris Fage 1937

ISOPODA

Stenasellidae

Stenasellus buili Remy 1949

Stenasellus racovitzai Razzauti 1925

Stenasellus virei angelieri Magniez 1968

Stenasellus virei boui Magniez 1968

Stenasellus virei hussoni Magniez 1968

Stenasellus virei virei Dollfus 1897

587

Appendix 1. (continued)

Asellidae

Gallasellus heilyi (Legrand 1956)

Proasellus albigensis (Magniez 1965)

Proasellus aquaecalidae (Racovitza 1922)

Proasellus beroni/ Henry & Magniez 1968

Proasellus boui Henry & Magniez 1969

Proasellus burgundus Henry & Magniez 1969

Proasellus cavaticus (Leydig 1871)

Proasellus chauvini Henry & Magniez 1978

Proasellus claudei Henry & Magniez 1996

Proasellus coiffaiti Henry & Magniez 1972

Proasellus coxalis (Dollfus 1892)

Proasellus meridianus (Racovitza 1919)

Proasellus nolli (Karaman 1952)

Proasellus racovitzai Henry & Magniez 1972

Proasellus rouchi Henry 1980

Proasellus spelaeus (Racovitza 1922)

Proasellus strouhali puteanus (Henry 1966)

Proasellus synaselloides (Henry 1963)

Proasellus valdensis (Chappuis 1948)

Proasellus vandeli Henry & Magniez 1969

Proasellus walteri (Chappuis 1948)

Proasellus sp. 1

Microparasellidae

Microcharon angelieri Coineau 1963

Microcharon boui Coineau 1968

Microcharon doueti Coineau 1968

Microcharon juberthiei juberthiei Coineau 1968

Microcharon juberthiei ramosus Coineau 1968

Microcharon reginae Dole & Coineau 1987

Microcharon rouchi Coineau 1968

Microcharon sisiphus Chappuis & Delamare 1954

Microcharon sp. 1

Microcharon sp. 2

Microcharon sp. 3

Microcharon sp. 4

Microcharon sp, 5

Microcharon sp. 6

Microcharon sp. 7

Sphaeromatidae

Caecosphaeroma burgundum burgundum Dollfus 1898

Caecosphaerorma burgundum rupisfucaldi Hubault 1934

Caecosphaeroma virei Dollfus 1896

Cirolanidae

Sphaeromides raymondi Dollfus 1897

Faucheria faucheri Dollfus & Vire 1900

Cirolanidae n. sp.

AMPHIPODA

Karstogiella lautieri

Bogidiellidae

Bogidiella albertimagni Hertzog 1933

588

Appendix 1. (continued)

Bogidiella nicolae Karaman 1988

Medigidiella chappuisi Ruffo & Delaimare 1952

Niphargidae

Niphargus angelieri Ruffo 1953

Niphargus aquilex Schiodte 1855

Niphargus balazuci Schellenberg 1951

Niphargus boulangei Wichers 1964

Niphargus burgundus Graf & Straskraba 1967

Niphargus ciliates Chevreux 1906

Niphargus corsicanus Schellenberg 1950

Niphargus delamarei Ruffo 1953

Niphargus fontanus Bate 1859

Niphargus foreli Humbert 1877

Niphargus gallicus Schellenberg 1935

Niphargus gineti Bou 1965

Niphargus gr.jovanovici S, Karaman 1931

Niphargus kieferi Schellenberg 1936

Niphargus kochianus kochianus Bate 1859

Niphargus ladmiraulti Chevreux 1901

Niphargus laisi Schellenberg 1936

Niphargus nicaensis Isnard 1916

Niphargus pachypus Schellenberg 1933

Niphargus plateaui Chevreux 1901

Niphargus renei G, Karaman 1986

Niphargus rhenorhodanensis Schellenberg 1937

Niphargus robustus Chevreux 1901

Niphargus schellenbergi S, Karaman 1932

Niphargus setiferus Schellenberg 1937

Niphargus thienemanni Schellenberg 1934

Niphargus vandeli Barbe 1961

Niphargus virei Chevreux 1896

Niphargopsis casparyi (Pratz 1866)

Crangonyctidae

Crangonyx subterraneaus Bate 1859

Salentinellidae

Parasalentinella rouchi Bou 1971

Salentinella angelieri Ruffo & Delamare 1952

Salentinella delamarei delamarei Coineau 1962

Salentinella delamarei macrocheles Coineau 1968

Salentinella gineti Balazuc 1957

Salentinella major Barbe 1965

Salentinella petiti Coineau 1968

Salentinella sp. 1

Pseudoniphargidae

Pseudoniphargus adriaticus S. Karaman 1955

Ingolfiellidae

Ingolfiella (Tyrrhenidiella) catalanensis Coineau 1963

Ingolfiella (Tyrrhenidiella) thibaudi Coineau 1968

OSTRACODA

Ostracoda n. sp.

Candonidae

589

Appendix 1. (continued)

Candonopsis boui Danielopol 1978

Cryptocandona kieferi (Klie 1938)

Fabaeformiscandona breuili (Paris 1920)

Fabaeformiscandona cf. breuili sp.1

Fabaeformiscandona cf breuili sp. 2

Fabaeformiscandona wegelini (Petkovski 1962)

Mixtocandona juberthieae Danielopol 1978

Mixtocandona laisi (Klie 1938)

Pseudocandona delamarei Danielopol 1978

Pseudocandona rouchi Danielopol 1973

Pseudocandona simililampadis Danielopol 1978

Pseudocandona zschokkei (Wolf 1920)

Schellencandona belgica (Klie 1937)

Schellencandona cf. schellenbergi sp. 1

Schellencandona cf. schellenbergi sp. 2

Schellencandona triquetra (Klie 1936)

Cyprididae

Cavernocypris subterranea (Wolf 1920)

Psychrodromus betharrarmi Baltanas, Danielopol, Roca & Marmonier 1993

Entocytheridae

Sphaeromicola cebennica cebennica Remy 1948

Sphaeromicola cebennica juberthiei Danielopol 1977

Sphaeromicola hamigera Remy 1948

Sphaeromicola topsenti Paris 1916

COPEPODA, CYCLOPOIDA

Cylopidae

Acanthocyclops hispanicus Kiefer 1937

Acanthocyclops rhenanus Kiefer 1936

Acanhocyclops sensitivus (Graeter & Chappuis 1914)

Diacyclops cf. paolae Pesce & Galassi 1987

Diacyclops clandestinus Kiefer 1933

Diacyclops zschokkei (Graeter 1910)

Eucyclops graeteri (Chappuis 1927)

Graeteriella (Paragraeteriella) bertrandi Lescher-Moutoue 1974

Graeteriella (Paragraeteriella) gelyensis Lescher-Moutoue1978

Graeteriella (Paragraeteriella) laisi (Kiefer 1936)

Graeteriella (Paragraeteriella) vandeli Lescher-Moutoue 1969

Graeteriella boui Lescher-Moutoue 1974

Graeteriella brehmi Lescher-Moutoue 1968

Graeteriella rouchi Lescher-Moutoue 1968

Graeteriella unisetigera (Graeter 1910)

Kieferiella delamarei (Lescher-Moutoue 1971)

Megacyclops brachypus Kiefer 1954

Speocyclops arregladensis Chappuis & Kiefer 1952

Speocyclops anomalus Chappuis & Kiefer 1952

Speocyclops castereti Lindberg 1954

Speocyclops demetiensis (Scourfield 1932)

Speocyclops gallicus Chappuis & Kiefer 1952

Speocyclops kieferi Lescher-Moutoue 1968

Speocyclops orcinus Kiefer 1937

Speocyclops proserpinae Kiefer 1937

590

Appendix 1. (continued)

Speocyclops racovitzai (Chappuis 1923)

Speocyclops racovitzai boscensis Kiefer 1954

Speocyclops racovitzai gouillounensis Kiefer 1954

Speocyclops racovitzai incerta Chappuis & Kiefer 1952

Speocyclops racovitzai liguensis Chappuis & Kiefer 1952

Speocyclops racovitzai noustensis Chappuis & Kiefer 1952

Speocyclops racovitzai peyortensis Chappuis & Kiefer 1952

Speocyclops racovitzai,sabartensis Kiefer 1954

Speocyclops racovitzai sandetsi Chappuis & Kiefer 1952

Speocyclops sisyphus Kiefer 1937

COPEPODA, HARPACTICOIDA

Ectinasomatidae

Pseudectinosoma vandeli (Rouch 1969)

Pseudectinosoma janineae Galassi, Dole-Olivier & De Laurentiis 1999

Ameiridae

Parapseudoleptomesochra subterranea subterranea (Chappuis 1928)

Parapseudoleptomesochra subterranea deminuta (Chappuis 1928)

Nitocrellopsis elegans (Chappuis & Rouch 1959)

Nitocrellopsis rouchi Galassi, De Laurentiis & Dole-Olivier 1999

Nitocrella beatricis Cottarelli & Bruno 1994

Nitocrella delayi Rouch 1970

Nitocrella gracilis Chappuis & Rouch 1959

Nitocrella dussarti Chappuis 1955

Nitocrella omega Hertzog 1936

Nitocrella sp.1 groupe hirta

Nitocrella sp, 2 groupe hitra

Canthocamptidae

Elaphoidella boui Rouch 1988

Elaphoidella bouilloni Rouch 1964

Elaphoidella brehieri Apostolov 2001

Elaphoidella calypsonis Chappuis & Rouch 1959

Elaphoidella cavatica Chappuis 1957

Elaphoidella coiffaiti Chappuis & Kiefer 1952

Elaphoidella elaphoides elaphoides (Chappuis 1923/124)

Elaphoidella federicae Pesce & Galassi 1988

Elapheidella garbeteneis Rouch 1980

Elaphoidella infernalis Rouch 1970

Elaphoidella leruthi leruthi/ Chappuis 1937

Elaphordella leruthi meridionalis Chappuis 1953

Elaphoidella longifurcata Chappuis & Kiefer 1952

Elaphoidella madiracensis Apostolov 1998

Elaphoidella mauro Chappuis 1956

Elaphoidella pyrenaica Rouch 1970

Elaphoidella reducta Rouch 1964

Elaphoidella vandeli Chappuis & Rouch 1958

Elaphoidella vasconica Rouch 1970

Ceuthonectes boui Apostolov 2002

Ceuthonectes bulbiseta Apostolov 2002

Ceuthonectes chappuisi Rouch 1980

Ceuthonectes gallicus Chappuis 1928

Ceuthonectes serbicus Chappuis 1923/24

591

Appendix 1. (continued)

Ceuthonectes vievilleae Rouch 1980

Moraria catalana Chappuis & Kiefer 1952

Moraria varica (Graeter 1911)

Bryocamptus (Articocamptus) vandowei (Kessler 1914)

Bryocamptus (Articocamptus) zschokkei triarticulata Kiefer 1929

Bryocamptus (Rheocamptus) alosensis Apostolov 1998

Bryocamptus (Rheocamptus) dentatus Chappuis 1937

Bryocamptus (Rheocamptus) pyrenaicus (Chappuis 1923)

Bryocamptus (Rheocamptus) unisaetosus Kiefer 1930

Antrocamptus catherinae Chappuis & Rouch 1960

Antrocamptus chappuisi Rouch 1970

Antrocamptus coffaiti Chappuis 1956

Antrocamptus longifurcatus Rouch 1970

Anlrocamptus stygius Rouch 1970

Parastenocarididae

Parastenocaris aedes Hertzog 1938

Parastenocaris corsica Cottarelli, Bruno & Berera 2000

Parastenocaris dentulatus Chappuis & Rouch 1959

Parastenocaris dianae Chappuis 1955

Parastenocaris fontinalis borea Kiefer 1960

Parastenocaris fontinalis fontinalis Schnitter & Chappuis 1915

Parastenocaris fontinalis meridionalis Hertzog 1936

Parastenocaris glareola Hertzog 1936

Parastenocaris hippuris Hertzog 1938

Parastenocaris mangini Rouch 1990

Parastenocaris micheli Chappuis & Rouch 1959

Parastenocar is nertensis Rouch 1990

Parastenocaris nicolasi Rouch 1996

Parastenocaris psammica Songeur 1961

Parastenocaris silvana Cottarelli, Bruno & Berera 2000

Parastenocaris stammeri gallicus Chappuis & Rouch 1959

Parastenocaris vandeli Rouch 1987

COPEPODA, GELYELLOIDA

Gelyellidae

Gelyella droguei Rouch & Lescher-Moutoue 1977

COPEPODA, CALANOIDA

Diaptomidae

Spelaeodiaptomus rouchi Dussart 1970

SYNCARIDA

Bathynellidae

Bathynella gallica Delamare & Chappuis 1954

Bathynella natans natans Vejdovsky 1882

Bathynella natans picardi Delamare 1961

Bathynella pyrenaica Delamare & Chappuis 1954

Delamareibathynella debouttevillei Serban 1989

Delamareibathynella motasi Serban 1992

Gallobathynella (Clamousella) delayi (Serban, Coineau & Delamare 1971)

Gallobathynella (Gallobathynella) boui Serban, Coineau & Delamare 1971

Gallobathynella (Gallobathynella) coiffaiti (Delamare 1963)

Gallobathynella (Gallobaihynella) juberthiae Serban, Coineau & Delamare 1971

Gallobathynella (Gallobathynella) tarissei Serban, Coineau & Delarnare 1971

592

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