International Symposium Jointly Organized By

Couverture 15/12/05 18:31 Page 1

International symposium jointly organized by the University Claude Bernard of Lyon 1 and CNRS within the frame of PASCALIS (Protocols for the ASsessment and Conservation

of Aquatic Life In the Subsurface) Editor : J. Gibert European research programme (contract EVK2-CT-2001-00121) http://www.pascalis-project.com

For ordering information, please contact :

Equipe Hydrobiologie et Ecologie Souterraines Janine Gibert - Université Claude Bernard Lyon 1 UMR-CNRS 5023 - Laboratoire des Hydrosystèmes Fluviaux 43 Bd du 11 Novembre 1918 - 69622 Villeurbanne cedex - France

E-mail : [email protected] Phone : +33 (0)4 72 44 82 59 Fax : +33 (0)4 72 43 15 23

HBES Subterranean BiodiversitySymposium on World - Proceedings HBES SYMPOSIUM on WORLD SUBTERRANEAN BIODIVERSITY

Held on 8 - 10 December 2004 University Claude Bernard of Lyon 1 - CNRS Villeurbanne, France

PROCEEDINGS

November 2005 Edited by Janine Gibert

HBES Published by : Equipe Hydrobiologie et Ecologie Souterraines Université Claude Bernard Lyon 1 UMR-CNRS 5023 Laboratoire des Hydrosystèmes Fluviaux 43, Bd du 11 Novembre 1918 69622 Villeurbanne cedex, France

For ordering information, please contact : J. Gibert Email : [email protected] Phone : +33 (0)4 72 44 82 59 Fax : +33 (0)4 72 43 15 23

The Subterranean Hydrobiology and Ecology research team (UMR/CNRS 5023, Laboratory of Fluvial Hydrosystems Ecology) is a University structure affiliated to the CNRS (French National Centre for Scientific Research); its activities include teaching, research, knowledge dissemination and protection of environment. The main research objective is understanding the adaptation to groundwater life. It also analyses the structure, role and determinants of groundwater biodiversity and the functioning of ecosystems. For further information : http://groundwater-ecology.univ-lyon1.fr/ http://www.pascalis-project.com/

Realised by A. Papin, Subterranean Hydrobiology and Ecology Research team, UCBL. Printed in France by Team Rush, Villeurbanne. Cover illustration by HBES. ORGANIZER

Conference co-ordinateur : Janine Gibert

Subterranean Hydrobiology and Ecology research team UMR-CNRS 5023 - Laboratory of Ecology of Fluvial Hydrosystems

University Claude Bernard of Lyon 1 43, Bd du 11 Novembre 1918 69422 VILLEURBANNE cedex, France.

SCIENTIFIC COMMITTEE

Anton Brancelj, National Institute of Biology, Ljubljana (Slovenia) Ana I. Camacho, National Museum of Natural Sciences, Madrid (Spain) David C. Culver, American University, Department of Biology, Washington DC (USA) Dan L. Danielopol, Institute of Limnology, Mondsee (Austria) Louis Deharveng, National Museum of Natural Sciences, Paris (France) Diana M.P. Galassi, Department of Environmental Sciences, L’Aquila (Italy) Janine Gibert, University Lyon 1, Fluvial Hydrosystems Ecology (France) William F. Humphreys, Western Australia Museum, Perth (Western Australia) Patrick Martin, Royal Belgian Institute of Natural Sciences, Brussels (Belgium) Guiseppe Messana, CNR, Florence (Italy) Boris Sket, Department of Biology, University of Ljubljana (Slovenia)

ORGANIZATION COMMITTEE

Janine Gibert Chairman (Professor, University Claude Bernard of Lyon 1, CNRS, AFL) Fabiana Castellarini (Post-doctoral fellow, University Claude Bernard of Lyon 1) Michel Creuzé des Châtelliers (Lecturer, University Claude Bernard of Lyon 1) Rita DeOliveira (PASCALIS coordination Assistant , University Claude Bernard of Lyon 1) Marie-José Dole-Olivier (Researcher, CNRS) Christophe Douady (Lecturer, University Claude Bernard of Lyon 1) Frédéric Hervant (Lecturer, University Claude Bernard of Lyon 1) Florian Malard (Researcher, CNRS) Dominique Martin (Engineer Assistant, CNRS) Jacques Mathieu (Professor University Claude Bernard of Lyon 1) Florian Mermillod-Blondin (Researcher, CNRS) Annick Papin (PASCALIS coordination Assistant , University Claude Bernard of Lyon 1) Jean-Louis Reygrobellet (Lecturer, University Claude Bernard of Lyon 1) SYMPOSIUM SUPPORTERS

The Subterranean Ecology and Hydrobiology research group greatly appreciates the following organizations and institutions : HBES

for their organizational support :

University Claude Bernard of Lyon 1 (UCBL), Laboratory of Fluvial Hydrosystems Ecology Conseil Scientifique

Centre National de la Recherche Scientifique (CNRS)

Ezus

the town council of Villeurbanne

for their financial support :

the European Community : PASCALIS research programme (contract : EVK2-CT-2001-00121)

IFR 41 Direction des Relations Internationales (DRI)

University Claude Bernard of Lyon 1 Institut Français de la Biodiversité (IFB)

Agence de l’Eau Rhône-Méditerranée-Corse

the Rhône-Alpes region

the Rhône Department

and their advertising sponsorship :

Association Française de Limnologie (AFL)

TABLE OF CONTENTS

SYMPOSIUM PRESENTATION Introduction J. Gibert ...... p 13 Programme ...... p 15

GENERAL INTRODUCTION TO BIODIVERSITY - Extended abstracts Global Change and biodiversity S. Morand ...... p 21 The different values of biodiversity and ethical consequences P. Joly ...... p 25 The Struggle to Measure Subterranean Biodiversity D.C. Culver ...... p 27 Why and how to take care of subterranean aquatic microcrustaceans ? D.L. Danielopol and P. Pospisil ...... p 29

PASCALIS GENERAL PRESENTATION - Extended abstracts GROUNDWATER BIODIVERSITY Protocols for the ASsessment and Conservation of Aquatic Life In the Subsurface (PASCALIS): overview and main results J. Gibert, A. Brancelj, A. Camacho, F. Castellarini, C. De Broyer, L. Deharveng, M.-J. Dole-Olivier, C. Douady, D.M.P. Galassi, F. Malard, P. Martin, G. Michel, B. Sket, F. Stoch, P. Trontelj and A.G. Valdecasas ...... p 39

SESSIONS - Extended abstracts Sessions Report D. Culver, L. Deharveng, A. Brancelj, B. Sket ...... p 55 Emerging knowledge of diversity, distribution and origins of some Australian stygofauna W.F. Humphreys, C.H.S. Watts and J.H. Bradbury ...... p 57 Assessment and conservation of aquatic life in the subsurface of the Pilbara region, Western Australia S.M. Eberhard, S.A. Halse, M.D. Scanlon, J.S. Cocking and H.J. Barron ...... p 61 Haplotype diversity in Pilbarus millsi, a widespread groundwater species of amphipod from the Pilbara, Western Australia T. Finston, M. Johnson, W. Humphreys, S. Eberhard, S. Halse ...... p 69 Phylogeography and taxonomic status of Niphargus virei (subterranean amphipod) T. Lefébure, C.J. Douady, M. Gouy, P. Trontelj, J. Briolay and J. Gibert ...... p 73 Hierarchical patterns of obligate groundwater biodiversity in France D. Ferreira, F. Malard, M.-J. Dole-Olivier and J. Gibert ...... p 75 Environmental gradients in ground waters. Main factors driving the composition of stygobiotic assemblages at a regional scale. M.-J. Dole-Olivier, F. Malard and J. Gibert ...... p 79 Improving the assessment of groundwater biodiversity by exploring environmental heterogeneity at a regional scale. F. Castellarini, M.-J. Dole-Olivier, F. Malard and J. Gibert ...... p 83 Mapping the stygofauna of the state of Baden-Württemberg, Southwest Germany H-.J. Hahn and A. Fuchs ...... p 89 Biodiversity of Belgian groundwaters : the Meuse basin P. Martin, C. De Broyer, F. Fiers, G. Michel, R. Sablon and K. Wouters ...... p 95 Distribution of groundwater invertebrates along an environmental gradient in a shallow water-table aquifer F. Paran, F. Malard, J. Mathieu, M. Lafont, D.M.P. Galassi and P. Marmonier ...... p 99 Does groundwater recharge stimulate biodiversity ? T. Datry, F. Malard and J. Gibert ...... p 107 Environmental quality of deep groundwater in the Lessinian Massif (Italy): signposts for sustainability T. Di Lorenzo, F. Stoch, B. Fiasca, E. Gattone, P. De Laurentiis, F. Ranalli and D.M.P. Galassi ... p 115

SESSIONS - Oral communication abstracts Distribution of the speciose genus Vestalenula, Rossetti & Martens, 1998 (Darwinulidae, Ostracoda), with the description of a new species from the Roussillon region (South-Eastern France) M. Artheau ...... p 129 Cladocera and Calanoida Ð Two interesting groups of stygobionts A. Brancelj and N. Mori ...... p 129 Use of groundwater copepods as a tool for environmental management : the case of Everglades National Park, Florida, USA M.C. Bruno and V. Cottarelli ...... p 130 COI data indicate a complex evolutionary history for Italian subterranean and surface populations of Proasellus (Crustacea, Isopoda) A. Campanaro, V. Ketmaier and R. Argano ...... p 131 Patterns of endemism of the Eastern North American cave fauna D.C. Culver, M.C. Christman, M. Madden, and D. White...... p 132 Are Bayesian inferences a useful tool for phylogeographers ? C.J. Douady and T. Lefébure ...... p 132 Hyporheic Copepod assemblages as a tool for detecting man-induced perturbation on GW/SW ecotone B. Fiasca, T. Di Lorenzo, P. De Laurentiis, D. M. P. Galassi ...... p 133 Population structure of stygobitic Isopods in the Pilbara, Western Australia C.J. Francis, M.S. Johnson and T. Finston...... p 133 Ectinosomatidae (Copepoda, Harpacticoida) in groundwater : radiation or multiple invasions ? R. Huys, D.M.P. Galassi and P. De Laurentiis ...... p 134 Ground-water flow and aquatic habitats W.K. Jones ...... p 134 Riverscape dynamics and the distribution of the hyporheos in a glacial river floodplain system F. Malard and M.Lafont ...... p 135 Epikarst fauna from North American caves T. Pipan and D. C. Culver ...... p 136 The relationship between Ostracod species diversity and water chemistry in the subsurface of the Pilbara region, Western Australia J. Reeves ...... p 137 The blind misleading the blind : false negatives and estimations of subterranean biodiversity K. Schneider, D.C. Culver, H.H. Hobbs III, D. Fong ...... p 138 Some auxiliary criteria for the selection of groundwater sites for biodiversity conservation B. Sket ...... p 138 Subterranean biodiversity in Great-Britain and Ireland : a predominantly post-glacial colonising fauna with possible trans-glacial elements P.J. Wood and G.S. Proudlove ...... p 139 POSTERS - Extended Abstracts A possible mechanism of participation of diapause in Cyclopid penetration into underground environments (Mongolian springs, well and in Sablinskije caves) V.R. Alekseev ...... p 143 Interstitial harpacticoids from volcanic lakes of Latium (Italy) : state of the art R. Berera, M.C. Bruno, L. Pariciani and V. Cottarelli ...... p 145 Distribution atlas of the stygobiont molluscs of France A. Bertrand ...... p 149 Hyporheic zone as refugium for the macroinvertebrate fauna in the Po river (NW Italy) : first data S. Fenoglio, T. Bo, M. Cucco and G. Malacarne ...... p 153 Microinvertebrates occurrence in ground-waters L. Mancini, D. Venanzi, L. Volterra, B. Pennelli and P. Formichetti ...... p 157 Diversity of subterranean fishes in Brazil E. Trajano and M. E. Bichuette ...... p 161

POSTERS Ð Abstracts A new subterranean catfish, genus Rhamdia, from Brazil (Siluriformes : Heptapteridae) with notes on ecology M.E. Bichuette and E. Trajano ...... p 165 Habitat determination and subterranean singularity A.I. Camacho, A.G. Valdecasas and J. Rodriguez ...... p 165 Effects of base flow reduction on aquatic ecosystems G. Cecchi, P. Formichetti and L. Mancini ...... p 166 Interstitial fauna dynamic in an island-braided flood plain (Tagliamento River, Italy) C. Claret and K. Tockner ...... p 166 Diversity and origin of the interstitial Isopod Microcharon (Crustacea, Microparasellidae) from the Roussillon and Western Languedoc regions (France) N. Coineau, M. Artheau, A. Bedos, M. Boulanouar, C. Boutin, F. Brehier, L. Deharveng, N. Giani ...... p 167 Obligate groundwater fauna of France : species diversity patterns and conservation implications D. Ferreira, F. Malard, M.-J. Dole-Olivier and J. Gibert ...... p 168 Diversity patterns of Copepod assemblages (Crustacea, Copepoda) of different caves from Southern and Central Italy E. Gattone, T. Di Lorenzo, B. Fiasca, P. De Laurentiis and D.M.P. Galassi ...... p 169 The stygobiontic fauna of Tunisia : preliminary results A. Ghlala, D. Della Vale, F. Charfi and G. Messana ...... p 169 Relationship between morphological taxonomy and molecular divergences : evidences from crustaceans T. Lefébure, C.J. Douady, M. Gouy and J. Gibert ...... p 170

SYMPOSIUM PARTICIPANTS ...... p 171

AUTHOR INDEX ...... p 179

SYMPOSIUM PICTURES ...... p 181

SYMPOSIUM PRESENTATION

SWSB December 2004 11

SYMPOSIUM PRESENTATION

INTRODUCTION

J. GIBERT

University Claude Bernard of Lyon 1, UMR/CNRS 5023, Laboratoire d’Ecologie des Hydrosystèmes Fluviaux, Equipe d’Hydrobiologie et Ecologie Souterraines, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France. ([email protected]). Groundwater biodiversity still remains largely than today, in particular in biodiversity-rich badly known compared to the biodiversity of regions. freshwater epigean habitats. Subterranean This international workshop was intended to : a) environment however, harbours a fauna of high mark the end of the European programme EVK2- originality, characterized by a moderate specific CT 2001-00121 PASCALIS (Protocols for the diversity and an extreme richness in endemics ASsessment and Conservation of Aquatic Life in and in phyletically isolated taxa within the the Subsurface) that ran from year 2002 to 2004; present fauna (Marmonier et al. 1993, Danielopol b) synthesize the expertise in the study of et al. 2000, Gibert & Deharveng 2002). terrestrial and aquatic subterranean communities Distribution of this diversity on the different and their dynamics in various countries of the continents is highly unequal, very changeable world, and c) open up new research and according to systematic groups and often highly cooperation ways in the field of underground unpredictable considering the current ecological biodiversity in relationship with the conservation conditions. Among the scientific fundamental and management policies on a regional, issues raised by the conservation of European and world scale. subterranean species, appear the questions relative to their origin and to the respective role of The scientific programme was composed of 3 historical, ecological and genetic factors in their parts, and included 37 oral communications in persistence during evolution. Our knowledge total according to the following breakdown : a) about the geographical distribution of these general introduction to Biodiversity presented by species is moreover very fragmentary; even in 3 invited speakers and 1 research team leader Europe, it is not possible at present to work out from our laboratory : 4 communications, b) efficient protection plans for this often threatened general presentation of the PASCALIS project by biological inheritance. the coordinator and partners or workpackage leaders : 7 communications, c) four scientific Groundwater represents the major source of sessions : Session 1: Australian stygofauna and drinking water for most Continents. Western Australian project (Pilbara region): 5 Nevertheless, most regulations pertaining communications; Session 2: Phylogeny, groundwater protection are inexorably linked with Phylogeography and Taxonomy : 7 the drinkableness of the water resource. In the communications; Session 3: Biodiversity patterns World, groundwater is severely threatened by : 9 communications; Session 4: Conservation human activities, with quantitative and qualitative and management : 5 communications rebound effects on both groundwater communities and ecological processes. The Symposium was successful since 88 Maintaining groundwater quality and conserving participants came from 14 different countries with its biodiversity are converging goals due to the following distribution: France: 44 participants, strong link existing among them (Gibert et al. Italy: 11, Slovenia: 6, Belgium: 4, UK: 2, Spain: 2, 1994, Griebler et al. 2001, Danielopol et al. Austria: 1, Russia: 1, Denmark: 1, USA: 3, 2003). Moreover, a rational management of Australia: 5, Brazil: 2 and Tunisia: 1. 37 oral groundwater resources cannot ignore the added communications have been presented and 20 value given by its unique biodiversity. posters have been exhibited on 8th and 9th Understanding the diversity of aquatic life in the December. subsurface is a necessary step for incorporating The Symposium enabled to study biodiversity current biological concepts within the framework under various aspects and to provide an of groundwater management. integrated approach of groundwater biodiversity. Finally, the strategies likely to maintain survival of It disclosed the latest scientific breakthroughs subterranean species and more specifically of revealed by PASCALIS. At present, it is possible endemics in the face of the regional socio- to map subterranean biodiversity in several economic requirements still remain to be defined. European countries outside the six partner At all these levels, a much more rigorous countries. Methods to rigorously sample a scientific basis would enable to target defined region, to analyze, predict and conserve management solutions much more efficiently biodiversity are now made available.

SWSB December 2004 13 SYMPOSIUM PRESENTATION

The wide range of countries represented by the Gibert J., D.L. Danielopol, and J.A. Stanford. 1994. participants’ ensured promotion and Groundwater ecology. Academic Press, San Diego. dissemination of the PASCALIS results on a Gibert, J. and L. Deharveng. 2002. Subterranean worldwide scale. Future collaborations with other ecosystems: a truncated functional biodiversity. countries such as the United States and Australia Bioscience 52:473-481. are being undertaken, as well as other European Griebler C., D.L. Danielopol, J. Gibert, H.P. Nachtnebel, and J. Notenboom. [Eds]. 2001. Groundwater countries (Germany and Austria). Ecology. A tool for management of water resources. EC Conference proceedings. European commission; REFERENCES Environment and climate programme, Bruxelles, 413 p. Danielopol, D.L., P. Pospisil and R. Rouch. 2000. Biodiversity in groundwater: a large-scale view. Marmonier, P., P.H. Vervier, J. Gibert and M.-J. Olivier. Trends in Ecology and Evolution 15:223-224. 1993. Biodiversity in ground waters. Trends in Ecology and Evolution 8:392-395. Danielopol, D.L., C. Griebler, A. Gunatilaka and J. Notenboom. 2003. Present state and future prospects for groundwater ecosystems. Environmental Conservation 30:104-130.

14 SWSB December 2004 SYMPOSIUM PRESENTATION

PROGRAMME

14.00-18.30 Registration session 18.30-19.00 Bus departure to Villeurbanne town hall 19.00-20.00 Welcoming reception by the mayor of Villeurbanne Wednesday 8th December 9.00-9.30 J. Gibert (organizer), P. Lantéri (University Lyon 1) Opening session and B. Andral (CNRS) in presence of institutional representatives General introduction to biodiversity Chairman : E. Pattee 9.30-10.00 S. Morand “Global change and biodiversity” CNRS-IRD Ð France 10.00-10.30 P. Joly “Values of biodiversity and motivation for its conservation” University Claude Bernard Lyon 1 - France Break 11.00-11.30 D. C. Culver “The struggle to measure subterranean biodiversity” American University Ð Washington DC Ð USA 11.30-12.00 D. L. Danielopol “Why and how to take care of subterranean aquatic Austrian Academy of Sciences - Austria microcrustaceans” General presentation of PASCALIS European research programme Chairman : J. Gibert 12.00-12.30 J. Gibert “How to assess and conserve groundwater biodiversity. Project coordinator The PASCALIS European project in the international context : main results” Lunch 14.00-14.30 L. Deharveng “The PASCALIS data base and the diversity of Natural History Museum Ð Paris - France stygobitic fauna in Europe.” 14.30-15.00 P. Trontelj, C. Douady “Cryptic diversity of selected groundwater taxa” University of Ljubljana Ð Slovenia 15.00-15.30 F. Stoch “Biodiversity indicators” University of L’Aquila Ð Italy Break 16.00-16.30 F. Malard “Additive partitioning of stygobiont species richness in CNRS-University Claude Bernard Lyon 1 Ð France Southern Europe” 16.30-17.00 F. Stoch “Assessing the conservation value of groundwater University of L’Aquila Ð Italy species” 17.00-17.30 G. Michel, F. Malard “Design of a network of priority sites for conserving Walloon Commission for Study and Protection of groundwater biodiversity in Europe” Subterranean Sites - Belgium 17.30-18.30 Poster session 20 posters Thursday 9th December Session 1 Australian Stygofauna and Western Chairman : Dan Danielopol Australian project (Pilbara region) Reporter : David Culver 9.00-9.20 W.F. Humphreys,C.H.S. Watts, J.H. Bradbury “Emerging knowledge of diversity, distribution and Western Australian Museum Ð South Australia origins of some Australian stygofauna” 9.20-9.50 S. Eberhard, S. Halse “Assessment and conservation of aquatic life in the Department of Conservation and Land Management - Australia subsurface of the Pilbara region in Western Australia.” 9.50-10.10 J. Reeves “The relationship between Ostracod species diversity and Australian National University - Australia water chemistry in the subsurface of the Pilbara region, Western Australia” 10.10-10.30 T. Finston, M.S. Johnson, S. Eberhard, S. Halse “Allozyme and haplotype diversity in Pilbarus Millsi, a The University of Western Australia Ð Crawley - Australia widespread groundwater species of amphipod from the Pilbara, Western Australia”

SWSB December 2004 15 SYMPOSIUM PRESENTATION

Break 11.00-11.20 C. J. Francis, M.S. Johnson, T. Finston “Population structure of stygobitic isopods in the Pilbara, The University of Western Australia Ð Perth - Australia Western Australia.” Session 2 Phylogeny, Phylogeography and Chairman : Ana Camacho Taxonomy Reporter : Louis Deharveng 11.20-11.40 A. Campanaro, V. Ketmaier, R. Argano “COI data indicate a complex evolutionary history for University of Rome “La Sapienza” - Italy Italian sub-terranean and surface populations of Proasellus (Crustacea, Isopoda)” 11.40-12.00 T. Lefébure, C.J. Douady, M. Gouy, P. Trontelj, “Phylogeography and taxonomic status of Niphargus virei J. Briolay, J. Gibert (subterranean amphipod)” University Claude Bernard Lyon 1 Ð France 12.00-12.20 C. J. Douady, T. Lefébure “Are Bayesian inferences a useful tool for University Claude Bernard Lyon 1 Ð France phylogeographers ?” Lunch 14.00-14.20 P.J. Wood, G. S. Proudlove “Subterranean biodiversity in Great-Britain and Ireland : Loughborough University - UK a predomi-nantly post-glacial colonising fauna with possi ble trans-glacial elements” 14.20- 14.40 R. Huys, D. Galassi, P. De Laurentiis “Ectinosomatidae (Copepoda, Harpacticoida) in The Natural History Museum Ð London - UK groundwater : radiation or multiple invasion ?” 14.40-15.00 A. Brancelj, N. Mori “Cladocera and Calanoida – two interesting groups of National Institute of Biology Ð Ljubljana - Slovenia stygobionts” 15.00-15.20 M. Artheau “Distribution of the speciose genus Vestalenula, Rosseti & University Paul Sabatier Ð Toulouse - France Martens, 1998 (Darwinulidae, Ostracoda), with the description of a new species from the Roussillon Region (South Eastern France)” Break Session 3 Biodiversity patterns Chairman : Diana Galassi Reporter : Anton Brancelj 16.00-16.20 D.C. Culver, M.C. Christman, M. Madden, “Patterns of endemism of the Eastern North American D. White cave fauna” American University Ð Washington DC Ð USA 16.20-16.40 D. Ferreira, F. Malard, M.-J. Dole-Olivier, “Hierarchical pattern of obligate groundwater biodiversity in J. Gibert France” University Claude Bernard Lyon 1 Ð France 16.40-17.00 H.J. Hahn, A. Fuchs “ Mapping the stygofauna of the state Baden-Württemberg, University of Koblenz-Landau - Germany Southwest Germany.” 17.00-17.20 K. Schneider, D.C. Culver, H.H. Hobbs, “The blind misleading the blind : false negatives and D. Fong estimations of subterranean biodiversity.” University of Maryland Ð College Park - USA

20.00 Bus departure from the University campus to the restaurant 20.15-23.00 Banquet dinner and return by bus

Friday 10th December Session 3 Biodiversity patterns : continuation 9.30-9.50 P. Martin, C. De Broyer, F. Fiers, G. Michel, “Biodiversity of Belgian groundwaters : the Meuse Basin” R. Sablon, K. Wouters Royal Belgian Institute of Natural Sciences Ð Brussels Ð Belgium 9.50-10.10 T. Pipan, D.C. Culver “Epikarst fauna from North American caves” Karst Research Institute Ð Postojna Ð Slovenia 10.10-10.30 F. Malard, M. Lafont, “Riverscape dynamics and the distribution of the Cemagref Ð Lyon Ð France hyporheos in a glacial river-floodplain system” Break

16 SWSB December 2004 SYMPOSIUM PRESENTATION

11.00-11.20 F. Paran, F. Malard, J. Mathieu, M. Lafont, “Distribution of groundwater invertebrates along an D. Galassi, P. Marmonier environmental gradient in a shallow water-table aquifer” Ecole National Supérieure des Mines – Saint-Etienne – France 11.20-11.40 W. K. Jones “Ground-water flow and aquatic habitats” Karst Waters Institute Ð Charles Town Ð USA

Session 4 Conservation and management Chairman : Patrick Martin Reporter : Boris Sket 11.40-12.00 B. Sket “Some auxiliary criteria for the selection of groundwater University of Ljubljana Ð Slovenia sites for biodiversity conservation” 12.00-12.20 M.C. Bruno, V. Cottarelli “Use of groundwater copepods as a tool for environmental University of La Tuscia Ð Viterbo- Italy management : the case of Everglades National Park, Florida, USA” Lunch 14.00-14.20 T. Datry, F. Malard, J. Gibert “Response of invertebrate assemblages to increased University Claude Bernard Lyon 1 Ð France groundwater recharge rates in a phreatic aquifer” 14.20-14.40 B. Fiasca, T. Di Lorenzo, P. De Laurentiis, “Hyporheic copepod assemblages as a tool for detecting D.M.P. Galassi man-induced perturbation on GW/SW ecotone.” University of L’Aquila Ð Italy 14.40-15.00 T. Di Lorenzo, B. Fiasca, F. Stoch, “Environmental quality of deep groundwater in the D.M.P. Galassi Lessinian mountains (Italy) : signposts for sustainability.” University of L’Aquila Ð Italy

15.00-15.50 Report on the sessions 5 reports 15.50-16.00 J. Gibert Closing of the Symposium Break

SWSB December 2004 17

GENERAL INTRODUCTION TO BIODIVERSITY

Extended abstracts

SWSB December 2004 19

GENERAL INTRODUCTION TO BIODIVERSITY

GLOBAL CHANGE AND BIODIVERSITY

S. MORAND

CNRS-IRD, Centre de Biologie et de Gestion des Populations (CBGP), Campus International de Baillarguet, CS 30 016, 34988 Montferrier-sur-Lez Cédex, France. ([email protected]).

KEYWORDS : CLIMATE CHANGE, PROJECTIONS

1. GLOBAL CHANGE increase in the earth mean temperature. Everything leads us to believe that due to the Aquatic and terrestrial ecosystems are being climatic systems inertia, an in-depth long-term subjected to multiple and synergetic impacts change of climate is to be expected. Climatic from human activities. These activities, that can change is a reality (IPPC, www.ipcc.ch). The be expressed by a measure as ecological print, increasing rate of CO and other greenhouse are characterized by increases and changes in 2 gases (N2O, CH4) leads to an increase in the the use of biological resources and ecosystems, global earth temperature. Everything indicates and by an impact on earth climate. that because of the inertia of the main world Ever more significant modifications in the uses climatic systems, the expansion of the impacts apply to all environments and are characterized should accelerate during the XXIth century with a by deforestations or reforestations, by global warming from 1.4 to 5.8¡ C and an expansions or abandonments of farmland, by increase in the sea level ranging from 20 cm to enlarging urban and peri-urban areas (increasing 80 cm. Important researches are carried out on urban sprawl), intensive exploitation of the climate dynamics in order to better predict renewable resources such as fishings, various future and anticipate adaptations of our societies. chronic pollutions, intensive water extraction…. All these research works propose more accurate The living itself is also directly submitted to new and localized projections. In addition to a global exploitations. Thus, the production of transgenic temperature increase, the first trends observed species (GMO) for agricultural purposes and for show a raise in the oceans level, a modification resistance to pesticides or bio-stressors is in the rain systems, a modification in soil immediately followed by their release in open humidity, a development of the ENSO et NAO fields, in several countries and on all continents. phases, and potentially, an impact on Economic globalization leads to important thermohaline circulations (the great conveyor commercial relationships and thus intensifies the belt). The effects of climatic change will also circulation of species on a worldwide scale, affect biogeochemical cycles as well as many either for agricultural and zootechnical purposes, ecosystems: coral reefs, forests and meadows, but also unintentionally. New commercial routes, mangrove swamps… and particularly South-to-South routes with new Interactions between use and climate changes, emerging economic powers, are going to joined under the global change concept, and increase introductions between distinct biodiversity cause a situation our planet has biogeographical zones but with comparable never experienced. The causes for this global climates. The massive use of antibiotics, animal change are thus linked to human activities and to and human vaccinations lead to select more and this tremendous human ecological print on more resistant and virulent pathogenic strains. energy resources, lands, oceans and living The same statement can be made for the case of resources. struggles against the spread of bio-stressors and pathogenic vectors showing multi-resistance 2. IMPACTS OBSERVED ON THE LIVING to various insecticides, anti-mollusc and 2.1 Distribution and phenology areas antifungal molecules… The impacts of global change can already be Ever more increasing energy needs useful to the observed on the living. Several researches world economy generate as well a significant indicate the effects of climatic change on the increase in the production and concentration of modifications of distribution areas, in latitude and greenhouse gases (nitrous oxide, carbon altitude, and for numerous plant and animal dioxide, methane) in the atmosphere, the species. In the same way, researches show the impacts of which can be already observed by the phenological changes (age at reproduction or at

SWSB December 2004 21 GENERAL INTRODUCTION TO BIODIVERSITY emergence) deeply affecting wild and cultivated 3.4 Projecting the impacts of biological plant species (for example wine grape harvest invasions date). Scenarii on biological invasions must be here again proposed, and following questions be 2.2 Biological resources : decline in answered : Which type of organism ? Which fishing areas at risk ? Which impacts in average ? Which Halieutic resources are influenced by climate future ? The answers are essential to manage variations (such as ENSO), but a decline in bioinvasions, upstream (quarantine fishing activities can be registered (such as cod mechanisms) and downstream (landscape in the North Atlantic) because of overfishing. ecology, eradication). Both combined phenomena increase the risk of collapse in the whole fishing activities. 3.5 Projecting the impacts on health (plants, animals, human) 2.3 Biological introductions, bio-invasions Climate has impacts on animal or human health. Introductions are more and more frequent and as Episodes such as El Nino type lead to an a consequence, the related problems more increase in coral bleaching and cholera acute. Furthermore, the interaction between epidemic. A raise in CO2 can favor the climatic change and biodiversity losses might development of plant pathogens. Climate change facilitate the settlement of invasive species and thus exerts a direct influence on the relationships reinforce their aggressive potential for the between pathogens and hosts. The fields of pathogenic species. human health, animal health and plant health are concerned. 3. WORK OUT PROJECTIONS FOR BIODIVERSITY 3.6 Projecting extinctions In biodiversity, projections should also be drawn The rate of extinctions would be currently up. They are essential to establish a dialogue accelerated by the impacts of global change. It between researchers, decision-makers and would be advisable to better quantify and qualify managers. They require to build multidisciplinary them : which diversity of the living ? in which research projects. regions, and on which biogeographical scales ?

3.1 Projecting future distributions 3.7 Projecting zones for biodiversity conservation One of the first challenges is to plan the future distribution of the living. Some models show that, The hotspots are areas defined upon their very in some cases, species must migrate at 1 high taxonomic richness and their high levels of km/year (equivalent to speeds of postglacial endemism. Efforts in simulation and projection of recolonisation) to reach a new suitable area. the impact from the climate change will allow to better predict the changes in the species These distributions concern Ç wild È species as distribution. However, and this represents quite a much as domesticated or cultivated species. gap with the past, the human ecological print Uses and further regulations for these new may be too much important to enable any distribution areas will have to be re-defined. movement of biomes because of urbanisation, or the use of some areas for agricultural production. 3.2 Projecting adaptive responses The protected areas, already often at stake in Species and their populations do not respond major issues and sustainability of which is only by a modification of their distribution areas. questioned, will have to anticipate impacts of The global change affects selection processes global change. and biotic interactions (predation, parasitism, symbiosis). The adaptive responses from 4. RESEARCH LINES populations et species and their abilities to 4.1 Role et fonction of biodiversity evolutive response will have to be better Biodiversity cannot be reduced to sheer specific approached. diversity, but this does not mean species diversity is not important per se. Biodiversity is dynamics 3.3 Projecting the evolution of living of interactions in which human is more and more resources involved. Human benefits from biodiversity and Fishing, forest exploitations, agriculture are all from goods and services provided by activities that are going to be impacted by global ecosystems. However, these human activities change but in which ways ? How to plan the generating a significant global change will adaptation of human activities related to the directly act upon the functioning of ecosystems exploitation of living resources to the impacts of and thus modify the advantages and benefits global change ? brought by the latter.

22 SWSB December 2004 GENERAL INTRODUCTION TO BIODIVERSITY

A first line of research aims thus to better won’t be able to develop empirical studies and to understand the functioning of ecosystems and exploit at best the data from observations. the relationship between biodiversity and this functioning, and to which extent this relationship 5. TOOLS FOR BIODIVERSITY can be modified by global change. As a consequence, we should also better understand Studies in biodiversity require some specific how the changes in biodiversity and in the tools: ecosystems functioning can have impacts on ¥ for the identification of organisms global change. (systematics) and references’ collections (Natural History Museum and Centres of Biological Resources) ; 4.2 Better understand interactions ¥ interconnected databases joining Biodiversity implies interactions. Theoretical as systematics, life- history traits, distribution well as empirical researches are still needed in areas, genetic information... ecology of communities and the interactions ¥ Biodiversity observatories in order to among species and populations and in conduct long-term monitoring, evolutionary biology. The interactions of ¥ for experimental testing : ecotrons, predators/preys, hosts/parasites or symbiosis manipulations of ecosystems… type or pollination type are and will become ¥ to model linkages of climatic, economical, impacted. One can wonder if the rate of global social, biotic variables… change does not overtake the potential rate of These studies require also the elaboration of new the adaptive change…. tools in order to facilitate interconnection and In terms of research, we have significant needs mutual development between life, environmental, for theoretical ecology. Without any theory, we human and social sciences.

SWSB December 2004 23

GENERAL INTRODUCTION TO BIODIVERSITY

THE DIFFERENT VALUES OF BIODIVERSITY AND ETHICAL CONSEQUENCES

P. JOLY

University Claude Bernard of Lyon 1, UMR CNRS 5023, Laboratoire d’Ecologie des Hydrosystèmes Fluviaux, Bât Darwin C, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne Cedex, France. ([email protected]).

The question “why preserve biodiversity?” results feelings, this cultural approach of Nature does more from philosophy than science because it not deeply differ from economic consumerism as deals with the conceptions of the relationships long as it does not break with duality (“Humans between Humans and Nature. These are evils who alter all things they touch”), it is conceptions are very diverse and, even within connected to economy (tourism, economic the present Western culture, several points of application of scientific knowledge), and it implies view are clearly perceptible, among which I will that Nature can be managed for producing such here consider anthropocentrism, ecocentrism cultural feelings (preservation of natural areas and biocentrism. Anthropocentrism can itself be from which humans are removed, creation of split between economic and cultural humanisms. gardens sensu lato). From this point of view, They mainly differ by the situation of Humans biodiversity takes an amenity value. with respect to Nature, and by the value humans Ecocentrism attempts to break with dualism in attach to biodiversity. setting Humans within Nature as a component of The roots of anthropocentrism are probably to be ecosystems. It considers that Nature is found in the skill of agriculture and food storage organised by regulating mechanisms that that released humans from the uncertainty of determine optimal conditions for life. Biodiversity day-to-day searching for food. The great plays an important role in such regulations efficiency of a technical approach of foraging (oxygen production, regulation of the water cycle, problems has opened the way for a radical elaboration of fertile soils, regulation of carbon dualism opposing the apparent chaos of wild dioxide, etc…). Because of resemblance with the Nature to the rationality of the human mind. This regulating mechanisms that ensure the dualism has first been reinforced by homeostasy of an organism, the Earth can be monotheisms that clearly make Humans special viewed as a superorganism for which it becomes creatures of divine essence facing a Nature that possible to establish health criteria. As a has to be dominated. Since the sixteenth century consequence, the future of Humans tightly it has also been reinforced by the emergence of depends on Earth health. The impact of certain modern science and the Cartesian conception human activities can then be considered as according to which Humans have received negative when such activities alter ecosystem science from God to take advantage of health. Continuing with the organism metaphor, understanding the mechanisms of Nature the exponential growth of human population (Humans have to focus on the “how” rather than might bring Humans to be assimilated with a on the “why” questions). Finally, the Age of cancer that can induce its own disappearance by Enlightenment confirmed the duality by killing its host. According to this conception, considering Humans as free and rational biodiversity has to be preserved because of its creatures extracted from naturalness, and who functional value. build themselves through experience and education (Rousseau, Kant). This laïc conception Biocentrism also breaks with dualism but on supports consumerist attitudes among which different grounds. From this point of view, liberalism and Marxism later became two biodiversity cannot be exploited as can be done alternatives. From this economic point of view, with inanimate things because i) of the role of biodiversity has a utility value (goods such as historical processes that forge its adaptive food, fuel, materials, energy, and services such nature, and ii) other living things share a similar as oxygen production, fertilisation, waste cycling, origin, nature and teleonomy with Humans. Both soil turbation, pollination). Humans can modify heritage nature and kinship with Humans give Nature for increasing the amount of such goods biodiversity an intrinsic value. and services they receive from it. Each of these points of view presents deep The cultural version of anthropocentrism caveats that impede any superiority of one of considers Nature as a source of personal them over the others. Because of individualistic blossoming, romantic wonder, knowledge egoism, anthropocentrism reveals its incapacity pleasure and mystical feeling. Although it is not to conceive a sustainable exploitation of Nature. easy to attribute an economic value to such The regulation of the exploitation of natural

SWSB December 2004 25 GENERAL INTRODUCTION TO BIODIVERSITY resources by market rules is an economic possible root for the conceptions of concept that is not adapted to the management Nature/Human relationships, and the only of most aspects of biodiversity. The philosophical reference of what they should be. But the foundations of anthropocentrism are themselves awareness of the recent biodiversity crisis shaken by the important weight modern biology implies the elaboration of new ethical pacts has given to genealogy in the determinism of between science, technology and economy. Like individual personality. Ecocentrism is burdened future generations and disadvantaged countries, with the difficulty of defining the healthy state of biodiversity is a low-power component of the an ecosystem otherwise than by referring to its political theatre at the global Earth level, and one utility, and by establishing a perspective to of the major externalities of human activities. human actions that would not be deeply Governance has to rely on regulatory constrained by a hypothetical common interest. mechanisms that would use biodiversity as an Biocentrism is burdened with the problem of indicator of sustainable economy. In this hierarchy of values between living things and by perspective, science has to get involved in a new attributing a negative value to enemies of human chain of interactions with technology and welfare such as pathogens. economy that should regulate the classical A critical appraisal of these positions leads us to runaway of sacrosanct growth and open the way think that anthropocentrism is unavoidable to sustainable development. because the human mind provides the only

26 SWSB December 2004 GENERAL INTRODUCTION TO BIODIVERSITY

THE STRUGGLE TO MEASURE SUBTERRANEAN BIODIVERSITY

D.C. CULVER

Department of Biology, American University, 4400 Massachusetts Ave. NW, Washington DC, 20016 USA. ([email protected]).

The subterranean fauna offers several for which there is an obvious confounding effect advantages to the biogeographer. First, it is a of area and sampling thoroughness. What is highly replicated environment in which many available is a list of relatively well-studied high successful independent evolutionary diversity caves developed by Culver and Sket experiments have occurred. Second, compared (2000) on a global basis and a list of high to most habitats, it is relatively easy to distinguish diversity tropical caves developed by Deharveng permanent residents (especially stygobionts and (2005). These studies of α-diversity suggest that troglobionts) from transients. Third, the high the highest diversity is a northern mid-latitude, levels of endemism at all scales make it an especially in Europe, and especially in the especially attractive model for the study of Dinaric Mountains. Within tropical latitudes, endemism. Fourth, the highly clustered diversity is highest in the Indo-Pacific region. geographic distribution make the subterranean realm a test case for the developing models of The causes of these patterns are not yet such distributions, such as zero-inflated Poisson completely understood, and in fact neither are distributions (Christman 2005). Finally, the large the patterns themselves. However, several points number of caves and other subterranean have emerged. The first is that cave density (and habitats results in a system that readily lends perhaps cave passage development) are an itself to quantification and statistical testing. Of important determinant of species numbers, course there are disadvantages as well. One especially troglobionts (Christman and Culver stands out. There is a perception that there is an 2001, Culver et al. 2003). The second is that apparent rarity of adaptation, especially new regional context matters. That is, caves in large character acquisition, compared to character loss regions of high cave density have more species (regressive evolution) that makes the than caves in small regions of high density, with subterranean biota a special rather than a a regional scales of tens and even hundreds of general case. Much research in the past several kilometers (Christman and Culver 2001, Culver decades has shown this to be wrong but the et al. 2004). Both of these factors point to the perception persists. importance of karst development in determining species number. I briefly touch on four topics—what are the biogeographic patterns, what are their causes, α how much confidence can we place in the If we base our results on -diversity, how much results, and where do we go from here. A key can we say about total diversity and with how advance was the summary of Gibert and much confidence? As (Gibert and Deharveng α Deharveng (2002), who pointed out that 2002) point out, -diversity and total diversity are subterranean communities share the following strongly correlated. And in fact, the most reliable features: information about regional diversity actually comes from a small number of caves, even in a ¥ High endemism result in low local diversity relatively well-studied area like Slovenia (Culver relative to regional diversity et al. 2004a). For example, the relationship ¥ A relative small number of subterranean between species number and number of caves is lineages resulting in taxonomic clearest when the 90th percentile of distribution disharmonies rather than the mean is regressed again number ¥ Many relicts, and of caves.ùù For two high diversity caves— ¥ Truncated food webs. Vjetrenica in Bosnia and Hercegovina (see Sket in Lucic (2003) and Mammoth Cave in Kentucky, To look for patterns, we must then move beyond USA—most of the species are not single cave the universals and look for factors that vary endemics but rather regional endemics, geographically. suggesting that these caves are representative of the region as a whole. Even for these two No summary of global subterranean biodiversity caves however, it is likely that new species will has been done beyond country lists of species continue to be discovered, as there is no sign

SWSB December 2004 27 GENERAL INTRODUCTION TO BIODIVERSITY that the accumulation curve of species number is Culver, D.C., M.C. Christman, W.R. Elliott, H.H. Hobbs III leveling off with time. The same holds true for and J.R. Reddell. 2003. The North American obligate small caves. Results of intensive sampling by cave fauna : regional patterns. Biodiversity and Conservation 12:441-468. Schneider and Culver (2004) of 65 caves in a 25 Culver, D.C. and Sket, B. 2000. Hotspots of subterranean km2 area in West Virginia indicated that only biodiversity in caves and wells. Journal of Cave and about half of the obligate cave fauna had been Karst Studies 62:11-17. found. In spite of these pessimistic findings there Culver, D.C., M.C. Christman, I. Sereg, P. Trontelj, and B. is reason for hope. The actual pattern of diversity Sket. 2004. The location of terrestrial species-rich seems to remain stable as more data are added caves in a cave-rich area. Subterranean Biology (Culver et al. 2004a). 2:27-32. Culver, D.C., M.C. Christman, B. Sket, and P.Trontelj. 2004a. Sampling adequacy in an extreme As we look to the future, we need to look to environment : Species richness patterns in Slovenian develop multiple sampling schemes as was caves. Biodiversity and Conservation 13:1209-1229. demonstrated by both Pipan and Schneider in Deharveng, L. 2005. Diversity patterns in the tropics, pp. this symposium, to develop predictive models as 166-170. In D.C. Culver and W.B. White [eds.], has begun to by Stoch and Malard in this Encyclopedia of caves. Elsevier Academic Press, symposium, using the PASCALIS results, and to Burlington, Mass. embed the results in conservation planning as Gibert, J. and Deharveng, L. 2002. Subterranean put forward by Michel in this symposium. ecosystems : a truncated functional biodiversity. Bioscience 52:473-481. Lucic, I. 2003. Vjetrenica. Pogled u dusu Zemlje. Zagreb- Ravno. BIBLIOGRAPHY Schneider, K. and Culver, D.C. 2004. Estimating Christman, M.C. 2005. Mapping subterranean subterranean species richness using intensive biodiversity, pp. 355-360. In D.C. Culver and W.B. sampling and rarefaction curves in a high density White [eds.], Encyclopedia of Caves. Elsevier cave region in West Virginia. Journal of Cave and Academic Press, Burlington, Mass., U.S.A. Karst Studies 66:39-45. Christman, M.C. and Culver, D.C. 2001. The relationship between cave biodiversity and available habitat. Journal of Biogeography 28:367-380.

28 SWSB December 2004 GENERAL INTRODUCTION TO BIODIVERSITY

WHY AND HOW TO TAKE CARE OF SUBTERRANEAN AQUATIC MICROCRUSTACEANS ?

1D.L. DANIELOPOL and 2P. POSPISIL

1. Institute of Limnology, Austrian Academy of Sciences, Mondseestr. 9, 5310 Mondsee, Austria. ([email protected])

2. Reichmangasse 3/6,1160 Wien, Austria.

ABSTRACT Subterranean habitats harbour a diversified fauna of microcrustaceans like copepods, ostracods, syncarids. There are scientific, moral and practical (economic) arguments to protect groundwater organisms and/or to prevent the deterioration of their environment. Many subterranean crustaceans are the product of unusual long evolutionary histories, an important aspect of their scientific interest. Microcrustaceans, e.g. the copepods, have also practical value, when used as ecological sentinels. Moral arguments to protect such organisms and their environment should be integrated within the general principles of human ethics. Hotspot areas for stygobitic Crustacea should be viewed as “natural museums” with cultural value that merits to be protected. The example of the “Austrian National Park Danube Wetlands” is presented.

KEYWORDS: AQUATIC MICROCRUSTACEANS, ENVIRONMENTAL PROTECTION, SUBTER- RANEAN HABITATS.

1. INTRODUCTION exist in a wide variety of aquatic subterranean The groundwater domain has an enormous habitats. In Europe more than two third of the extension below the soil-surface. Note that 30 exclusively dwelling subterranean crustaceans percentages of the total earth’s freshwater (cf. Botosaneanu 1986) belong to groups with reserves are located in the subsurface, either in miniaturised body shape, like the Syncarida large spaces of consolidated rocks like in karstic Bathynellacea (fig. 1), the Copepoda and areas or in porous voids of non-consolidated Ostracoda. sediments. The biological exploration of this Sket (1999) compared the species richness of “subterranean ocean” existing below our feet hypogean crustacean fauna of Europe with the was intensified during last years. Gibert and epigean one. He noted for instance that from a Culver (2004) point out that about 7,700 aquatic total of 245 species of Harpacticoida and 145 stygobitic species were known by year 2000 species of Cyclopoida mentioned in the Illies’ world-wide and their number increases steadily (1978) compendium “Limnofauna Europaea” with new exploration of various geographic areas 70% of the former group and 60% of the latter

Figure 1 : Antrobathynella stammeri stammeri (Jakobi), body approx. 1.5mm length (del. E. Serban, in Coineau 1998) in Europe, Australia, South America, Asia. The one were stygobites. There are other recent European Community project PASCALIS microcrustacean groups like Bathynelacea and (cf. this volume) documents well this positive Thermosbaenacea which are represented trend. practically only by subterranean species. The We know now that microcrustaceans (animals importance of these minute crustaceans is not with body length in the range of millimetres size) only scientific (they offer us new insight into the

SWSB December 2004 29 GENERAL INTRODUCTION TO BIODIVERSITY diversity of life within one of the major segments groundwater specimens. The genus of the earth crust) but also have cultural and in a Microceratina Swanson contains marine benthic lesser extent practical (economic) value. ostracod species known from the Upper With the increasingly high anthropogenic stress Cretaceous (about 70 My years) to the Recent. In exerted on subterranean aquatic systems, like all cases specialists never could observe living the over-extraction of water for human needs, specimens, only valves were recorded (Namiotko chronic pollution, e.g. with various chemicals of et al. 2004). The unique species for which the both porous and karstic aquifers it becomes limbs were described is M. martensi Namiotko et more and more common to see the local al. a stygobiont recently discovered in an disappearance of subterranean dwelling animals anchialine cave in the Indian Ocean, on the and/or their specific habitats. This situation offers Christmas Island (Namiotko et al. 2004). arguments for the necessity to view the Within the EC project PASCALIS a new ostracod environmental protection of subterranean genus and species were discovered which has organisms as a human ethical problem. In the closely affinities with a Tertiary ostracod group following, we will offer examples for the why and from China (Z. Gído, J-P. Colin, M. Artheau, D.L. how to take care of the subterranean life Danielopol and P. Marmonier, in prep.). This new especially those of microcrustaceans, a group of stygobitic ostracod found in southern France can animals that we observed intensively during be considered a “living fossil”. Such examples many years. We will use not only already known represent an important progress in our arguments (cf. Sket 1999, Elliott 2004) but also knowledge on the subterranean organismic our own experience derived from groundwater diversity. It produces intellectual stimulation ecology projects in Austria. when thinking about the origin and the evolution of the lineages to which these species belong. It 2. WHY TO TAKE CARE OF produces also for the scientists who study them SUBTERRANEAN FAUNA ? a pleasant feeling, a kind of excitement once we better understand or arrive to reconstruct a tiny 2.1 Cultural (scientific) motivation aspect of the earth’s history. Therefore, caves or We use the term Culture in the every-day even porous aquifers containing a high number communication without difficulty. However when of rare species are considered a kind of “natural we intend to use it precisely, in our case related museums” for which a wide spectrum of people to scientific topics and/or the scientific display interest and admiration (Danielopol endeavour, we discover that it has different 1998). meanings. D’Andrade, (1984): mentions the following possible definitions: (1) accumulation 2.2 Practical (economic) aspects and transmission of knowledge, (2) producer of Groundwater dwelling microcrustaceans conceptual structures which help humans to especially copepods can be used as understand the reality of the world they inhabit, environmental descriptors. They are sometimes (3) the extension of social institutions and human called “sentinels” because they offer information traditions, an important factor for the acquisition about the state of groundwater ecosystems and preservation of knowledge. (Dole-Olivier et al. 1993). When successful, this Subterranean hydroscapes are replete with rare approach can offer an economic advantage to or unique stygobitic microcrustaceans. Within the the environmental monitoring programmes. Copepoda for instance one distinguishes nine Rouch pioneered this approach using the orders with about 200 families and several harpacticoid fauna of the Baget karstic system in thousands of species. The order Gellyelloida southern France. Using the taxonomic diversity Huys has only one family erected for a unique of harpacticoids he could document the origin genus Gelyella Rouch & Lescher-Moutoué; This and the dynamics of the water within the flooded latter contains only two stygobitic species, G. karst, with the possibility to differentiate between droguei Rouch & Lescher- Moutoué and G. the water coming from the main drains and those monardi Moeschler & Rouch localised in of lateral reserves (Rouch 1986). This approach restricted karstic areas in Europe and for which was successfully used in other projects too we know very little about their ecology and their (Gibert 1986). The presence of epigean phylogenetical affinities (Boxshall and Halsey microcrustaceans especially of cyclopoids in the 2003). In Austria a small cyclopoid Austrocyclops deep groundwater karstic systems are in many vindobonae Kiefer was described from a unique cases related to rapid infiltration of water loaded specimen caught in a well at Vienna (Kiefer with organic matter, hence the use of these 1964). Its morphology was so strange that it was crustaceans as ecological indicators (cf. Malard difficult to find the taxonomic position within the 2001). order. Only 30 years later Pospisil and Stoch Microcrustaceans can be used also for toxicity (1997) could solve the problem of this unique monitoring. Notenboom and Boessenkool (1992) stygobitic crustacean using additional experimented with the harpacticoid

30 SWSB December 2004 GENERAL INTRODUCTION TO BIODIVERSITY

Parastenocaris germanica for testing computer techniques (Fig. 2). The example toxicological effects of pesticides in groundwater. below is an expression of this kind of aesthetics. Mösslacher (2000) showed that the sensitivity of stygobitic microcrustaceans species were higher 3. HOW TO TAKE CARE OF THE SUBTER- than those of epigean related species when RANEAN FAUNA ? tested for the survival in high concentrations of Kalium Chloride. 3.1 The danger of local extinction for valuable stygobitic species Groundwater organisms especially crustaceans displayed during exhibitions organised by natural There are a multitude of causes which could history museums attract the interest of visitors, eliminate various stygobitic species which closely fit to the more or less stable environment of hence they play a practical role to the success of groundwater aquifers. Organic pollution along such cultural projects (P. Moeschler, pers. comm. the rivers eliminate many hyporheic to D.L.D. about the presentation of model assemblages, including stygobitic stygobitic crustaceans in an exhibition organised microcrustaceans. Danielopol (1976) showed by the Natural History Museum in Geneva). such an effect at Vienna below a sewage outlet 2.3 Ethic and aesthetic values where stygobitic ostracods were present only Pospisil and Danielopol (1990) using the with the rest of their carapaces. Chronic chemical philosophical views of W. Schröder consider that pollution of rivers achieve the same effect animals in their natural environment have to be (Danielopol et al. 2003). Local extinction of protected following the rules we apply to stygobitic fauna in some cases become a real humans. Respectively, we should not accept the tragedy because of impossibility to recover or systematic elimination of subterranean compensate for their disappearance. This is the organisms that since innumerable generations case of the minute isopod Microcerberus plesai live in equilibrium with their environment. Chappuis and Delamare Deboutteville (Fig. 3) Additionally, if one considers that he/she is an known only from a unique site, a sandy-gravel bank in the cave Vadu Crisului (Pestera lui Zichy) educated person who is able to value potential near Cluj, in Romania (cf. Botosaneanu 1986). life on earth than it is important to answer a The interest of this species is related to the fact question advanced by P. Dasgupta (2001) in that the Microcerberoidea represents an isopod “Human well-being and the natural environment”: group of marine origin spread around the can we live comfortably with the idea of the Mediterranean as well as in tropical and exhaustion of natural resources ? subtropical areas world-wide (Hobbs 2000). M. Groundwater and the organisms which inhabit it plesai is a freshwater cavernicolous species exerted since many centuries on the human mind found only in Transylvania far away from the a strong fascination combining feelings of typical biogeographic distribution that we know pleasure with fear ones. One should remember nowadays. It is apparently a relict of a Tertiary how Antoine de Saint-Exupery (1939) warm fauna. During many years this species characterised the subterranean water: “the could always be collected in its unique interstial world’s greatest treasure, and also the most habitat from the Vadu Crisului cave (Fig. 3). delicate, so pure, deep within the earth...”. During the last 10-15 years the cave became a During our long-year research in the Danube tourist attraction and the sandy-gravel bank was wetland Lobau, at Vienna, we were fascinated by completely deteriorated by the passage of the beauty of crustaceans when observing them visitors. Repeated investigations of this site in their natural environment with videometry showed the disappearance of this unique techniques. It stimulated one of us (P.P.) to endemic species (S. Iepure, pers. comm. to recreate the mysterious atmosphere of their life- D.L.D.). As the stygobitic fauna in Romania is habits within tiny porous space with virtual very poor, respectively there are few species of

Figure 2 : Virtual electronic image of life in a groundwater porous habitat (Artistic representation realised by P. Pospisil)

SWSB December 2004 31 GENERAL INTRODUCTION TO BIODIVERSITY

Figure 3 : Microcerberus plesai (left panel, del. C. Plesa) from the cave Vadu Crisului where Dr. C. Plesa (right panel, from Danielopol 1965) sampled it during many years

Isopoda, mainly Aselloidea. The loss of the restricted wetland area within the national park unique Microcerberoidea representative known and was already declared during the 70ies an in Romania has a special significance. It is UNESCO biosphere reserve. Long-term equivalent to the loss of a highly precious object investigations of the groundwater fauna within an of art in a museum. area of about 0.8 Km2 in the Lobau allowed to discover 35 stygobitic species, from which 2/3rd 3.2 Environmental protection strategies are represented by microcrustaceans, viz. One of the first measure we have to do is to Copepoda and Ostracoda (Danielopol and proceed to map the subterranean biodiversity at Pospisil 2001). There are endemic species various spatial scales. Efforts for the known only from this area like the ostracod development of effective algorithms are Mixtacandona spandli Rogulj & Danielopol, proposed in Culver et al. (2001). This was also others which are very rare but occur also in other one of the aims of the EC project PASCALIS. parts of the park and/or occasionally also in Juberthie (2000) reviewed protection strategies Lower Austria, like Austriocyclops vindobonae for the caves and their environment, mainly Kiefer,Acanthocy-clops gmeineri Pospisil, under the European Community legislation. Diacyclops felix Pospisil and Stoch (Fig.4). Generally it is recommended to avoid pollution of Finally, there are stygobitic microcrustaceans groundwater by keeping clean the surrounding which even if they have a wider geographic environment. Further it is advisable to distribution in Europe, they occur also in low concentrate the protection effort on sites with a numbers in completely isolated places. Such a special interest, e.g. those displaying a high case are those of the cyclopoid Graeteriella species richness of stygobites. Culver and Sket unisetigera Graeter (Fig. 4) and of the ostracod (2000) provided a rule of thumb for such a Cryptocandona kieferi Klie (Namiotko et al. measure, respectively they defined hotspot sites 2005). The comparative study of individuals of those areas like cave systems which have at this latter species from France, Germany, Austria least 20 stygobitic species. Worldwide only 20 and Romania showed that the local populations karstic systems and two sites in porous aquifers found within the National Park, belong to a new can be termed as hotspot areas (cf. Culver and subspecies (Namiotko et al. 2005). Especially Sket 2000, Danielopol and Pospisil 2001). One of interesting are the species of cyclopoids which this sites is the Lobau wetland at Vienna, in the differ not only through their morphology (Fig. 4) National Park. Its interest for the protection of but also through their ecological and micro- groundwater fauna is briefly reviewed below. geographical distribution (Pospisil and 3.3 The “National Park Danube Danielopol 2000). Wetlands”, a natural reserve for Because many habitats where earlier we could groundwater fauna sample cyclopoids and other groundwater An area of about 9,300 ha of wetlands were microcrustaceans disappeared or became declared national park in Lower Austria at the altered by various types of pollution, this is the end of 1996. The park is located east of Vienna, case especially along large rivers and their down to the boarder with Slovakia, within the floodplains like the Danube and/or the Rhine Danube flood plain, hence also the name rivers, the possibility to keep protected rare or “Danube Flood Plain National Park” (cf. unique groundwater microcrustaceans in a Danielopol and Pospisil 2001, 2002). It protects national park is of extreme importance. It offers one of largest natural floodplain areas in Central the chance to the future generation of Europe and it has a wide variety of terrestrial and groundwater specialists to continue studies on aquatic habitats which support a high number of the ecology of these poorly known species. species both plants and animals. The Lobau is a Presenting the data to lay-people who visit the

32 SWSB December 2004 GENERAL INTRODUCTION TO BIODIVERSITY

Figure 4 : The surprising rich diversity of Cyclopoid fauna in the “National Park Danube Wetlands” park allows to improve their interest and There are more and more educational acceptance for the necessity to keep alive the programmes organised by various institutions sustainability of groundwater systems, including especially within university departments dealing their life forms. Hence, the example of the with groundwater ecology and the management “National Park Danube Wetlands”, as a natural and/or protection of groundwater resources. As reserve for groundwater fauna should be an example one one should mention an followed when possible in other countries too. advanced study course on groundwater ecology organised by the Institute of Limnology at the Austrian Academy of Sciences, within the 4. NEEDS FOR MORE INFORMATION AND framework of the European Commission’s EDUCATION Environment and Climate Research programme For the sustainability of groundwater resources (Griebler et al. 2001). Additionally, we have to including the groundwater fauna, like the become also activists for the protection of the microcrustaceans and their habitats, one needs groundwater environment making clear to more education and information. We have to politicians and/or decision makers the necessity make understandable to a wide spectrum of to integrate within their policies also ecological humans the ecological research we are carrying views related to groundwater ecosystems on the groundwater environment and the interest (Danielopol et al. 2003, 2005). for groundwater organisms. The publication of Finally, as environmentalists it appeared us that the first textbook on groundwater ecology edited our duty is in a long-term to contribute to a switch by Gibert et al. (1994) is in retrospect in the human perception for groundwater toward appreciated as a success if one considers that more hydrosolidarity, toward better protection not during its ten year existence more than 1,500 only of the subsurface water reserves but also of copies were distributed al other the world. A these invaluable small organisms. similar success seems to enjoy the first textbook on groundwater ecology written for German- CONCLUSION speaking readers by Griebler and Mösslacher (2003). Within one year more than 500 copies of Paraphrasing ideas expressed by the Austrian historian of art E.H. Gombrich (cf. “Ideals and this book were distributed. We need to intensify idols”, 1979) we will close this essay with the other forms of communication too. For instance following statements: one of us (P.P.) produced for lay-people an interactive play distributed on CD-ROM and on 1. The knowledge of the subterranean diversity which one finds the multiple aspects existing in of microcrustaceans yields pleasure and the underground of the city of Vienna from the life enrichment. in groundwater to historical relics and the artistic 2. What makes subsurface systems so attractive production related to these aspects (Pospisil is the awareness that there are things which 2003). Several encyclopaedic books deal with are not only simply rare but unique. topics related to caves and their environment. 3. We have faith in the power of a new creative Such compendia have a high educational impact generation of groundwater ecologists which if well produced. The most recent one edited by espoused cultural values and will successful Culver and White (2004) represents a superb invest time and energy for taking care of the achievement of this goal. groundwater fauna and their environment.

SWSB December 2004 33 GENERAL INTRODUCTION TO BIODIVERSITY

4. As ecologists we have to communicate also Danielopol, D.L. and Pospsil, P. 2002. Taxonomic the idea of the development of extended diversity of Crustacea Cyclopoida in the Austrian hydrosolidarity for groundwater sustainability. “Danube Floodplain” National Park. Vie Milieu 52:67- 75. This should encompass the broad spectrum of ecological problems, from which we selected Danielopol, D.L., C.Griebler, A.Gunatilaka, and J.Notenboom 2003. Present state and future here a tiny fraction, those of the groundwater prospects for groundwater ecosystems. organisms. Environmental Conservation 30:104-130. Danielopol, D.L., J. Gibert, C. Griebler, A. Gunatilaka, H.J. Hahn, G. Messana, J. Notenboom, and B. Sket. 2005. ACKNOWLEDGEMENTS Incorporating ecological perspectives in European This essay benefited from long-term cooperation groundwater management policy. Environmental Conservation 31:185-189. with Janine Gibert and her research group (Lyon) as well as with Raymond Rouch (formerly in Dasgupta, P. 2001. Human well-being and the natural environment. Oxford Univ. Press, Oxford. Moulis). The support received by one of us (D.L.D.) from the EC-Project PASCALIS is much Dole-Olivier, M.-J., M. Creuzé des Châtelliers, and P.Marmonier. 1993. Repeated gradients in appreciated. The Austrian Fund for Scientific subterranean landscape Ð Example of the stygofauna Research (FWF) is acknowledged for the long- in the alluvial floodplain of the Rhône river (France). term support of the investigations done by D.L.D. Arch. Hydrobiol. 127:451-471. and P.P. in the Lobau at Vienna. We are also Elliott, W.R. 2004. Protecting caves and cave life, pp. 458- much indebted to our colleagues who provided 467. In D.C. Culver and W.B. White [eds.], data and information: Malvina Artheau Encyclopedia of caves. Elsevier Academic Press, San (Toulouse), Sanda Iepure and Cornel Plesa Diego. (Cluj), Ionel Tabacaru (Bucharest), (Cluj) Zolt Gibert, J. 1986. Ecologie d’un système karstique Gidó (Debrecen), Christian Griebler (München), jurassien : Hydrogéologie, dérive animale, transits de Pascal Moeschler and Ilinca Juvara-Bals matières, dynamique de la population de Nyphargus (Crustacé, Amphipode). Mém. Spéléol. 13:1-380. (Geneva), David J. Horne (London), Jean-Paul Gibert, J. and Culver, D.C. 2004. Diversity patterns in Colin (Bordeaux), Nicole Coineau (Banyuls Europe, pp. 196-201. In D.C. Culver and W.B. White s/Mer). [eds.], Encyclopedia of caves. Elsevier Academic Press, San Diego. Gibert, J., D.L. Danielopol, and Stanford, J.A. 1994. BIBLIOGRAPHY Groundwater Ecology. Academic Press, San Diego. Botosaneanu, L. 1986. Stygofauna mundi. E.J. Brill, Gombrich, E.H. 1979. Ideals and idols. Phaidon, Oxford. Leiden. Griebler, C. and Mösslacher, F. 2003. Grundwasser Boxshall, G.A. and Halsey, S.A. 2003. An introduction to Ökologie. Facultas, Wien. copepod diversity. The Ray Society, London. Griebler, C., D.L. Danielopol, J.Gibert, H.P.,Nachtnebel, Coineau, N. 1998. Syncarida, pp. 863-876. In C. and J. Notenboom. 2001. Groundwater ecology Ð A Juberthie and V. Decu [eds.], Encyclopaedia tool for management of water resources. Office Publ. biospeologica. Soc. de Biospeologie, Moulis. Of EC, Luxembourg. Culver, D.C. and Sket, B. 2000. Hotspots of subterranean Hobbs III, H.H. 2000. Crustacea, pp.95-107. In H. biodiversity in caves and well. J. Cave and Karst Res. Wilkens, D.C. Culver and W.F. Humphreys [eds.], 62:11-17. Subterranean ecosystems. Ecosystems of the World, Culver, D.C. and White, W.B. 2004. Encyclopedia of 30. Elsevier, Amsterdam. caves. Elsevier Academic Press, San Diego. Illies, J. 1978. Limnofauna Europaea. G. Fischer, Culver, D.C., L. Deharveng, J. Gibert, and I. Sasowsky. Stuttgart. 2001. Mapping subterranean biodiversity. Karst Juberthie, C. 2000. Conservation of subterranean Waters Inst., Special Publ. 6., Charles Town, WVa. habitats and species, pp. 691-700. In H. Wilkens, D.C. D’Andrade. 1984. Cultural meaning systems, pp. 88-119. Culver and W.F. Humphreys [eds.], Subterranean In R.A. Shweder and R.A. LeVine [eds.], Culture ecosystems. Ecosystems of the World, 30. Elsevier, Theory, Cambridge Univ. Press, Cambridge. Amsterdam. Danielopol, D.L. 1965. Nouvelles données sur les Kiefer, F. 1964. Zur Kenntnis der subterranene Ostracodes d’eau douce de Roumanie. Ann. Limnol. Copepoden (Crustacea) Österreichs. Ann. Nathist. 1:443-463. Mus, Wien 67:477-485. Danielopol, D.L. 1976. The Distribution of the Fauna in the Malard, F. 2001. Groundwater contamination and Interstitial Habitats of Riverine Sediments of the ecological monitoring in a Mediterranean karst Danube and the Piesting (Austria). Int. J. Speleol. ecosystem in Southern France, pp 183-194. In C. 8:23- 51. Griebler, D.L. Danielopol, J. Gibert, H.P. Nachtnebel Danielopol, D.L. 1998. Conservation and protection of the and J. Notenboom [eds.], Groundwater ecology Ð A biota of karst: assimilation of scientific ideas through tool for management of water resources. Office Publ. artistic perception. J. Cave and Karst Res. 60:67. Of EC, Luxembourg. Danielopol, D.L. and Pospisil, P. 2001. Hidden biodiversity Mösslacher, F. 2000. Sensitivity of groundwater and in the groundwater of the Danube Flood Plain surface water crustaceans to chemical pollutants and National Park (Austria). Biodiversity and Conservation hypoxia : implications for pollution management. Arch. 10:1711-1721. Hydrobiol.149:51-66.

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Namiotko, T., K. Wouters, D.L. Danielopol, and Pospisil, P. and Danielopol, D.L. 2000. Diversity of W.F.Humphreys. 2004. On the origin and evolution of groundwater dwelling Cyclopoida (Crustacea, a new anchialine stygobitic Microceratina species Copepoda) in a Danube wetland in Austria. Vie Milieu (Crustacea, Ostracoda) from Christmas Island (Indian 50:137-150. Ocean). J. Micropal. 23:49-59. Pospisil, P. and Stoch, F. 1997. Rediscovery and Namiotko, T., P. Marmonier, and Danielopol, D.L. 2005. redescription of Austriocyclops vindobonae Kiefer, Cryptocandona kieferi (Crustacea, Ostracoda): 1964 (Copepoda, Cyclopoida) with remarks on the redescription, morphological variability, geographic subfamily Eucyclopinae Kiefer. Crustaceana 70:901- distribution. Vie Milieu 55:(in press). 910. Notenboom, J. and Boessenkol, J.-J. 1992. Acute toxicity Rouch, R. 1986. Sur l’écologie des eaux souterraines testing with the groundwater copepod Parastenocaris dans le karst. Stygologia 2:352-398. germanica (Crustacea), pp. 311-317. In J.A. Stanford Saint-Exupéry, A. de 1939. Terre des hommes. Gallimard and J.J. Simons [eds.], Proc. 1st, Int. Conf. on (Oeuvres, nrf, bibliothèque de la Pléïade, 1959), Groundwater Ecology, AWRA, Bethesda, MD. Paris. Pospisil, P. 2003. Unter der Stadt, Kultur und Sket, B. 1999. High biodiversity in hypogean waters and Naturgeschichte des Wiener Untergrundes, CD-ROM. its endangerment Ð the situation in Slovenia, the Brüder Hollinek, Purkersdorf. Dinaric Karst and Europe. Crustaceana 72:767-779. Pospisil, P. and Danielopol, D.L. 1990. Vorschläge für den Schutz der Grundwasserfauna im geplanten Nationalpark “Donauauen” östlich von Wien, Österreich. Stygologia 5:75- 85.

SWSB December 2004 35

PASCALIS GENERAL PRESENTATION

SWSB December 2004 37

PASCALIS GENERAL PRESENTATION

GROUNDWATER BIODIVERSITY

Protocols for the ASsessment and Conservation of Aquatic Life In the Subsurface (PASCALIS): overview and main results

1J. GIBERT, 2A. BRANCELJ, 3A. CAMACHO, 1F. CASTELLARINI, 4C. DE BROYER, 6L. DEHARVENG, 1M.-J. DOLE-OLIVIER, 1C. DOUADY, 6D.M.P. GALASSI, 1F. MALARD, 4P. MARTIN, 5G. MICHEL, 7B. SKET, 6F. STOCH, 7P. TRONTELJ and 3A.G. VALDECASAS.

1. University Claude Bernard of Lyon 1, UMR/CNRS 5023, Laboratoire d’Ecologie des Hydrosystèmes Fluviaux, Equipe d’Hydrobiologie et Ecologie Souterraines, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France. ([email protected]), ([email protected]), ([email protected]), ([email protected]), ([email protected]).

2. National Institute of Biology, Vecnaù pot 111, 1000 Ljubljana, Slovenia. ([email protected]). 3. Museo Nacional de Ciencias Naturales, Dpto. de Biodiversidad y Biología Evolutiva, C/ José Gutiérrez Abascal 2, 28006, Madrid, Spain. ([email protected]), ([email protected]).

4. Royal Belgian Institute of Natural Sciences, 29 rue Vautier, 1000 Brussels, Belgium. (Claude.De [email protected]), ([email protected]).

5. Commission Wallonne d’Etude et de Protection des Sites Souterrains (CWEPSS), 21 avenue Auguste Rodin, 1050 Brussels, Belgium. ([email protected]). 6. National Museum of Natural History, Origine, Structure et évolution de la biodiversité, Département Systématique et Evolution, 45 rue Buffon, 75005 Paris, France. ([email protected]). 7. University of L’Aquila, Dipartimento di Scienze Ambientali, Via Vetoio, Coppito, 67100 L’Aquila, Italy. ([email protected]), ([email protected]).

7. University of Ljubljana, Department of Biology, Biotechnical Faculty, Vecnaù pot 111, P.P. 2995, 1001 Ljubljana, Slovenia. ([email protected]), ([email protected]).

ABSTRACT The PASCALIS project (EVK2-CT-2001-00121) (2002-2004) was the first comprehensive proposal that specifically addresses the groundwater biodiversity issue at the European scale and represents also an important backbone for research development in groundwater. The main objective was to establish a rigorous and detailed protocol for assessing groundwater biodiversity and to develop operational tools for its conservation. More precisely, a solid unmatched piece of work was provided by the PASCALIS database that reflects our present- day knowledge on stygobiotic biodiversity distribution in six European countries. The cryptic diversity has been explored by using molecular approach on reference taxa and revealed to be a useful tool in the implementation and extension of our knowledge of groundwater biodiversity. Furthermore, a sampling protocol has been assessed, allowing reliable results on species distribution within selected hierarchical units. Results show that species distribution differs from one region to another, and that species similarity between regions is very low due to the high level of endemicity observed. In addition, biodiversity indicators among stygobionts resulted to be a very useful tool to build up a network of reference sites at European scale with reduced costs and sampling effort. Significant improvements have been introduced in the groundwater biodiversity conservation strategy by proposing new methodologies relying on sound scientific results. Finally, the ways to effectively implement a coherent, sustainable and scientific solid conservation strategy have been explored in the framework of EU policies related to biodiversity and sustainable management of groundwater resources.

SWSB December 2004 39 PASCALIS GENERAL PRESENTATION

1. INTRODUCTION 2. MAIN RESULTS Groundwater plays an important role for the 2.1 Insight on the diversity of stygobio- sustainability of many earth ecosystems and a tic species and mapping in southern crucial role in human life and socio-economic Europe development (Danielopol et al. 2003, Gibert et al. Knowledge of the spatial dimension of in press). Because there are continuously biodiversity is the basic need to any conservation evolving economic, social and cultural pressures strategy. Consequently, mapping biodiversity is on groundwater resources, it is becoming urgent the fundamental starting point to understand to provide an updated picture of groundwater spatial patterns of biodiversity (Gaston and biodiversity and to propose a global strategy for Spicer 2004). Little has been actually achieved groundwater management and conservation. In on groundwater biodiversity at the European this framework, the main objective of the scale (Danielopol et al. 2000; Gibert and Culver proposal was to establish a strict and 2004). Only a few maps for some given comprehensive protocol to estimate groundwater taxonomic groups are presently available at biodiversity and to elaborate operational tools for national levels in sparse European countries its conservation. Knowledge of groundwater (Juberthie and Decu 1994, 1998, Stoch 2001) biodiversity stands behind that of all other but no global representation of national aquatic ecosystems. This puts a major constraint groundwater biodiversity exists. The PASCALIS on a successful implementation of its project attempted to characterize distribution conservation. In this context, the specific patterns of biodiversity at operational scales for scientific objectives of the PASCALIS project biogeography and conservation. were (Gibert 2001): A flexible biodiversity storage tool has been ¥ to establish the general patterns of groundwater developed, for the six European countries biodiversity at the European scale in creating a (Belgium, France, Italy, Portugal, Spain, and large database, firstly to store all information Slovenia), as a basis for a mapping strategy to be available on subterranean taxa distribution, used for groundwater habitats. Based on the secondly by incorporating also the new records Endemism project (Deharveng 2001), a central obtained following the PASCALIS sampling database structure has been produced campaigns, to map the groundwater (Appendix 1), comprising 5 tables (Reference, biodiversity at the European scale; Distribution, Species, Genera, Co-ordinates) of 10 to 25 fields each, and 22 procedures written in ¥ to improve biodiversity evaluation by exploring the 4D language. The reference table contains genetic diversity of taxa in order to estimate the 1,756 records. The distribution table has 17,367 cryptic biodiversity within selected groups; records of which 14,780 concern stygobionts. The Species table contains 2,346 species or ¥ to set up a stratified sampling scheme in order infraspecific taxa, of which 1,264 are stygobionts to obtain an optimum field sampling strategy (Table 1). The Genera table has 662 genera, and an unbiased estimate of biodiversity at the most of which are not stygobionts. The Co- regional scale; ordinate table has 3,971 records. The dataset ¥ to analyse the patterns of biodiversity and covers most (likely over 90%) of the published identify potential biodiversity indicators at the records of stygobiotic taxa in the 6 selected regional scale; countries. Different distributional maps were produced: ¥ to develop strategies for conservation of species richness (Fig.1), hotspots of species groundwater biodiversity in the European socio- richness, strict endemism, endemicity scores, economic context by identifying some regions range-size rarity scores, taxonomic isolation of special biological interest, the spatial scale of scores. A map of the residuals of the regression relevance for conserving biodiversity within between species richness and site numbers was these regions, in order to propose a series of produced and used to assess the level of appropriate measures for maintaining their knowledge of groundwater stygofauna in biodiversity. southern Europe.

40 SWSB December 2004 PASCALIS GENERAL PRESENTATION

Table 1: Taxonomic distribution of main stygobiotic taxa in Europe (Belgium, Spain, France, Italy, Portugal and Slovenia)

Figure 1 : Species richness in the PASCALIS consortium countries

SWSB December 2004 41 PASCALIS GENERAL PRESENTATION

Present biodiversity distribution patterns are still subterranean environment. Not less important is largely impacted by uneven sampling effort the role played by vicariance at different spatial among regions and countries. However, the and temporal scales, which may allow the analysis of these patterns already reveals splitting up of an ancestral population and its hotspots and coldspots of highly isolated taxa, diversification in different, sometimes using LISA models (Anselin 1995). The highest morphologically indistinguishable, cryptic values are concentrated in a longitudinal area species. It is in this general framework, that in the extending from Slovenia to the Mediterranean PASCALIS project, four representative taxa have French coast following the Italian Pre-Alps, while been selected for molecular analyses (Trontelj et areas of very low taxonomically isolated al. 2004). assemblages are found in the northern part of ¥ The amphipod Niphargus is the most widely France. Most of Paleothyrrhenian areas distributed and diverse in Europe, comprising (Sardinia, Corsica and coastal areas of Italy) and about 300 species/subspecies. The preliminary part of the Iberian Peninsula show stygobiotic phylogenetic hypotheses based on 28S rDNA assemblages, poor in terms of number of species sequences seem to argue for a paraphyly of the but conversely rich in endemics. genus. The overall structure of the tree is rather The present-day knowledge of the groundwater bush-like, with many short internal branches. It biodiversity distribution patterns has been deeply is possible that few clades are oversplit. On the enlarged in all the PASCALIS countries, even if other hand, a single species may be composed further field work is still needed to obtain more of distinct, well separated lineages. For information in some poorly sampled regions, example, Niphargus virei that shows a weak such as the Iberian Peninsula, southern Italy and morphological differentiation among different north-western France. populations (Ginet 1960), presents an unpredicted molecular tripartite structure based on two independent genes (28S and COI). This 2.2 Hidden diversity in groundwater suggests that the history of this species might In the PASCALIS project, biodiversity have been eventful, involving phenomena of assessment essentially relies upon dispersal and vicariance at various stages of its morphologically based identification and evolutionary history (Lefébure et al. in press). traditional taxonomic criteria. However, previous ¥ The isopod Asellus aquaticus is an expansive studies showed that groundwater and Western Palaearctic species, which has subterranean habitats as a whole harbour successfully dispersed across Europe (Henry remarkably high numbers of sibling or cryptic and Magniez 1983). It has a prominent genetic species (Stoch 1995). These species cannot be population structure at small geographic identified by standard morphological analyses, distances (<100 km). A. aquaticus can be identification keys and only sporadically considered a target “species-group” for morphological microcharacters may be used to clarifying the processes leading to high lineage distinguish closely related species. The diversity in restricted areas in the groundwater awareness about this high cryptic diversity is network (habitat and microhabitat scales). based on a number of studies on genetic Discrepancies between the morphological- differentiation and population structure (e.g., based identification versus the genetic results Avise and Selander 1972, Laing et al. 1976, placed in a phylogeographic scenario were Sbordoni et al. 1979, Caccone et al. 1986, Sket found. and Arntzen 1994, Gentile and Sbordoni 1998, Wiens et al. 2003, Lefébure et al. submitted). ¥ Hirudinea belonging to the genera Dina and Most of these studies used allozymes as genetic Trocheta are frequently found in various markers. These markers are suitable for groundwater habitats, mainly in karstic aquifers. detecting genetic population structure, gene flow, Their systematics is controversial and the or boundaries to gene flow, but are less reliable specific and generic attributions open to in terms of phylogenetic accuracy, evolutionary question. Phylogenetic analyses of ITS and rates, and discerning historical events from COI gene sequences reveal in some cases a ongoing processes. Many evolutionary models strong discrepancy between morphologically- try to explain the high degree of hidden diversity based systematics and phylogenetic in subterranean systems (reviewed in Wilkens et relationships among taxa. The cryptic diversity al. 2001) and the overall diversity of groundwater observed in both genera could be the fauna (Stoch 1995). The commonly recognized consequence of convergent or parallel models to explain the high level of differentiation evolutionary changes in non-sister groundwater in the hypogean environment argue a taxa. combination of radiation and multiple invasions. ¥ Proteus anguinus, the blind cave salamander Homoplasy by means of convergence and is distributed along the Dinaric karst of the parallelism seems to be widespread in the western Balkan Peninsula in several

42 SWSB December 2004 PASCALIS GENERAL PRESENTATION

hydrographically isolated areas (Sket 1997). in particular habitats, such as those by Deep splits between some groups of Scarsbrook and Halliday (2002) and Boulton et populations were found in the phylogenetic al. (2003, 2004) in the porous aquifer, no analysis, indicating the presence of cryptic sampling strategies have been proposed nor species. tested up to now, to solve such a basic question. The molecular approach of selected taxa A rigorous sampling protocol has been applied in revealed that cryptic diversity may represent a 6 selected regions (Walloon karst in Belgium, significant part of groundwater biodiversity at the Meridional Jura and Roussillon in France, European scale. The genetic variation can be Cantabria in Spain, Lessinia in Italy and Krim explained by the multiple cave invasions model massif in Slovenia) (Malard et al. 2002). It is or can be the consequence of convergent or based on a stratified sampling strategy, following parallel evolutionary changes in non-sister a hierarchical scheme, including: region, basin, groundwater taxa. Convergence may stem from aquifer type (karst and alluvial), zone in each the conservative nature of groundwater in aquifer (unsaturated zone and saturated zone in respect to the more changeable epigean karst, hyporheic zone and phreatic zone in freshwater, and may be the result of adaptation unconsolidated sediments) (Fig.2). In each to the major environmental describers (Galassi region, 192 sampling sites were selected to 2001). Convergence toward similar assess the groundwater biodiversity. morphological traits may be the consequence of The main results indicate that the high sampling the same response to similar groundwater effort performed during the PASCALIS surveys environments. For this reason, genetically was not satisfactory in some areas (192 sites per distinct phylogenetic lineages of stygobiotic taxa, region) due to the high amount of rare species that have ceased to exchange genes, may and strict endemics. The stratification escape clear taxonomic distinction. These results considering karstic areas and unconsolidated give a good idea about the extent of diversity and sediments as different sampling units should be endemism that may be overlooked when relying used in every sampling design, being statistically upon traditional taxonomy. From data of the significant in discriminating different species present study, the cryptic biodiversity should be assemblages (Stoch et al. 2004). considered as a priority tool for assessing overall In order to explore the relationship between the groundwater biodiversity. structure of obligate-groundwater assemblages and environmental gradients, multivariate 2.3 Sampling protocols and biodiversity statistical analyses were applied to the assessment environmental and species data sets. Trends of Due to the inaccessibility of the subterranean biodiversity were highlighted by the statistical environment, planning an efficient groundwater analyses, suggesting the dominant role of sampling strategy is a difficult task (Gibert et al. geography, paleogeography, habitat structure 1994; Griebler et al. 2001). It may not be possible and water chemistry. to distribute sampling sites wherever necessary Additive partitioning of groundwater species because access is only possible through a diversity across nested spatial scales - aquifers, limited number of outputs (springs, resurgences) basins, and regions - using species-richness or windows (caves, wells). Apart a few attempts data, was performed; the between-region

Figure 2 : Hierarchical sampling scheme.

SWSB December 2004 43 PASCALIS GENERAL PRESENTATION component made by far the highest contribution relictuality) were used to assign a cumulative to the stygobiotic species richness, e.g. conservation value to each of the species community composition varied most importantly included in the database. Limitations to this over broader spatial scales. The results clearly procedure are linked to the high level of indicate that the most effective way to preserve endemism of stygobiotic species (more than 83 stygobiotic diversity in southern Europe is to % of them can be defined as strict endemics) protect multiple aquifers within different regions which prevents the selection of a threshold useful and with different environmental features. for discriminating priority species (Stoch et al. 2004).

2.4 Identification of potential biodiversity indicators and conservation 2.5 Elaboration of an operational indicators conservation strategy Species richness is a simple measure of In the European policies, most of the attention biodiversity and a widely used criterion for regarding groundwater has understandably biodiversity assessment and conservation. concerned its use as a safe source of drinking However, data on species richness are reliable water of primary importance for a large majority only if derived from exhaustive inventories; of the EU inhabitants. However, it has become unfortunately, inventories over large spatial increasingly obvious that groundwater should not scales are expensive and time-consuming. only be viewed as a drinking water reservoir but Predictive models of species richness have been also as critical aquatic ecosystems that must be used as an alternative to conduct extensive field taken into account in the European legislation. studies. These models rely on environmental The Habitat Directive (European Commission variables, named “environmental surrogates” of 2003) has so far mostly neglected groundwater species richness (Araujo et al. 2001) or the habitats and species (Juberthie 1995, IUCN identification of a representative species group, 2004) and the Water Framework Directive able to reflect the overall species richness of an (European Union 2001) did not take groundwater entire biota (Pearson, 1994; Pimm et al. 2001). biodiversity into consideration and did not These species are named “biodiversity expressly perceive the “good ecological status of indicators” (Mac Nally and Fleishman 2004). ground water” as for surface waters. Sets of environmental parameters, species and As GW ecosystems were never considered when higher level taxa were selected as indicators of defining priority sites and areas for conservation, groundwater biodiversity; multiple regression there is an evident missing point in excluding models and statistically sound information criteria stygofauna in the establishment of natural were used to select the indicators and assess reserves. In order to fill this gap, different their predictive power of species richness. methods to select the most efficient network of Unfortunately, stygobiotic species vary from one region to another due to the high degree of GW biodiversity reserves, priority species and endemism in groundwater taxa. However, at both habitats were tested (De Broyer et al. 2004). European and regional scales, the higher taxa like Gastropoda, Harpacticoida and Amphipoda 2.5.1 The designation of “Groundwater Biological appear to be significantly correlated with total Reserves” species richness (Stoch 2004). Different methods were tested to design a Thus, biodiversity indicators are a very useful network of aquifers (as biological groundwater tool to detect spatial patterns of species richness reserves) containing a significant proportion of at large scale, with reduced costs and sampling European groundwater biodiversity (including effort. However, the model should be applied with species of high conservation value) with low-cost caution, as the results cannot be extended procedures. In order to maximize the outside the study area, given the large amount of conservation effectiveness of the network fine-scale autocorrelation. composed of a fixed amount of grid cells, we A standard method to build conservation indices, tested and compared the conservation effects of based on the information stored in the different cell assemblages, taking into account PASCALIS database and on the grid cells used different parameters, such as species richness, to map the distribution of species over Europe, number of endemics, relictuality and has been developed. The degree of endemicity, conservation value of species contained in the range-size rarity, habitat selection and taxonomic selected cells. The construction of the isolation (included phylogenetic relictuality) were conservation network, based on complementarity incorporated after normalization in a test (between grid cells) offers a more effective conservation index. Mean values of endemicity, approach instead of selection methods based on rarity, and taxonomic isolation (weighted for hotspots of biodiversity and endemism.

44 SWSB December 2004 PASCALIS GENERAL PRESENTATION

A simple method of risk assessment was methods. An innovative approach has been proposed which combined criteria of aquifer proposed as follows: the reciprocal discrimination vulnerability and human activities. Vulnerability of correspondence analysis and hierarchical and human pressure indexes were combined in cluster analysis were used to distinguish order to distinguish between 3 distinct types of between distinct groundwater habitats based on biological groundwater reserves: healthy, under biological data collected (as an example) from 55 threat, highly anthropised. Despite several aquifers in France. Results show that a spatial methodological difficulties linked to the paucity of hierarchical approach provides a consistent information available for some aquifers, the theoretical framework for identifying groundwater method proves to be a valuable tool for habitats comprising dissimilar set of species. prioritizing conservation efforts among aquifers Alpha and beta diversity can be used to prioritize of biological interest. conservation efforts among groundwater habitats (Ferreira et al. submitted). The PASCALIS project provides a rare 2.5.2 Identification of priority species for groundwater biodiversity conservation opportunity to propose to European stakeholders a specific Action Plan for the conservation of a Umbrella species have been selected (Andelman unique and neglected part of the European and Fagan 2000) for groundwater biodiversity aquatic biodiversity (De Broyer et al. 2004). This conservation, according to the basic principle Action Plan recommends to integrate the that they are species with ecological groundwater biodiversity protection concern in all requirements encompassing those of the relevant European policies, legislation and remaining members of a community. By instruments dealing with conservation and recommending some conservation measures for sustainable management of natural resources - aquifers hosting a limited number of umbrella in particular water resources (WFD and GW species, the conservation effects do benefit to all Daughter Directive), nature conservation (Habitat close-by species. Different ways of selecting Directive, Natura 2000), - as well as in relevant umbrella species have been tested aiming at horizontal legislation and instruments concerning defining a limited number of species covering in particular environment, land use, and priority biodiversity hotspots and associated to a agriculture. high proportion of priority species. A list of 30 taxa that should receive a legal status in order to Relying on the scientific outputs of the contribute to a better protection of GW PASCALIS project and the expertise of the biodiversity has been proposed. PASCALIS contributing teams, the following general recommendations are made for the implementation of an Action Plan for the 2.5.3 Definition of a list of priority habitats for conservation of groundwater (GW) biodiversity in groundwater conservation Europe: The selection of priority groundwater sites was mostly based on traditional expert selection

REC. 1: TO INTEGRATE THE GW BIODIVERSITY AND ECOSYSTEMS CONSERVATION CONCERN IN ALL RELEVANT EUROPEAN POLICIES

¥ Considering that GW is not only an exploitable resource but also an ecosystem as all other water bodies; ¥ Considering the important scientific, heritage and eco-functional value of the European GW biodiversity and ecosystems; ¥ Noting the present and potential threats they face or may face; ¥ Considering that GW biodiversity, habitats and ecosystems have been largely overlooked in the current policies implementing the EU biodiversity strategy; It is recommended:

¥ to integrate the GW biodiversity and ecosystems concerns in all relevant European policies (legislation and instruments) dealing with biodiversity and sustainable management of natural resources, in particular water resources (Water Framework Directive and GW Directive) and biodiversity conservation (Habitat Directive, Natura 2000), as well as in horizontal environmental legislation and instruments regarding in particular land use, agriculture, environmental impact assessment, and environmental monitoring.

SWSB December 2004 45 PASCALIS GENERAL PRESENTATION

REC. 2: TO COMPLETE THE HABITAT DIRECTIVE POLICY

¥ Considering the diversity of GW habitats of special conservation interest;

¥ Noting the high conservation value of most GW species;

¥ Considering the lack or very limited inclusion of priority GW habitats and species in the Habitat Directive Annexes 1 and 2;

It is recommended:

¥ To formally include the priority GW species and habitats in the respective Annexes of the Habitat Directive at the next revision opportunity.

Rec. 2.1 : To establish a list of priority GW species for conservation

¥ Considering that the establishment of a list of priority stygobiotic species at European level as well as at national and regional levels should follow a sound selection protocol taking into account the conservation value of the stygobiotic species (in particular on the basis of criteria of endemism, rarity, and phylogenetic relictuality) and their endangered status;

¥ Noting that, given the high conservation value of most stygobiotic species and the (potential) endangered status of many of them, a comprehensive list of European GW priority species for conservation does not appear operational for complete inclusion in Annex 2 and that alternative limited selection of representative GW species has to be defined;

It is recommended:

¥ To rely on a surrogate species approach (such as umbrella species and biodiversity indicators) as an operational conservation strategy to establish a limited list of representative priority species whose habitat conservation allow covering a significant part of the European GW biodiversity.

¥ To explore new ways of defining species value and of integrating it into site selection process, for a better estimate of site biological value.

Rec. 2.2: To establish a list of priority GW habitats

¥ Considering that the existing habitat categories relevant for GW in Annex 1 of the Habitat Directive do not acknowledge the diversity of groundwater habitats;

¥ Considering the complexity of the procedures involved to modify the Habitat Directive and to add some species and habitats to its Annexes;

¥ Considering that the subdivision of the subterranean hydrosphere in various habitats must be based on sound hydrogeological units taking into account the physico-chemical conditions, and, on the other hand, on the biogeographic spatial and temporal context;

It is recommended:

¥ To rely on a pragmatic strategy to include priority GW habitats in Annex 1 as it stands by broadening the interpretation of the existing habitat categories relevant for GW and completing the “Interpretation Manual of European Union Habitats” accordingly, before the formal recognition by a revision of the Directive.

¥ To define the GW (macro)habitats in a regional framework on the basis of the aquifer unit or assemblages of aquifers having identical hydrogeological characteristics and similar palaeoclimatic or palaeogeographic histories, and sharing a suite of characteristic species.

Rec. 2.3 : To develop a European network of priority sites (GW natural reserves) for GWB conservation

¥ Considering that the appropriate spatial scale to take into account for defining GW natural reserves on ecological and hydrological basis is the aquifer;

¥ Considering that the most effective way to preserve stygobiotic biodiversity in Europe is to protect multiple aquifers presenting high biodiversity complementarity and different

46 SWSB December 2004 PASCALIS GENERAL PRESENTATION

environmental features within different regions and, there, by maintaining regionally distinctive species-rich assemblages;

¥ Considering that the European biogeographic regions as referred to by the Habitat Directive do not apply to the GW fauna for which specific biogeographic stygoregions have to be defined;

It is recommended:

¥ To establish a network of priority aquifers as natural reserves for GWB conservation at European scale but also at national and regional scale, by relying on a sound selection protocol taking into account the intrinsic scientific and heritage value of their biodiversity and by properly assessing (on the basis of the aquifer(s) vulnerability and intensity of human pressure) the risk they are facing in order to define priorities in terms of conservation.

¥ To attempt to include the maximum number of GW taxa in the priority aquifers at the minimal cost in socio-economical terms by using complementarity approaches for site selection.

¥ To regularly update, under control of recognized experts, the priority sites network based on a complementarity approach, in order to integrate progress of knowledge on GWB.

¥ To define biogeographic stygoregions to allow assessing the representativeness of GW habitats and species and to integrate this information into priority species and habitat selection protocols.

¥ To develop biodiversity modelling in order to compensate the lack of information on biodiversity distribution in many areas that handicaps the habitat selection protocol.

REC. 3: TO INTRODUCE BIODIVERSITY CONCERN AND GOOD ECOLOGICAL STATUS OF GW IN THE WATER FRAMEWORK DIRECTIVE AND THE GW DIRECTIVE POLICY

¥ Noting that the WFD and the proposed GW Directive consider GW only as a resource and not as an ecosystem, contrary to the surface water systems for which the ecological dimension is duly recognized;

¥ Recalling that, like surface water systems, GW systems undergo to hydrological, chemical and biological processes which collectively define their ecological status;

¥ Noting that in the proposed GW Directive the GW protection measures are mainly based on the maintenance of a quantitative equilibrium (balance between GW abstraction and recharge), and on chemical quality (based on threshold concentrations of some GW chemical constituents);

¥ Considering that the maintenance of a high diversity of GW micro-organisms is crucial to the self-purification processes of GW systems, and that beside micro-organisms, stygobiotic invertebrate and vertebrates can be selectively used as bioindicators of the structural, functional and healthy state of GW ecosystems;

It is recommended

¥ To elaborate the instruments (including the required expertise and knowledge background) necessary to accurately define the “good ecological status” of GW in the regional context and take it into account in GW management.

¥ To develop efficient monitoring strategies for GWB and ecosystems (selective biological and environmental indicators, reference sites) in order to ensure a follow-up of the GW populations in selected priority aquifers and of the functional characteristics of the GW ecosystems.

¥ To consider developing potential monitoring synergies by relying - where relevant - to the reference sites network required by the WFD for monitoring GW resources.

¥ To improve by appropriate investigations the indicator potential of the GW biodiversity in the characterization of the connectivity with surface waters (and therefore the GW vulnerability) and of the qualitative state of GW.

SWSB December 2004 47 PASCALIS GENERAL PRESENTATION

REC. 4 : TO DEVELOP SCIENTIFIC KNOWLEDGE ON GW BIODIVERSITY AND ECOSYSTEMS

¥ Considering the large gaps in the scientific knowledge of the nature and distribution of GW biodiversity, its habitats and communities as well as its eco-functional roles;

¥ Noting that these limitations hamper the development of comprehensive conservation initiatives in favour of GW ecosystems and biodiversity in the framework of European, national or regional policies, as well as the development of a sound ecological approach of groundwater management;

It is recommended:

¥ To make efficiently available the scientific information on European GW biodiversity and distribution by developing and maintaining a centralized European GWB database.

¥ To support the establishment of a European platform for GW biodiversity combining the expertise of ecologists, biologists and hydrologists to manage the European GWB database and advise on the implementation of relevant European environmental policies and of a GWB conservation strategy along the lines recommended in this Action Plan.

¥ Among the priority research topics to focus on, it is suggested:

¥ To plan large scale sampling surveys to complete the knowledge of the nature and distribution of European groundwater biodiversity at regional and aquifer scale.

¥ To plan, for reference purposes, in depth all-taxa surveys of selected representative aquifers of different characteristics, which include the assessment of cryptic biodiversity by molecular methods.

¥ To stimulate the taxonomic study of GW fauna and the development of efficient taxonomic identification tools usable by non-taxonomists.

¥ To define the European biogeographic stygoregions.

¥ To develop population dynamics studies of reference GW species in reference sites in order to accurately assess their endangered status.

¥ To improve the understanding of the structure and function of GW ecosystems in relation to various environmental conditions and anthropogenic influences.

¥ To assess the role of micro-, meio- and macro-organisms in GW self- purification and biological remediation, as well as their potential as GW quality indicators.

¥ To strengthen the ecological basis of groundwater management by conducting studies on the sensitivity of GW species to pollution and physical disturbance; the effects of pollutants on biodegradation processes; the ecological recovery of groundwater ecosystem after remediation.

¥ To improve the assessment of human impact and land-use practices on structural and functional aspects of phreatic, hyporheic, and karstic ecosystems; and the impact of water mining on the functioning of GW ecosystems.

REC. 5 : TO RAISE PUBLIC AWARENESS

¥ Recalling that the recent Malahide Conference identified the promotion of broader public awareness, understanding and support for the conservation and sustainable use of biodiversity as a critical overarching issue, and recognized that there is an urgent need to mobilise public opinion in support of biodiversity;

¥ Considering the general lack of knowledge by decision-makers as well as by the general public of the existence, interests and roles of the GW ecosystems;

¥ Recognising the high heritage, scientific and ecological values of their biodiversity;

¥ Stressing that an effective conservation policy requires the acceptance, support, and understanding of the socio-economic actors;

48 SWSB December 2004 PASCALIS GENERAL PRESENTATION

It is recommended:

¥ To undertake systematic campaigns of public awareness targeted towards stakeholders and policy makers in charge of biodiversity conservation, management of natural resources and the environment at European, national, regional and local scale, the water basin agencies and water supply companies, the education- (at all pertinent levels), scientific- and media communities as privileged information relays, the civil society organizations, the general public.

¥ In these campaigns, to present groundwater fauna and its conservation in the more global framework of groundwater protection, as an essential resource for human welfare.

¥ To develop educational supports dealing not only with GW biodiversity but with more general aspects of groundwater (geological features of aquifers, water cycle, origin of drinking water, etc.).

¥ To attempt to designate some GW flagship species which can help in arising people interest on GW ecosystems and biodiversity.

3. CONCLUSIONS structure, elevation gradient, historical factors, etc, have been strongly recommended. PASCALIS is the first research Programme on groundwater ecology and conservation, funded ¥ Cryptic diversity appears very high in under the “Global Change, Climate and subterranean environments. Cryptic diversity biodiversity” within the Energy, Environment and due to cryptic speciation and between non- Sustainable Development framework monophyletic lineages may significantly Programme of the European Commission. The increase alpha and beta diversity at multiple study of groundwater biodiversity in 6 countries different spatial scales. (Belgium, France, Italy, Portugal, Slovenia and ¥ Considering the biodiversity partitioning, Spain) and in 6 selected regions more between-region richness made by far the specifically investigated (Walloon karst, highest contribution (80%) to total richness in Meridional Jura, Roussillon, Cantabria, Lessinia Europe. The contribution of alpha and beta and Krim massif) allows to draw the following diversity to regional species richness probably conclusions: changes as a function of spatial scale. ¥ Synthetic information on groundwater ¥ Environmental parameters and higher taxa biodiversity becomes available through richness can be used as good indicators and PASCALIS and constitutes an important predictors of species richness at regional scale. backbone for further scientific research. It may However, indicators of biodiversity at the also serve as a strong basis for communication species level were different in each region; for about groundwater biodiversity in Europe this reason, higher taxa were used to predict towards policy makers and other disciplines. species richness at the European scale. ¥ The subterranean environment is unique if ¥ PASCALIS helps to identify regions and compared to surface environments because it aquifers harbouring high biodiversity. The includes a large number of endemic and rare construction of the conservation network, species. Over 83% of the stygobiotic species in based on complementarity test offers a more the PASCALIS countries can be classified as effective approach than selection methods strict endemics and over 69% are rare. based on hotspots of biodiversity and ¥ At regional scale, for the first time, a rigorous endemism. Reserves should be preferentially stratified sampling scheme based on a defined using aquifers as reference units. hierarchical approach by examining biodiversity Moreover, PASCALIS can help to answer the in 6 regions has been applied. General question to what extent Natura 2000 covers guidelines have been proposed to improve also aquifers of high subterranean biodiversity. sampling protocols at regional scale, Overlay analysis may help to clarify where demonstrating the efficiency of a stratification surface and subsurface biodiversity protection sampling based on aquifer type (karstic vs. go together and where the most important gaps porous aquifers). Nevertheless, due to the low are traceable. similarity between regions and the high level of Follow-up research should focus on further endemism at sub-regional scale, a stronger spatial analysis and identification of main sampling effort and a finer stratification, which pressures for groundwater biodiversity, linking takes also into account basins, habitat structural and functional aspects, and modelling

SWSB December 2004 49 PASCALIS GENERAL PRESENTATION the most important pressure-response biodiversity. Cartographie de la biodiversité souterraine. relationships. Better link is needed between Culver D.C., Deharveng L., Gibert J. & I. Sasowsky [Eds.], Karst Waters Institute, Special Publication 6:12- hydrological, biogeochemical and biodiversity 15. information of aquifers placed in a land-use De Broyer, C, G. Michel, F. Malard, P. Martin, and J. Gibert. scenario, water management and agriculture 2004. PASCALIS D10 Deliverable for Workpackage 9: policies. Action Plan for conservation of groundwater biodiversity. European Project Protocols for the Assessment and Conservation of Aquatic Life in the Subsurface (PASCALIS No EVK-CT-2001-00121), (Unpublished ACKNOWLEDGEMENTS report). This work was financially supported by the European Commission. 2003. Interpretation manual of European Programme PASCALIS (Protocols for European Union Habitats. Eur 25 DG ENVIRONMENT. Nature and biodiversity. the ASsessment and Conservation of Aquatic http://europa.eu.int/comm/environment/nature/nature_co Life In the Subsurface) (no EVK2-CT-2001- nservation/eu_enlargement/2004/pdf/habitats_im_en.pdf 00121) under the EU 5th Framework European Union. 2001. Common strategy on the Programme : Global Change, Climate and implementation of the Water Framework Directive. Biodiversity. Annick PAPIN (Lyon1 University) is www.europa.eu.int/comm/environment/water/water- framework. greatly acknowledged for her invaluable Ferreira, D., F. Malard, M.-J. Dole-Olivier, and J. Gibert. assistance and co-ordination along the entire Hierarchical patterns of obligate groundwater biodiversity duration of the project. Authors wish also to thank in France (in press). all the PASCALIS participants for their Galassi, D.M.P. 2001. Groundwater copepods: diversity involvement and cooperation and all the external patterns over ecological and evolutionary scales. experts, who in different ways made this project Hydrobiologia 453/454:227-253. feasible, and ameliorated the working plans with Gaston, K.J. and Spicer, J.I. 2004. Mapping biodiversity. In criticism, suggestions, collection and Gaston K.J. & J.I. Spicer [eds.], Biodiversity: an Introduction (second edition). Blackwell Science, identification of groundwater fauna. Oxford,191p. Gentile, G. and Sbordoni, V. 1998. Indirect methods to estimate gene flow in cave and surface populations of BIBLIOGRAPHY Androniscus dentiger (Isopoda: Oniscidea). Evolution 52:432Ð442. Andelman, S. and Fagan, W. 2000. Umbrellas and flagships: Gibert, J. (Coord.) 2001. Protocols for the ASsessment and Efficient conservation surrogates or expensive mistakes? Conservation of Aquatic Life In the Subsurface Proc. Natl. Acad. Sci., USA 97:5954-5959. (PASCALIS): a European project. 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Juberthie, C. 1995. Underground habitats and their Sket, B. and Arntzen, J.W. 1994. A black, non-troglomorphic protection. Council of Europe, Strasbourg, Nature and amphibian from the karst of Slovenia: Proteus anguinus Environment paper: 72. parkelj n. ssp. (Urodela: Proteidae). Contrib. Zool. 64:33- Laing, C., R.G. Carmody, and S.B. Peck. 1976. How common 53. are sibling species in cave-inhabiting invertebrates? Am. Stoch, F. 1995. The ecological and historical determinants of Nat. 110:184-189. crustacean diversity in groundwaters, or : why are there Lefébure, T., C.J. Douady, M. Gouy, P. Trontelj, J. Briolay, and so many species ? Mém. Biospéol. 23:139-160. J. Gibert. (submitted): Molecular phylogeography of Stoch, F. 2001. Mapping subterranean biodiversity: structure Niphargus virei (Crustacea: Amphipoda) reveals of the database, mapping software (CKMAP), and a evolutionary patterns in groundwater. Molecular Ecology. report of status for Italy. In Mapping subterranean Mac Nally, R. and Fleishman, E. 2004. A successful predictive biodiversity. Cartographie de la biodiversité souterraine. model of species richness based on indicator species. Culver D.C., Deharveng L., Gibert J. & I. Sasowsky [Eds], Conservation Biology 18(3):646Ð654. Karst Waters Institute, Special Publication 66:29-36. Malard, F., M.-J. Dole-Olivier, J. Mathieu, and F. Stoch. 2002. Stoch, F., F. Malard, F. Castellarini, M.-J. Dole-Olivier, and J. PASCALIS D4 Deliverable for Workpackage 4: Sampling Gibert. 2004. PASCALIS D8 Deliverable for manual for the assessment of regional groundwater Workpackage 7: Statistical analyses and identification of biodiversity. European Project Protocols for the indicators. European Project Protocols for the Assessment and Conservation of Aquatic Life in the Assessment and Conservation of Aquatic Life in the Subsurface (PASCALIS; No EVK-CT-2001-00121), Subsurface (PASCALIS; No EVK-CT-2001-00121) (Unpublished report). (unpublished report). Pearson, D.L. 1994. Selecting indicator taxa for the Stoch, F. 2004. PASCALIS D9 Deliverable for Workpackage quantitative assessment of biodiversity. Philosophical 8: Maps of measured and predicted patterns of Transactions of the Royal Society of London, Series B biodiversity and hotspots of species richness and 345:75Ð79. endemism. European Project Protocols for the Assessment and Conservation of Aquatic Life in the Pimm, S.L., M. Ayres, A. Balmford, G. Branch, K. Brandon, T. Subsurface (PASCALIS; No EVK-CT-2001-00121). Brooks, R. Bustamente, R. Costanza, R. Cowling, L.M. (unpublished report). Curran, A. Dobson, S. Farber, G.A.B. da Fonseca, C. Gascon, R. Kitching, J. McNeely, T. Lovejoy, R.A. Trontelj, P., C.,Douady, C. Fiser, S. Goricki, T. Lefébure, and Mittermeier, N. Myers, J.A. Patz, B. Raffle, D. Rapport, P. R. Verovnik. 2004. PASCALIS D7 Deliverable for Ravens, C. Roberts, J.P. Rodriguez, A.B. Rylands, C. Workpackage WP6B: Cryptic diversity of selected Tucker, C. Safina, C. Samper, M.L.J. Stiassny, J. groundwater taxa. European Project Protocols for the Supriatna, D.H. Wall and D. Wilcove. 2001. Can we defy Assessment and Conservation of Aquatic Life in the nature’s end? Science 293:2207-2208. Subsurface (PASCALIS; No EVK-CT-2001-00121). (unpublished report). Sbordoni, V., M. Cobolli Sbordoni, and De Matthaeis, E. 1979. Divergenza genetica tra popolazioni e specie Wiens, J.J., P.T. Chippindale, and D.M. Hillis. 2003. When are ipogee ed epigee di Niphargus (Crustacea, Amphipoda). phylogenetic analyses misled by convergence? A case Lav. Soc. Ital. Biogeogr. N.S. 6:1-23. study in Texas cave salamanders. Systematic Biology 52:501-514. Scarsbrook, M.R. and Halliday, J. 2002. Detecting patterns in hyporheic community structure: does sampling method Wilkens, H., D.C. Culver, and W.F. Humphreys [Eds]. 2001. alter the story. New Zeal. J. Mar. Fresh. 36:443-453. Subterranean ecosystems. Ecosystems of the World 30. Elsevier, Amsterdam. Sket, B. 1997. Distribution of Proteus (Amphibia: Urodela: Proteidae) and its possible explanation. J. Biogeogr. 24:263-280.

SWSB December 2004 51 PASCALIS GENERAL PRESENTATION

Appendix 1: The PASCALIS central database Structure

52 SWSB December 2004 SESSIONS

SESSIONS

Extended abstracts

SWSB December 2004 53

SESSIONS

SESSIONS REPORT

SESSION 1 Chairman : Dan L. Danielopol Reporter : David C. Culver

AUSTRALIAN STYGOFAUNA AND WESTERN AUSTRALIAN PROJECT (PILBARA REGION)

Western Australia is an emerging hotspot of diverse. Unlike most other areas, it appears that aquatic subterranean biodiversity and presents aridity, especially in the Miocene, has been the both recurring themes and new challenges to our driving force for the colonization of the understanding of the subterranean biota. A series subterranean realm. In common with other karst of five presentations was devoted to the results areas, subterranean dispersal is highly limited, of an ongoing regional-scale survey of the based on the distribution of species and Western Australia stygofauna. Most of the molecular analysis of cryptic variation in sampling was performed in wells and boreholes. apparently widespread species. Hydrologic Ultimately, there may be 200 known stygobionts basins act as barriers with little dispersal beyond from this region. In common with other karst areas, subterranean species in Australia are a scale of tens of kilometres. Dispersal is further “disharmonious” in the sense that, compared to restricted by important differences in water surface-dwelling species, some groups are over- chemistry, chiefly resulting from the extreme represented and some are under-represented. In aridity. Even in this sparsely inhabited region, the the case of the Western Australian stygofauna, human ecological footprint is growing large as a the Dytiscidae (Coleoptera) are particularly result of extensive mining operations.

SESSION 2 Chairman : Ana I.Camacho Reporter : Louis Deharveng

PHYLOGENY, PHYLOGEOGRAPHY AND TAXONOMY

Aquatic subterranean biodiversity remains poorly information on stygobiotic biodiversity is terribly known. More than in any other habitat in Europe, poor. new taxa are described every year, and the It has been repeatedly said that good taxonomy contribution of the PASCALIS project is already is the basis of any further analysis of biodiversity. outstanding in this respect. Diversity patterns of Traditional morphological approaches remain the this subterranean aquatic fauna are nevertheless only operational tool for analyzing biodiversity at slowly emerging. They have been shaped by a wide taxonomic regional scale or for large number of factors of which three are of groups. But the most promising remain molecular overwhelming importance : the ability of lineages tools. Molecular approaches can disclose more to speciate and to adapt, the opportunity to of diversity (i.e. cryptic diversity). They can also colonize underground habitats given to them by help understanding evolutionary history of geological and palaeoclimatic changes, and the colonization or population fragmentations, by possibility that populations have persisted dating population divergences. This justifies their underground during the last glacial periods. rapidly increasing use by the scientific Like for stygobiont-rich versus stygobiont-poor community. These methodological aspects groups, we are primarily interested in hotspots, deserve certainly more in-depth consideration in and we often forget how much can be drawn from taxonomy, phylogeny and phylogeography, as considering coldspots. Hotspots can only be progress is extremely rapid in these fields. defined in contrast to colder spots, where

SWSB December 2004 55 SESSIONS

SESSIONS REPORT

SESSION 3 Chairman : Diana M.P. Galassi Reporter : Anton Brancelj

BIODIVERSITY PATTERNS

New highlights on the major patterns affecting patterns are : a) geology, fossil river basins and biodiversity problems in ground water and covering vegetation, b) fragmentation of habitats related ecotones have been presented. and hierarchical structure i.e. karstic vs. Some groups (Syncarida and Isopoda) reflect a interstitial and saturated / un-saturated or restricted regional distribution, while others hyporheic / phreatic systems, c) migration (Copepoda) have wide distribution pattern. Beta between systems and the phenomenon of diversity is considered as the main contributor to speciation increase the number of single-site the overall biodiversity. With increasing number endemics. of different habitats, areas with increasing size Main problems faced for successful estimation should become increasingly dissimilar. The best and evaluation of biodiversity patterns are in : a) approach to study groundwater biodiversity un-sufficient and/or non-standardized sampling should be based on aquifer scale. methods and b) relation between sampling effort The main factors, influencing biodiversity and number of taxa recorded = cost / benefit ratio

SESSION 4 Chairman : Patrick Martin Reporter : Boris Sket

CONSERVATION AND MANAGEMENT

Different aspects of the groundwater fauna troglobionts prefer deeper layers, the troglophiles ecological distribution as well as interrelations the upper ones. between the human impacts and the Possible criteria for the groundwater sites groundwater habitat or/and fauna have been evaluation have been discussed. Besides the discussed. Copepod faunas have been generally number of troglobiotic species, some other used as model. The influence of man faunistic and ecological data can be used. Some

(eutrophization -C, NO3 contents) can modify non-biological (historic, cultural etc) data may be subterranean assemblages. The accessibility of of particular importance when the conservation organic matter appeared to be the primary factor measures are difficult to be accepted by the determining the biodiversity patterns ; society.

56 SWSB December 2004 SESSION 1 : AUSTRALIAN STYGOFAUNA AND WA PROJECT (PILBARA)

EMERGING KNOWLEDGE OF DIVERSITY, DISTRIBUTION AND ORIGINS OF SOME AUSTRALIAN STYGOFAUNA

1W.F. HUMPHREYS, 2C.H.S. WATTS and 3J.H. BRADBURY.

1. Western Australian Museum, Locked Bag 49, Welshpool DC, WA 6986, Australia. ([email protected]). 2. South Australian Museum, North Terrace Adelaide, SA 5000, Australia. ([email protected]). 3. University of Adelaide, SA 5005, Australia. ([email protected]).

KEYWORDS : ARID ZONE, CALCRETE, REFUGIA, STYGOFAUNA, WESTERN SHIELD.

The last decade has transformed understanding two-thirds of the land mass lies within the arid of the diversity, distribution and affinities of zone, and it is in the arid zone where much of the Australian stygofauna, especially in the more arid stygal diversity occurs. Examination of the parts. Western Australia, in particular, is proportional distribution of eight higher emerging as having an exceptional diversity of taxonomic groups within Australia compared with often unexpected lineages. These studies are the world average proportional distribution of still in their early stages, in terms of both those taxa (from Scarsbrook et al. 2003) reveals geographic coverage, and taxonomic and that the Australian fauna is probably deficient in systematic knowledge. However, published isopods and molluscs (9% and 18% world species plus information from specialists in average) but that beetles are greatly over certain taxa places the total recognised number represented (3,100% world average) amongst of stygal taxa recognised in Australia at 515 the Australian stygofauna. The remaining groups species (Dytiscidae, 80; Ostracoda, 29; considered (Acari, Amphipoda, Syncarida, Copepoda, 109; Isopoda, 40; Amphipoda, 65; Ostracoda, Copepoda) have between 61% and Syncarida, 32; Acarina, 150; Gastropoda, 10). 159% world average). Further research may alter This is known to be a considerable this picture, as has occurred recently with the underestimate as in numerous higher taxa many Dytiscidae. undescribed species, not included in figure 1 are Australian amphipods are diverse and known to occur from molecular (R. Leys, pers. predominently stygal, containing representatives comm.) or morphological evidence, especially of ancient freshwater lineages, of which the from the Copepoda (T. Karanovic, pers. comm. crangonyctoids are the most ubiquitous 2004), Ostacoda (I. Karanovic, pers. comm. (Bradbury 1999), and those of relatively recent 2004), Bathynellacea (J-L Cho, pers. comm. marine origin (Hadziidae, Melitidae). The 2004) and Amphipoda. Stygobiont amphipod distributions match that of land areas supposedly species yet to be described are known from not submerged—as also does the distribution of Queensland (Chillagoe sp.), New South Wales other freshwater lineages, the Phreatoicidea and (Neoniphargidae), South Australia (north- Tainisopidea (Humphreys 2004)—and western, east-central, and south-eastern - submerged, respectively, in the ocean since the Melitidae and Paramelitidae), Tasmania Cretaceous. Crangonyctoid amphipoda have (Paramelitidae), and Western Australia hitherto, with few exceptions, been regarded as (Ashburton River, Pilbara region, Yilgarn region - confined by the cool fresh waters of the south Bogidiellidae, Melitidae and Paramelitidae).The east and south west of the continent and state of flux of this estimate can be gained when Tasmania (Barnard and Williams 1995). Calcrete it is realised that 54% of this number (including deposits have developed extensively in the both described and recognised species) have Pilbara and Yilgarn Cratons, tectonically stable been formally described in the last five years. regions which have been emergent above the This value represents 15% of the 3,410 sea since the Precambrian. Where these ground described species in 13 large higher taxa of waters have remained fresh, especially in the freshwater stygobites that were enumerated by extensive palaeochannels (Humphreys 1999, Scarsbrook et al. (2003) from the world synopsis 2001), they have been found to contain diverse (Botosaneanu 1986). While these world figures crangonyctoid species as well as bogidiellids; are now dated, Australia, which comprises ~7% where more saline, to contain hadziid species. It of the Earth’s land area, would appear to greatly is already clear the knowledge of the diversity of exceed the world average of stygal species per hypogean amphipods in Australia confirms area. This is unexpected in a continent where Holsinger’s (1993) view that southern Australia is

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Figure 1 : Venn diagrams showing the number of species (large numerals) and percentage overlap between adjacent regions of Western Australia for stygal Copepoda (upper) and and Ostracoda (lower) species (left) and genera (right). Regions (denoted by number or writing on map): 1, Kimberley; 2, Pilbara; 3,Yilgarn; 4, Leeuwin-Naturaliste, 5, Cape Range/Barrow I.. Date from Ivana Karanovic (pers. comm. 2004) and Tomislav Karanovic (pers. comm. 2004). a region of significant diversity for this group. The ostracod fauna of Western Australian However, description of Paramelitidae from the continental subterranean waters almost Pilbara region of Western Australia and work exclusively comprises representatives of the underway on species from the Yilgarn Craton subfamily Candonidae (Podocopida). A total of contradicts findings elsewhere which predict 29 species has been recorded, 28 of which have “only in subtropical areas where freshwater been described recently (Karanovic & Marmonier stygobionts have been derived from marine 2002, 2003; I. Karanovic 2003a, b; 2004, 2005, in ancestors by stranding ..... have significant press). They comprise 11 genera in three distinct stygobiont amphipod faunas evolved in limnic tribes: Humphreyscandonini Karanovic, in press; waters” (Holsinger 1993). Candonopsini Karanovic, 2004 and a third yet to Stygal copepods have been recorded from about be formalized (Karanovic in prep). All these tribes 500 sites, 112 species of representing four are very old lineages within the subfamily orders, 15 families and subfamilies, and 44 Candoninae. The first tribe has eight genera in genera. These are distributed amongst regions Australia (19 species) found only in the Pilbara as follows: in Western Australia, Margaret River, region of Western Australia, and three genera 12; Yilgarn, 31 (T. Karanovic 2004); Pilbara, 43; elsewhere (subterranean waters of India and Kimberley, 8; Queensland: Pioneer Valley, 15; Europe). The Candonopsini contains four New South Wales, 3 stygoxenes, mainly from genera, two in Australia and the other two in caves. Generally, most forms of ancient Central and South America, and the third tribe freshwater origin show Gondwana or Eastern will contains two genera. In addition to the Gondwana connections on global scale, while Candoninae, two species of the family those of marine origin have Tethyan connections. Darwinulidae occcur in the interstitial waters and Of the 112 species, 84% have been described wells (Martens & Rossetti, 2002). Stygal species from aquifers in Western Australia (T. Karanovic in anchialine waters are known from the pers. comm. 2004). Cytheruridae (Namiotko et al. 2004) and

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Thaumatocypridae (Myodocopida) (Danielopol et 2004); Oniscidea, Haloniscus: south of arid zone, al., 2000) (I. Karanovic, pers. comm. 2004). Taiti & Humphreys 2001, S. Taiti et al. There is an exceptional number of Dytiscidae unpublished; Dysticidae: widely outside arid species in the stygofauna of the Australian arid zone, CHS Watts, data), troglobitic and stygobitic zone, with so far ~80 species known, an order of species are confined to the arid zone; there is magnitude greater than elsewhere in the world. It evidence, also, of a southward progression includes stygal species in both the tribes Bidissini through time of the isolation of dytiscids in (Limbodessus and Bidessodes) and Hydroporini groundwater calcrete aquifers (Leys et al. 2003), (Nirripirti which may include Kintingka) within the mapping the southward spread of aridity. subfamily Hydroporinae (Watts and Humphreys The Western Shield contains a number of ancient 2004), as well as the first stygal diving beetle not (crangonyctoid amphipods, bathynellacean belonging to the subfamily Hydroporinae, namely syncarids, phreatoicidean isopoda) as well as in the Copelatinae (Copelatus: Balke et al. 2004). Tertiary stygal lineages (Dytiscidae, Oniscidea) The Hyphydrinae and family Noteridae, which but the northern region (Pilbara) has a have stygal representatives in other parts of the stygofauna distinct from the southern part world, are not known as stygofauna in Australia. (Yilgarn) despite having supposedly been a They occur in the northern and central Yilgarn of single land mass since at least the Palaeozoic. Western Australia, and in the Ngalia Basin in the This distinction applies to those presumably Northern Territory. Up to four species are found in ancient lineages above, as well as to Tertiary the same calcrete (13 cases of sympatric sister invaders such as the dytiscids and oniscideans. species pairs; Leys et al. 2003; Leys et al. pers. In Figure 1 the diversity (to 2004) of stygal comm.) and are well separated by size. They Copepoda and Ostracoda species in Australia exhibit a small range of stygomorphies (reduced and the general lack of overlap, in the ostracods or absent eyes, wings, colour and increased a total lack of overlap between adjacent regions mobility of prothorax) which stands in contrast to even at the generic level. It is more remarkable the large range of unusual morphologies in that this distinction holds even in the contiguous comparison to epigean close relatives that seem and long emergent parts of the Western Shield, a to be a result of relaxation of selection pressures. distinction that remains enigmatic. The chains of salt lakes (playas) in arid Australia are the base level of groundwater flow along the course of palaeodrainages constrained by ACKNOWLEDGEMENTS Permian palaeovalleys. Groundwater calcretes We appreciate information and/or discussion with form near salt lakes and contain diverse stygal Drs J-L. Cho, M.S. Harvey, W. Ponder, S. Taiti,. communities (Dytiscidae, Amphipoda, Oniscidea, G.D.F. Wilson, and especially, for the detailed Gastropoda, Ostracoda, Copepoda, summaries of Drs Tom Karanovic and Ivana Bathynellacea: Humphreys 2001) endemic to a Karanovic. given deposit. The fauna is best known from the Oniscidea (Scyphacidae: Haloniscus: Taiti and Humphreys 2001; S. Taiti et al. unpublished) and BIBLIOGRAPHY Dytiscidae (80 species of Copelatinae, Hydroporinae: Bidessini, Hydroporini: e.g. Watts Balke, M., C.H.S. Watts, S.J.B. Cooper, W.F. Humphreys, and Humphreys 2004) for which both and A.P. Vogler. 2004. A highly modified stygobitic diving beetle of the genus Copelatus (Coleoptera, morphological and molecular data are available Dytiscidae): Taxonomy and cladistic analysis based (Cooper et al. 2002; Leys et al. 2003) and, on mtDNA sequences. Systematic Entomology contrary to the initial hypothesis about their origin 29:59-67. (Humphreys 2001), their phylogenies do not map Barnard, J.L. and Williams, W.D. 1995. The taxonomy of the palaeodrainage channels. Conversely, freshwater Amphipoda (Crustacea) from Australian ancient lineages, such as crangonyctoids, also freshwaters: Part 2. Records of the Australian represented by discrete populations in the Museum 47:161-201. calcretes (R. Leys, pers. comm.), do tend to map Botosaneanu, L. 1986. Stygofauna Mundi: A faunistic, distributional and ecological synthesis of the world the palaeodrainages in the Pilbara (T. Finston fauna inhabiting subterranean waters (including the and M. Johnson pers. comm. 2004). marine interstitial). Brill and Backhuys, Leiden. There is increasing evidence from arid Australia Bradbury, J. H. 1999. The systematics and distribution of that elements of both terrestrial and aquatic Australian freshwater amphipods: a review. In F.R. subterranean fauna are refugial in origin Schram and J.C. von Vaupel Klein [eds.], (troglobitic Schizomida, stygal Oniscidea, stygal Crustaceans and the Biodiversity Crisis. Proceedings of the Fourth International Crustacean Congress, Dytiscidae). In each case, despite wide Amsterdam, The Netherlands, July 20-24, 1998. Brill: distribution of close relatives outside the arid Leiden. zone (Schizomida: northern monsoonal Cooper, S.J.B., S. Hinze, R. Leys, C.H.S. Watts, and W.F. rainforest, Harvey 1992, 2000, pers. comm. Humphreys. 2002. Islands under the desert:

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molecular systematics and evolutionary origins of Karanovic, I. 2005. Towards a revision of Candoninae stygobitic water beetles (Coleoptera: Dytiscidae) from (Crustacea, Ostracoda): Australian representatives of central Western Australia. Invertebrate Systematics the subfamily, with descriptions of three new genera 16:589-598. and seven new species. New Zealand Journal of Marine and Freshwater Research 39:29-75. Danielopol, D.L., A. Baltanás, and W.F. Humphreys. 2000. Danielopolina kornickeri sp. n. (Ostracoda, Karanovic, I. (in press). A new Candoninae genus Thaumatocypridoidea) from a Western Australian (Crustacea, Ostracoda) from subterranean watres of anchialine cave: morphology and evolution. Zoologica Queensland, with cladistic analysis of the tribe Scripta 29:1-16. Candonopsini. Memoirs of the Queensland Museum. Harvey, M.S. 1992. The Schizomida (Chelicerata) of Karanovic, I. and Marmonier, P. 2002. On the genus Candonopsis (Crustace: Ostracoda: Candoninae) in Australia. Invertebrate Taxonomy 6:77-129. Australia, with key to the world recent species. Harvey, M.S. 2000. A review of the Australian schizomid Annales de Limnologie, 38:199-240. genus Notozomus (Hubbardiidae). Memoirs of the Karanovic, I. and Marmonier, P. 2003. Three new genera Queensland Museum 46:161-174. and nine new species of the subfamily Candoninae Holsinger, J.R. 1993. Biodiversity of subterranean (Crustacea, Ostracoda, Podocopida) from the Pilbara amphipods crustaceans: global patterns and Region (Western Australia). Beuafortia 53:1-51. zoogeographic implications. Journal of Natural History Karanovic, T. 2004. Subterranean copepods (Crustacea: 27:821-835. Copepoda) from arid Western Australia. Crustaceana Humphreys, W.F. 1999. Relict stygofaunas living in sea Supplement 3: 1-366. salt, karst and calcrete habitats in arid northwestern Leys, R., C.H.S. Watts, S.J.B. Cooper, and W.F. Australia contain many ancient lineages. 219-227. In Humphreys. 2003. Evolution of subterranean diving W. Ponder and D. Lunney [eds.], The Other 99%. The beetles (Coleoptera: Dytiscidae: Hydroporini, Conservation and Biodiversity of Invertebrates. Bidessini) in the arid zone of Australia. Evolution Transactions of the Royal Zoological Society of New 57:2819-2834. South Wales, Mosman 2088. Martens, K. and Rossetti, G. 2002. On the Darwinulidae Humphreys, W.F. 2001. Groundwater calcrete aquifers in (Crustacea, Ostracoda) from Oceania. Invertebrate the Australian arid zone: the context to an unfolding Systematics 16: 195-208. plethora of stygal biodiversity, pp 63-83. In: Namiotko, T., K. Wouters, D.L. Danielopol, and W.F. Subterranean Biology in Australia 2000, W.F. Humphreys. 2004. On the origin and evolution of a Humphreys and M.S. Harvey (Eds). Records of the new anchialine stygobitic Microceratina species Western Australian Museum, Supplement No. 64. (Crustacea, Ostracoda) from Christmas Island (Indian Humphreys, W.F. 2004. Diversity patterns in Australia, Ocean). Journal of Micropaleontology 23:49-59. pp183-196. In D.C. Culver and W. White [eds.], Scarsbrook, M.R., G.D. Fenwick, , I.C. Duggan, and M. Encyclopedia of Caves. Academic Press, San Diego. Haase. 2003. A guide to the groundwater invertebrates of New Zealand. NIWA Science and Karanovic, I. 2003a. Towards a revision of Candoninae Technology Series, Wellington. (Crustacea, Ostracoda): Description of two new genera from Australian groundwaters. Species Taiti, S. and Humphreys, W.F. 2001. New aquatic Diversity 8:353-383. Oniscidea (Crustacea, Isopoda) from groundwater calcretes of Western Australia. In: Subterranean Karanovic, I. 2003b. A new genus of Candoninae Biology in Australia 2000, W.F. Humphreys and M.S. (Crustacea, Ostracoda, Candonidae) from the Harvey [eds.], Records of the Western Australian subterranean waters of southwestern Western Museum, Supplement No. 64:133-151. Australia. Records of the Western Australian Museum Watts, C.H.S. and Humphreys, W.F. 2004. Thirteen new 21:315-332. Dytiscidae (Coleoptera) of the genera Boongurrus Karanovic, I. 2004. Towards a revision of Candoninae Larson, Tjirtudessus Watts and Humphreys and (Crustacea, Ostracoda): On the genus Candonopsis Nirripirti Watts and Humphreys, from underground Vavra, with description of new taxa. Subterranean waters in Australia. Transactions of the Royal Society Biology 2:91-108. of South Australia 128:99-129.

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ASSESSMENT AND CONSERVATION OF AQUATIC LIFE IN THE SUBSURFACE OF THE PILBARA REGION, WESTERN AUSTRALIA

S.M. EBERHARD, S.A. HALSE, M.D. SCANLON, J.S. COCKING and H.J. BARRON

Department of Conservation and Land Management, Science Division, PO Box 51, Wanneroo, WA 6065, Australia. ([email protected]), ([email protected]), ([email protected]), ([email protected]), ([email protected]).

ABSTRACT To provide a framework for assessment of mine development projects in the Pilbara, as well to plan conservation of groundwater biodiversity, the Department of Conservation and Land Management (CALM) is undertaking a five-year survey (2002 to 2007) of Pilbara stygofauna. The project aims to map regional patterns in subterranean biodiversity by sampling the range of groundwater environments that can be accessed via bores, wells, caves, springs, spring- brooks and hyporheic habitats. The first year of sampling has revealed many new stygal species, and indicate that stygofauna is abundant and widely distributed across the region, and occurs in several different aquifer types. Seventy-one per cent of 355 bore/well samples yielded stygofauna. Modifications to sampling equipment and sampling protocol may have contributed to higher recovery rates of stygofauna than reported in other survey work. The Pilbara stygofauna comprises at least 150 species belonging to 77 genera and 39 families. The Pilbara is an important region for subterranean biodiversity.

KEYWORDS : AUSTRALIA, PILBARA, STYGOFAUNA, SURVEY, CONSERVATION, GROUNDWATER.

1. INTRODUCTION (Department of Mineral and Petroleum The Pilbara region (20-24o S and 115-122o E) Resources 2001). Groundwater is a critical issue covers an area of approximately 178,000 km2 in in the Pilbara because mining below the north-west Western Australia (Figure 1). The watertable in large open-cut pits requires Pilbara lies adjacent to Cape Range and Barrow dewatering of surrounding aquifers. Other major Island, two areas well known for containing rich impacts on groundwater resources include stygofaunas (Humphreys 2000). Work in the abstraction for water supplies to towns, industry, Pilbara during the 1990s, spearheaded by W.F. and agriculture (Water and Rivers Commission Humphreys and colleagues, established the 1996). In recent years, there has been some presence of a diverse and scientifically conflict between groundwater abstraction or mine interesting stygofauna there as well (Humphreys dewatering operations, and the protection of 2001). To date about 50 stygal species (nearly all groundwater-dependent ecosystems, including Crustacea) have been described or recorded, stygofauna (Water and Rivers Commission 1996; including: Amphipoda (Barnard and Williams Johnson and Wright 2001; Playford 2001). 1995; Bradbury 2000), Spelaeogriphacea (Poore and Humphreys 1998, 2003), Isopoda (Knott and In the most extreme cases, mine development Halse 1999; Wilson 2003), Copepoda (Pesce et proposals have been halted or delayed because al. 1996; Laurentiis et al. 1999, 2001; Lee and it was considered they posed a threat to the Huys 2002; Karanovic 2004), Ostracoda conservation of stygofauna (Playford 2001; (Martens and Rossetti 2002; Karanovic 2003; Finston et al. 2004). In Western Australia, all Karanovic and Marmonier 2003), Acarina native fauna species (though not necessarily (Harvey 1998), and Oligochaeta (Pinder 2001). individual animals) are protected under the Wildlife Conservation Act 1950 and environmental impact assessment of The Pilbara has a sparse human population but development proposals requires that proponents is economically important because of its mineral demonstrate that no species are placed at risk of wealth. In 2000, Western Australia produced 158 extinction (Playford 2001; Department of million tonnes of iron ore representing 15% of Conservation and Land Management 2004). This world production with forecasts of expanded requires accurate recognition of species and future markets and new mine developments knowledge of their distributions.

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To provide a framework for assessment of and Port Hedland Coast Basins respectively development projects in the Pilbara, as well to (Figure 2). Each basin drains to the Indian Ocean plan conservation of groundwater biodiversity, via a series of river systems characterised by the Department of Conservation and Land highly sporadic flow regimes. The major Management (CALM) is undertaking a five-year topographic features of the Pilbara are the survey (2002 to 2007) of Pilbara stygofauna. The Hamersley Plateau, which coincides with the project aims to map regional patterns in Central Pilbara iron ore region and contains the subterranean biodiversity by sampling the range Hamersley Range that reaches an elevation of of groundwater environments that can be 1,250 m above sea level (asl), and which is accessed via bores, wells, caves, springs, separated by the Fortescue Valley from the spring-brooks, and hyporheic habitats. This short Chichester Plateau and Range of more subdued paper describes the approach and methods used relief (618 m asl). The river drainage systems in this survey, and presents some preliminary have deeply dissected the margins of the results from the first year of sampling. Hamersley and Chichester Plateaus, but then

Figure 1. Map of Australia showing the Pilbara region in Western Australia.

1.1 Climate and physiography follow broad low-gradient valleys across The Pilbara experiences extreme climatic extensive lowlands to wide coastal plains. conditions characterised by high daytime Groundwater occurs throughout the Pilbara temperatures in summer (average maximum 36- region in Precambrian basement rocks, 39¡C), low winter minima (average minimum 6- Phanerozoic sedimentary basins and Cainozoic 12¡C), and high evaporation (average annual deposits. Total stored fresh-saline groundwater potential evaporation 3200-4000 mm). Annual resources are estimated at 67,140 GL with average rainfall is very low (200-350 mm) but renewable fresh-marginal resources of 291 with high inter-annual variation associated with GL/year (Allen 1997). The aquifers have been irregular tropical cyclones (Gentilli 1972; Bureau classified into three types by Johnson and Wright of Meteorology 1977). (2001): (1) Unconsolidated sedimentary aquifers; The Pilbara region coincides with the emergent (2) Chemically-deposited aquifers; (3) Fractured- part of the Pilbara Craton, which has remained rock aquifers. Unconsolidated sedimentary more or less continually above sea level since aquifers comprise Cainozoic valley and coastal the Proterozoic (> 545 Ma), bordered by marine plain alluvium and colluvium, while the environments (including the Tethys Sea) from the chemically-deposited aquifers consist of calcrete Devonian (410-354 Ma) until the fragmentation of or pisolitic limonite formed within the valley-fill Gondwana in the Cretaceous (141-65 Ma) sequences. Fractured rock aquifers are (Cockbain and Hocking 1990). The Pilbara developed in Proterozoic and Archaean encompasses five major drainage basins, the sedimentary and volcanic rocks including Ashburton, Fortescue, De Grey, Onslow Coast dolomite, sandstone, shale, chert, banded-iron

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Figure 2 : Pilbara region showing the five major hydrographic basins and the physiographic features mentioned in the text. formation and basalt. The calcretes and spring-brooks, and > 3,700 wells and bores dolomites are commonly karstic, and vuggy associated with pastoral and town water supply, porosity is developed within the limonites road and railway construction, and mine (Johnson and Wright 2001). dewatering operations (Allen 1997). However, most wells and bores were concentrated either in high-yield valley-fill aquifers used for water supply, or local aquifers where mine dewatering, 2. METHODS road and railway construction occurs, rather than 2.1 Survey constraints, limitations and being evenly distributed across the landscape. approach An important subsidary aim of the survey was to The scope of the survey was constrained by collaborate with taxonomists to support several factors, including: (1) Large land area to description of new taxa. Groups in which be covered with relatively few access points for particular efforts were made are Copepoda, sampling groundwater; (2) Limited existing Ostracoda, Isopoda and Amphipoda. The survey knowledge of the taxonomy and biology of has also collaborated with geneticists using DNA Pilbara species and their distributions; (3) techniques to define species boundaries and to Taxonomic impediment in that most Western reconcile genetic data with morphological Australian invertebrates are undescribed with characters. The latter have been found to be few taxonomists working on them and a poor variable in some Amphipoda (see Bradbury framework for morpho-species identifications; (4) 2000; Finston et al. 2004). Logistical constraints imposed by field operations 2.2 Sample sites in a remote and rugged location with extreme Sample sites were selected to provide broad climate. With these constraints in mind, the geographic coverage and to encompass the overall aim was to obtain an overview of range of geologic, topographic, and biodiversity patterns at a regional scale. physiographic environments across the Pilbara Achieving broad spatial coverage in a restricted region. Sites were also selected to cover different time frame (4 years for fieldwork) reduced the aquifer types (porous sedimentary, chemically- capacity for repeated sampling at individual sites deposited, fractured-rock, karstic and vuggy and for investigation of distribution patterns at aquifers). Approximately 450 (eventually 550) smaller spatial, or longer temporal, scales. bores and wells were selected (Figure 3). Parallel studies (described below) evaluated Resource and time constraints prevented issues such as sampling efficiency, and comparable sampling coverage of springs, stratification in the water column. spring-brooks and the hyporheos, so about 40 of Despite large deposits of karstified carbonate these groundwater habitats will be sampled. In rocks (Proterozoic dolomites and Cainozoic conjunction with the stygofauna survey, there is a calcretes), there were very few caves accessible parallel survey of aquatic fauna in Pilbara surface for sampling. There were many springs and waterbodies, including springs, brooks and rivers

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(A.M. Pinder, S.A. Halse and J.M. McRae water sample was refrigerated for laboratory unpublished data). Where possible, both surveys determination of solute concentrations (Ca2+, 2+ 3+ + 2+ 2+ + - - 2- will sample the same springs, spring-brooks and Fe /Fe , K , Mg , Mn , Na , CL , NO3 , CO3 , - 2- groundwater-fed river pools to evaluate the fauna HCO3 , SO4 , SiO2, Si, Sr), alkalinity, hardness, in groundwater/surface water ecotones. colour, turbidity, electrical conductivity, pH, and total dissolved solids. Laboratory methods 2.3 Sampling protocol and equipment followed APHA (1995). 2.3.1 Bores and wells Bores and wells were sampled for stygofauna Environmental attributes recorded for each site using a plankton net of suitable diameter (47 mm, include latitude, longitude, altitude, bore/well 97 mm, 147 mm, or 197 mm) to match the construction details (including where available, bore/well. The net, with a weighted McCartney the depth and geology at the slotted interval), vial attached, was lowered to the base of the surface geology, vegetation, land-use, and bore/well then agitated up and down (±1 m, 6 impacts. times) to disturb the bottom sediment. Six hauls

Figure 3 : Pilbara region showing the five major hydrographic basins and location of 450 bores and wells selected for sampling stygofauna. The clustered and linear distribution patterns reflect, respectively, the concentration of bores in water supply borefields within high-yield valley-fill aquifers and alignments of bores sunk for construction of roads and railways. The spatial coverage will be improved by sampling an additional 100 bores/wells.

Standing water level (SWL, in metres below of the entire water column were made, the first ground level) and the maximum depth of each three hauls used a 150 µm net to capture bore was measured to the nearest 0.05 m with a macrofauna, the second three hauls used a 50 Richter Electronic Depth Gauge or weighted µm net to capture microfauna such as rotifers. To Lufkin tape measure. Temperature, pH, electrical minimize loss of fauna through bow-wave effects conductivity, dissolved oxygen, turbidity and during hauling, the McCartney vial had the redox were measured at—1 m SWL using a bottom removed and replaced with 50 µm mesh. calibrated Yeo-Kal 611 water quality analyser. The entire net haul sample was transferred to a Water samples for laboratory analysis 120 ml labelled polycarbonate container and (undertaken at Chemistry Centre, Perth) were preserved in 100% ethanol. To maximize collected from < 1 m SWL using a sterile bailer preservation for possible DNA analysis, the (Clearwater PVC disposable 38 x 914 mm), and ethanol was replaced after a few hours by stored in sterile, rinsed 250 ml plastic bottles. decanting the sample through a 50 µm net and One 250 ml water sample was filtered through a refilling the sample bottle with fresh 100% 45 µm membrane and frozen for analysis of ethanol. To eliminate the possibility of faunal nutrients (total soluble N, total soluble P). Highly contamination between sites, the nets were turbid samples were pre-filtered though a glass- sterilized by washing in a decontaminant (5% fibre filter using a hand vacuum pump (Millipore solution of Decon 90), then rinsed in distilled Sterifil Aseptic 47 mm OM041). A second 250 ml water and air-dried.

64 SWSB December 2004 SESSION 1 : AUSTRALIAN STYGOFAUNA AND WA PROJECT (PILBARA)

2.4 Targeted field studies maximum depth 100 m using the Yeo-Kal 611 2.4.1 Repeated sampling water quality analyser. A down-hole video camera (Underwater Video Systems UVS-VDS- The very large land area to be surveyed meant 2001 connected with a JVC GR-DV900AA cam that each bore/well was sampled on only two occasions, while springs, spring-brooks and the recorder) will be used to relate any physico- hyporheos were sampled only once. Each chemical stratification to the distribution and bore/well was sampled once in late autumn-early abundances of animals within the bore water winter (April to July) and once in late winter-early column. spring (August to October) to capture possible seasonal influences on species occurrence 2.5 Springs, spring-brooks and hyporheos (extremely hot summers preclude field sampling Preliminary sampling of interstitial habitats in at that time of year). While preliminary data on springs, spring-brooks and the hyporheos of sampling efficiency suggested two sample semi-permanent river pools was undertaken events per site was less than optimal for using a Bou Rouch pump. Three subsamples, documenting the stygofaunal assemblage of between 2 and 10 m apart, were collected at individual bores/wells, whether there are each site to account for small-scale implications for description of regional patterns heterogeneity. Unless shallow bedrock will be evaluated by repeated sampling of a small underlying the sediments prevented it, sampled subset of the sites. The subset consisted of 14 depth was generally 0.5-1 m below the ground spatially dispersed sites located in both coastal surface. A diaphragm bilge pump (Munster and inland settings. The sites exhibited a range Simms Eng. Ltd.) with 25 mm inlet/outlet was in species richness (from zero to > 20 taxa used to pump a 10 L volume of sample as fast as recorded during the initial two sampling possible into a 10 L bucket with 1 L graduations events).These sites will be sampled twice each marked. Pump discharge rate (L/sec) was year for 2 to 3 years, with > 3 months separating recorded and the physico-chemical parameters each sample event, and species accumulation (temperature, pH, electrical conductivity, curves will be generated. dissolved oxygen) of the sample in the bucket 2.4.2 Sampling efficiency were measured with a WTW Multi 340i meter, The sampling efficiency of the net haul method and 2 x 250 ml water samples were collected for (cf. Malard et al. 1997) is being evaluated in five laboratory analysis of nutrients and ionic bores in alluvial aquifers. A standard set of six net concentrations. The bucket sample was then hauls is taken and then the bore is purged. elutriated to separate water and animals from Purged water is passed through a 50 µm sediment, and filtered through a plankton net (50 plankton net over the pump outlet and all animals µm mesh). After elutriation, the volume of retained. The volume purged is three times the sediment remaining in the bucket was estimated bore volume, using a centrifugal pump (Grundfos to nearest 0.1 L. Between sites, the Bou Rouch MP-1) with maximum pump rate of 3 m3/hour. equipment was thoroughly rinsed in filtered After the bore refills, it is re-sampled with six net water, and the net was decontaminated. hauls. Sampling efficiency is calculated by Samples were preserved in 100% ethanol. comparing the fauna collected pre-purging with Future sampling will utilise a Grillot pump as the total fauna collected pre-, during and post- recommended in Malard (2003). pur-ging. Except after heavy rainfall, Pilbara rivers consist 2.4.3 Water column profiling and down-hole video of isolated pools with either no, or very limited, At a few selected sites, physico-chemical flow, which precludes detection of upwelling and profiling of the water column in bores and wells downwelling zones by pressure differential using will be undertaken to investigate possible effects the T-bar method (see Malard et al. 2003). from stratification, which may influence the Instead, potential zones of groundwater distribution of animals or the interpretation of upwelling were inferred from geomorphic water chemistry data collected at—1 m SWL. In structures such as bedrock outcrops which cased bores where the slotted interval occurs frequently form barrages to hyporheic flow in the deeper than the SWL, a ‘deadwater zone’ of alluvium of river channels (eg. Davidson 1975). uncharacteristic water chemistry may develop Environmental attributes recorded at each Bou (Malard et al. 2003). Additionally, stratified Rouch sampling site were the same as for anchialine habitats may be present in coastal bores/wells. The following additional details were areas such as the lower Fortescue River where also recorded: channel width, alluvium thickness, there is a saltwater interface (Commander 1993) estimated discharge (L/sec), flow regime and elements of an anchialine fauna occur (perennial, semi-permanent, intermittent), spring (Humphreys 2001). Accordingly, the physico- type (limnocrene, rheocrene, helocrene); type of chemistry of the water column will be profiled to hyporheic zone (1, 2, or 3 sensu Malard (2003)).

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2.6 Identification and taxonomy with zero taxa). Across all samples, the mean Prior to sorting, samples were first separated into number of animals per sample was 23.2 (range 0 three size fractions by sieving through 250, 90, Ð 250). For the 97 sites sampled twice, a mean of and 53 µm Endicott sieves. Sorting occurred 5.4 taxa per site was recorded. On average (104 under Leica MZ3, 6, 8 or 12 dissecting comparisons), two sampling events had 24% microscopes with 0.6x or 10x eyepiece and 10x species in common, while each sample or 16x objective. Each taxon was identified to the comprised 38% common species. lowest taxonomic rank possible using published 3.2 Systematic diversity keys and descriptions, and the numbers of each Initial sampling has revealed the presence of taxon were recorded. Identification of microfauna many new species of Crustacea and Acarina, in used a Zeiss Axioskop 2 compound microscope addition to undescribed Mollusca, Oligochaeta, with 10x eyepiece and 5x, 10x, 20x, 40x and Polychaeta, Tricladida, and Nematoda. Thus far, 100x objective. Identifications were confirmed by the Pilbara stygofauna comprises, minimally, > specialist taxonomists as necessary. Many taxa 150 species belonging to 77 genera and 39 were new and undescribed so examples of each families. The classes/orders and families new (morpho-) species collected were retained in a voucher collection and used for checking recorded to date include: Acarina (Arrenuridae, identifications and designating new species. In Halacaridae, Limnesiidae, Mideopsidae, collaboration with molecular geneticists (at the Pezidae, Oribatida), Amphipoda (Bogidiellidae, South Australian Museum and the University of Melitidae, Neoniphargidae, Paramelitidae), Western Australia, see for example Finston et al. Aphonaneura, Bathynellacea (Bathynellidae, this volume), DNA techniques were used to Parabathynellidae), Copepoda (Ameiridae, assist in defining species boundaries, particularly Canthocamptidae, Cyclopidae, Diosaccidae, in groups such as Amphipoda that appear to Ectinosomatidae, Parastenocaridae), Decapoda have few reliable morphological characters for (Atyidae), Gastropoda (Hydrobiidae), Isopoda distinguishing species. Publication of (Amphisopidae, Cirolanidae, Microcerberidae, descriptions of new species collected during the Phyloscidae, Tainisopidae), Oligochaeta survey will be supported. (Enchytraeidae, Naididae, Phreodrilidae, Tubificidae), Polychaeta (Neiridae), Ostracoda 2.7 Database, analysis, and reporting (Candonidae, Darwinulidae, Limnocytheridae), All survey data are stored in a relational Rotifera (Philodinidae), Speleaogriphacea database using MS Access. The database (Spelaeogriphidae), Thermosbaenacea structure comprises several linked tables that (Halsobaenidae), Tricladida (family encompass taxa, site, and environmental indeterminate), Nematoda (family attribute data. Site x species and site x indeterminate). environmental attribute matrices will be compiled for analysis of regional-scale biodiversity patterns using multivariate analysis software 4. DISCUSSION AND CONCLUSION such as PATN (Belbin 1993). Analysis will seek to The Pilbara is an important region for determine inter alia, groups of species with subterranean biodiversity. It contains a similar patterns of occurrence, groups of sites comparatively rich systematic diversity of with similar stygofauna and the environmental stygofauna (cf. Botosaneanu 1986). The current attributes that influence biodiversity patterns. A survey has uncovered many new stygal species. parallel study will examine groundwater Preliminary results from the first year of sampling chemistry and its influence on the distribution of indicate that stygofauna is abundant and widely ostracods (J. Reeves and P. De Deckker, distributed across the region, and occurs in Australian National University, unpublished several different aquifer types. The 71% recovery data). The survey results are intended to be rate of stygofauna from 355 samples was higher published in a series of papers in refereed than anticipated given the generally lower rates scientific journals. previously reported in other survey work, both in the Pilbara and elsewhere in Western Australia. 3. PRELIMINARY RESULTS Recovery rates from other surveys in the Pilbara averaged 38% (range 0 to 69%, n = 355 samples 3.1 General sampling statistics from 311 bores/wells in 14 aquifers and 15 In the first year of sampling (November 2002 to surveys; data extracted from Eberhard 1998; November 2003), 355 samples were collected Eberhard and Humphreys 1999 ; Biota from 253 bores, of which 97 bores were sampled Environmental Services 2002, 2003, 2004; Knott at least twice including five bores that were and Goater 2004). Lower recovery rates were purged (three samples per event). Fauna was also reported from porous and limestone aquifers detected in 252 of 355 samples (71%), with a in south-west Western Australia, a region where mean of 3.8 taxa per sample (excluding samples stygofauna appear to be less diverse. One

66 SWSB December 2004 SESSION 1 : AUSTRALIAN STYGOFAUNA AND WA PROJECT (PILBARA) survey (150 bores) in the Perth Basin yielded De Laurentiis, P., G.L. Pesce, and Humphreys, W.F. 1999. stygofauna in 2% of samples (reported in Knott Copepods from ground waters of Western Australia. and Goater 2004), while another survey (33 IV. Cyclopids from basin and craton aquifers (Crustacea: Copepoda: Cyclopidae). Records of the bores) yielded 15% (Eberhard 2003). The higher Western Australian museum 19:243-257. recovery rates in the CALM Pilbara survey may De Laurentiis, P., G.L. Pesce, and Humphreys, W.F. 2001. be the result of: (1) Use of smaller net mesh Copepods from ground waters of Western Australia. sizes (50 and 150 µm) to capture microfauna, V1. Cyclopidae (Crustacea: Copepoda) from the whereas other sampling has used 200, 250 or Yilgarn Region and the Swan Coastal Plain. Records 350 µm mesh; (2) Modifications to net hauling of the Western Australian Museum, Supplement equipment and sampling protocol (x 6 net hauls 64:243-257. with vigorous agitation of bottom sediments) to Department of Conservation and Land Management. maximize capture of animals from sediment in 2004. Towards a biodiversity conservation strategy for Western Australia. Government of Western Australia. the bottom of bores, and to minimize fauna loss 79 pp. through bow-wave effects during hauling. Some Department of Mineral and Petroleum Resources. 2001. other surveys employed only 2 or 3 net hauls, Western Australian Iron Ore Industry. July 2001. and less vigorous agitation of bottom sediments; Department of Mineral and Petroleum Resources, (3) Sorting in the laboratory under relatively high Perth. magnification, rather than field sorting with lower Eberhard, S. 1998. Groundwater fauna in the West magnification microscopes. Angelis project area. Western Australian Museum report to Ecologia Environmental Consultants, Perth. BIBLIOGRAPHY 1-19 pp. Eberhard, S. 2003. Preliminary investigation of Allen, A.D. 1997. Groundwater: the strategic resource - a stygofauna in the Blackwood Groundwater Area. geological perspective of groundwater occurrence Report prepared as part of establishment of interim and importance in Western Australia. Western ecological water requirements for the Blackwood Australia Geological Survey, Report 50:1-61. Groundwater Area, Western Australia. Murdoch APHA. 1995. Standard methods for the examination of Environment report to URS Australia Pty Ltd., Perth. water and wastewater. American Public Health 14 pp. Association, Washington. Eberhard, S.M. and Humphreys, W.F. 1999. Stygofauna Barnard, J.L. and Williams, W.D. 1995. The taxonomy of survey - Ore Body 23 (Newman) and Mine Area C. crangonyctoid Amphipoda (Crustacea) from Western Australian Museum report prepared for BHP Australian freshwaters: part 2. Records of the Iron Ore Pty Ltd, Perth. 57 pp. Australian Museum 47:161-201. Finston, T.L., J.H. Bradbury, M.S. Johnson, and B. Knott. Belbin, L. 1993. PATN v. 3.5: pattern analysis package. 2004. When morphology and molecular markers Commonwealth Scientific and Industrial Research conflict: a case history of subterranean amphipods Organisation (CSIRO), Canberra. 108 pp. from the Pilbara, Western Australia. Animal Biota Environmental Services. 2002. Hamersley Iron Biodiversity and Conservation 27:83-94. stygofauna sampling - 2002. Interim Summary report Gentilli, J. 1972. Australian climate patterns. Thomas to Hamersley Iron Pty Ltd, Perth. 28 pp. Nelson and Sons, Melbourne. 285 pp. Biota Environmental Services. 2003. West Angelas Harvey, M. 1998. Unusual new water mites (Acari: stygofauna survey. Report to Robe River Mining Co Hydracarina) from Australia, Part 1. Records of the Pty Ltd, Perth. 30 pp. Western Australian Museum 19:91-106. Biota Environmental Services. 2004. Sherlock Bay Nickel Humphreys, W.F. 2000. The hypogean fauna of the Cape Stygofauna Survey. Report to Sherlock Bay Nickel Range peninsula and Barrow Island, northwestern Corporation Ltd, Perth. 20 pp. Australia. In: H. Wilkens, D.C. Culver and W.F. Botosaneanu, L. 1986. Stygofauna Mundi: A faunistic, Humphreys [eds.], Ecosystems of the world, Vol. 30 - distributional and ecological synthesis of the world Subterranean ecosystems, 581-601. Elsevier, fauna inhabiting subterranean waters (including the Amsterdam. marine interstitial). Brill and Backhuys, Leiden. Humphreys, W.F. 2001. Groundwater calcrete aquifers in Bradbury, J.H. 2000. Western Australian stygobiont the Australian arid zone: the context to an unfolding amphipods (Crustacea: Paramelitidae) from the Mt plethora of stygal biodiversity. Records of the Western Newman and Millstream regions. Records of the Australian Museum, Supplement 64:63-83. Western Australian Museum, Supplement 60:1-102. Johnson, S.L. and Wright, A.H. 2001. Central Pilbara Bureau of Meteorology. 1977. Climatic atlas of Australia. groundwater study. Water and Rivers Commission. Australian Government Publishing Service, Canberra. 102 pp. Cockbain, A.E. and Hocking, R.M. 1990. Phanerozoic. In, Karanovic, I. 2003. Towards a revision of Candoninae Geology and Mineral resources of Western Australia: (Crustacea: Ostracoda): description of two new Western Australia Geological Survey, Memoir 3, 750- genera from Australian groundwaters. Species 755. Department of Mines, Perth. Diversity 8:353-383. Commander, P.D. 1993. Hydrogeology of the Fortescue Karanovic, I. and Marmonier, P. 2003. Three new genera River alluvium. Geological Survey of Western and nine new species of the subfamily Candoninae Australia Hydrogeology, Report No. 1993/14: 45. (Crustacea: Ostracoda: Podocopida) from the Pilbara Davidson, W.A. 1975. Hydrogeological reconnaissance of region (Western Australia). Beaufortia 53:1-51. the northwest Pilbara region. Western Australia Karanovic, T. 2004. The genus Metacyclops Kiefer in Geological Survey Record. Australia (Crustacea: Copepoda: Cyclopoida), with

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description of two new species. Records of the Pesce, G.L., P. De Laurentiis, and Humphreys, W.F. 1996. Western Australian Museum 22:193-212. Copepods from groundwaters of Western Australia, I. Knott, B. and Goater, S. 2004. Draft report for Fortescue The genera Metacyclops, Mesocyclops, Microcyclops Metals Group, Pilbara iron ore mine sites. Zoology and Apocyclops (Crustacea: Copepoda: Cyclopidae). Department, University of Western Australia, Perth. 11 Records of the Western Australian Museum 18:77-85. pp. Pinder, A.M. 2001. Notes on the diversity and distribution Knott, B. and Halse, S.A. 1999. Pilbarophreatoicus of Australian Naididae and Phreodrilidae platyarthricus n.gen., n.sp. (Isopoda: Phreatoicidae: (Oligochaeta: Annelida). Hydrobiologia 463:49-64. Amphisopodidae) from the Pilbara region of Western Playford, P.E. 2001. Subterranean Biotas in Western Australia. Records of the Australian Museum 51:33- Australia. Report for the Environmental Protection 42. Authority, Perth. 17 pp. Lee, W. and Huys, R. 2002. A new genus of groundwater Poore, G.C.B. and Humphreys, W.F. 1998. The first Ameiridae (Copepoda, Harpacticoida) from boreholes record of the Spelaeogriphacea (Crustacea) from in Western Australia and the artificial status of Australasia: a new genus and species from an aquifer Stygonitocrella Petkovski, 1976. Bulletin of the in the arid Pilbara of Western Australia. Crustaceana, Natural History Museum, London (Zoology) 68:39-50. 71:721-742. Malard, F., M.-J. Dole-Oliver, J. Mathieu, and Stoch, F. Poore, G.C.B. and Humphreys, W.F. 2003. Second 2002. Sampling manual for the assessment of species of Mangkurtu (Spelaeogriphacea) from north- regional groundwater biodiversity. PASCALIS western Australia. Records of the Western Australian European Project: Protocols for the assessment and Museum :67-74. conservation of aquatic life in the subsurface. 74 pp. Water and Rivers Commission. 1996. Pilbara region (unpublished report). water resources review and development plan, Malard, F., J.-L. Reygrobellet, R. Laurent, and Mathieu, J. summary report 4. Water and Rivers Commission, 1997. Developments in sampling the fauna of deep Perth. 26 pp. water-table aquifers. Archiv für Hydrobiologie Wilson, G.D.F. 2003. A new genus of Tainisopidae fam. 138:401-432. nov. (Crustacea: Isopoda) from the Pilbara, Western Martens, K. and Rossetti, G. 2002. On the Darwinulidae Australia. Zootaxa, 245:1-20. (Crustacea, Ostracoda) from Oceania. Invertebrate Systematics 16: 195-208.

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HAPLOTYPE DIVERSITY IN PILBARUS MILLSI, A WIDESPREAD GROUNDWATER SPECIES OF AMPHIPOD FROM THE PILBARA, WESTERN AUSTRALIA

1T. FINSTON, 1M. JOHNSON, 2W. HUMPHREYS, 3S. EBERHARD and 3S. HALSE.

1. School of Animal Biology M092, The University of Western Australia, 35 Stirling Hwy., Crawley, WA 6009, Australia ([email protected]). 2. Western Australian Museum, Locked Bag 49, Welshpool DC, Western Australia 6986, Australia. ([email protected]). 3. Department of Conservation and Land Management, P.O. Box 51, Wanneroo, WA6946. Australia. ([email protected]), ([email protected]).

ABSTRACT The amphipod Pilbarus millsi, has a widespread distribution in northwestern Western Australia. It is unclear whether ongoing gene flow or historical processes account for this pattern. Analysis of mtDNA sequence divergence between samples from ten catchments showed no evidence of contemporary gene flow. Instead, amphipods from each catchment had unique and highly distinct haplotypes, indicating that the distribution is ancient. Phylogenetic analysis revealed clades containing haplotypes from only a single catchment, suggesting past fragmentation events.

KEYWORDS : AMPHIPODS, CO1, MTDNA, PHYLOGEOGRAPHY, STYGOFAUNA.

1. INTRODUCTION Johnson, 2004).The present study examined Twenty-six species of stygobitic amphipods have specimens of P. millsi from ten catchments, using been described from the Pilbara, an arid region in a partial sequence of the mitochondrial COI the northwest of Western Australia (Bradbury and gene, to test hypotheses about the origin and Williams, 1997; Bradbury, 2000). Many are maintenance of a widespread species in a highly known from single bores. One species, Pilbarus subdivided habitat. The species may be millsi, is widespread, occurring in multiple widespread due to its dispersal capabilities, catchments throughout the Pilbara. However, its perhaps linked to its occasional occupation of distribution appears to be limited to areas surface waters and flooding events associated containing calcrete formations (Finston and with the area. Alternately, historical processes

Figure 1 : Map of the Pilbara, Western Australia, showing sample sites within the Ashburton and Fortescue River basins.

SWSB December 2004 69 SESSION 1 : AUSTRALIAN STYGOFAUNA AND WA PROJECT (PILBARA) may have involved a past fragmentation event of calculated using Kimura’s two-parameter a once widespread ancestor, perhaps linked to distance model in the DNADIST module. A the onset of aridity in Australia. maximum likelihood phylogeny was produced using the module DNAML on 100 multiple data 2. METHODS sets (generated in SEQBOOT). Specimens of P. millsi were collected from ten creek catchments located within two major river 3. RESULTS basins, the Ashburton, and the Fortescue (Fig. The number of haplotypes detected varied 1). With some exceptions, between five and ten among catchments, ranging from a single individuals per catchment were sequenced. haplotype at Caves Creek, Roy Hill and Whole genomic DNA was extracted from Spearhole Creek, to six at Weeli Wolli Creek. specimens in 50µl of a proteinase K extraction Importantly, haplotypes were not shared among buffer for between 10 and 20 hours. A 710 base catchments. Sequence divergence between pair (bp) fragment of the 3’ end of the cytochrome haplotypes within the same catchment did not oxidase subunit 1 (COI) gene was amplified, exceed 2.3%. Within the Ashburton, sequence using the primers LCO1490 (forward, 5’ GGT divergence ranged from 8.9 to 13.1% between CAA CAA ATC ATA AAG ATA TTG G 3’) and catchments and within the Fortescue, divergence HCO2198 (reverse, 5’ TAA ACT TCA GGG TGA ranged from 16.4 to 26.5% between catchments. CCA AAA AAT CA 3’; Folmer et al., 1994). The 25 Sequence divergence between haplotypes in µl PCR reactions used 0.2mM dNTPs, 4.0mM different basins ranged from 20.1 to 26.8%. The

MgCl2, 1x buffer, 12.5 pmoles of each primer, 1 phylogenetic analysis showed that genetic unit Taq, and 2.5 µl template. Sequences were structure within P. millsi reflected hydrological cleaned using the UltraCleanª PCR Clean-up structure. Each mtDNA lineage contained DNA purification kit (MoBio Laboratories, Inc.) haplotypes from only a single catchment (Fig. 2). prior to sequencing. The sequencing reaction While the use of a molecular clock is fraught with was carried out using the BigDye V3 Ready assumptions that are often violated, it can give a Reaction Mix (ABI Prism) and the products were rough estimate of the time frame associated with sequenced using both primers on an ABI 373 phylogenetic events. Using an estimate of the automated sequencer (Applied Biosystems). evolutionary rate of the COI gene of Sequences were aligned and edited by eye with approximately 2% divergence per million years, GeneDoc, version 2.6.002, using default haplotypes from the Ashburton and Fortescue settings. Relationships among the sequences River basins last shared a common ancestor were analysed using PHYLIP, version 3.57c. between 10 and 13 mya. Pair-wise nucleotide sequence divergence was

Figure 2 : CO1 maximum likelihood tree. The tree is rooted using A. subtenuis, (family Ceinidae) and Nedsia sp. (family Melitidae). Confidence levels > 50%, assessed by 100 replicate bootstraps are shown on branches. Number of individuals represented by each haplotype is shown for n>1.

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4. DISCUSSION BIBLIOGRAPHY Genetic structure within P. millsi reflected the Bradbury, J.H. 2000. Western Australian stygobiont hydrological structure of the region. There was amphipods (Crustacea: Paramelitidae) from the Mt no apparent gene flow between catchments, and Newman and Millstream regions. Records of the each lineage corresponded to individual creek Western Australian Museum Supplement, No. 60. catchments. Strong geographical associations of Bradbury, J.H., and Williams, W.D. 1997. The amphipod haplotypes may be evidence of a past (Crustacea) stygofauna of Australia: Description of new taxa (Melitidae, Neoniphargidae, Paramelitidae), fragmentation event (Templeton, 1998). This and a synopsis of known species. Records of the pattern of diversity suggests that the current Australian Museum 49:249-341. distribution of P. millsi is not due to gene flow, but Finston, T. and Johnson, M.S. 2004. Geographic patterns supports the hypothesis that a historical of genetic diversity in subterranean amphipods of the fragmentation event isolated populations of the Pilbara, Western Australia. Marine and Freshwater species within catchments and basins. One Research 55:619-628. explanation is that a surface ancestor sought Folmer, O., M. Black, W. Hoeh, R. Lutz, and R. refuge in subterranean waters when the climate Vrijenhoek. 1994. DNA primers for amplification of became more arid during the Miocene mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine (Humphreys, 2001). In support of this Biology and Biotechnology 3:294-299. hypothesis, the molecular clock estimate places Humphreys, W.F. 2001. Groundwater calcrete aquifers in divergence between the two basins at 10 Ð 13 the Australian arid zone: the context to an unfolding my, corresponding to the late Miocene or early plethora of stygal biodiversity. Records of the Western Pliocene. Pilbarus millsi may comprise multiple Australian Museum Supplement, No. 64, 63-83. morphologically cryptic species, or may be an Templeton, A.R. 1998. Nested clade analysis of example of morphological stasis in a genetically phylogeographic data: testing hypotheses about gene diverse, widespread species. flow and population history. Molecular Ecology 7:381- 397.

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SESSION 2 : PHYLOGENY, PHYLOGEOGRAPHY AND TAXONOMY

PHYLOGEOGRAPHY AND TAXONOMIC STATUS OF NIPHARGUS VIREI (SUBTERRANEAN AMPHIPOD).

1T. LEFÉBURE, 1C.J. DOUADY, 2M. GOUY, 3P. TRONTELJ, 4J. BRIOLAY and 1J. GIBERT.

1. University Claude Bernard of Lyon 1, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne Cedex, France. Laboratoire d’Ecologie des Hydrosystèmes Fluviaux, UMR-CNRS 5023. ([email protected]), ([email protected]), ([email protected]). 2. University Claude Bernard of Lyon 1, 43, Bd du 11 Novembre 1918, 69622 Villeurbanne Cedex, France. Laboratoire de Biométrie et Biologie Evolutive, UMR CNRS 5558. ([email protected]). 3. Department of Biology, Biotechnical Faculty, Univ. of Ljubljana, P.O. Box 2995, 1001 Ljubljana, Slovenia. ([email protected]). 4. University Claude Bernard of Lyon 1, 43, Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France. Développement des Techniques d’Analyse Moléculaire de la Biodiversité (DTAMB). ([email protected]).

Extreme conditions are suspected to be Likelihood analyses, based on two independent responsible for morphological convergence, and genes (28S and COI), revealed the same so to bias the assessment of biodiversity in tripartite structure (Fig. 1). N. virei populations groundwater. This hypothesis was tested within from Benelux (C in Fig.1), Eastern (B) and the well-defined subterranean amphipod Western France (A) appeared as independent Niphargus virei. All our Bayesian and Maximum evolutionary units. Molecular rates estimated via

Figure 1 : Most likely tree from the COI gene (left panel), 28S gene (upper right panel), their concatenation (lower right panel) and distribution of A, B and C groups based on the COI (middle panel). Numbers on the tree correspond to bootstrap support (Bayesian bootstrap for COI and ML bootstrap for the others, at least 100 replicates). On the middle panel white circles represent locations sampled and black circles distinct COI haplotypes.

SWSB December 2004 73 SESSION 12 : AUSTRALIANPHYLOGENY, STYGOFAUNAPHYLOGEOGRAPHY AND WA AND PROJECT TAXONOMY (PILBARA) global or Bayesian relaxed clock suggest that Protocols for the ASsessment and Conservation this split is at least 16 my old and accredit the of Aquatic Life In the Subsurface, (contract no cryptic diversity hypothesis. Surprisingly, the EVK2-CT-2001-00121) of the Fifth Research and geographical distribution of these lineages Technological Development Framework Program (particularly the group A, Fig. 1) pointed to of the European Community. evidence of recent dispersal through apparent vicariant barriers. Comparison of the molecular divergences BIBLIOGRAPHY between these lineages and between several Lefébure, T., J.C. Douady, M. Gouy, P. Trontelj, J. Briolay, thousand of different species of Crustacean give and J. Gibert. 2005. Molecular phylogeography of evidence that the three lineages of N. virei should Niphargus virei (Crustacea: Amphipoda) reveals evolutionary patterns in groundwater. Submitted to be considered as different species. In Molecular Ecology. consequence, we argue that future analyses of Lefébure, T., J.C. Douady, M. Gouy, and J. Gibert. 2005. subterranean biodiversity, or more generally of Relationship between morphological taxonomy and biodiversity in extreme environments, should molecular divergences: evidences from crustacean. consider morphology as a potentially biased Submitted to Molecular Ecology. marker and DNA of promising help.

ACKNOWLEDGEMENTS This work was financially supported by the European research project PASCALIS :

74 SWSB December 2004 SESSION 3 : BIODIVERSITY PATTERNS

HIERARCHICAL PATTERNS OF OBLIGATE GROUNDWATER BIODIVERSITY IN FRANCE

D. FERREIRA, F. MALARD, M.-J. DOLE-OLIVIER and J. GIBERT

University Claude Bernard of Lyon 1, UMR CNRS 5023, Ecologie des Hydrosystèmes Fluviaux, Equipe Hydrobiologie et Ecologie Souterraines, 43, Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France. ([email protected]), ([email protected]), ([email protected]), ([email protected]).

ABSTRACT

Groundwater biodiversity is expected to follow a spatialized hierarchical pattern so that nested areas of increasing size should become increasingly dissimilar in term of species composition. We selected a subset of 55 aquifers that had received comparable sampling efforts. Reciprocal scaling and a UPGMA dendrogram (unweighted pair-group method using arithmetic averages) were used to examine differences in species composition among aquifers. Both methods distinguished between 2 clusters of aquifers belonging to the Rhône River and Garonne River basins that contained distinct sets of species (beta diversity > 90%). The dendrogram divided these basins into smaller spatial units of increasing species similarity, thereby supporting the hypothesis of a spatial hierarchization of groundwater biodiversity. We argue that the most efficient strategy of data acquisition for identifying groundwater biodiversity regions would consist in determining the species composition of selected aquifers in territories that have been poorly investigated.

KEYWORDS : BETA DIVERSITY, PATTERNS, HIERARCHIZATION, STYGOFAUNA, FRANCE

1. INTRODUCTION and subspecies (Ferreira et al. 2005). The The delineation at different spatial scales of analysis of species occurrence data in 20 x 20 areas having dissimilar sets of species (i.e. grid cells did not enable to reveal the real spatial biodiversity regions) is of paramount importance pattern of biodiversity in France because of for prioritizing conservation efforts (Gering et al. taxonomic and sampling biases among cells. We 2003, Summerville et al. 2003). The diversity of therefore selected a subset data of aquifers that obligate-groundwater species (i.e. stygobionts) had received comparable sampling effort. Moreover in order to avoid taxonomic biases assemblages is presumably low at a local scale between selected aquifers, we only retained the but diversity is expected to increase markedly group of Crustacea (except Ostracoda and with increasing spatial scales because of the Syncarida) that is a well studied group both in importance of groundwater system fragmentation terms of taxonomy and geographic distribution. in generating species. We therefore The final subset data extracted from the full data hypothesized that biodiversity is expected to base was a presence - absence matrix of 117 follow a spatialized hierarchical pattern so that species distributed among 55 aquifers (Figure 1). nested areas of increasing size should become increasingly dissimilar in terms of species composition. For testing this hypothesis, we have 2.2 Data analysis examined the spatial variation of stygobiotic For the analysis, we used two types of methods species composition (beta diversity) among that have been classically used in the different selected aquifers in France that have examination of beta diversity patterns (Magurran received a comparable sampling effort. 1988). First, in order to investigate the overall similarity 2. MATERIAL AND METHODS of aquifers and to pick out major groupings, we used an ordination method called reciprocal 2.1. Data set scaling of the correspondence analysis that Within the framework of the PASCALIS program displays visually a simultaneous ordination of (Gibert 2001), a data base has been established species and aquifers in a common factorial plane at France scale that assembled a total of 380 (Thioulouse and Chessel 1992). Each species obligate groundwater (i.e. stygobiotic) species and each aquifer is represented by one ellipse.

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Figure 1 : Distribution of 55 selected aquifers (from Ferreira et al. in preparation).

Analyses were performed using ADE-4 software factorial scores revealed a strong regional (Thioulouse et al. 1997). clustering in the composition of aquifers Second, we used hierarchical cluster analysis assemblages by separating aquifers located in β the Rhône-Rhine region (negative scores) from involving the beta index sim (Lennon et al. 2001) considered as a narrow sense measure of beta those located in the Adour-Garonne region diversity in that it focuses more on difference in (positive scores). species composition than on difference in These two regions shared a limited number of species richness (see review in Koleff et al. species clearly identified by their bigger ellipses 2003). This index has been calculated for all such as Graeteriella unisetigera and Niphargus possible pairs of aquifers using the following kochianus kochianus (Figure 2). formula: In addition, five different clusters of aquifers β sim = (min (b,c)) / (min (b,c) + a) could be distinguished, (Figure 2, left panel), each of them being characterised by a distinct where a = number of species in common set of species (Figure 2, right panel): aquifers of between aquifers B and C, b = number of species the Jura Moutains characterized by the presence that occurs in the aquifer B but not in the aquifer of the isopods Caecosphaeroma virei and C, and c = number of species that occurs in the aquifer C but not in the aquifer B. Proasellus valdensis, alluvial aquifers of the Rhône River and its tributaries (e.g. Ain) The UPGMA linkage method (unweighted pair- characterised by species including the isopods group method using arithmetic averages) was Microcharon reginae and Proasellus walteri, used to compute a hierarchical tree in R software aquifers of the Corbières and North-Montpellier (Version 1.7.1, R Development Core Team, areas harboring several characteristic species http://www.R-project.org). including the isopods Stenasellus builli and Faucheria faucheri, aquifers of the Garonne 3. RESULTS River Basin characterised by putative species The reciprocal scaling analysis clearly identified including the polytypic isopod species two clusters of regions. The mapping of the Stenasellus virei, and the Arbailles Mountain

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Figure 2 : Reciprocal scaling of stygobiotic species tolerance and aquifer diversity in France. Left panel: the 55 aquifer ellipses with clusters corresponding to distinct regions. Right panel: the 117 species ellipses. Species lists correspond to species occurring preferentially in each cluster of aquifers. The scale (valid for both panels) is given by the diagram in the left bottom corner of the figure. (Modified from Ferreira et al. in preparation).

comprising distinct set of species including the by the highly fragmented nature of groundwater isopods Proasellus vandeli and P. spelaeus. systems. Because the reciprocal scaling analysis An aquifer-level approach is particularly supported that between-aquifers diversity is amenable to determine areas of dissimilar larger than the within aquifers-diversity, the use species composition and identify groundwater of a classification method appears to be of a habitats to be included in the Habitat Directive. great of interest. Indeed, the cluster analysis Our results suggest that sampling a limited clearly identified the same two clusters of regions number of aquifers in poorly studied areas would between which there is nearly 90% of average be the most effective strategy for obtaining a dissimilarity (Fig. 3). The dendrogram further valuable assessment of stygobiotic diversity divided these two regions into smaller spatial heterogeneity. Our results further provide units of increasing species composition similarity. guidelines for the implementation of ecological The Garonne River Basin and Jura Mountains background in the management of groundwater were further subdivided into 2 clusters of resources in France. First, to use aquifer as a aquifers. reference unit for identifying priority areas for conservation since aquifers have already used 4. DISCUSSION by the French Water Agency as spatial units for Both ordination and classification methods groundwater management. Second, to identify clearly distinguished between two clusters of aquifers of high biological value (i.e. species regions characterized by distinct sets of species. richness, number of endemics) and provide a list Following the definition of Olson and Dinerstein of characteristic species for each aquifer. (1998), these two regions can be considered as distinct stygofaunistic regions. The dendrogram divided these two regions into smaller spatial ACKNOWLEDGEMENTS units of increasing species composition similarity, This work was financially supported by the thereby supporting the hypothesis of a spatial European research project PASCALIS: Protocols hierarchization of groundwater species diversity. for the ASsessment and Conservation of Aquatic This spatialized hierarchical pattern is probably Life In the Subsurface, (contract no EVK2-CT- the result of multiple speciation events induced 2001-00121) of the Fifth Research and

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Figure 3 : Unweighted pair-group method using arithmetic averages (UPGMA) dendrogram of 55 aquifers based on presence - absence data for 117 crustacean species showing determined cluster of aquifers (modified from Ferreira et al. in preparation).

Technological Development Framework Program Koleff, P., K.J. Gaston, and J.L. Lennon. 2003. Measuring of the European Community. beta diversity for presence-absence data. Journal of Animal Ecology 72:367-382. Lennon, J. L., P. Koleff, J.J.D. Greenwood, and K.J. Gaston. 2001. The geographical structure of British BIBLIOGRAPHY bird distributions : diversity, spatial turnover and scale. Ferreira, D., M.-J. Dole-Olivier, F. Malard, and J. Gibert. Journal of Animal Ecology 70:966-979. 2003. Faune aquatique souterraine de France : base Magurran, A. 1988. Ecological diversity and its de données et éléments de biogéographie. measurement. Princeton University Press, Princeton. Karstologia 42:15-22. Olson, D.M. and Dinerstein, E. 1998. The global 200 : a Ferreira, D., F. Malard, M.-J. Dole-Olivier, and J. Gibert. representation approach to conserving the Earth's 2005. Obligate groundwater fauna of France : most biologically valuable ecoregions. Conservation diversity patterns and conservation implications. Biology 12(3):502-515. Biodiversity and Conservation (in submission). Summerville, K.S., M.J. Boulware, J.A. Veech, and T.O. Gering, J.C., T.O. Crist, and J.A. Veech. 2003. Additive Crist. 2003. Spatial variation in species diversity and partitioning of species diversity across multiple spatial composition of forest Lepidoptera in Eastern scales : implications for regional conservation of Deciduous Forests of North America. Conservation biodiversity. Conservation Biology, 17:488-499. Biology 17:1045-1057. Gibert, J. 2001. Protocols for the assessment and Thioulouse, J. and Chessel, D. 1992. A method for conservation of aquatic life in the subsurface reciprocal scaling of species tolerance and sample (PASCALIS) : a European project. In : Mapping diversity. Ecology 73(2): 670-680. Subterrenean Biodiversity : Proceedings of the International Workshop held, 18-20 March 2001, Thioulouse, J., D. Chessel, S. Dolédec, and J.-M. Olivier. Laboratoire Souterrain du CNRS, Moulis, Ariège, 1997. ADE 4 : a multivariate analysis and graphical France. D.C. Culver, L. Deharveng and J. Gibert and display software. Statistics and Computing 7:75-83. I.D. Sasowsky [Eds.], Karst Waters Institute, Special Publication 6, Charles Town, West Virginia, 19-21.

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ENVIRONMENTAL GRADIENTS IN GROUND WATERS. MAIN FACTORS DRIVING THE COMPOSITION OF STYGOBIOTIC ASSEMBLAGES AT A REGIONAL SCALE.

M.-J. DOLE-OLIVIER, F. MALARD and J. GIBERT

University Claude Bernard of Lyon 1, UMR CNRS 5023, Laboratoire d’Ecologie des Hydrosystèmes Fluviaux 43, Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France. ([email protected]), ([email protected]), ([email protected]).

ABSTRACT

The relationships between the structure of stygobiotic assemblages and environmental / palaeogeographic attributes of ground waters were explored by using biological (presence Ð absence of 62 species) and environmental data (16 variables) collected in the Meridional Jura (269 sites distributed among 4 hydrogeographic basins). An OMI (Outlying Mean Index) analysis was performed to identify the main factors driving community structure and to evidenciate the ecological preferences of taxa along environmental gradients. The average marginality of all taxa was highly significant, indicating the strong influence of pore size and historical determinants (i.e., distance to the Würm glacier). Almost all taxa had their centroids displaced towards the most permeable geological formations (first axis of the analysis), i.e., the karst aquifer and coarse alluvium, also characterized by a high oxygen content. Stygobiotic assemblages were enriched both with increased distance to the Würm glacier and decreasing elevation (second axis of the analysis). Land use variables and physicochemical

parameters (Mg, Ca, Specific conductance, PO4, NO3-N) had low influence on the composition of the stygobiotic assemblages of the Meridional Jura.

KEYWORDS : ENVIRONMENTAL GRADIENTS, STYGOBIONTS, NICHE ANALYSIS, PRESENCE- ABSENCE DATA, MERIDIONAL JURA

1. INTRODUCTION simultaneous collection of stygobiotic species in One important issue for groundwater biodiversity the same set of sites. Sites were selected and assessment and the subsequent research in spatially distributed following the standardized optimizing sampling strategies, is the knowledge sampling protocol described within the of environmental variables driving the PASCALIS framework (Malard et al. 2002). The composition of stygobiotic assemblages. For initial protocol (192 sites) was extended to a total historical reasons, this question was studied and of 296 sites, which were distributed among 4 documented separately in karst and porous different habitats, (1-unsaturated zone and 2- habitats so that we are unaware of any complete saturated zone in the karst, 3-hyporheic zone investigation comprising both habitats and and 4-phreatic zone in porous habitats) within reporting on the main environmental variables at four hydrogeographic basins (Suran, Albarine, a regional scale (Gibert et al., 1994). The Oignin and Valouse rivers). The faunal data set objective of the work was to determine the main contained presence/absence of species within environmental factors driving the stygobiotic each site. The environmental data set gathered assemblages in the Meridional Jura, and to sixteen variables related to habitat (1- elevation, 2- geology, 3- hydraulic conductivity), water identify the ecological preferences of species. mineralisation (4- pH, 5- specific conductance, 6- This observational approach was developed Dissolved oxygen, 7- Calcium, 8- Magnesium, 9- within the framework of the European PASCALIS Nitrates, 10- Phosphates), land cover (11- program (Protocols for the ASessment and artificial lands, 12- coniferous and mixed forests, Conservation of Aquatic LIfe in the Subsurface, 13- deciduous forests, 14- meadows, 15- Gibert, 2001) in the Meridional Jura. agriculture) and history (16- distance to the Würm glacier). 2. METHODS The relationships between environmental The sampling strategy involved the variables and the composition of stygobiotic measurement of environmental variables and the assemblages were investigated by matching

SWSB December 2004 79 SESSION 3 : BIODIVERSITY PATTERNS these two tables through a niche analysis (OMI: representing the distribution of sites according to Outlying Mean Index, Doledec et al., 2000). In their coordinates on F1 and F2 (Figure 1B). On this OMI analysis the position of each species this graph, karstic sites have positive values on depends on their niche deviation (marginality) the axis 1, when clay, glacial till and colluvium from a reference, which corresponds to a have negative values. Medium alluvium and theoretical ubiquitous species, i.e., a species coarse alluvium have an intermediate position. uniformly distributed along the environmental Thus, the first axis described a gradient linked to gradient. This analysis is suited for the geology, from highly permeable aquifers that investigation of multidimensional niche breadths, drain well oxygenated water to very low in the case of strong driving forces and in finding permeable formations containing poorly the most important ecological factors. A Monte- oxygenated water. The second axis of the OMI Carlo permutation test was used to check the analysis represented 20% of the total variability. statistical significance of the marginality for each On this axis, the distance to the Würm glacier, taxon as well as the average marginality of all which is negatively correlated with elevation is taxa. Analysis and graphical displays were also a significant parameter (Figure 1A). This is performed using the ADE software (Thioulouse clearly illustrated by the factorial graph et al., 1997). representing the distribution of sites according to their coordinates on F1 and F2 (Figure 1C). On 3. RESULTS AND DISCUSSION this graph, low elevation sites (200-400 m) have A total of 62 species were collected through this positive values, when high elevation sites (600- sampling strategy (Table 1), from which fifteen 800 m) have negative values. Medium elevation species were probably new to science (labelled sites (400-600 m) have an intermediate position. “sp.J1, sp.J2…”). Six others were also new for Thus, the second axis described a gradient this region (Parabathynella cf. stygia, Crangonyx linked to distance to the Würm glacier, from high sp., Bogidiella albertimagni, Proasellus altitude sites situated within the glacier, and to synaselloides, Microcharon reginae, Siettitia low elevation sites situated far from the limits of avenionensis), underlying the efficiency of the ancient glaciated areas. sampling protocol that involved all kinds of Land use variables (Coniferous and mixed groundwater habitats. forests, deciduous forests, Meadow, Agriculture, 3.1 Environmental variables Artificial lands) and physicochemical parameters

The first axis of the OMI analysis represented (Mg, Ca, Specific conductance, PO4, NO3-N) 52% of the total variability. On this axis, geology, were unimportant for the definition of the which is negatively correlated with dissolved environmental gradient also indicating their low oxygen, is the most significant parameter (Figure influence on the composition of stygobiotic 1A). This result is illustrated by the factorial graph assemblages at this scale.

Table 1 : List of species collected in the Meridional Jura, following the sampling protocol defined in PASCALIS framework. The abbreviations assigned to the species is used in Fig 1D.

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Figure 1 : Main results of the OMI analysis. A- canonical weights of the 16 environmental variables along axes 1 and 2. B & C- Distribution of sites according to their coordinates on F1 and F2: B- sites are classified within 5 classes of geology; C- sites are classified within 6 classes of elevation. D- Distribution of species according to their coordinates on F1 and F2; the significance of abbreviations is given in Table 1.

3.2 Ecological gradient exclusively in low permeable formations and The great majority of species were displaced poorly oxygenated waters. In the contrary Niphargus foreli (NpFor), Bryocamptus sp. towards the positive values on F1 (Figure 1D), (BrJ1), Speocyclops sp. (SpJ2 and Sp.J3), corresponding to highly permeable geological Elaphoidella phreatica (ElPh) and Moraria formations (limestone and coarse alluvium) and (Moraria) sp. (MMJ1) were only found in well well oxygenated water. The most significant oxygenated habitats and highly permeable species for defining this gradient are the aquifers. Copepoda Ceuthonectes serbicus (CeSe), Most of the species were also preferentially Eucyclops graeteri (EuGr) and Speocyclops collected in low elevation sites situated far from sp.J3 (SpJ3), the Ostracoda the Würm glacier (positive values on F2, Figure Fabaeformiscandona breuili (FaBr) and 1D). The most significant species to define this Schellencandona sp.J1 (ScJ1), the Isopod altitudinal gradient were the Syncarida, Proasellus cavaticus (PrCa) and the Amphipod Parabathynella cf. stygia (Pab2S), the copepoda Niphargus rhenorhodanensis (NpRh). One Attheyella sp.J1 (AAJ1), the Amphipoda species, Niphargus kochianus (NpK), occurred Salentinella juberthieae (SaJu), Crangonyx sp.

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(Cr1) and Niphargopsis casparyi (NpCa), the Protocols for the ASsessment and Conservation Isopoda Proasellus walteri (PrWa) and the of Aquatic Life In the Subsurface, (contract no Oligochaeta Trichodrilus cf. serei (Tr2S). None of EVK2-CT-2001-00121) of the Fifth Research and the species was restricted to high elevation sites Technological Development Framework Program but a set of species associated with long of the European Community. distances to the glacier limits and to low altitudes. In this case were found Proasellus walteri, Haber BIBLIOGRAPHY turquini, Salentinella juberthieae, Siettitia Gibert, J. (Coordinator) 2001. Protocols for the avenionensis, Parabathynella cf. stygia, and ASsessment and Conservation of Aquatic Life In the Microcharon reginae. These species are Subsurface (PASCALIS): a European project. In common-dwellers of the Rhône river aquifer Mapping subterranean biodiversity. Cartographie de (Dole-Olivier et al., 1994) and its main tributaries. la biodiversité souterraine. Culver D.C., Deharveng L., Gibert J. & Sasowsky, I. [Eds.], Karst Waters Institute, Special Publication 6:19-21. 4. CONCLUSION Gibert, J., J.A. Stanford, M.-J. Dole-Olivier, and J.V. Ward. The major environmental parameters driving 1994. Basic Attributes of groundwater ecosystems stygobiotic assemblages at the scale of the and prospects for research. Groundwater Ecology, J. Meridional Jura were geological attributes versus Gibert, D.L. Danielopol & J.A. Stanford [Eds.], pp. 7- dissolved oxygen and elevation versus distance 40. to the Würm glacier. The great majority of Dole-Olivier, M.-J., P. Marmonier, M. Creuzé des species were displaced towards highly Châtelliers, and D. Martin. 1994. Interstitial fauna permeable geological formations (karst and associated with the alluvial floodplains of the Rhône river. Groundwater Ecology. J. Gibert, Stanford, J.A., coarse alluvium) draining well oxygenated Danielopol, D.L., Academic Press: 313-346. waters. Species were successively added as Dolédec, S., Chessel, D., and C. Gimaret-Carpentier. elevation decreases and distance to the maximal 2000. “Niche separation in community analysis: a new extension of the glacier during the Würm, method.” Ecology 81(10):2914-2927. increases. These results would be helpful for the Malard, F., Dole-Olivier, M.-J., J. Mathieu, and F. Stoch. spatial modelisation and prediction of occurrence 2002. Sampling manual for the assessment of of the stygobiotic species at a regional scale and regional groundwater biodiversity. European Project also for improving the existing sampling PASCALIS, available at http://www;pascalis- strategies. project.com. Thioulouse, J., S. Dolédec, D. Chessel, and J.-M. Dole- Olivier. 1997. ADE-4: a multivariate analysis and graphical display software. Statistics and Computing ACKNOWLEDGEMENTS 7:75-83. This work was financially supported by the European research project PASCALIS :

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IMPROVING THE ASSESSMENT OF GROUNDWATER BIODIVERSITY BY EXPLORING ENVIRONMENTAL HETEROGENEITY AT A REGIONAL SCALE.

F. CASTELLARINI, M.-J. DOLE-OLIVIER, F. MALARD and J. GIBERT University Claude Bernard of Lyon 1, UMR CNRS 5023, Equipe Hydrobiologie et Ecologie Souterraines, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France. ([email protected]), ([email protected]), ([email protected]), ([email protected]).

ABSTRACT The assessment of groundwater biodiversity has the inherent and recurrent problem related to the accessibility of the groundwater environment. This study explored potential sources of environmental heterogeneity to improve sampling strategies developed for the assessment of the stygobiotic species richness. Thus random and stratified strategies were analyzed for 247 sites in the southern Jura (France). Decentred Correspondence Analysis (DCOA), Principal Component Analysis (PCA) and correlation analysis showed significant relationships between the species richness patterns and the main sources of heterogeneity (habitat structure, elevation and historic events).

1. INTRODUCTION glaciers, Buoncristiani & Campy, 2004). At present it is an upland forested massif of Although ground waters are widely exploited for karstified Jurassic limestone. Four surface human use, they are amongst the most poorly hydrogeographic basins, the Suran, Albarine, known ecosystems because of their hidden Oignin and Valouse rivers were selected for this character and their relative inaccessibility study. Each basin contained several groundwater (Marmonier et al. 1993, Malard et al. 1994). In recent years, efforts have been done to improve aquifers occurring in two general geologic the sampling techniques for collecting settings: the karstic-consolidated (K) and the groundwater fauna (Malard et al. 1994, 1997; porous-unconsolidated (P) sediments. Samples Mauclaire et al. 1998, Scarsbrook and Halliday, were taken following a hierarchical stratified 2002) or to determine a sampling strategy sampling (Malard et al., 2002), which was carried (sample volume, number of samples…) for its out using three strata according to habitat evaluation at a local scale (Hunt and Stanley structure. Here habitat structure refers to the size 2000; Boulton et al. 2003, 2004). Thus, the of the voids and their hydrological search for a methodology to assess groundwater connectedness. The first stratum contained the biodiversity at a larger scale becomes the basins; the second, the aquifer types: the karst following and challenging task, especially to (K) and the porous (P) aquifers and the third, the define conservation priorities. hydrogeologic zones: the unsaturated karst (uk, n=49), the saturated karst (sk, n=54), the The main goal of this study was to provide some hyporheic (h, n=44) and the phreatic zones (p, general indications for optimal sampling for the n=100). Samples were taken wherever it was assessment of ground water biodiversity on a accessible, according to the methodology regional scale. For this, the heterogeneity of the described in Malard et al. (2002). habitat structure was explored as a starting point Presence/absence data were used as estimate for the stratification scheme and definition of the hierarchical units to be sampled. of the species richness. We considered that species richness roughly represents the exhaustive biodiversity of the area because the 2. MATERIAL AND METHODS number of sampling sites (N=247) represents an 2.1 Study area intensive effort. The sampling area for this study, which covered A Decentred Correspondence Analysis (DCoA, a surface of 1,272 km2, was located in the Dolédec et al. 1995) was used to assess the southern Jura, eastern France. The Jura is a low effect of habitat structure on the species perialpine mountain range which was covered in richness. Although DCoA allows both the spatial- its eastern and southern parts by a local ice temporal heterogeneity and the heterogeneity of sheet during the Quaternary (Riss / Würm sampling effort to be taken account, the analysis

SWSB December 2004 83 SESSION 3 : BIODIVERSITY PATTERNS was done by sub-sampling (N total=160). For potential and additional sources of variation in this, 10 replicates, the most spatially separated species richness (i.e. altitudinal gradient, last sites within each stratum, were taken. The main glacial event), Principal Component Analysis purpose of this selection was to reduce the (PCA) and correlation were calculated. These spatial autocorrelation between sites, which is two analyses were carried out on the sub- imposed by the accessibility to the groundwater sampling above, correlating the position of each environment, and to cover the maximum area. To site in one dimension of PCA with the altitudinal optimize the separation between groups a gradient and the last glacial event. All analyses Between and Within analysis (Dolédec and were done using ADE4 software (Thioulouse et Chessel 1989) was computed. To explore al. 1997).

Niphargus rhenorhodanensis (1) Diacyclops cf. belgicus (2) Avenionia (3) Bythiospeum (4) Pseudocandona zschok k ei (5) Fabaeformiscandona breuili (6) Ceuthonectes serbicus (7) Islamia sp. (8) Eucyclops graeteri (9) Niphargus virei (10) Niphargus indet. (11) Attheyella (A) sp.J1 (12) Schellencandona spJ1 (13) Elaphoidella phreatica (14) Parabathynella cf. stygia (15) Proasellus cavaticus (16) Salentinella juberthieae (17) Speocyclops spJ.3 (18) Niphargopsis casparyi (19) Proasellus "non walteri" (20) Diacyclops languidoides (21) Rhyacodrilus balmensis (22) Cryptocandona k ieferi (23) Moraria (M.) sp.J1 (24) Graeteriella unisetigera (25) Fabaeformiscandona wegelini (26) Parastenocaris sp J1 (27) Spiralix sp. (28) Cavernocypris subterranea (29) Niphargus k ochianus (30) Islamia minuta (31) Proasellus valdensis (32) Niphargus fontanus (33) Schellencandona triquetra (34) Parastenocaris glareola (35) Alona phreatica (36) Caecosphaeroma virei (37) Microcharon reginae (38) Schellencandona insueta (39) Parastenocaris sp. J2 (40) Schellencandona sp J3 (41) Nitocrella gr. hirta sp. J1 (42) Acanthocyclops sensitivus (43) Bryocamptus sp. J1 (44) Speocyclops sp.J1 (45) Pseudobathynella sp.J1 (46) Proasellus walteri (47) Bogidiella albertimagni (48l) Niphargus forelii (49) Graeteriella cf. boui (50) Speocyclops sp.J2 (51) Gianus cavealis (52) Siettitia avenionensis (53) Proasellus synaselloides (54) Crangonyx indet. (55) Schellencandona spJ4 (56) Speocyclops indet. (57) Speocyclops sp.J4 (58) Bythinella (59) Spiralix vitrea (60) Haber turquini (61) Trichodrilus cf. serei (62) 0102030405060708090100 110 120 Absolute frequency

Figure 1 : Stygobiont absolute frequency of occurrence in the southern Jura, France (N=247). J* and letters in brackets mean that they are new or undetermined species.

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(a) 60

0 Karst Porous (b) 45

Number of Species 40 35 30 25 20 15 10 5 0 sk uk h p Total Exclusive

Figure 2 : Occurrence of stygobionts in the different strata of the study area. Ntotal = 62 species. sk: saturated karst, uk: unsaturated karst, h: hyporheic and p: phreatic zone.

3. RESULTS stratum, on the other hand, shows a spatial A total of 62 stygobionts (strict groundwater heterogeneity along the second factorial axis species) were identified (Fig.1). The two most (F2) with three communities, the first belonging frequent species (Niphargus rhenorhodanensis to the Suran basin, the second to the Valouse and Diacyclops cf. belgicus) were collected basin and the third to the Albarine and Oignin within 95% and 60% of the sites, 50% of species basins. had a very low frequency of occurrence (< 5%, Species composition of the clusters shows that hereafter called rare species) and 16% of this heterogeneity is mainly due to low-frequent species were found only once. Stygobionts are and exclusive species in the case of G1 and G3 totally absent in 4 sites only. No marked clusters, exclusive in the case of G4 and rare, differences in species richness were observed high-frequent and exclusive in G2 and G5 between karstic and porous strata (43 and 45 (Figures 2 and 3b). species respectively). More than 60% of species Although less conspicuous, the same were shared between both of them but more than configuration was found in the PCA. Correlation 24% and 29% of the species were exclusive to between values of each site along the axis 2 of the karstic and porous strata respectively. Less PCA and the last glacial event (Würm glacier), than 16% were exclusive species of the uk, sk, h and between axis 2 and the altitudinal gradient and p strata (Fig.2). were significant but low (r = —0.54 and r = 0.43, 3.1 Sampling strategy and effort P< 0.05 respectively). Total inertia of the DCoA (11.16) was explained 4. DISCUSSION by the “basin effect” linked with the effect of the Karstic and Porous aquifers, hereafter called Successional communities usually change from Habitat-Basin effect (Hab-Bas). The result was a earlier stages (i.e., pronounced dominance and configuration of five clusters (G1-5, Fig.3a), in low species richness) to later stages (reduced which 61% of total inertia was explained by the dominance, high species richness, and a large between groups effect (clusters) and the rest by number of rare species, May 1981). The the within groups effect (uk, sk, h and p stygobiotic species richness pattern of the hydrogeologic zones). Regarding the gradient southern Jura suggest that the community is in a along the first factorial axis (F1), three clusters mature stage with rare species (<5% of (G1, G2 and G4) represent communities occurrence) related to their habitat specificity belonging to the karstic strata and the other two (“exclusive species”, Figures 1, 2 and 3b) but not (G3 and G5) to the porous strata. The porous necessarily to their endemism (see concepts in strata shows moreover, two clusters, one Gaston 1997). Such is the case, for example, of clustering stygobionts of Oignin, Suran and Haber turquini (61) and Proasellus synaselloides Valouse basins and the other representing (32) which are rare and exclusive to the uk and p stygobionts of the Albarine basin. The karst habitats respectively, or Niphargus fontanus (3)

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a b F2 51 57 F2 58 61 uk 45 46

G1 10 37 Suran

15 35 22 12 53 54 34 16 59 4 2340 52 47 sk 39 17 60 50 2 11 30 55 19 2736 33 38 7 246 31 3 48 62 5 26 13 21 41 1 F1 18 8 43 9 25 32 28 2029

44 14 G2 56 49 sk Valouse h h h p uk Valouse Albarine Suran F1 p p G3 h p Oignin

G4 G5 uk sk

Albarine Oignin

uk sk

Figure 3 : Results of the Decentred Correspondance Analysis. a) Spatial configuration of clusters (G1-G5) within karstic and porous strata,

Nsites= 160. b) Distribution of Stygobiotic species in the strata, (see code of species in Fig. 1).

which is rare and exclusive to the porous assumption that habitat structure could be a stratum. Nevertheless all of them are not source of heterogeneity, at least as regards endemic and may be also found in other basins differences in the species composition between like the lowest part of the Rhône river basin (P. the karst and porous strata. Nevertheless, they synaselloides Botosaneanu, 1986), high Rhône also suggested that other sources of variation basin (H. turquini, Dole 1994) or with a wide and within the karst and the porous strata may cause dispersed distribution in France (N. fontanus, environmental heterogeneity. In this way, within Ginet 1996). At the same time the species the porous stratum, along the positive side of F1 richness pattern also shows rare and endemic axis (Fig 3a), the cluster G5 results from the species as Schellencandona sp J4(56), isolation of three regional basins, which contain Speocyclops spp J1(45), J2(51), J3(18) and the fauna of the Jura region (hereafter called J4(58) and Pseudobathynella sp J1(46) which “Jurassic fauna”), while the cluster G3 results are still not described. Although all of these rare from the co-occurrence of two different regional species (50% of the total) may not be very fauna. This co-occurrence may be explained by important for studies of functional ecology, they the connection between the Albarine basin, become an important component in studies of belonging to the Jura region, and the Rhône and biodiversity because rare species often Ain basins, belonging to the Mediterranean determine the differences between communities fluvial corridor which allows the colonization of (Cao et al. 1998). Mediterranean species such as Microcharon Results of the DCoA give insights as much on the reginae (38), Schellencandona triquetra (34), sources of heterogeneity acting on groundwater Salentinella juberthieae (17) and Niphargus communities as the similarities and differences fontanus (33) towards the Jura region giving thus between these communities. In fact, the relative a cluster with fauna of two different regions (Fig position of clusters confirmed our starting 3b).

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On the other hand, the position of clusters within the manuscript. Taxonomical experts identified the Karst stratum along the F2 axis (Fig 3a), and stygobiotic species, N. Coineau (Syncarida), M. results of PCA and correlation analysis suggest & G. Falkner (Mollusca), F. Fiers (Cyclopoida), D. the karstic habitats are affected by two negatively Galassi (Harpacticoida), R. Ginet (Amphipoda), correlated variables, the altitude and the distance J. Juget (Oligochaeta), G. Magniez (Isopoda), P. to the limit of the Würm glacier (last glacial Marmonier (Ostracoda) and B. Sket (Hirudinae). event). In this way the G1 cluster includes sites that have not been affected by the ice sheet, REFERENCES most of them also being in altitudes lower than Addo-Bediako, A., S. Chown, and K. Gaston. 2000. 400 m; G2 includes sites located at intermediate Thermal tolerance, climatic variability and latitude. altitudes (300-600 m) mostly without effect of the Proc. R. Soc,. London, 267:739-745. glacier and finally, G4 includes sites mostly Botosaneanu, L. 1986. Stygofauna mundi, E.J. Brill/Dr W. located in the higher altitudes (> 400 m) and Backhuys, Leiden.Brose, U, N. mostly affected by the glacial event. Apart from Boulton, A., M.-J. Dole-Olivier, and P. Marmonier. 2003. the well-known relationships between altitude Optimizing a sampling strategy for assessing and other variables, i.e. oxygen and temperature, hyporheic invertebrate biodiversity using the Bou- which can play an important role in the Rouch method: within-site replication and sample volume. Archiv für Hydrobiologie 156: 431-456 physiology of organisms and thus, in their limits Boulton, A., M.-J. Dole-Olivier, and P. Marmonier. 2004. of distribution (Addo-Bediako et al. 2000; Issartel Effects of sample volume and taxonomic resolution on et al. 2005), the distance to the last glacial event assessment of hyporheic assemblage composition introduces biogeographical hypothesis to explain sampled using Bou-Rouch pump. Archiv für the distribution of the karstic groundwater fauna Hydrobiologie 159:327-355 in the Jura (Ginet and Juberthie 1987). In this Buoncristiani, J. and Campy, M. 2004. Expansion and way it is possible to explain the relative position retreat of the Jura ice sheet (France) during the last of clusters along the second axis in Fig 3a and glacial maximum. Sedimen Geol, 165:253-264. the occurrence of endemic species such as Cao, Y., D. Williams, and N. Williams. 1998. How Speocyclops spp J1(45), J2(51) and J4(58) and important are rare species in aquatic community Pseudobathynella sp J1 (46) in the G1 cluster; ecology and bioassessment Limno Oceanogr, 43:1403-1409. species in the G2, represented by the “Jurassic Dolédec, S. and Chessel, D. 1989 Rythmes saisonniers fauna” similar to that found in the porous strata et composantes stationnelles en milieu aquatique II- (G5), and finally the G4 cluster with exclusive Prise en compte et élimination d’effets dans un species such as N. foreli (49), Cavernocypris tableau faunistique. Acta Îcologica, Îcologia subterranea (29) and Elaphoidella phreatica (14), Generalis 10, 3, 207-232. (Figures 1 and 3b). Dolédec, S., D. Chessel, and J.-M. Olivier. 1995. l’Analyse des correspondances décentrée: application aux 5. CONCLUSIONS peuplements ichtyologiques du Haut-Rhône. Bull Fr Pêche Piscic 336:29-40. The 62 stygobionts collected in the southern Jura Dole-Olivier, M.-J., P. Marmonier, M. Creuzé des represent a rich set of species for such a Châtelliers, and D. Martin. 1994 Interstitial fauna constraining environment, being larger than associated with the alluvial floodplains of the Rhône previously sampled in the alluvial sediments of river. Groundwater Ecology. J. Gibert, Stanford, J.A., the Upper Rhône river (Dole-Olivier et al., 1994) Danielopol, D.L., Academic Press: 313-346. or in the specious karstic area of the northern Gaston, K. 1997. What is rarity? In The Biology of Rarity. region of Montpellier (Malard et al., 1997). Edited by W. Kunin and K. Gaston. Chapman and Secondly, on a regional scale we suggest the use Hall, London. pp 30-47. of habitat structure, with the dichotomy Ginet, R., and C. Juberthie. 1987. Le peuplement animal karst/porous as a stratum plus the elevation des karsts de France. Eléments de biogéographie souterraine pour les invertébrés. Première partie: la gradient and a biogeographical factor as major fauna aquatique. Karstologia 10:43-52. sources of additional heterogeneity. Ginet, R. 1996. Bilan systématique du genre Niphargus en France, Société Linéenne de Lyon. ACKNOWLEDGMENTS Hunt, G.W. and Stanley, E.H. 2000. “An evaluation of This work and the fellowship of FC were alternative procedures using the Bou-Rouch method financially supported by the European for sampling hyporheic invertebrates.” Can J Fish Commission’s PASCALIS (Protocols for the Aquat Sci 57:1545-1550. Assessment and Conservation of Aquatic Life In Issartel, J., F. Hervant, Y. Voituron, D. Renault, and P. the Subsurface) project (Contract: EVK2-CT- Vernon. (in press) Behavioural, ventilatory and respiratory responses of epigean and hypogean 2001-00121), under the EU 5th Framework crustaceans to different temperatures. Comp Biochem Programme: Global Change, Climate and Phys A. Biodiversity. Authors wish thank T. Lefébure, D. Malard, F., J. Gibert, R. Laurent, and J.-L. Reygrobellet. Ferreira, D. Martin and L. Vitry for Field 1994. A new method for sampling the fauna of deep assistance; and J.-M. Olivier for comments on karstic aquifers. C. R. Acad. Sc. 317:955-966.

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Malard, F, J. Gibert and R. Laurent. 1997. L’aquifère de la comparison of coring and pumping techniques. Arch. source du Lez: un réservoir d’eau et de biodiversité. Hydrobiol. 142:111-123. Karstologia 30:49-54. May, R.M. 1981. Patterns in multi-species communities. Malard, F., J.-L. Reygrobellet, R. Laurent, and J. Mathieu. In Theoretical Ecology. Edited by R.M. Blackwell 1997. Developments in sampling the fauna of deep Scientific Publications, Oxford, UK. pp. 197-227 water-table aquifers. Arch. Hydrobiol. 138, 3: 401- Orghidan, T. 1955. Un nou domeniu de viata acvatica 432. subterana : “biotopul hiporeic”. Buletin Stiintific sectia Malard, F., M.-J. Dole-Olivier, J. Mathieu, and F Stoch. de Biologie si stiinte Agricole si sectia de Geologie si 2002. Protocols for the Assessment and Conservation Geografie VO VII(3): 657-676. of Aquatic Life In the Subsurface. Sampling Manual. Scarsbrook, M.R. and Halliday, J. 2002. “Detecting published within the framework of the EU Project patterns in hyporheic community structure: does PASCALIS. Available from (http://www.pascalis- sampling method alter the story.” New Zeal J Mar project.com/home/innovation.html). Fresh 36:443-453. Marmonier, P., P. Vervier, J. Gibert, and M.-J. Dole-Olivier. Thioulouse, J, D. Chessel, S. Dolédec, and J.-M. Olivier 1993. Biodiversity in ground waters. Tree 8, 11, 392- 1997. ADE-4: a multivariate analysis and graphical 395. display software. Stat Comput 7:75-83. Mauclaire, L, P. Marmonier, and J. Gibert. 1998. Sampling water and sediment in interstitial habitats: a

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MAPPING THE STYGOFAUNA OF THE STATE OF BADEN-WÜRTTEMBERG, SOUTHWEST GERMANY

H.J. HAHN and A. FUCHS

Universität Koblenz-Landau, Campus Landau, Abt. Biologie, Arbeitsgruppe GrundwasserökologieIm Fort 7, 76829 Landau, Germany. ([email protected]), ([email protected]).

ABSTRACT

The stygofauna of the state of Baden-Württemberg was mapped by sampling 304 bores, which were representative of the state`s geography, hydrogeology and hydrography ; 106 species were reported, among them 74 stygobiontic and stygophilous ones. Some species of stygofauna were found to be relict forms, which reflect the Pliocene river systems. Many of these species are seldom and due to their rareness and isolation potentially endangered - an aspect that should be considered for future protective measures. The hydraulic conductivity of the aquifer was identified as the main factor shaping stygofaunal communities. The fauna of non-alluvial aquifers is considered to be more suitable for the characterisation of regional stygofaunal communities than the fauna of alluvial or karstic aquifers. The large scale distribution of the stygofauna was in accordance with the geographic situation, but was supposed to be mainly the result of the different proportions of the different hydraulic groups within the groups of natural geographic regions (naturraums).

KEYWORDS : BADEN-WÜRTTEMBERG, LARGE SCALE DISTRIBUTION, NATURRAUM, REGIONAL MODELS OF STYGOFAUNA

1. INTRODUCTION communities, and spring fauna seem to be not representative of the fauna of the adjacent Information on large scale distribution of aquifers (Hahn & Matzke 2005). stygofauna is scant, and data on large scale distribution patterns are nearly lacking. With the However, the Landesanstalt für Umweltschutz exception of Karst, most studies on stygofauna Baden-Württemberg (LfU, State Institute for are focused on alluvial aquifers, while nearly no Environmental Protection Baden-Württemberg) information is available on non-alluvial and supported this study to describe regional fractured rock aquifers. Furthermore, regional stygofauna communities, and to find out by which models of stygofauna communities, analogous to factors (e.g. geography, hydrogeology, surface waters (s. EU Water Framework hydrography and hydraulic conductivity of the Directive), are yet lacking. Although such aquifer) influence the patterns found. Preliminary regional models will probably not be required by results are presented here. the new EU Groundwater directive, which is still under discussion, they can help to assess 2. STUDY AREA AND METHODS groundwater habitats on an ecological basis and The state of Baden-Württemberg covers an area provide information for future protective of 37,750 km2 and consists of 11 hydrogeological measures. units and 66 naturraums, which are aggregated In the state of Baden-Württemberg, South- to 13 groups of naturraums (Fig. 1) like the Black Western Germany, only one study (Steenken Forest (6 naturraums) or the Southern Upper 1998) tried to describe the stygofauna of two Rhine Plateau (5 naturraums). natural geographical regions, the so-called Baden-Württemberg belongs to two major river naturraums, by sampling 23 bores and 11 basins: The Rhine system covers most of the springs. Given a number of 66 naturraums on the area, while the River Danube and its tributaries state`s territory and the high heterogenity of are restricted on the south eastern part of the groundwater habitats, such a small number of state). In the Pliocene period, the catchment of sites does not provide sufficient data to the River Rhine was smaller than today with its characterize typical regional stygofaunal origin situated around 150 km north of its present

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Figure 1 : Map of the state of Baden-Württemberg. Dots indicate the bores sampled, shading patterns the different groups of naturraums.

source, while the size of the River Danube basin 3. RESULTS AND DISCUSSION was larger than today (Fig. 2). It stretched much 3.1 Distribution of selected species more westwards until the Lake Léman comprising the uppermost parts of the present In the study area, 106 species were found, Rhône River catchment, and northward. The among them 74 stybiontic and stygophilous present Rhine catchment north of Strasbourg, ones. Most of these species occurred in the the present Upper Neckar River catchment and whole area, but around 15 stygobites were restricted in their spatial distribution. significant parts of the upper Main catchment also belonged to the Pliocene River Danube Due to the climatic situation in the quarternary in basin. So, in the Quartenary, the River Rhine the mountainous areas (“Mittelgebirge”) of took over large parts of the former upper River Central Europe, we supposed most groundwater Danube system. species to be widely spread. In the ice age, the mountainous areas were not covered by glaciers, In the study area, 304 monitoring bores (Fig. 1), but the climate was very cold and dry. This lead which were supposed to reflect the geographical, to the extinction of many species including many hydro geological and hydrographical conditions, endemites, even in the groundwater. As a result were sampled twice for fauna and water between of recolonisation at the end of the ice age most November 2001 and December 2002. Sampling species are widely spread, today. For details see of the fauna was carried out by using a so-called Thienemann (1950). But it seems that some of net sampler, a modified phreatic net. All the old Tertiary stygofauna have survived, and metazoan taxa except nematodes were that their large scale distribution reflects the determined on species level. Pliocene hydrography.

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Figure 2 : River sytems of Rhine and Danube in the Pliocene and today.

All species with a restricted spatial distribution, Parastenocaris psammica and Nitocrella omega. with the exception of the cyclopoid Other species of the Pliocene Rhine catchment Acanthocyclops gmeineri, were found in the are Niphargellus nolli and Bathynella Rhine catchment. Acanthocyclops gmeineri freiburgensis (not displayed here), which were occurred exclusively in the Danube catchment also found mainly in the central and northern part and was considered formerly as an endemic of the Upper Rhine Plateau and in the lower species of the alluvial groundwater of the River Neckar catchment. Danube near Vienna. This finding obviously The Pliocene Danube system comprised parts of reflects the fact, that the River Rhine took over the present upper Rhône system, the upper large parts of the upper Danube catchment Ð Rhine system, the upper Neckar system and the together with most of its stygofauna. upper Main system. As a consequence, we Furthermore, most of the species with a expected and found a Danube-Rhine-Rhône restricted distribution are somewhat rare and fauna as described firstly for some fish and must be considered as relictual forms of the old groundwater fauna by Thienemann (1950): The Pliocene river systems. Both rareness and amphipod Niphargopsis casparyi was found in relictual character are indicative of their potential the present Danube catchment, in the endangerment. As a consequence, it is urgently groundwater of the present southern Upper required to handle such species as subjects of Rhine Plateau, of the upper Neckar River protective measures. catchment and of the Main River catchment. It is In the Pliocene, the spring of the Rhine river was also reported from the upper Rhône catchment situated near todays Strasbourg, 150 km north of (Fig. 3 B). The same applies with both the its present position. Species of the old Rhine cyclopoid Acanthocyclops sensitivus and the system were thus expected, and found, in the isopod Proasellus slavus. These three species groundwater of the central and northern part of were also found at a few sites in the lower the Upper Rhine Plateau and in the lower Neckar Neckar catchment and in the central part of the catchment (Fig. 3 A). These species are the Upper Rhine catchment, indicating a moderate harpacticoids Parastenocaris germanica, migration since the Pliocene.

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Figure 3 : Distribution of selected species in the study araea. A : Species of the Pliocene Rhine system : Nitocrella omega, Parastenocaris germanica, Parastenocaris psammica. B : Species of the Pliocene Danube system : Acanthocyclops sensitivus, Niphargopsis casparyi, Proasellus slavus. Black lines indicate the Pliocene watershed between the Danube and the Rhine River. Grey design indicates the present Rhine catchment, dotted the present Danube catchment.

Figure 4 : MDS of the groups of naturraums of Baden-Württemberg using the species abundances aggregated by means. The designs of the dots correspond with the designs of the map, which indicate the different groups of naturraums.

3.2 Large scale distribution patterns of communities. To test this hypothesis, faunal stygofauna communities abundance data were aggregated by means over Corresponding to aquatic surface fauna, we the groups of naturraums and ordered by an expected for the stygofauna regional MDS using Bray-Curtis dissimilarities. The MDS

92 SWSB December 2004 SESSION 3 : BIODIVERSITY PATTERNS plot (Fig. 4) reflects well the geographical characterisation of regional stygofaunal situation of the groups of naturraums: The communities the fauna of non-alluvial aquifers is northern, central and southern Upper Rhine more suitable than the fauna of alluvial or karstic Lowlands and the Hochrhein Area were ordered aquifers. in accordance with the map from North to South There could be two reasons why the fauna of as well as the Swabian Alb, the Donau-Iller-Lech alluvial and karstic aquifers was so similar: Both Area and the Prealpine Hill and Swamp Area. types of aquifers are open systems allowing for a Between them, and corresponding to the map, migration between different aquifers and thereby the Neckar-Tauber-Gäu Area and the Swebian mixing their fauna, and both aquifer types are Keuper-Lias Country were ordered. The Black strongly influenced by surface water. Forest was situated in the Southwest and the It seems that the reflection of the geographical Odenwald Mountains in the North Ð also in situation by the fauna is mainly the result of the congruence with the map. different proportions of the different hydraulic As the main factor influencing the distribution of groups within the groups of naturraums, and this stygofauna, the hydraulic conductivity of the is again an argument to focus more on the fauna aquifer was indentified by using the PRIMER procedure of BIOENV. Four groups of aquifers BIBLIOGRAPHY could be distinguished: Aquifers with low Hahn, H.J. and Matzke, D. 2005. A comparison of (compact sediments), moderate stygofauna communities inside and outside (unconsolidated/alluvial sediments), high groundwater bores. Limnologica (in press). (fractured rock) and very high (karst) hydraulic Steenken, B. 1998. Die Grundwasserfauna - Ein conductivity. The fauna of all unconsolidated Vergleich zweier Grundwasserlandschaften in Baden- Württemberg. Ecomed Verlag, Landsberg am Lech. sediments and the karstic aquifers was very Thienemann, A. 1950. Die Verbreitungsgeschichte der similar, while the communities of the fractured Süßwassertierwelt Europas. Versuch einer rock and compact aquifers were heterogenous in historischen Tiergeographie. Die Binnengewässer 18, both groups. This finding implies that for the XVI, 1 - 809

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BIODIVERSITY OF BELGIAN GROUNDWATERS : THE MEUSE BASIN

1P. MARTIN, 1C. DE BROYER, 1F. FIERS, 2G. MICHEL, 1R. SABLON and 1K. WOUTERS.

1. Royal Belgian Institute of Natural Sciences, 29 rue Vautier, 1000 Brussels, Belgium. ([email protected]), ([email protected]), ([email protected]), ([email protected]), ([email protected]). 2. Commission Wallonne d’Etude et de Protection des Sites Souterrains (CWEPSS), 21 avenue Auguste Rodin, 1050 Brussels, Belgium. ([email protected]).

KEYWORDS : GROUND WATER, BIODIVERSITY, BELGIUM, MEUSE

The “Walloon karst” is a lowland area made of Science. This is a minimal amount since some limestone, which bisects Belgium in a west- taxa, such as the cyclopoid Acanthocyclops cf. eastern direction and belongs to the catchment venustus, clearly belong to a species complex basin of the Meuse River. This rock-stratigraphic which requires further studies. To date, the total formation contains the most important aquifers in number of stygobitic species amounts to 40, of Belgium. Leruth (1939) was the first to initiate a which 14 species are new for the Belgian fauna. systematic exploration of this subterranean Interestingly, 9 species known from the literature domain and to compile a list of Belgian were not found again in PASCALIS samples. groundwater species. Since then, the knowledge As a rule, the number of occurrences for of the groundwater fauna was not significantly stygobiont species is always exceedingly low. improved so that only 25 stygobitic species are Stygobionts are absent from 37% of sampled known in Belgium to date (Delhez et al., 1999). sites. In addition, more than 50% of species are The launching of the PASCALIS project, of which present in less than, or equal to, 3% of sites. No the “Walloon karst” is a component, offered a statistically significant differences were unique opportunity to fulfil gaps in this knowledge observable in species richness whatever the and to give an overview of groundwater stratification scheme considered. A few species biodiversity in the Meuse basin. are exclusive to one of the considered zone, in The Walloon karst was intensively sampled accordance with their known biology. The same during a field campaign in 2002-2003 according amount of species (26), while of a different to a stratified sampling scheme and a composition, was present in the karst and porous standardized protocol (See this issue, J. Gibert et strata, respectively, and 9 species, in both cases, al.)(Malard et al., 2002). A total of 202 sampling were exclusive to each of the two strata (Fig. 2). sites were selected in four hydrogeographic Most statistical analyses are not significant as a basins belonging to the Meuse Basin, equally possible result of exceedingly low number of shared out among the saturated and unsaturated occurrences for stygobite species. This suggests zones of the fractured medium (karst) and that, in the Walloon karst, the best estimate of among the hyporheic and the phreatic zones of stygobitic species richness can be obtained the porous medium (Fig. 1). Site-specific using a random sampling strategy instead of a environmental attributes were measured in stratified one. However, the possibility that a parallel, in order to test the relationship between stratification pattern could emerge as a result of fauna and environmental parameters and to an increased sampling effort cannot be ruled out. characterize the stygobitic fauna from a historical Species accumulation curves do not reach a and ecological perspective. plateau, suggesting that the total number of More than 173 species were found inhabiting this sampled stations is not satisfactory as regards environment, representative of the Amphipoda, the best estimate of species richness (Fig. 3) ; Cladocera, Copepoda, Hydrachnidia, Isopoda, more samples have to be taken into account in Nematoda, Oligochaeta, Ostracoda and future studies. The total number of stygobiont Tardigrada. Thirty-two stygobite species were species can be estimated as amounting to 36 to identified, of which three, Pseudocandona leruthi 40 species. As such a figure is in good (Klie, 1936) (Crustacea, Ostracoda) Proasellus accordance with the total number of stygobitic hermallensis (Arcangeli, 1938) and Speocyclops species known so far in Belgium, including fontinalis Fiers (2005) (Crustacea, Copepoda) literature data, this suggests that this fauna is are endemic to Belgium, the latter being new for nearly completely known in the Meuse basin. If

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Figure 1 : Map of Walloonia (Belgium) showing the location of sites sampled during the PASCALIS project (circles) and sites mentioned in the literature (triangles). The four hydrogeographic basins considered in this study are surrounded by a quadrilateral. (A): Haute-Meuse; (B): Lesse; (C): Ourthe; (D): Amblève. future studies should provide a better overview of Most taxa were typically more frequent in highly the stygobiont distribution in this basin, the permeable formations, with hard waters, probability of finding new stygobiont species is characterized by moderate to low hydrological low. connectivity. The bulk of stygobite taxa can be Species-environment relationships were considered as ubiquitous species which seem to examined by means of an OMI analysis (Outlying have wide ranges of tolerance for most mean index ; Dolédec et al., 2000). This environmental factors. These ecological multivariate method addresses the question of dispositions probably facilitated recolonization of niche separation and niche breadth and helps in the Walloon karst, following the eradication of its finding which ecological factors are important for fauna during the Quaternary (Ice Age). If the community structure. The average marginality of maximum ice extension never reached the taxa was slightly significant, indicating a Walloon area, the permafrost penetrated several moderate influence of environmental and dozen of meters below the surface. historical variables on the distribution of taxa.

Figure 2 : Species occurrence in the unsaturated (US), saturated (S), hyporheic (H) and phreatic (P) zones of the “Walloon karst”.

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Figure 3 : Species richness accumulation curves in the Walloon region (karst and porous strata pooled together) according to various richness estimators (Sobs : number of species observed in the pooled samples ; ACE : Abundance-based Coverage Estimator; Jack 1-2: First and second-order Jackknife ; MMMean : Michaelis-Menten - Colwell, 2000).

ACKNOWLEDGEMENTS grottes de Wallonie. Bulletin des Chercheurs de la Wallonie 39:27-54. This work was financially supported by the European research project PASCALIS : Dolédec, S., D. Chessel, and C. Gimaret-Carpentier. 2000. Niche separation in community analysis : a new Protocols for the ASsessment and Conservation method. Ecology 81:2914-2927. of Aquatic Life In the Subsurface, (contract no Fiers, F. 2005. Speocyclops fontinalis spec. nov. EVK2-CT-2001-00121) of the Fifth Research and (Crustacea, Copepoda, Cyclopoida) from the Han Technological Development Framework Program Cave, a well known and popular cavern, in south of the European Community. This manuscript eastern Belgium. Bull. Inst. R. Sc. Nat. Belg., Biologie greatly benefited of useful comments and 75: in press. improvements by Georges Michel (CWEPSS, Leruth, R. 1939. La biologie du domaine souterrain et la Belgium). faune cavernicole de la Belgique. Mém. Mus. r. His. Nat. Belgique 87: 1-506. Malard, F., M.-J. Dole-Olivier, J. Mathieu, and F. Stoch. 2002. Sampling Manual for the assessment of BIBLIOGRAPHY Regional Groundwater Biodiversity. European project Colwell, R. K. 2000. EstimateS : Statistical estimation of : Protocols for the Assessment and Conservation of species richness and shared species from samples. Aquatic Life in the Subsurface (PASCALIS ; EVK2- Version 6b1. User’s guide and application published CT-2001-00121 ; http://www.PASCALIS- at: http://viceroy.eeb.uconn.edu/estimates. project.com/results/samplingmanual.html). Delhez, F., M. Dethier and J.-M. Hubart. 1999. Contribution à la connaissance de la faune des

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DISTRIBUTION OF GROUNDWATER INVERTEBRATES ALONG AN ENVIRONMENTAL GRADIENT IN A SHALLOW WATER-TABLE AQUIFER

1F. PARAN, 2F. MALARD, 2J. MATHIEU, 3M. LAFONT, 4D.M.P. GALASSI and 5P. MARMONIER.

1. Ecole Nationale Supérieure des Mines de Saint-Etienne, Centre SITE, 158, Cours Fauriel, 42023 Saint-Etienne Cedex 2, France. ([email protected]). 2. University Claude Bernard of Lyon 1, UMR CNRS 5023, Ecologie des Hydrosystèmes Fluviaux, 43, Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France. ([email protected]), ([email protected]). 3. UR Hydrobiologie, CEMAGREF, 3bis, Quai Chauveau, CP. 220, 69336 Lyon, France. ([email protected]). 4. Dipartimento di Scienze Ambientali, Università di l’Aquila, Via Vetoio, Coppito, 67100 L’Aquila, Italy. ([email protected]). 5. Université de Rennes 1, UMR CNRS 6553, Ecobio, Campus de Beaulieu, Bâtiment 14A, 263, Avenue du Général Leclerc, CS 74205, 35042 Rennes, France. ([email protected]).

ABSTRACT Hydrogeological attributes such as permeability are expected to drive biodiversity patterns in groundwater not only because they exert a direct influence on organisms but also control fluxes of nutrients and organic matter. We used OMI analysis to examine responses of multiple species to hydrogeological and physico-chemical variables in a shallow water-table aquifer along the Loire River, France, in which the water chemistry has been severely affected by gravel extraction and refilling with demolition materials. Hydrogeological attributes were obtained from a finite-difference groundwater flow model. Physico-chemical variables and biological data were collected during a single survey of 29 wells. Groundwater had elevated concentrations of dissolved organic carbon (DOC) and contained a high number of species, among which the majority were epigean. Spatial variation in the physico-chemistry of groundwater induced by the weathering of buried demolition materials, had no detectable influence on the distribution pattern of fauna. Rather, species were distributed along an environmental gradient defined primarily by permeability, aquifer thickness, and vadose zone thickness. Our results emphasize the importance of including hydrogeological measurements within the framework of ecological studies for understanding invertebrate assemblage distribution patterns in groundwater.

KEYWORDS : GROUNDWATER FLOW MODEL, PERMEABILITY, VADOSE ZONE THICKNESS, GROUNDWATER CHEMISTRY, GRAVEL EXTRACTION, GROUNDWATER CONTAMINATION, HYPOGEAN FAUNA.

Distinct factors are expected to drive zone thickness (VZT) are thought to exert a groundwater biodiversity at different spatial and major control on the distribution of groundwater temporal scales (Gibert et al. 1994). At the invertebrates because they influence fluxes of megascale (i.e. continent), differences in species matter and energy among spatial units of the composition among ecoregions reflect events aquifer (Gibert et al. 1994; Datry et al. in press). such as the formation of geological deposits, Human activity including groundwater drawdown, climate change (Quaternary glaciations), and the gravel extraction, and water pollution is another alternance of marine transgressions and potential source of biotic heterogeneity at the regressions. At a macroscale, the ecoregion is aquifer scale because it generates a mosaic of made of a mosaic of aquifers having distinct patches differing in their degree of alteration hydrogeological features such as land cover, (Mösslacher and Notenboom 1999). This pore size, and substrate permeability. At a anthropogenic source of heterogeneity is mesoscale, the aquifer comprises distinct spatial potentially superimposed on natural processes to units that differ in their disturbance regime and generate complex patterns of biodiversity. the degree of connectivity with the surface Variation in species composition between spatial environment. Hydrogeological features such as units differing in their flow conditions were permeability, groundwater recharge, and vadose already examined in a number of carbonated

SWSB December 2004 99 SESSION 3 : BIODIVERSITY PATTERNS aquifers including the Baget karst (Rouch et al. plain has an area of 750 km2. Average altitude is 1993), the Dorvan system (Gibert 1986), the 340 m and the plain is crossed from south to Organ cave basin (Culver et al. 1994), and the north by the Loire River (Duclos 1967). The Mammoth cave system (Poulson 1992). Forez aquifer is essentially fed by rainwater and Ecologists however, have only recently begun to is drained by the Loire River (figure 1). The explore the relationships between the general groundwater flow direction is from composition of invertebrate assemblages and southwest to northeast. The aquifer consists of environmental gradients in groundwater of quaternary alluvial deposits (gravels, sands, unconsolidated sediments (Danielopol, 1989; clays) and the surface area and volume of the Mauclaire and Gibert 2001; Dumas et al. 2001). aquifer are 10 km2 and 18 km3, respectively. The We examined invertebrate responses to lower confining bed which consists of Oligo- hydrogeological and physico-chemical gradients Miocene impermeable claystone is located at a in a shallow water-table alluvial aquifer along the depth of 6 m below the soil surface. Although the Loire River, France, in which the water chemistry Loire River is generally considered as one of the had been severely altered by gravel extraction last semi-natural large rivers in France, the Forez and refilling with demolition materials. Our plain has been highly modified by human objective was to determine whether the species activities including river embankment, intensive distribution patterns were controlled primarily by agriculture, and in particular gravel extraction hydrogeological features or by spatial variation in (Ulmer 1997). Gravel pits either formed artificial groundwater chemistry caused by gravel ponds fed by groundwater or they were filled with extraction and refilling with demolition materials. demo-lition materials. 1.2 Groundwater flow model 1. MATERIALS AND METHODS A finite-difference groundwater flow model 1.1 Study area : the Forez aquifer (NewSam type) was used with data from the The study area was an alluvial aquifer located in following parameters; water level measurements the sedimentary Forez plain which is part of the in multiple wells, permeability values obtained Loire River catchment, France. This sedimentary from resistivity measurements and pumping

Figure 1 : Map of the study aquifer showing piezometric head contours in January 2003.

100 SWSB December 2004 SESSION 3 : BIODIVERSITY PATTERNS tests, and estimates of rainfall, evaporation, and physicochemical variables. This two-table infiltration. The model provided piezometric ordination method calculates the marginality of levels for cells in a grid every 25, 50 or 100 m. species. Species marginality measures the The model was calibrated by modifying distance between the mean habitat conditions permeability until the calculated piezometric used by a species (species centroid) and the levels and the measured piezometric levels were mean habitat conditions in groundwater. A similar. Details of the groundwater flow model Monte-Carlo permutation test was used to check were provided by Mimoun (2004). the statistical significance of the marginality of each species. 1.3 Physico-chemical sampling The environmental data consisted of 3 Groundwater samples were collected from 29 hydrogeological features (aquifer thickness, wells in January 2003 to test for the influence of VDZ, and permeability) and 8 physicochemical gravel pits on groundwater chemistry (figure 1). 3- - variables (temperature, DO, DOC, PO4 , NO3 , In addition, 3 water samples were collected for + 2+ 2+ - NH4 , SiO2, and the sum of Ca , Mg , HCO3 , analysis from groundwater flowing through 2- - SO4 , and Cl expressed as meq/L). The faunal demolition materials. Groundwater was pumped data were expressed as presence / absence. from each well using a Bou-Rouch pump Four wells (coded as 35, C, C2 and P9 in figure (discharge rate ~10 L/min) (Bou and Rouch 1) were excluded from the analysis because they 1967). The first 50 L of pumped water were used had very low temperature and/or specific for sampling invertebrates (see below). conductance, suggesting direct infiltration of Groundwater was collected in burned glass surface runoff water into the well. The OMI bottles to determine dissolved organic carbon analysis positioned species along an (DOC) and in polypropylene bottles for analysis environmental gradient based on a maximization of Ca2+, Mg2+, HCO -, SO 2-, Cl-, PO 3-, NO -, NH +, 3 4 4 3 4 of their average marginality. The statistical 2 and SiO . Water samples were stored at 4¡C, significance of the average marginality of all returned to the laboratory within 4 h of collection, species was tested using a Monte-Carlo and filtered through a 0.45-µm membrane filter. permutation test. Multivariate analyses and Water analyses were performed following graphical displays were performed using ADE-4 standard methods (Clesceri et al. 1998). Specific software (Thioulouse et al. 1997; software conductance (WTW LF 330), pH (WTW pH 330) available on http://pbil.univ-lyon1.fr/ADE-4/). and dissolved oxygen (DO) (WTW OXI 330) were measured on the field. 2. RESULTS 1.4 Faunal sampling Invertebrate assemblages were collected from all 2.1 Hydrogeological features of the 29 wells in January 2003. The first 50 L of water aquifer extracted with the Bou-Rouch pump were filtered Permeability decreased from the margin to the through a 100-µm mesh net. Invertebrates were centre of the alluvial plain but increased close to preserved in 4% formaldehyde before processing the Loire River (figure 2). Aquifer thickness under a dissecting microscope. The most decreased from approximately 6 m at the margin abundant groups including Amphipoda, of the plain to 1 m near the Loire River. VZT Cyclopoida, Harpacticoida, and Oligochaeta increased from 1 m at the margin of the alluvial were counted and identified to genus or species plain to 6 m near the Loire River. where possible. Invertebrates were identified as 2.2 Groundwater chemistry hypogean or epigean depending on their degree of adaptation to the groundwater environment. Groundwater flowing through buried demolition Hypogean invertebrates are obligate materials (n=3 samples) was suboxic (i.e. DO < groundwater taxa that complete their entire life 0.3 mg/L) and had high DOC concentrations cycle exclusively in subsurface water (i.e., (mean ± standard deviation : 11 ±3.7 mg/L), stygobites in Gibert et al. 1994). Epigean specific conductance (1,221 ±477 µS/cm), and invertebrates are taxa that have no affinities for sulphate concentrations (295 ±306 mg/L). Nitrate groundwater but occur accidentally in alluvial concentration was distinctly lower in groundwater 3 sediments (i.e., stygoxenes in Gibert et al. 1994) flowing through demolition materials (NO < 0.1 or taxa that exploit groundwater resources during mg/L) than in groundwater flowing through 3 at least a part of their life cycle (i.e., stygophiles alluvial deposits (35.5 ±34.3 mg/L NO , n=29). 2+ 2+ - in Gibert et al. 1994). Specific conductance and Ca , Mg , HCO3 , 2- SO4 , and Cl- concentrations in groundwater 1.5 Data analysis were positively correlated (0.41

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Figure 2 : Distribution map of permeability after model calibration. up to 1315 µS/cm) and concentrations of major which only one taxon was hypogean, showed a ions were measured in most wells located significant deviation (p<0.1) of their habitat downstream of the gravel pits. DOC preference from a uniform distribution (figure 4). concentration in groundwater averaged 3.8±3.5 Eight of these 9 non-uniformly distributed taxa mg/L and tended to increase with increasing favoured the permeable side of the distance from the Loire River. Mean DO environmental gradient. Most hypogean taxa concentration in groundwater was 2.6 ±1.5 mg/L occupied a central position along the O2 (n=29). environmental gradient (figure 4). 2.3 Species-environment relationships 3. DISCUSSION A total of 15,458 invertebrates were collected This study is the first on groundwater from the groundwater wells. Fifty taxa were invertebrate assemblage composition in an identified of which 15 were hypogean (Appendix aquifer of the Loire River catchment (Ferreira et 1). Mean density was 533 invertebrates/50 L of al. in press). Despite severe alteration by gravel pumped water (SD = 1414, range: 9Ð7,709 n = mining, the Forez aquifer harbours a rich 29). Axis 1 of the OMI analysis explained 52.8% groundwater fauna including 15 hypogean taxa. of the variability and arranged the sampling wells Average invertebrate density was high and many along a gradient of increasing permeability, epigean taxa were present. Mean densities in the increasing aquifer thickness (from bottom to top alluvial aquifer of the Ariège River and the glacio- in figure 4) and decreasing VDZ (17.4, 20.2, and fluvial aquifer of the Lyon metropolitan area, 18.8% of the formation of axis 1, respectively). France, were 1,000 invertebrates per 1,000 L of There was no significant correlation (p>0.10, pumped water and 44 ±117 invertebrates per 50 n=25) between hydrogeological attributes and Lof pumped water, respectively (Dumas and groundwater mineralization. Hydrogeological Fontani 2001, Datry et al. in press). Groundwater features had a moderate but significant influence fauna in these two aquifers was dominated by on the distribution of taxa (p=0.10, global Monte- obligate groundwater taxa. High faunal densities Carlo permutation test). Nine of 43 taxa, among and the presence of high numbers of epigean

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Figure 3 : Distribution map of specific conductance in groundwater. invertebrates in shallow groundwater of the relative to the permeability of the aquifer (Peffer Forez plain are likely to reflect a high food supply. 1982). Groundwater flow would tend to Reduced thickness of the vadose zone has been circumvent compacted wastes, thereby isolating shown to increase the amount of soil-derived low-permeability demolition materials and labile DOC reaching the groundwater table reducing their impact on groundwater (Pabich et al. 2001). The average DOC mineralization. DOC enrichment and subsequent concentration in the shallow water-table aquifer depletion of DO and NO3 within groundwater of the Forez plain was 7 times higher than that flowing through buried demolition materials reported by Datry et al. (in press) for the deep suggest that reduction processes may be a water-table aquifer of the Lyon metropolitan area, source of dissolved organic matter (organic France. High DOC input at the water table of the matter degradation) but a sink of nitrogen Forez aquifer probably stimulated microbial (denitrification). Additional studies are needed to production, thereby providing sufficient food determine the potential for groundwater resources for epigean species. contamination resulting from the leachate of Refilling gravel pits with demolition materials heavy metals and/or organic compounds. strongly affected the physico-chemistry of The analysis of species-environment groundwater. The dissolution of buried materials relationships in this highly-altered aquifer such as plaster, concrete, or asphalt suggests that spatial variation in physico- considerably increased the concentrations of chemistry of groundwater induced by the major ions, particularly sulphates, calcium, and dissolution of buried demolition materials was not chloride. The increase in total dissolved solids the primary factor controlling the distribution was observed at considerable distance pattern of fauna. Rather, species were distributed downstream of the gravel pits indicating that the along an environmental gradient defined dissolution processes had a global effect on primarily by hydrogeological features including groundwater quality. The influence of gravel pits permeability, aquifer thickness and VDZ. Dumas on groundwater chemistry is likely to depend et al. (2001) demonstrated that the distribution of upon the permeability of the demolition materials groundwater macrocrustaceans in the alluvial

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Figure 4 : Outlying mean index (OMI) analysis of groundwater invertebrate assemblages. Axis 1 representing 52.8% of the variability was used for graphical representation. Left panel: canonical weights of environmental variables (DOC: dissolved organic carbon; mineralization: 2+ 2+ - 2- - sum of Ca , Mg , HCO3 , SO4 , and Cl concentrations expressed as meq/L). Middle panel: scores of sampling wells. Right panel: distribution of taxa along the environmental gradient as a function of their weighted average position along well scores. The sizes of black circles are proportional to the total occurrence of taxa. Vertical lines are standard deviations. Asterisks indicate taxa whose distribution deviates significantly from a uniform distribution along the environmental gradient (Stmi: Stylodrilus mirus, Acro: Acanthocyclops robustus, Paff: Parastenocaris fontinalis fontinalis, Rhfa: Rhyacodrilus falciformis, Tbtu: Tubifex tubifex, Luva: Lumbriculus variegatus, Deoo: Dero sp., Frga: Fridericia galba, Psal: Pseudocandona albicans) See appendix 1 for the codes of taxa (Caju: juveniles of Candoninae, Ostracoda). aquifer of the Ariège River in France was driven reaching the groundwater. Links between by hydrogeological features, particularly aquifer hydrogeological features, fluxes of organic transmissivity, rather than groundwater quality matter, and competition processes among taxa caused by agriculture. A number of recent potentially result in complex biodiversity patterns studies emphasize the importance of integrating in groundwater. High fluxes of dissolved organic hydrogeological measurements within the matter into permeable and shallow water-table framework of ecological studies for areas of the Forez plain facilitate the colonization understanding distribution patterns of by epigean invertebrates. These epigean invertebrate assemblages in the groundwater of organisms may compete with obligate unconsolidated sediments (Mösslacher 1998, groundwater taxa that are relegated to areas less Mauclaire and Gibert, 2001, Datry et al. in press). rich in organic matter. This model of community Permeability, i.e. the degree of organization provides a working hypothesis for interconnectedness between the pores of an future ecological investigations in this alluvial aquifer, has a direct influence on groundwater aquifer. fauna because it is likely to determine the spatial extent of available habitats as well as the amount of energy spent by organisms to explore their ACKNOWLEDGEMENTS environment. Hydrogeological factors also This work was a part of the ADNT program indirectly control biodiversity patterns by (Territorial Decision and Negotiation Maker influencing fluxes of nutrients, organic matter, according to governance principles, leader : D. and heat from the surface environment. Datry et Graillot) funded by the École des Mines de Saint- al. (in press) demonstrated that VDZ was a Etienne and the Rhône-Alpes Région and was primary factor determining biodiversity patterns supported by the FRAPNA Loire and Morillon in groundwater of the Lyon metropolitan area Corvol. We are indebted to T. Datry for his because it controlled the amount of DOC valuable help during sampling, J. Gibert and her

104 SWSB December 2004 SESSION 3 : BIODIVERSITY PATTERNS research group for their support and most Gibert, J. 1986. Ecologie d’un système karstique dedicated help with the laboratory work, R. Ginet jurassien. Hydrogéologie, dérive animale, transits de for his valuable help in the identification of matières, dynamique de la population de Niphargus (Crustacé Amphipode). Mémoires de Biospéologie Niphargus species (Amphipoda), and D. Mimoun 13:1-379. for the implementation of the groundwater flow Gibert, J., J.A. Stanford, M.-J. Dole-Olivier, and J.V. Ward. model. We thank A. Wright-Stow (NIWA, New 1994. Basic attributes of groundwater ecosystems Zealand) for comments that improved an earlier and prospects for research. In: Groundwater ecology, version of this manuscript. J. Gibert, D.L. Danielopol and A. Stanford [Eds.], Academic Press, San Diego, 7-40. Mauclaire, L. and Gibert, J. 2001. Environmental determinants of bacterial activity and faunal BIBLIOGRAPHY assemblages in alluvial riverbank aquifers. Archiv für Bou, C. and Rouch, R. 1967. Un nouveau champ de Hydrobiologie 152:469Ð487. recherche sur la faune aquatique souterraine. Mimoun, D. 2004. Spatialisation de l’information : une Compte-Rendus de l’Académie des Sciences de aide à l’analyse hydraulique et paysagère développée Paris 265:369Ð370. lors de la réhabilitation de sites post-industriels. Cas Clesceri, L.S., A.E. Greenberg, and A.D. Eaton. 1998. des réaménagements des gravières en eau en milieu Standard methods for the examination of water and alluvionnaire. Thèse de doctorat, Ecole nationale wastewater. American Public Health Association, supérieure des mines, Saint-Etienne. Washington. Mösslacher, F. and Notenboom, J. 1999. Groundwater Culver, D.C., W.K. Jones, D.W. Fong, and T.C. Kane. biomonitoring. In: Biomonitoring of polluted water, A. 1994. Organ cave karst basin. In: Groundwater Gerhardt [Ed.], Trans Tech Publications, Zürich, ecology, J. Gibert, D.L. Danielopol and A. Stanford Switzerland, 119Ð140. [Eds.], Academic Press, San Diego, 451-473. Mösslacher, F. 1998. Subsurface dwelling crustaceans as Danielopol, D.L. 1989. Groundwater fauna associated indicators of hydrological conditions, oxygen with riverine aquifers. Journal of the North American concentration and sediment structure in an alluvial Benthological Society 8:18Ð35. aquifer. International Review of Hydrobiology 83:349- Datry, T., F. Malard, and J. Gibert. 2005. Response of 364. invertebrate assemblages to increased groundwater Pabich, W.J., I. Valiela, and H.F. Hemond. 2001. recharge rate in a phreatic aquifer. Journal of the Relationship between DOC concentrations and North American Benthological Society, in press. vadose zone thickness and depth below the water Dolédec, S., D. Chessel, and C. Gimaret-Carpentier. table in ground water of Cape Cod, U.S.A. 2000. Niche separation in community analysis: a new Biogeochemistry 55:247Ð268. method. Ecology 81(10), 2914-2927. Peffer, J.R. 1982. Fly ash disposal in a limestone quarry. Duclos, P. 1967. Géologie et minéralisation uranifères de Ground Water 20:267-273. la Plaine tertiaire du Forez. Mémoire de thèse, Centre Poulson, T.L. 1992. The Mammoth Cave ecosystem. In: à l’Energie Atomique, rapport 3117, Fontenay-aux- The Natural History of Biospeology, A.I. Camacho Roses. [Ed.], National Museum of Natural Sciences, Madrid, Dumas, P., C. Bou, and J. Gibert. 2001. Groundwater 568-611. macrocrustaceans as natural indicators of the Ariège. Rouch, R., A. Pitzalis, and A. Descouens. 1993. Effets International Review of Hydrobiology 86:619Ð633. d’un pompage à gros débit sur le peuplement des Dumas, P. and Fontani, G. 2001. Sampling fauna in Crustacés d’un aquifère karstique. Annales de aquifers: a comparison of net-sampling and pumping. Limnologie 29:15-29. Archiv für Hydrobiologie 150:661-676. Thioulouse, J., D. Chessel, S. Dolédec, and J.-M. Olivier. Ferreira, D., F. Malard, M.-J. Dole-Olivier, and J. Gibert. 1997. ADE-4: a multivariate analysis and graphical 2005. Obligate groundwater fauna of France: diversity display software. Statistics and Computing 7:75Ð83. patterns and conservation implications. Biodiversity Ulmer, A. 1997. Expertise écologique de l’Ecozone du and Conservation, in press. Forez. Rapport interne, FRAPNA Loire.

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Appendix 1 : List of taxa collected in 29 wells of the Forez aquifer, France. Hypogean taxa are indicated with an asterisk. Codes of taxa are used in figure 4.

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DOES GROUNDWATER RECHARGE STIMULATE BIODIVERSITY ?

T. DATRY, F. MALARD and J. GIBERT

University Claude Bernard of Lyon 1, UMR CNRS 5023, Ecologie des Hydrosystèmes Fluviaux, 43 Bd 11 Novembre 1918, 69622 Villeurbanne Cedex, France. ([email protected]), ([email protected]), ([email protected]).

ABSTRACT In groundwater ecosystems, DOC supply and spatio-temporal variability in environmental conditions are likely to be the most important parameters driving the distribution and the composition of invertebrates assemblages. Because areas with higher groundwater recharge rates may present both higher DOC supply and higher spatio-temporal variability, we expected groundwater biodiversity to peak in recharge areas of aquifers. We took advantage of the increasing use of artificial infiltration of stormwater into groundwater to test this hypothesis by comparing the density and species richness of invertebrate assemblages among reference sites and sites artificially recharged with stormwater. Because we suspected the vadose zone thickness (VZT) to play an important role in controlling DOC supply and damping spatio-temporal variability in groundwater, we selected sites with a large range of VZT. The vertical distribution of invertebrate assemblages at a shallow water-table recharge site and a nearby reference site was also examined. Results showed that artificial recharge of groundwater induced elevated dissolved organic carbon (DOC) concentrations in groundwater and increased spatio-temporal physico-chemistry variability. However, DOC enrichment was controlled by the vadose zone thickness (VZT), whereas spatio-temporal variability was not. Artificially recharged groundwater presented higher invertebrate assemblage density and richness, the amount of which was also controlled by VZT. This suggested that organic matter supply, rather than spatio-temporal variability, was a primary factor determining biodiversity patterns in groundwater. At shallow water-table sites, invertebrate density increased and species composition shifted with increasing depth below the groundwater table. Some taxa, including several stygophiles, preferentially occurred near the water table, whereas others, including several stygobites, colonized deeper groundwater layers. This could be the result of biotic interactions between stygophiles and stygobites, the latter, although found limited by food supply, being outcompeted in the uppermost layer of groundwater.

KEYWORDS : STORMWATER, DISSOLVED ORGANIC CARBON, DISSOLVED OXYGEN, ENVIRONMENTAL VARIABILITY, HYPOGEAN INVERTEBRATES, FAUNAL GRADIENT

1. INTRODUCTION 1998, Baker et al. 2000). DOC input into In groundwater ecosystems, DOC supply is likely groundwater is also expected to stimulate the to be the most important parameter driving production of microbial biofilms, which are then biodiversity. Indeed, the absence of light grazed by invertebrates (Cummins and Klug prevents photosynthesis, and organic carbon 1979, Bärlocher and Murdoch 1989). standing stocks are generally very low. Typically, Surprisingly, only few studies have focused on groundwater dissolved organic carbon (DOC) the role of DOC supply in driving groundwater concentrations are lower than 1 mg/L, and biodiversity (Holsinger 1966, Sinton 1984). sediment organic matter does not exceed 0.5% Following the predictions of the habitat template by dry weight (Leehneer et al. 1974, Wedepohl model developed for surface ecosystems 1978, Malard & Hervant 1999). Surface-derived (Townsend 1989), spatio-temporal variability of DOC supply has already been showed to limit habitat could also be an important factor affecting some of the most important biogeochemical biodiversity in groundwater. However, overall processes in groundwater, including aerobic spatio-temporal variability is highly reduced in respiration, denitrification or even groundwater ecosystems, in terms of both methanogenesis (Jones et al. 1995, Hedin et al. physical habitat and environmental conditions

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(Ginet & Decu, 1977, Gibert et al. 1994). with stormwater in addition to rain water, and 11 Previous studies addressing the question of does reference sites recharged only by direct spatio-temporal variability, in terms of infiltration of rain water at the land surface. Sites environmental conditions, affect the distribution were selected to constitute 2 categories of of invertebrates in groundwater did not provide vadose zone thickness (VZT): < 10 m and > 10 m strongly consistent results (Pospisil 1999). for both recharge and reference sites. Each site Higher DOC supply and higher spatio-temporal was equipped with a monitoring well with a variability in groundwater are expected in perforated casing intersecting the water-table recharge areas of aquifers, where surface water region of the aquifer. Additionally, for each VZT infiltrates downward to the water table. Several category, a recharge and a nearby reference studies demonstrated that DOC concentrations sites were equipped with well clusters to sample in groundwater were distinctly higher in recharge groundwater each meter below the water table areas of shallow groundwater because the and examine differences in the vertical residence time of water in the vadose zone was distribution of invertebrate assemblages (Datry et insufficient to allow microbial degradation of al. 2004, Malard et al. 2004). surface-derived DOC (Starr and Gillham 1993, 2.2 Methods Pabich et al. 2001). Areas of groundwater recharge should also exhibit a higher spatio- 2.2.1 Spatio-temporal variability in groundwater temporal variability in physico-chemistry because physico-chemistry the infiltration of surface water induces strong We used temperature and specific conductance temporal fluctuations in temperature and water as surrogates of overall spatio-temporal chemistry, which are attenuated with increasing variability of groundwater. Differences in the depth and distance from the recharge area spatio-temporal variability of groundwater (Datry et al. 2004, Malard et al. 2004). between recharge and reference sites were thus The increasing use of artificial infiltration basins assessed by recording specific conductance and to dispose of stormwater and tertiary effluent into temperature in well clusters of the shallow and the subsurface is likely to induce locally high deep water-table sites for 10 months with YSI groundwater recharge rates, and subsequent 600 XLM multi-parameter loggers (Yellow Spring changes in groundwater DOC supply and spatio- Instrument Co., Yellow Springs, Ohio, USA). temporal variability of groundwater physico- 2.2.2 Groundwater DOC, DO and phosphates and chemistry (Pitt et al. 1999, Datry et al. 2004). biodiversity at recharge and reference sites Taking advantage of that, we addressed the role Groundwater was collected from each well on 4 of increased groundwater recharge rates in dates with a pneumatic piston pump equipped sustaining biodiversity in a regional phreatic with 2 inflatable packers (pumping rate ~ 10 aquifer by examining differences in the density L/min) (UWITEC Company, Mondsee, Austria). and species richness of invertebrate Groundwater was also collected from the well assemblages among reference sites and sites clusters at the recharge and reference sites (i.e. artificially recharged with stormwater. We also each m below the water table) on 5 dates with a examined differences in the vertical distribution Bou-Rouch pump (Bou & Rouch 1967). DOC, of invertebrate assemblages between a dissolved oxygen (DO) and phosphate reference site and a shallow water-table concentrations were determined using standard recharge site. We expected that : methods (Clesceri et al. 1998). Invertebrate ¥ organic matter supply and spatio-temporal assemblages were analysed in 50 L of pumped heterogeneity would be higher in recharge groundwater. Two-way repeated measures areas; ANOVAs and Scheffé tests were used to compare concentrations of DOC, DO, and ¥ increased DOC inputs and spatio-temporal phosphate, and invertebrate density and diversity heterogeneity at recharge sites would lead to at recharge and reference sites. higher invertebrate assemblage density and species richness;

¥ the effect of recharge would be mediated by the 3. RESULTS vadose zone thickness (VZT). 3.1 Differences in spatio-temporal variability among recharge and reference sites 2. MATERIAL & METHODS For both VZT categories, artificial recharge 2.1 Study sites increased the temporal variability of groundwater The study area was a glaciofuvial aquifer 10 km temperature and specific conductance (Fig. 1). east from Lyon, France. A total of 24 sites were This induced temporal variability was drastically studied; 13 recharge sites artificially recharged damped with depth below the water table.

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Figure 1 : Time series (1-h intervals) of temperature and specific conductance measured at depths of 1.2 and 4 m (black, dark grey and light grey patterns) below the groundwater table at the shallow water-table sites (vadose zone thickness < 10 m, left panel) and deep water-table sites (vadose zone thickness > 10 m, right panel), and rainfall. Arrows indicate reference-time and recharge-time series. Modified after Datry et al. 2005.

3.2 Differences in DOC, DO and 3.3 Vertical patterns of DOC and phosphate concentrations among phosphate concentrations at the recharge and reference sites shallow water-table site There were no statistical differences between Groundwater DOC and phosphate DOC, DO and phosphate concentrations at concentrations were higher at the recharge site recharge and reference sites in deep (> 10 m) compared to the reference site (Fig. 3). At the groundwater (p > 0.05) (Fig. 2). In contrast, DOC recharge site, vertical gradients in DOC and and phosphate concentrations were higher phosphates developed below the water table (Scheffé test, p < 0.001 and p = 0.012, immediately after rainfall events (Fig. 3). respectively) and DO lower (Scheffé test, p < 0.001) at recharge sites than at reference sites in shallow groundwater (VZT < 10m) (Fig. 2).

3- Figure 2 : Differences in dissolved organic carbon (DOC), dissolved oxygen (DO), and phosphate (PO4 ) between recharge sites (black triangles, n ≥ 5) and reference sites (white circles; n ≥ 5) for vadose zone thickness < 10 m (left grey panel) and > 10 m (right white panel). Modified after Datry et al. 2005.

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Figure 3 : Vertical changes in DOC and phosphates in ground water at the shallow water-table recharge site (black triangles) and shallow water-table reference site (white circles) during dry weather (upper panels, mean + standard deviation ; n = 3) and rainfall events (lower panels, mean + standard deviation ; n = 2). Modified after Datry et al. 2005.

3.4 Differences in biodiversity among pumped groundwater at recharge sites (n = 20 recharge and reference sites samples) against 9 + 13 invertebrates per 50 L at Density and species richness were higher at references sites (n = 24 samples) (Fig. 4). recharge sites for VZT < 10 m (Scheffé test, p < However, there were no statistical differences for 0.001), with 62 + 100 invertebrates per 50 L of VZT > 10 m (Fig. 4).

Figure 4 : Differences in density and richness of invertebrate assemblages between recharge sites (black triangles) and reference sites (white circles) for vadose zone thickness < 10 m (left grey panel) and > 10 m (right white panel). Modified after Datry et al. 2005.

Figure 5 : Vertical changes in density and richness of invertebrate assemblages in ground water at the shallow water-table recharge site (black triangles) and shallow water-table reference site (white circles) during dry weather (upper panels, mean + standard deviation; n = 3) and rainfall events (lower panels, mean + standard deviation; n = 2). Modified after Datry et al. 2005.

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3.5 Vertical patterns of invertebrate distributed along the 1st axis (18.4% of the assemblages at the shallow water- variance) following a gradient of density, and table site along the 2nd axis (10.6% of the variance) The density of invertebrate assemblages according to their habitat affinities (stygophiles- decreased with increasing depth at shallow stygobites). Ordination of species according to water-table sites, regardless of recharge or their factorial scores along the second axis of a weather conditions (Fig. 5). PCA showed that several taxa, including a majority of stygophile invertebrates (e.g., E. Average density decreased from 411 + 378 serrulatus, P. fimbriatus, M. albidus, A. robustus, invertebrates per 50 L of water at a depth of 1 m and D. languidoides ssp.), occurred near the to 33 + 16 invertebrates at a depth of 4 m at the water table (Fig. 6). Other taxa, including most of recharge site and from 32 + 8 to 5 + 4 the stygobite species, either occurred at multiple invertebrates, respectively, at the reference site. depths (e.g., P. zschokkei, S. juberthiae) or were In contrast, richness did not show any vertical restricted to deeper depths (e.g., M. reginae, S. pattern. lescherae, and A. rhenanus) (Fig. 6). This shift in The first two axes of a principal component species composition with depth was observed analysis (PCA) explained altogether 29% of the both at the recharge and reference site. variance in the distribution of taxa. Taxa were

Figure 6 : Vertical distribution of selected species at the shallow water-table sites. Species were ordinated according to their factorial scores along axis 2 of the principal component analysis. Asterisks indicate stygophile species. Diameters of circles are proportional to the mean number of individuals collected per 50 L of pumped water (n = 5 dates). Crosses correspond to sites where no individuals were collected. Modified after Datry et al. 2005.

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4. DISCUSSION ACKNOWLEDGEMENTS As predicted, increased infiltration rates led to 1) This work was part of the OTHU project higher DOC supply, and 2) higher spatio- (Experimental Observatory for Urban Hydrology) temporal variability in groundwater. Results funded by the urban community of Lyon clearly indicated that VZT influenced DOC supply (COURLY) and the Rhône-Alpes Region. We are but did not attenuate heterogeneity. This role of indebted to G. Bouger and L. Vitry for dedicated the vadose zone in retaining/degrading DOC assistance with field and laboratory work. We from infiltrating water was consistent with thank to the following specialists for their previous studies showing that shallow water- valuable help in the identification of taxa : M.-J. table aquifers presented higher DOC Dole-Olivier (Salentinella), R. Ginet (Niphargus), concentrations than deep water-table aquifer F. Lescher-Moutoué and J. Mathieu (Cyclopoida) (Starr & Gillham 1993, Pabich et al. 2001). and P. Marmonier (Ostracoda). Density and richness of invertebrate assemblages were higher at recharge sites for VZT < 10 m. However, despite a strong increase BIBLIOGRAPHY in spatio-temporal variability at recharge sites Baker, M.A., H.M. Valett, and C.N. Dahm. 2000. Organic with a VZT > 10 m, density and richness were not carbon supply and metabolism in a shallow different from reference sites. These results groundwater ecosystem. Ecology 81(11):3133-3148. provided strong evidences that organic matter Bärlocher, F. and Murdoch, J.H. 1989. Hyporheic biofilms- supply, rather than heterogeneity, was the a potential food source for interstitial animals. Hydrobiologia 184:61-67. primary driver of biodiversity in groundwater. Bou, C. and Rouch, R. 1967. Un nouveau champ de Invertebrate density decreased with increasing recherche sur la faune aquatique souterraine. depth below the water table. This indicated a Compte-Rendus de l’Académie des Sciences de tendency among groundwater invertebrates to Paris 265:369-370. develop abundant populations in resource-rich Brunke, M. and Gonser, T. 1999. Hyporheic invertebrates patches located in the vicinity of surface - the clinal nature of interstitial communities structured production sites of organic matter. This trend was by hydrological exchange and environmental consistent with patterns observed in the gradients. Journal of the North American Benthological Society 18:344-362. hyporheic zone of rivers, where invertebrates were generally more abundant in downwelling Clesceri, L.S., A.E. Greenberg, and A.D. Eaton. [eds.], 1998. Standard methods for the examination of water sites (Marmonier et al. 1992, Fowler et al. 2002). and wastewater. American Public Health Association, By contrast, richness did not show any pattern Washington. with depth, supposedly because food Cummins, K.W. and M.J. Klug. 1979. Feeding ecology of competition among taxa induced a shift in stream invertebrates. Annual Review of Ecology and species composition (see below). Systematics 10:147-172. The vertical shift in species composition was Datry, T., F. Malard, and J. Gibert. 2004. Dynamics of observed at both recharge and reference sites solutes and dissolved oxygen in shallow urban groundwater below a stormwater infiltration basin. despite marked differences in DOC input and Science of the Total Environment 329:121-229. spatio-temporal heterogeneity between sites. Datry, T., F. Malard, and J. Gibert. 2005. In press. This shift in invertebrate assemblages could Response of invertebrate assemblages to increased reflect the outcome of biotic interactions among groundwater recharge rates in a phreatic aquifer. taxa. Some taxa, including several stygobites Journal of the North American Benthological Society. could be found limited by food supply but they Fowler, R.T. and M.R. Scarsbrook. 2002. Influence of were out-competed in the uppermost layer of hydrologic exchange patterns on water chemistry and groundwater by other taxa, including several hyporheic invertebrate communities in three gravel- stygophiles. Brunke and Gonser (1999) bed rivers.New Zealand Journal of Marine and Freshwater Research 36:471Ð482. suggested that the depth distribution of stygoxen, stygophile and stygobite taxa within the Gibert, J., J.A. Stanford, M.-J. Dole-Olivier, and J.V. Ward. 1994. Basic attributes of groundwater ecosystems hyporheic zone of the Töss River, Switzerland, and prospects for research. pp. 7-40 in: Groundwater was governed by trade-offs. According to these Ecology, J. Gibert, D.L. Danielopol and J.A. Stanford. authors, depth penetration within the sediment [Eds.], Academic Press, San Diego. by stygoxen and stygophile taxa was limited by Ginet, R., and Decu, V. 1977. Initiation à la biologie et à food supply, whereas colonization of the upper l’écologie souterraines. J.-P. Delarge [Ed.], Paris, (ou carbon-rich layers by stygobite taxa was Edit. Univers., 400 p.), 345 pp. restricted by interference competition. This Hedin, L.O., J.C. Von Fischer, N.E. Ostrom, M.G. Brown, model of community organization within the and G.P. Robertson. 1998. Thermodynamic hyporheic zone of streams might also be valid for constraints on nitrogen transformation and other biogeochemical processes at soil-stream interfaces. explaining the distribution pattern of invertebrate Ecology 79:684-703. assemblages at the water-table interface of Holsinger, J.R. 1966. A preliminary study on the effects of phreatic aquifers. organic pollution of Banners Corner Cave, Virginia.

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International Journal of Speleology 2:75-89. Pabich, W.J., I. Valiela, and H.F. Hemond. 2001. Jones, J.B., R.M. Holmes, S.G. Fisher, N.B. Grimm, and Relationship between DOC concentrations and D.M. Green. 1995. Methanogenesis in Arizona, USA, vadose zone thickness and depth below the water dryland streams. Biogeochemistry 31:155-173. table in groundwater of Cape Cod, U.S.A. Biogeochemistry 55:247-268. Leehneer, J.A., R.L. Malcolm, P.W. McKinley, and L.A. Eccles. 1974. Occurrence of dissolved organic carbon Pitt, R., S. Clark, and R. Field. 1999. Groundwater in selected groundwater samples in the United States. contamination potential from stormwater infiltration Journal Research U.S. Geol. Survey 2:361-369. practices. Urban Water 1:217-236. Malard, F., T. Datry, and J. Gibert. 2004. The use of Pospisil, P. 1999. The Composition of Cyclopoid multilevel wells and multiparameter loggers for Assemblages in Ecologically Different Groundwater monitoring groundwater quality below stormwater Habitats of a Danube Riverine Wetland in Austria. infiltration basins. p.p. 713-720 In: GRAIE [Ed.], Crustaceana 72(8):883-892. Sustainable techniques and strategies in urban water Sinton, L.W. 1984. The macroinvertebrates in a sewage- management. Novatech 2001 Proceedings, Delta polluted aquifer. Hydrobiologia 119:161-169. Imprimerie, Lyon, France. Starr, R.C. and Gillham, R.W. 1993. Denitrification and Malard, F. and Hervant, F. 1999. Oxygen supply and the organic carbon availability in two aquifers Journal of adaptations of animals in groundwater. Freshwater the North American Benthological Society Biology 41:1-30. Townsend, C.R. 1989. The patch dynamic concept of Marmonier, P., M.-J., Dole-Olivier, and M. Creuzé des stream community ecology. Journal of the North Châtelliers. 1992. Spatial distribution of interstitial American Benthological Society 8(1):36-50. assemblages in the floodplain of the Rhône River : Wedepohl, K.H. [ed.], 1978. Handbook of Geochemistry, Regulated Rivers. Research and Management Vol II- 1, Springer Verlag, Berlin. 7:75Ð82.

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ENVIRONMENTAL QUALITY OF DEEP GROUNDWATER IN THE LESSINIAN MASSIF (ITALY): SIGNPOSTS FOR SUSTAINABILITY

T. DI LORENZO, F. STOCH, B. FIASCA, E. GATTONE, P. DE LAURENTIIS, F. RANALLI and D.M.P. GALASSI

Dipartimento di Scienze Ambientali, University of L’Aquila, Via Vetoio, Coppito, 67100, L’Aquila, Italy. ([email protected]), ([email protected]), ([email protected]), ([email protected]), ([email protected]).

ABSTRACT Groundwater conservation and management planning in the Lessinian Massif (Veneto, Italy) is herein discussed following an ecological approach dealing with stygobiont species. The aim was to provide baseline signposts for the sustainable management of groundwater sites in order to guarantee the preservation of the stygobiota. The approach is organised in the following steps: 1) groundwater biodiversity assessment; 2) selection of priority sites for conservation; 3) selection of respect zones for the priority sites; 4) evaluation of the main anthropogenic risk drivers; 5) assessment of aquifer intrinsic vulnerability; 6) evaluation of risk; 7) proposal of sustainability signposts.

KEYWORDS : AQUIFER VULNERABILITY, COPEPODS, RISK, STYGOBIONT, SUSTAINABILITY.

1. INTRODUCTION In the document about the sustainable use of Consequently, a possible strategy for obtaining groundwater worked out by RIZA/RIVM (1991) sustainability is the use of a general protection for the EU Minister, groundwater sustainability framework, which takes into account the was hung on two conditions: 1) no loss of multifunctional features of the groundwater groundwater potential functions; 2) preservation resources, as supplies for anthropic uses and of ecosystem diversity and maintenance of repository of groundwater biodiversity. Moreover, species richness. However, the conception that the protection framework should foresee a logical groundwater is not only important in supporting approach in accordance with different human welfare, but also in harbouring different environmental pressures and/or sensitivities per taxa, belonging to several phylogenetic lineages country and/or region (Notenboom, 2001). The (Danielopol et al., 2004), is not yet well present contribution falls in this context and the recognised at political level. The EU enforced the final objective is to suggest some sustainable protection of groundwater environments via a management practices of groundwater resources number of Directives which mainly give in the Lessinian area (northern Italy). The recognition to anthropic aspects (91/271/EEC; approach allows an analysis of risk that 91/676/EEC; 2000/60/EC). The 2000/60/EC Lessinian aquifers will be likely to experience. Directive and the proposed EU Groundwater The study was organised in the following steps: Directive 2003 both require the achievement of a 1) groundwater biodiversity assessment; 2) good “groundwater status” and the selection of priority sites for conservation; 3) protection/restoration of groundwater bodies with selection of respect zones for the priority sites; 4) extensive monitoring programmes, completely evaluation of the main risk drivers; 5) ignoring their ecological status (Danielopol et al., assessment of aquifer intrinsic vulnerability; 6) 2004). Hence, although the concept of evaluation of risk; 7) proposal of sustainability sustainability has been introduced in the signposts. European Directives, the misconception about the meaning of sustainability still remains. What 2. MATERIALS AND METHODS is needed is the overall awareness that a sustainable groundwater use can only be 2.1 Study area achieved if groundwater management is part of The Lessinian Mountains are a 472.14 km2 an integrated approach, directed toward the trapezium-shaped massif, located in the maintenance of groundwater potential functions Venetian Pre-Alps (Veneto, Italy). The massif and biodiversity (Notenboom, 2001). extends southward in divergent finger-like ridges,

SWSB December 2004 115 SESSION 4 : CONSERVATION AND MANAGEMENT branching and entering in contact with the alluvial The survey was carried out from June to July plane of River Adige. Minor valleys between the 2002 during the low hydrogeological regime ridges are narrow and steep in the northern when the annual precipitation is about 100 mm sector and, in general, wide and smoothly vs. 300 mm in October-November (Patrizi and sloping in the southern one (Fig. 1). The Lavagnoli, 2001). Two saturated strata were composition of the massif is dominated by selected according to the aquifer typology: a limestones of Mesozoic and Cenozoic ages, karstic stratum (K) and a porous stratum (P). interspersed by Cenozoic volcanic rocks and Spatial distribution of fauna and environmental Eocenic limestone outcrops (Sauro, 1973). parameters were investigated from a network of Quaternary alluvial deposits fill up the valleys 76 cased wells, following a standardized and cover the whole plane. The alluvial thickness methodology (Malard et al., 2002). Wells were is variable, ranging from 100 to 200 m. The flow sorted out randomly in each stratum: 29 wells in the karstic network is characterised by high were selected in the karstic stratum and 47 wells speed and short travel-time and the hydrological in the porous one which is more widely

Figure 1 : Map of the Lessinian Mountains (Italy), showing in detail the study area. supply is exclusively vertical, deriving from represented in the lower part of the massif. Wells rainfalls which tend to infiltrate vertically in few depth ranged from 4 to 375 m and water level hours to a maximum of few days. The fast from 0 to 230 m below ground surface. The subterranean karstic circuit presents a renewal of investigated wells were sealed and cased, being about 2-4 months. The subterranean flow provided of a steel, PVC or hydraulic cement direction of the whole Lessinian basin is from casing perforated in the saturated zone with slots north to south and parallel to the minor valleys of at least 3 mm, closed on the top and equipped direction. The total subterranean discharge is with a submerged pump connected to an engine. quantifiable in about 50 m3/s: 15 m3/s flow at the The wells were sampled once; before pumping, contact between limestones and the alluvial the water level was measured by an electrical sediments; 30 m3/s through the streambeds and water level meter. A quantitative volume of 500 l 5 m3/s at the karstic outlet of Montorio (Pasa, of water was pumped and filtered through a 60- 1954; Sorbini, 1993; Patrizi and Lavagnoli, µm mesh net at the outlet of the discharge pipes: 2001). the filtered sediment was collected in plastic vials and the biological component fixed with 7% 2.2 Sampling strategy formaldehyde solution. Per each sampling site, A stratified random sampling was performed in a the following chemico-physical analyses were macro-scale approach in the lower part of the carried out: temperature, pH, dissolved oxygen, Lessinian Massif covering an area of about 150 saturation oxygen, electrical conductivity, 2 km . While some water from the regional aquifer calcium (mg/l) and total hardness (mg/l) were is used for livestock watering, domestic supply measured in the field; magnesium (mg/l) and industrial purposes, the main water use, in concentration was determined by difference by the investigated area, is for irrigation. standard methods. Water samples for the

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Figure 2 : Abundance of taxa collected in karstic (left) and porous (right) aquifers.

determination of nitrate and phosphate were Focusing on the frequency of occurrence, poured in plastic bottles, twice washed with copepods showed the highest incidences in both deionised water and rinsed with pumped water. karstic and porous aquifers (Fig. 3). Nitrate (mg/l) and total phosphate (mg/l) According to these observations, copepods were concentrations were measured by a HACH DR selected as focal group, sensu Hammond 2000 spectrophotometer. Biological samples (1995), of the investigated communities and the were hand-sorted under a stereomicroscope and following analyses were focused on this taxon the collected speciments belonging to different only. According to the degree of adaptation to taxa (Nematoda, Oligochaeta, Mollusca, groundwater life and by integrating the definitions Copepoda, Ostracoda, Amphipoda, Isopoda, given by Notenboom (1991), Gibert et al. (1994) Syncarida, Insecta) were stored in separated and Galassi (2001), copepods were classified as vials and preserved in 70% alcohol solution and stygobionts (Sb), stygophiles (Sp) and identified at species or subspecies level, stygoxenes (Sx). In order to minimise whenever possible. misinterpretation errors, the use of the wider stygophile concept, considered as potential 3. DATA ANALYSIS AND RESULTS intermediate evolutionary step in the “stygobitization” process (Stoch, 1995), was 3.1 Groundwater biodiversity assessment avoided and the Sp attribute was limited to the The composition of faunal assemblages was eustygophile species. As expected, stygobiont analysed in order to assess the number of species were dominant in both karstic and stygobiont species in the saturated Lessinian porous aquifers, representing more than 60% of aquifers. As frequently observed in groundwater the taxocoenosis and occurring with 12 species habitats, Crustacea Copepoda were the most vs. a total of 19. abundant and species richest taxon. In the investigated groundwater sites, copepods 3.2 Selection of priority sites represented the 90% and 84% of the biological The selection of priority sites/aquifers took into samples collected respectively in karstic and account the Intrinsic Biological Value (I.B.V.) of porous aquifers (Fig. 2). the investigated sites independently from other

Figure 3 : Frequency of occurrence of taxa collected in karstic (left) and porous aquifers (right).

SWSB December 2004 117 SESSION 4 : CONSERVATION AND MANAGEMENT considerations (De Broyer et al., 2004). The preservation is here defined as the maintenance I.B.V. of a site was computed by summing the of the chemico-physical and structural features, Conservation Value (C.V.) of each collected through the application of management actions stygobiont species/subspecies. The C.V. was including interdictions, limitations and duties. As computed through an algorithm which averages the groundwater systems (both karstic and the value of the following stygobiont species porous) are characterised by a physical and features: endemism, rarity, habitat selection, hydrogeological continuity, the best choice in taxonomic isolation (Stoch et al., 2004). Twelve order to preserve groundwater habitats would be stygobiont copepod species were collected in the to safeguard the whole recharge basin. This Lessinian aquifers. In Appendix, the C.V. of each option may be excessive for porous aquifers species is reported together with the I.B.V of the which are self-protected by low permeable sites in which at least one copepod species was sediments, while it seems more appropriate for found. In order to preserve the 12 Sb species, a the karstic ones. As a consequence, two different network of priority sites was identified. The criteria are hereafter proposed. criterium utilised for detecting the priority sites The criterium selected for porous sites P6s and forming the network consisted in: 1) ordering the P9p consisted in the identification of a Restricted priority sites in function of decreasing values of Respect Zone (R.R.Z.) as defined by the Italian the I.B.V. (as in Appendix); 2) selecting the normative (Repubblica Italiana, 1999: D.Lgs. priority site with the highest I.B.V. value; 3) 152/99). Following the law recommendation, the selecting a second priority site with the highest isochrone of 30 days was considered the most I.B.V. value and harbouring one or more sustainable one for delimiting the R.R.Z. The stygobiont species not harboured in the previous sites P6s and P9p are located in two different selected site; 4) selecting a third, a forth, etc. alluvial valleys of the Lessinian area priority site until completing the entrance of all (respectively Squaranto and Valpantena valleys) stygobiont species. In the study case a network at 57 and 88 m a.s.l. and are utilised for irrigation. of 9 priority sites was selected (Appendix). The depth of the wells is 18 m (P6s) and 42 m Results hereafter illustrated deal with three of the (P9p) respectively. According to the priority sites (P6s, P9p, K13p). hydrogeological parameters, the permeability of the porous medium was estimated about 10-4 m/s 3.3 Selection of respect zones around the for both sites. According to permeability and priority sites transit-time values (30 days), the radius of the Following the Directive 92/43/EEC, the protection R.R.Z. resulted to be 259 m and the area about of a species may be realized by the preservation 10 km2 for both P6s and P9p. Indeed, the R.R.Z. of the habitat in which it lives. Habitat was thought as a semi-circle oriented along with

Figure 4 : Topographic map indicating the location of P9p (dot on the left) and K13p (dot on the right) sites. Black arrows indicate the subterranean flow direction. The R.R.Z. (Restricted Respect Zone) and the recharge area of the karstic site are reported too. The thick white arrow indicates the Pantena stream (scale 1:10000).

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P6s P9p K13p PPP

Water exploitation (agriculture) 1 1 1 Nitrates leaching (agriculture) 1 1 1 Pesticides leaching (agriculture) 1 1 1 Sewage leakage (urbanisation) 0.5 0 0 Motorways oil seeping (urbanisation) 1 1 0.5

Table 1 : Risk drivers and P (impact probability) of the selected priority sites. the main subterranean flow direction (the R.R.Z In Table 1, the risk drivers are reported together of the P9p site is represented in Fig. 4). The with the probability of the correlated impacts. The karstic site K13p (depth = 50 m) is located at an probability (P) assumed the values 0, 0.5 and 1, altitude of 76 m a.s.l., at the base of a Cenozoic which respectively correspond to a low, medium calcareous hill of about 400 m of height and is or high probability of the impact. utilised for irrigation. The recharge area to be 3.5 Assessment of aquifer intrinsic protected was estimated to be about 4 km2 and vulnerability no preferential infiltration pathways (eg., sinkholes) were identified for 2 km around the The Intrinsic Vulnerability (I.V.) refers to the site (Fig. 4). intrinsic characteristics of the aquifer and in particular to its own aptitude of swallowing and 3.4 Evaluation of the main risk drivers spreading a contaminant infiltrated from the Risk drivers are the underlying causes which surface (Civita, 1994). Thus defined, I.V. is lead to environmental negative impacts. In this distinct from pollution risk which depends on I.V. section a check-list of drivers is provided and also on the existence of a significant together with the probability of occurrence of the pollutant loading entering the subsurface negative impacts which may be produced. The environment. More in detail, I.V. increases when drivers were identified through information which the aquifer is interested by the presence of resulted from field recognitions and local surface water fast-infiltration pathways (as authorities database. Five risk drivers were sinkholes or gravel layers) which allow the identified, due to agriculture (water exploitation, entrance of pollutants in the saturated zone and nitrates leaching, pesticides leaching) and to act as preferential pathways for the epigean urbanisation (sewage leakage and motorways oil fauna, as well. As a consequence, stygoxene seeping). species may be collected at very elevated depth

Figure 5. : Abundance of stygoxenes (Sx) along with aquifer depth. On the horizontal axis, sites are ordered from the shallowest one to the deepest one.

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Table 2 : Mean and standard deviation of chemico-physical parameters in karstic (K) and porous (P) aquifers. T = temperature;

Cond. = electrical conductivity; O2 = dissolved oxygen; O2% = saturation oxygen; PO4 = total phosphates; NO3 = nitrates; Ca = calcium; Mg = magnesium. as in the investigated Lessinian sites. In the 29 concentrations (t = —3.37; p = 0.0011) and pH karstic sites, stygoxene species were distributed (t = 2.42; p = 0.01). at various depths reaching 240 m. On the On the basis of the above observations, the contrary, in the 47 porous media, the stygoxenes presence of stygoxenes in the 9 saturated sites is seemed to be confined to the uppermost layers likely to be indicative of the presence of of the aquifers, occurring frequently at low depths hydrological connections between groundwater (4-16 m) and sporadically and with low and surface water. More in particular, the abundances at more consistent depths (23 and presence of stygoxenes in deep, oligotrophic 120 m) (Fig. 5). groundwater environment is to be viewed as the The observed distribution of stygoxenes in the result of a fast passive transport operated by two aquifer typologies resulted not to be linked to infiltrating surface water. As far as the Crustacea chemico-physical factors, as karstic and porous Copepoda are concerned, the passive aquifers showed similar behaviour from this point displacement is facilitated by the presence of of view (Table 2). In regard to the 9 priority sites larval stages (nauplia and copepodids), prone to of the network of conservation, neither stygoxene be transported. Consequently, the occurrence of nor stygobiont species seemed to be affected by epigean copepods in deep aquifers may be high concentration of nitrate and/or phosphate or frequently interpreted as a result of hydro- by the oxygen depletion or by other chemical transport (Rouch, 1980). By the way, stygoxenes parameters (Table 3). may play the role of Active Exchange Describers The t-test for temperature (t = —1.14; p = 0.25), (A.E.D.), that is indicators of hydraulic exchange dissolved oxygen (t = 1.66; p = 0.1), saturation zones (surface water vs. groundwater), which oxygen (t = 1.60; p = 0.11), phosphate (t = —0.8; represent also the main infiltration pathways for p = 0.42), nitrate (t = —0.66; p = 0.5) and calcium pollutants (Lafont et al., 1992). The presence of concentrations (t = —1.74; p = 0.08) confirmed stygoxenes in shallow aquifers (both karstic and that there is no statistically significant difference porous) of the Lessinian area was somewhat between these parameter values in karstic and expected, due to the physical closeness of these porous aquifers at a α-probability = 0.05. The two aquifers to the surface. On the contrary, the aquifers typologies differ only for magnesium collection of alive stygoxenes in deep,

Table 3 : Abundances of Sx (stygoxene), Sp (stygophile), Sb (stygobiont) copepods and chemico-physical parameters of the 9 priority sites

forming the network for conservation (T = temperature; Cond = electrical conductivity; O2 = dissolved oxygen; O2% = saturation oxygen;

PO4 = total phosphates; NO3 = nitrate; Ca = calcium; Mg = magnesium).

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Table 4 : I.V. (Intrinsic Vulnerability) of the selected priority sites (I.V.m = I.V. from the hydrogeological map; Sx = presence of stygoxenes; I.V.i = implemented I.V. values). oligotrophic aquifers (both porous and karstic heavy metals, chloride and some pesticides ones) was taken as a warning of intrinsic seriously affect the survival of stygotaxa (among vulnerability as already pointed out by Malard et others, Mathews et al., 1977; Bosnak and al. (1994, 1998). Morgan, 1981a, b; Mösslacher and Notenboom As a result of this, an eco-hydrogeological 1999). On the other hand, the lowering of the approach, which combines the Sx-presence and water-table, due to an over-exploitation, may hydrogeological data for the assessment of produce reduction in both habitat dimension and aquifer intrinsic vulnerability, was applied. The heterogeneity with the consequent disturbance of I.V. values given by the I.V. hydrogeological map the communities. Rouch (1980, 1986) observed were increased of one unit if stygoxene that the recovery of karstic communities after an copepods were collected in the priority site under over-exploitation is slow but no more detailed consideration. The I.V. from the hydrogeological information is available on this topic. Similarly, map was already classified into 5 classes: 1 clear evidences about the effect of nitrates and (low); 2 (medium); 3 (high); 4 (very high); 5 sulphurous oxides on stygo-communities did not (elevated). The I.V. values derived by the eco- receive the necessary attention. In regard to the hydrological approach were therefore rescaled study area, high concentrations of nitrates (as from 1 to 6: 1 (low); 2 (medium); 3 (high); 4 (very observed in P20a where [NO3] = 56.8 mg/l) high); 5 (elevated); 6 (very elevated). With definitely did not affect the presence of regards to K13p site (3 Sx vs. 4 Sb), the Sx- stygobionts while it was observed that very low correction factor increased the I.V. from 1 to 2. In concentrations of heavy metals (few mg/l) have a Table 4 the I.V. values of P6s, P9p and K13p are strong effect on stygobiont species survival reported. (Mösslacher and Notenboom (1999) for a review). According to this, D values were 3.6 Computation of the risk assigned on the basis of the amount and The risk faced by an aquifer depends primarily on reliability of data from literature. Moreover, D of its natural defences rather than on the intensity demonstrated alteration effects should be a and type of the pollution source. According to multiple of the value assigned to the supposed this, the risk to which the priority sites are ones. The values proposed herein are D = 1 for exposed was computed as the product of three the damages derived from nitrate leaching, factors: Intrinsic Vulnerability (I.V.), Probability of sewage leakage and oil seeping, and a higher occurrence of the potential negative impacts (P) value (D = 3) for the damage caused by and intensity of the impact Damage (D). In this pesticides leaching. In the Lessinian area, the approach the impact Damage (D) is supposed to over-exploitation of the groundwater resources be a parameter which can assume only two occurs from July to September during the values: 1 (low damage) and 3 (high damage). vegetative period of the crops; in the other The damage intensity was defined on the basis months the wells are not utilised and the of the degree of alteration experienced by resource has the time to recover. For this reason, groundwater communities/habitats after the D = 1 was assigned to water exploitation, as well impact. Eco-toxicological tests demonstrated that (Table 5).

Table 5 : D (damage intensity) of the selected risk drivers.

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The Risk (R) a site experiences was computed enough oxygenated (Tab. 3). As a result of this, by the arithmetic average of the partial risks (D x the present management of the area is thought Px I.V.) multiplied to a factor 10 to avoid not to affect the groundwater biota and therefore decimals. In the Lessinian area R ranges from 0 the strategy proposed is conservative. to 84. Three levels of risk were selected dividing the risk range in three equivalent classes of 4. CONCLUSION values: low (0-28); moderate (29-56) and high One of the main purposes of the European and (57-84). The risk values for the selected priority Italian water policies is the pursuing of the sites are the following :

P6s: R = {[(1x1x3)+(1x1x3)+(3x1x3)+(1x0.5x3)+(1x1x3)]/5}x10 = 39

P9p: R = {[(1x1x4)+(1x1x4)+(3x1x4)+(1x0x4)+(1x1x4)]/5}x10 = 48

K13p: R = {[(1x1x2)+(1x1x2)+(3x1x2)+(1x0x2)+(1x0.5x2)]/5}x10 = 22

which correspond respectively to moderate (P6s sustainable use of the water resources and P9p) and low (K13p) levels of risk. (2000/60/EC; D.Lgs. 152/99). Similarly, it is also stated that the status of aquatic ecosystems 3.7 Proposal of signposts for (including surface waters, costal waters and sustainability groundwaters) has to be enhanced and The conservation actions a site requires were preserved from further deterioration estimated on the basis of the risk it faces. The (2000/60/EC, art. 1) through management purpose at the basis of the approach is to practices which guarantee the water bodies to suggest conservation actions which are support wide and well diversified faunistic and sustainable in the Lessinian area which is low- floristic communities (D.lgs 152/99, art. 1). On industrialized and low-densely populated. the other hand, in both normatives, neither the According to this, feasible and low-cost ecological status of groundwater nor the management procedures were suggested for the importance of groundwater biota as indicators of moderate-risked sites (P6s and P9p), while groundwater status have been yet recognised. conservative actions were proposed for the low- As a matter of fact, although the knowledge risked aquifer (K13p). Into the R.R.Z. of the about the groundwater ecosystem has greatly porous sites and in the recharge area of the increased in the last years, the dissemination of karstic one, some interdictions are information about groundwater biota and recommended, such as on gravel extraction, ecosystems is still scarce. Particularly in Italy, dumps and mine openings (Sket, 1999). More in these topics rarely overcome the boundaries of particular, the utilization of more efficient the scientific academic world which should face irrigation techniques, such as drip irrigation, is the challenge of widespreading information. The suggested in the R.R.Z. of P9p, in order to approach here proposed applies to the Italian reduce the water loss with the consequent context the procedure developed by De Broyer et decreasing of the water-table. Furthermore, P9p al. (2004) within the framework of the PASCALIS is located very close to Pantena stream (2002) project. The procedure was focused on (indicated by the white thick arrow in Fig. 4) so the paradigm of sustainability for groundwater that it would be possible to utilize also stream through the endorsement of the biota water for irrigation; however, because of the low conservation in the use of the resources. The aim discharge of the stream, this practice should be of the approach was to preserve groundwater pursued only in the first weeks of the vegetative species through the conservation of their natural period when the water need of the crops is low. habitats. An index of conservation values (C.V.) The priority site P6s is located downstream the for stygobiont species (Stoch et al., 2004) was urbanised area of Montorio (Verona Province) utilised in order to identify a network of and its R.R.Z. includes part of the urban groundwater sites priority from a biological point agglomerate. The conservation measures for this of view. According to the economic context of the site should foresee the sewage pipes Lessinian area, some management actions were maintenance in order to avoid the leakage of proposed with the aim to avoid further contaminants. The recharge area of K13p is groundwater habitat reductions and interested by agriculture and small manures, contaminations. The proposed conservation however, the nitrate and phosphate procedures were applied to limited areas such as concentrations are low and the groundwater is R.R.Z. (Restricted Respect Zones) and recharge

122 SWSB December 2004 SESSION 4 : CONSERVATION AND MANAGEMENT areas around the priority sites in order to spare Council of 21st May 1992. Conservation of natural revisions of regional planning in terms of soil habitats and wild fauna and flora. Official Journal of uses and management practices. The use of the European Communities L 206:1-57. A.E.D. (Active Exchange Describers) in the European Community. 2000. Directive 2000/60/EC of the rd assessment of the intrinsic vulnerability of European Parliament and of the Council of 23 October 2000. Establishing framework for community aquifers was usefully proposed in this approach. action in the field of water policy. Official Journal of the On the other hand, the procedure requires further European Communities L 327:1-7. standardization. As a matter of fact, the list of risk European Community. 2003. Proposal for a Directive of drivers is far from being complete and some the European Parliament and of the Council on the other drivers, occurring in other areas (e.g., salt protection of groundwater against pollution. Brussels, water infiltration), should be included. 19th September 2003, COM 2003/0210 (COD) [www Accordingly to this, the risk range is subjected to document]. URL http://europa.eu.int/eur - further revision. Hence, the proposed procedure, lex/en/com/pdf/2003/com2003_0550en01.pdf. which fitted to the Lessinian sites, is in need of Galassi, D.M.P. 2001. Groundwater copepods: diversity validation and adjustments in order to be usefully patterns over ecological and evolutionary scales. Hydrobiologia 453/454:227Ð 253. applied at national and European level. Gibert, J., J.A. Stanford, M.-J. Dole-Olivier, and J.V. Ward. 1994. Basic attributes of groundwater ecosystems and prospects for research. In: J. Gibert , D. L. ACKNOWLEDGEMENTS Danielopol & J.A. 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Comparison of describe groundwater flow and contaminant transport acute toxicity for Cd, Cr, Cu between two distinct populations of aquatic hypogean isopods (Caecidotea in a fractured rock aquifer. Arch. Hydrobiol. 131 sp.). 8th International Congress of Speleology 1:72- (1):93Ð 110. 74. Malard, F., J.-L. Reygrobellet, and Laurent, R. 1998. Bosnak, A.D. and Morgan, E.L. 1981b. Acute toxicity of Spatial distribution of epigean invertebrates in an cadmium, zinc and total residual chlorine to epigean alluvial aquifer polluted by iron and manganese. and hypogean isopods (Asellidae). NSS Bulletin Rhône River, France. Verh. Int. Ver. Limnol. 26:1590- 43:12-18. 1594. Civita, M. 1994. Le carte di vulnerabilità degli acquiferi Malard, F., M.-J. Dole-Olivier, J. Mathieu, and Stoch, F. all’inquinamento: teoria e pratica. Ð Quaderni di 2002. Sampling manual for the assessment of Tecniche di protezione ambientale. Pitagora [Ed.], regional groundwater biodiversity. European project Bologna. PASCALIS, available at http://www.pascalis- project.com. Danielopol, D.L., J. Gibert, C. Griebler, A. Gunatilaka, H.J. Hahn, G. Messana, J. Notenboom, and B. Sket. 2004. Mathews, R.C., A.D. Bosnak, D.S. Tennant, and E.L. Incorporating ecological perspectives in European Morgan. 1977. Mortality curves of blind cave crayfish groundwater management policy. Environ. Conserv. (Orconectes australis australis) exposed to 31(3):1Ð5. chlorinated stream water. Hydrobiologia 53 (2):107- 111. De Broyer, C., G. Michel, P. Martin, F. Malard, F. Stoch, and J. Gibert. 2004. Action plan for conservation of Mösslacher, F. and Notenboom, J. 1999. Groundwater groundwater biodiversity. PASCALIS D9 for Biomonitoring. In: A. Gerhardt [ed.], Biomonitoring of Workpackage 9. European project PASCALIS polluted waters. Trans Tech Publ., Zurich. 117-139. (Protocols for the Assessment and Conservation of Notenboom, J. 1991. Marine regression and the evolution Aquatic Life in the Subsurface). (unpublished report). of groundwater dwelling amphipods (Crustacea). J. European Community. 1991. Directive 91/271/EEC of the Biogeogr. 18:41-65. European Parliament and of the Council of 21st May Notenboom, J. 2001. Managing ecological risks of 1991. Urban water treatment. Official Journal of the groundwater pollution. In: C. Griebler, D.L. European Communities L 135:40-52. Danielopol, J. Gibert, H.P. Natchtnebel, and J. European Community WD. 1991. Directive 91/676/EEC of Notenboom [eds.], Groundwater ecology. A tool for the European Parliament and of the Council of 12th management of water resources. Austrian Academy December 1991. Protection of waters against of Sciences, Institute of Limnology Vienna, Mondsee. pollution caused by nitrates from agricultural sources. pp. 183Ð196. Official Journal of the European Communities L Pasa, A. 1954. Carsismo ed idrografia carsica nel gruppo 375:1-8. del M. Baldo e nei Lessini veronesi. Consiglio European Community. 1992. Directive 92/43/EEC of the Nazionale delle Ricerche, Bologna, pp. 1Ð137.

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PASCALIS. 2002. Protocols for the Assessment and Sauro, U. 1973. Il paesaggio degli alti Lessini. Studio Conservation of the Aquatic Life In the Subsurface. geomorfologico. Mus. Civ. St. Nat. Verona. Mem. F.S., Contract no EVK2-CT-2001-00121 - PASCALIS of the 6:1-161. Fifth Research and Technological Development Sket, B. 1999. The nature of biodiversity in hypogean Framework Program supported by the European waters and how it is endangered. Biodiversity and Community. Conservation 8:1319Ð 1338. Patrizi, G. and M. Lavagnoli. 2001. Indagine Sorbini, L. 1993. Geologia, idrogeologia e qualità dei idrogeologica, geochimica e geochimica-isotopica principali acquiferi veronesi. Mem. Mus. Civ. St. Nat. sugli acquiferi della Lessinia. Rapporto finale. Verona (II serie). Sezione Scienze della Terra, 4: Provincia di Verona, Settore Ecologia, pp.1Ð96 . 1Ð150. Repubblica Italiana. 1999. Decreto Legislativo 11 Maggio Stoch, F. 1995. The ecological and historical determinants 1999, n. 152. Disposizioni sulla tutela delle acque of crustacean diversity in groundwaters, or: why are dall’inquinamento e recepimento della direttiva there so many species? Mém. Biospéol. 22:139Ð160. 91/271/EEC concernente il trattamento delle acque reflue urbane e della direttiva 91/676/EEC relativa alla Stoch, F., F. Malard, , F. Castellarini, and M.-J. Dole- protezione delle acque dall’inquinamento provocato Olivier. 2004. Proposal of method for assessing the dai nitrati provocati da fonti agricole. G.U. 29 Maggio conservation value of species. In: Statistical analyses 1999, n. 124, S.O. and identification of indicators. PASCALIS D8 Deliverable for Workpackage 7:121-149. European RIVM/RIZA. 1991. Sustainable use of groundwater: project PASCALIS : Protocols for the Assessment and problems and threats in the European Communities. Conservation of the Aquatic Life In the Subsurface. Report prepared for Ministersseminar Groundwater, (unpublished report). th 26-27 November 1991. RIVM-report 600025001. Vial, G. and O. Tondelli. 1997. Mappa della vulnerabilità Rouch, R. 1980. Le système karstique du Baget. X. La idrogeologica intrinseca, Tav. 2 del Piano Territoriale communauté des Harpacticides. Richesse spécifique, Provinciale, Amministrazione della Provincia di diversité et structure d’abondance de la nomocénose Verona. hypogée. Ann. Limnol. 16 (1):1Ð20. Rouch, R. 1986. Sur l’écologie des eaux souterraines dans les karst. Stygologia 2:352-398.

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Appendix. The C.V. (Conservation Value) of each stygobiont species is reported together with the I.B.V. (Intrinsic Biological Value) of the 42 sites in which at least one copepod species was found. The 9 priority sites forming the network are in bold and the new-entered species are underlined

SWSB December 2004 125

SESSIONS ABSTRACTS

SESSIONS

Oral communication abstracts

SWSB December 2004 127

SESSIONS ABSTRACTS

DISTRIBUTION OF THE SPECIOSE GENUS VESTALENULA, ROSSETTI & MARTENS, 1998 (DARWINULIDAE, OSTRACODA), WITH THE DESCRIPTION OF A NEW SPECIES FROM THE ROUSSILLON REGION (SOUTH-EASTERN FRANCE).

M. ARTHEAU

Laboratoire Dynamique de la Biodiversité, UMR/CNRS 5271, UPS, 118 route de Narbonne, 31062 Toulouse cedex 4, France. ([email protected]).

A new species of Vestalenula Rosseti & Martens, 1998, belonging to the danielopoli-lineage is described. It was found during the PASCALIS project when sampling 5 hyporheic sites on the Aude and Tech rivers (Roussillon region, South Eastern France). This new record of the genus brings to 20 the number of Vestalenula species worldwide. Vestalenula representatives are known from the Lower Miocene to Present. The biogeography of this group will be reviewed in order to stress out the interest of the discovery of a new species in Southern France. Most living species of Vestalenula occur in subtropical regions. V. cylindrica, V. boteai and V. danielopoli lived or are living in the Palearctic. V. pagliolli occurs in both hemispheres. The other 16 are Recent species, 12 of them are living in the southern hemisphere. The Recent species occur in restricted areas and many live in semi-terrestrial habitats and/or interstitial habitats. The morphological characters which enable the Vestalenula species to be successful in colonizing groundwater habitats will be discussed. This work has been done during a stay in June 2004 at the Limnological Institute of the Austrian Academy of Sciences in Mondsee, Austria.

CLADOCERA AND CALANOIDA TWO INTERESTING GROUPS OF STYGOBIONTS

A. BRANCELJ and N. MORI

National Institute of Biology, Vecna pot 111, 1000 Ljubljana, Slovenia. ([email protected]), ([email protected]).

Crustacea are well represented in subterranean environment in terms of number of taxa, with Copepoda (Harpacticoida, Cyclopoida), Amphipoda and Isopoda being the most numerous. Exceptions among Crustacea are Anostraca, Conhostraca and Notostraca with no known stygobionts. Cladocera and Calanoida (Copepoda) are very well represented in most types of fresh-water bodies, each with about 500 taxa all over the world, but they have only few representatives in subterranean environment. Members of both groups were found relatively late in different types of groundwater. The first cladoceran recognised as stygobiont was discovered in 1973 (Alona smirnovi) and the first stygobiotic member of Calanoida was found in 1942 (Microdiaptomus cokeri). So far, only six stygobionts and additional two might-stygobionts are known among Cladocera and eight stygobionts and two might-stygobionts among Calanoida. The number of new stygobionts from both groups increased in the last few years due to intensive patterns of geographical distribution of representatives of both groups, their morphological and ecological adaptations to subterranean environment as well as appropriate sampling techniques will be discussed.

SWSB December 2004 129 SESSIONS ABSTRACTS

USE OF GROUNDWATER COPEPODS AS A TOOL FOR ENVIRONMENTAL MANAGEMENT : THE CASE OF EVERGLADES NATIONAL PARK, FLORIDA, USA

M.C. BRUNO and V. COTTARELLI

Università della Tuscia, Dipartimento di Scienze Ambientali, Largo dell’Università, 01100 Viterbo, Italy. ([email protected]), ([email protected]).

Everglades National Park (ENP), is an extensive subtropical wetland ecosystem marsh ; its hydrology changed radically from the 1950s as a result of the construction of a canal system, which drained water eastwards and southwards. The Rocky Glades is an area of higher elevation in the Eastern part of ENP, with short hydroperiod, characterized by a karstic landscape with thousands of solution holes, which are filled with water during the wet season (Jun.-Oct.), and are dry in the dry season (Nov.-May). We collected copepods from different depths in the superficial aquifer in 2 sets of wells in the Rocky Glades; and in a third set of wells along the canals bordering ENP to the east. Objectives of the three studies were to assess : 1) stygoxenic and stygophilic species and their spatial and temporal distribution ; 2) how seasonal changes in groundwater levels influence the ability of Everglades surface copepods to disperse into groundwater to survive the drought ; 3) how local geological features influence faunal exchange between surface and groundwater. Groundwater communities were dominated by surface copepods ; abundance and diversity decreased with depth, due to different limestone permeability, and copepods did not disperse far from the input location. In the Rocky Glades, surface copepods colonized groundwater during the dry season, with peaks at the beginning and at the end of the wet season.. In canals bordering ENP, copepods colonized groundwater following groundwater seepage along canals. Recommendation for the management included : 1) solution holes in the Rocky Glades act as an exchange area between surface and groundwaters : over drainage should be avoided to preserve ENP biodiversity; 2) the use of copepods as indicators of seepage could be a tool to assess the direction and the duration and the direction of surface and ground water exchange ; 3) the minimum target water levels for the Rocky Glades had been set at 45 cm below the surface. This level might be too low for copepods to migrate vertically into the aquifer, or to survive adverse physical and chemical conditions in groundwater.

130 SWSB December 2004 SESSIONS ABSTRACTS

COI DATA INDICATE A COMPLEX EVOLUTIONARY HISTORY FOR ITALIAN SUBTERRANEAN AND SURFACE POPULATIONS OF PROASELLUS (CRUSTACEA, ISOPODA).

A. CAMPANARO, V. KETMAIER and R. ARGANO

Department of animal and Human Biology, University of Rome “La Sapienza”, V.le dell’Università 32, 00185 Rome, Italy. ([email protected]), ([email protected]), ([email protected]).

The water louse species of Proasellus are widely distributed in the surface and subterranean freshwaters of the Mediterranean region. We investigated patterns of genetic divergence and phyletic relationships in 40 populations of Proasellus, sampled in continental Italy and in Sardinia, sequencing a fragment of the mitochondrial gene coding for the cytochrome oxidase I (COI; 411 bp). The study includes 26 surface populations from continental Italy, two subterranean species from Central Italy, P. acutianus and a new (still non described) species (P.n.sp.), and 12 both surface and subterranean populations from Sardinia. Phylogenetic and network analyses revealed the existence of two well-supported clades. The first clade is characterised by a relatively low degree of genetic heterogeneity and groups all the surface populations from continental Italy and three surface populations from Sardinia. The second clade includes all the remaining populations from Sardinia, P. acutianus and P.n.sp. This group shows a high level of genetic heterogeneity, suggesting several different cladogenetic events and multiple independent invasions of subterranean environments. The placement in the same clade of all the subterranean populations, independently from their geographic origin, suggests that these taxa could be extant representatives of an ancient (Miocenic) wave of colonisation. Divergence time estimates, calculated using published COI rate for Stenasellus, indicate that the split between the two clades occurred around 30 myr ago and should be related to the tectonic events leading to the displacement of the Sardinia-Corsica micro plate from southern continental Europe. Previous morphological and allozyme data on the P. coxalis-complex are also discussed.

SWSB December 2004 131 SESSIONS ABSTRACTS

PATTERNS OF ENDEMISM OF THE EASTERN NORTH AMERICAN CAVE FAUNA

1D.C. CULVER, 2M.C. CHRISTMAN, 3M. MADDEN, and 4D. WHITE.

1. Department of Biology, American University, Washington DC 20016 USA. ([email protected]) 2. Department of Animal and Avian Sciences, University of Maryland, College Park MD 20742 USA. 3. Environmental Studies Program, American University, Washington DC 20016 USA 4. Environmental Protection Agency, Corvallis OR 97333 USA

Over 250 species of obligate terrestrial cave-dwelling animals (troglobionts) are known from single caves in the eastern United States. We investigated their geographic distribution especially in relation to non- single cave endemic troglobionts. We related these patterns to taxonomic group, opportunities for dispersal, and geographic location. We associated over 3000 records of over 450 troglobiotic species and subspecies with hexagons of 1,000, 5,000, and 10,000 km2 in size. We found Moran’s I, black-white joins, and cubic regression of endemics on non-endemics at all three spatial scales. For 5,000 km2 hexagons, we found the spatial autocorrelation of the residuals of cubic regression of endemics on non-endemics. Differences among orders in percent single cave endemism were not significant except for Pseudoscorpionida which was higher (67%) than any other order. At all three scales, Moran’s I and black- white joins were significant, indicating a clumped distribution of both single-cave endemics and other troglobionts. Spatial patterns were similar at all three scales and Moran’s I was highest at 5000 km2. The cubic fit of endemics to nonendemics was consistently better with less systematic error or residuals than was linear or quadratic models. Residuals showed a significant geographic pattern with excess endemics in more southerly locations. There was both a non-spatial and spatial component to the pattern of single cave endemism. The non- spatial component was the association of high levels of single cave endemism with areas of high diversity of non-endemics. It may be that both are high because of high secondary productivity. Spatially, single- cave endemism is high in central rather than peripheral areas and in the southern part of the range. It is not higher in areas of more dissected limestone which would reduce migration rates. Regional spatial effects are important indicating that cave communities cannot be understood (or protected) in isolation.

ARE BAYESIAN INFERENCES A USEFUL TOOL FOR PHYLOGEOGRAPHERS ?

C.J. DOUADY and T. LEFÉBURE

University Claude Bernard of Lyon 1, UMR CNRS 5023, Laboratoire d’Ecologie des Hydrosystèmes Fluviaux, Plate-forme d’Ecologie Moléculaire des Hydrosystèmes Fluviaux & Equipe d’Hydrobiologie et Ecologie Souterraines, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France. ([email protected]), ([email protected]).

Usage of Bayesian inferences to reconstruct phylogenetic relationships between organisms is exponentially increasing. This method has been wildly accepted for both theoretical and pragmatic reasons. It is indeed quite satisfying to be able to take into account the prior knowledge on the probability of a change, to obtain in a single analysis an estimation of reconstruction reliability and to do this in a fraction of the time necessary to frequentist methods (i.e. Maximum likelihood inferences). Surprisingly, most phylogeographic analyses realized in the course of PASCALIS program revealed an important discrepancy in branch lengths between Bayesian and Maximum Likelihood methods. Branch lengths being one of the most important phylogenetic / phylogeographic information, after the branching pattern itself, we here investigate this behavior using 38 real data sets and 100 simulated ones. We report our first observations and discuss theirs consequences for phylogeographic usage of Bayesian inferences. Finally a new bioinformatic tool developed to allow user-friendly bootstrapping of Bayesian inferences is presented.

132 SWSB December 2004 SESSIONS ABSTRACTS

HYPORHEIC COPEPOD ASSEMBLAGES AS A TOOL FOR DETECTING MAN-INDUCED PERTURBATION ON GW/SW ECOTONE

B. FIASCA, T. DI LORENZO, P. DE LAURENTIIS and D.M.P. GALASSI

Dipartimento di Scienze Ambientali, University of L’Aquila, Via Vetoio, Coppito, 67100 L’Aquila, Italy. ([email protected]), ([email protected]), ([email protected]).

The comparative analysis of hyporheic copepod assemblages was carried out in order to focus on the effects of surface anthropogenic activities on groundwater/surface water ecotone. The streams under study are tributaries of the River Adige and are fed by the Lessinian regional aquifer (Verona, northern Italy). The hydrology is strongly influenced by meteoric events and the valleys under study show quite marked differences in land use and resources exploitation. The sampling sites were selected and sampled once according to the PASCALIS standardized sampling procedure. Each sampling site consisted of four biological and four chemico-physical replicates. Specific conductivity, temperature, dissolved oxygen, pH, calcium, nitrates and phosphates were measured. Moreover, the sediment volume per each sample was assessed and the granulometric composition of the substrate was estimated. Most of the species showing high ecological plasticity co-occurred in the hyporheic zone of all streams. The remaining ones were clearly partitioned. Descriptive analyses and multivariate statistical methodologies were used in order to compare chemistry, physical parameters and geomorphological conditions of hyporheic zones, to check the occurrence of differences in composition and structure of the taxocoenoses and possibly assess correlations between environmental features and copepod assemblages. Temperature, pH and concentration of nitrates, phosphates and dissolved oxygen seemed to play an important role in structuring the hyporheic copepod assemblages; hydraulic conditions and substratum properties, on the contrary, could be helpful to describe site-specific attributes, just related to a chemico-physical gradient along each altitudinal stream profile.

POPULATION STRUCTURE OF STYGOBITIC ISOPODS IN THE PILBARA, WESTERN AUSTRALIA

C.J. FRANCIS, M.S. JOHNSON and T. FINSTON

School of Animal Biology, The University of Western Australia, Stirling Hwy, Perth, WA 6014, Australia. ([email protected]), ([email protected]), ([email protected]).

The arid northern region of Western Australia contains one of the world’s most diverse and unique assemblages of stygofauna. In the Pilbara, the occurrence of locally restricted fauna has become a major environmental issue, as it coincides with extensive mining industries. Conservation of this endemic and relictual fauna is hampered by limited knowledge of species diversity and distributions. An understanding of the connectivity of populations and mechanisms generating genetic diversity is urgently needed for the management of stygofauna in this region. The application of molecular markers may be the only way to reveal the true level of biodiversity in stygobitic fauna. Therefore, I am using a molecular phylogenetic approach to establish conservation priorities in the Pilbara region, focussing on isopod, copepod and ostracod species. As the first stage of this study, the population structure of stygobitic isopods within and between three catchments was examined using mtDNA cytochrome oxidase I (COI) sequences. Genetic variation was assessed within the context of local hydrogeological structure.

SWSB December 2004 133 SESSIONS ABSTRACTS

ECTINOSOMATIDAE (COPEPODA, HARPACTICOIDA) IN GROUNDWATER: RADIATION OR MULTIPLE INVASIONS?

1R. HUYS, 2D.M.P. GALASSI and 2P. DE LAURENTIIS

1. The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. ([email protected]) 2. Dipartimento di Scienze Ambientali, University of L’Aquila, Via Vetoio, Coppito, 67100 L’Aquila, Italy. ([email protected])

The family Ectinosomatidae currently contains 20 genera, showing high radiation in the marine environment. Only secondarily, members of the family, belonging to the genera Ectinosoma, Halectinosoma, Arenosetella and Pseudectinosoma, entered true freshwater habitats and, to less extent, groundwater. A recent discovery from groundwater in India added the new genus Rangabradya to the present-day known genera of the family. A sampling campaign in the porous aquifers of the Lessinian Mountains (Italy) revealed the presence of an additional member of the family, whose taxonomic position required a wider analysis on phylogenetic informative morphological characters, and a revision of the family in order to evaluate the monophyly of some questionable genera. Based on that, the new groundwater population is attributable to an undescribed genus of the family, rising the question whether the colonization of groundwater by ectinosomatids occurred only once in the evolutionary history of the family or alternatively by multiple invasions which took place independently, in different times and by different ways. A phylogenetic background is presented in order to evaluate the most plausible colonization patterns in groundwater of different members of the family. Three different lineages at least have been envisaged, which colonized groundwater following different ways: the Arenosetella-lineage by marine psammic environment, the Bradya-Ectinosoma-Halectinosoma-Rangabradya-lineage which probably passed the limnicoid surface habitat before entering in groundwater and the Sigmatidium- Noodtiella-Lineosoma-Pseudectinosoma-lineage which entered fresh groundwater through crevicular karstic habitats from epibenthic/epiphytic brackish environment.

GROUND-WATER FLOW AND AQUATIC HABITATS

W.K. JONES

Karst Waters Institute, P.O. Box 537, Charles Town, WV 25414, USA. ([email protected]).

Description of ground-water habitats are typically based on physical and ecological characteristics of the sample location where certain animals are collected. The most important ecological variables controlled by the nature of the subsurface pore spaces are limits to animal size, the residence time of the water, and the flux of nutrients. Most aquifers are a mixture of different size pore spaces and water chemistry and therefore contain different potential habitats so sampling results will be influenced by the scale of the sampling or capture method employed. For consolidated rock aquifers, most of the total volume of water in storage is in the less permeable but high porosity intergranular pore space, but most of the flow is through fractures or conduits. These higher permeability flow routes also represent preferential routes for the flux of nutrients or energy through the aquifer. Wells or bore holes will generally have a restricted capture zone and may not intercept the more permeable but highly localized flow paths that actually control much of the ground water flow. The results from sampling the interstitial habitats at wells should give variable capture counts, but the population assemblages will be somewhat limited, at least with respect to physical size. At the other extreme of sampling scale, collecting at a large karst spring may represent a “lumped parameter” model ranging from laminar flow through the epikarst zone to open channel flow through caves and covering a potential capture area of tens of square kilometers. In general, collecting at areas of concentrated ground-water flow will yield higher animal counts and greater species diversity. Flow is concentrated at zones of higher permeability and along stratigraphic contacts with less permeable units. Soil moisture is stored and held under tension if below saturation. Sampling in the unsaturated zone is often controlled by precipitation events and may result is a sort of “piston” flow where old water is displaced by new event water.

134 SWSB December 2004 SESSIONS ABSTRACTS

RIVERSCAPE DYNAMICS AND THE DISTRIBUTION OF THE HYPORHEOS IN A GLACIAL RIVER-FLOODPLAIN SYSTEM

1F. MALARD and 2M. LAFONT

1. CNRS-University Claude Bernard of Lyon 1, Laboratoire d’Ecologie des Hydrosystèmes Fluviaux, 43, Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France. ([email protected]). 2. UR Hydrobiologie, Cemagref, 3 bis Quai Chauveau, C.P. 220, 69336 Lyon, France. ([email protected]).

River-floodplain systems are dynamic riverscapes, the extent, composition, and configuration of which vary at several temporal scales. The biodiversity of a habitat not only depends on its environmental attributes but also on its age, shape, and position within the riverscape that controls the exchange of organisms with neighboring habitat patches. We developed a functional habitat classification that integrated the suitability, life span, and position of water bodies within the riverscape for testing hypotheses on biodiversity patterns in a pulsing glacial river-floodplain system. We expected that the pattern of biodiversity in the hyporheic zone of the floodplain would reflect the intersection between dissimilar life cycles and dispersal capacities among insects and permanent aquatic taxa and the suitability, life span, and position of habitats. Flooding resulted in a highly fragmented mosaic of habitats, so that newly created suitable habitats (i.e. clear water channels) were separated from source areas of colonizers by inhospitable habitats (i.e. turbid water channels). Because of their high dispersal capacities, insects colonized equally the hyporheic zone of suitable habitats that differed, however, in their life span and position within the riverscape. As turbid water expanded throughout the floodplain, insects became increasingly more abundant in newly created suitable habitat patches. Permanent aquatic taxa preferentially colonized hyporheic sediments of permanent suitable habitats but they failed to develop abundant populations in newly created suitable habitats, some of which had continuous flow for 8 months. The dispersal of permanent aquatic taxa within the riverscape was apparently constrained by the spatial arrangement of suitable and inhospitable habitats and the weakness of hydrological connections between hyporheic sediments and underlying ground water.

SWSB December 2004 135 SESSIONS ABSTRACTS

EPIKARST FAUNA FROM NORTH AMERICAN CAVES

1T. PIPAN and 2D.C. CULVER

1. Karst Research Institute ZRC SAZU, Titov trg 2, 6230 Postojna, Slovenia. ([email protected]). 2. American University, Department of Biology, 4400 Massachusetts Ave. NW, Washington DC 20016, USA. ([email protected]).

Drip pools and trickles of percolation water in 17 caves in West Virginia were sampled for epikarstic species, using aspirators and fine-mesh nets designed to capture copepods and other minute invertebrates. A total of 20 genera of copepods were found. 11 different genera of harpacticoids and 9 of cyclopoids were recognized in the samples collected from pools. In addition to copepods there were also specimens of Nematoda, Gastropoda, Troglochaetus among Archiannelida, Oligochaeta, Acarina, Ostracoda, Bathynellacea, Isopoda, Amphipoda, Millipedia, Collembola, Coleoptera and insects larvae. Results of the correlation analyses indicate that there was correlation between copepod abundance and number of species on the one hand and the quantity of pumped water on the other hand. For understanding of the structure and physico-chemistry of epikarst, the focus was specially on trickles rather than pools. We choose three sampling sites of the Organ cave system: the Lipps, Sively 2 and Sively 3 streams. In Lipps streams we had collection of 5 drips within 4 m of each other. In passages Sively 2 and Sively 3 we found 4 trickles at distances of up to 200 m. Sampling and water chemistry measurements were undertaken at the sites at approximately 10-day intervals. All together in drips we found 13 genera of copepods, from which 8 of harpacticoids and 5 of cyclopoids. Monitoring of trickles tens or hundreds of meters apart showed that there is statistically significant negative correlation between the distance between the drips and the similarity of the fauna. Due to horizontal movement of water in epikarst it is hard to say if each drip represents an isolated habitat or nearby drips come from the same habitat. The positive correlation of abundance and taxonomic richness in drips is consistent with faunistic exchanges between microhabitats in epikarst. The epikarst fauna will add significantly to the overall species richness found in Appalachian caves, and likely elsewhere in the U.S. as well.

136 SWSB December 2004 SESSIONS ABSTRACTS

THE RELATIONSHIP BETWEEN OSTRACOD SPECIES DIVERSITY AND WATER CHEMISTRY IN THE SUBSURFACE OF THE PILBARA REGION, WESTERN AUSTRALIA

J. REEVES

The Australian National University, Department of Earth and Marine Sciences, Canberra, 0200 ACT, Australia. ([email protected]).

The Pilbara region, north-western Australia provides an eclectic array of environments for the study of subterranean fauna. Although climatically regarded as semi-arid with annual evaporation outweighing preci-pitation by around 10:1, the Pilbara is located at the tropical fringe and receives rainfall from cyclonic events during the summer months. At such times, the strength of the water inundates roads throughout the floodplains and commonly causes damage to bridges.

The region may be divided broadly into the ranges, floodplain and coastal zones. The ranges date to the Early Proterozoic Ð Archaean and include the metasedimentary Hamersley Ranges in the central Pilbara, and the predominantly volcanic Chichester Ranges to the north. These units overlie the Archaean greenstone and granite, which outcrop to the northeast of the region. The Fortescue and Ashburton Rivers form floodplains either side of the Hamersley’s, and the Robe, Yule and De Grey Rivers extend as broad deltas toward the coast. The significant aquifers in the region include recent valley-fill alluvium and colluvium, chemically deposited calcretes and pisolites associated with former watercourses, and fractured-rock dolomite, banded-iron formations and granite. Groundwater in the region is typically fresh to brackish and bicarbonate-dominant, although Na-Cl-rich waters are common in both the coastal and arid eastern margins.

This paper presents analyses from the first two seasons of the Pilbara Stygofaunal survey (see Eberhard and Halse, this volume). The distribution of ostracods is highlighted as an example of the faunal diversity. Around 20 species of the family Candoninae have recently been identified from this region, and at least a further 20 taxa are yet to be described. The evolution of the groundwater through the aquifer is detailed in terms of major-ion and stable-isotope geochemistry of the waters, in conjunction with measured physical parameters, and related to processes of weathering, evaporation and solute evolution. The relationship between these patterns and species distribution is discussed.

SWSB December 2004 137 SESSIONS ABSTRACTS

THE BLIND MISLEADING THE BLIND : FALSE NEGATIVES AND ESTIMATIONS OF SUBTERRANEAN BIODIVERSITY

1, 2K. SCHNEIDER, 1D.C. CULVER, 3H.H. HOBBS III, 1D. FONG

1. American University, 4400 Massachusetts Avenue NW, Washington DC 20016, USA. ([email protected]), ([email protected]). 2. University of Maryland, 1205 Biology-Psychology Building, College Park MD 20742, USA. ([email protected]). 3.Wittenberg University, PO Box 720, Springfield, OH 45501, USA. ([email protected]).

As the cave habitat is difficult (and expensive) to sample, most of what is known about cave biodiversity is based on a fraction of the world’s caves, and at that, predominately large, easily accessible caves. But how representative are these caves of local or regional biodiversity? How many species are missed by only focusing on a small subset of the underground realm? How inaccurate and/or misleading is our knowledge of subterranean biodiversity and biogeography? We set out to answer these questions by thoroughly investigating every cave in a high cave density and species rich area of West Virginia, U.S.A. We sampled 65 caves within a 20 km2 study site. As a result of intensive sampling that included visual censuses and trapping procedures, 18 obligate stygobionts and troglobionts were collected. Only 7 caves were needed to collect 95% of the species, and by sampling only the largest 7 caves, 89% of the species could be recovered. However, the species accumulation curve did not reach an asymptote, and estimates of species richness based on frequency of rare species predicted that nearly half of the species were not collected at all. Two years later (2004), these caves were revisited and resampled. Preliminary results show that many groups (isopods, pseudo scorpions, beetles, and diplurans, for example) were encountered in more caves than in the previous study. At the same time, they were often not recollected from previously recorded caves. Beetles (Pseudanopthalmus sp.), for example, were only collected from half of the 13 caves recorded in the previous study; yet they were collected from 9 additional caves. Clearly, these repeated visits reveal that false negatives are in fact an issue when quantifying subterranean biodiversity. This complete dataset, based on two years of intensive sampling, contains information across many caves and entire communities, and enables a powerful investigation of biases associated with estimations of species richness.

SOME AUXILIARY CRITERIA FOR THE SELECTION OF GROUNDWATER SITES FOR BIODIVERSITY CONSERVATION

B. SKET

University of Ljubljana, Department of Biology, Biotechnical Faculty, Vecna pot 111, P.P. 2995, 1001 Ljubljana, SLOVENIA. ([email protected]).

If we are selecting a groundwater site for the biodiversity conservation, the main criterion will be the richness and uniqueness (i.e. endemism or phylogenetic and taxonomic singularity) of its fauna. This may be connected either by the site’s biogeographical position or by a diversity of habitats included. However, considering other aspects will on the one hand bring additional benefits to the science and/or to the local culture, and on the other hand, it will facilitate the achievement of the legal protection. If the site (e.g. a cave system, a site of gravel deposits) is the type locality of some taxa, it is scientifically more important; the same if it is at the same time a paleontological or archaeological site. If it is particularly appropriate for scientific study, for the public education, if it is a historical place of any kind, or of a particular beauty, the need for its protection will be locally and nationally easier recognisable. Postojna-Planina Cave System will be used to demonstrate these principles.

138 SWSB December 2004 SESSIONS ABSTRACTS

SUBTERRANEAN BIODIVERSITY IN GREAT BRITAIN AND IRELAND: A PREDOMINANTLY POST-GLACIAL COLONISING FAUNA WITH POSSIBLE TRANS-GLACIAL ELEMENTS

1P.J. WOOD and 2G.S. PROUDLOVE

1. Department of Geography, Loughborough University, Loughborough, LE11 3TU, UK. ([email protected]). 2. Department of Zoology, Manchester Museum, Manchester, M13 9PL, UK. ([email protected]).

Most of the land area of Great Britain and Ireland was covered by ice during several of the Pleistocene glaciations. Only small areas to the south of these islands remained ice free but they would have been subject to permafrost conditions. If there were any subterranean biota before the Pleistocene they would have been made extinct during this time. .Samples collected over the past 60 years show that there are few obvious troglobites and stygobites. Terrestrial communities are dominated by Collembola, Acari and Diptera. Aquatic communities are dominated by Copepoda, Amphipoda and Isopoda. Most subterranean species appear to be morphologically similar to epigean species and we class them as troglophiles and stygophiles. However it is possible that some species found in caves are ecologically and evolutionarily troglobitic and stygobitic, with no genetic exchange between epigean and hypogean populations. In general the fauna appears to be colonising subterranean habitats in these areas. There is a strong possibility that the stygobitic Amphipoda (Niphargus species and Crangonyx subterraneus) and Isopoda (Proasellus cavaticus), together with a syncarid crustacea Antrobathynella stammeri, survived the glacial periods in sub-glacial refugia such as caves and deep groundwater. The syncarid is found in central Scotland many hundreds of km north of the last (Devensian) glacial maximum. The distribution abilities of these animals is very poor and a sub-glacial refugium model is the best explanation for current distribution.

We will provide distributional and genetic data for the current distribution of subterranean animals in Great Britain and Ireland.

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A POSSIBLE MECHANISM OF PARTICIPATION OF DIAPAUSE IN CYCLOPID PENETRATION INTO UNDERGROUND ENVIRONMENTS (MONGOLIAN SPRINGS, WELLS AND IN SABLINSKIJE CAVES).

V.R. ALEKSEEV

Zoological Institute of the Russian Academy of Science, University emb., 1, 199034, St. Petersburg, Russia ([email protected]).

ABSTRACT The main adaptive change in cyclopid species recently penetrating into underground environment was transformation of population structure resulting in accumulation of diapausing copepodid stages. Such recent invaders had no other clear seen adaptation to life in this environment that was confirmed also by morphological and genetical comparison with species from surface populations. Species with much longer history of inhabiting underground environment (that was indicated by both DNA and morphological differences from surface relatives) had normal population structure and clear seen morphological and biological adaptations to life in such conditions. An hypothesis on a possible mechanism of participation of diapause in Cyclopid penetration into underground environments is suggested.

KEYWORDS : DIAPAUSE, MONGOLIAN DESERT, WELLS, SPRINGS, CYCLOPID PENETRATION, UNDERGROUND ENVIRONMENT.

1. INTRODUCTION to study adaptations in crustaceans to survive in The Goby is a high-mountain stone desert with periodically drying up waterbodies. hundreds coldwater streams and karstic zones. In 1992, some materials on diversity among In the north part of Goby, there is the so named cyclopids in recently created human-made Great Lake Valley Ð an area with several large Sablynskuje caves (Ladoga Lake area, lakes periodically almost drying up that tightly Leningrad district) were obtained from Dr. Peter follows to the Solar activity. Krylov, Zoological Institute, RAS. Traditional methods of sampling (filtration of 50- Organisms surviving in these lakes should 200 L of water via a plankton net, mesh size 75 possess some special adaptations to these mcm), conservation (4% formalin) and harsh conditions. For true aquatic fauna, we observation (a compond Zeiss microscope with found three main strategies to survive when 1000 times resolution) were used in this study. lakes almost dry up for more than one season : ¥ Producing of well protected resting eggs 3. RESULTS AND DISCUSSION (Calanids; Cladocerans; Fireshrims; Rotifers); It was shown that the Copepodid 4 stages of the ¥ Going up-stream to the never drying up streams small cyclopid Microcyclops afganicus Lindberg in mountains (Fishes, Amphibians); were the most common copepod specimens in ¥ Underground water penetrating via springs Mongolian streams and desert wells (Alekseev, (Amphipods, cyclopids). The last adaptation 1996). This copepodis stage is also known as a could be regarded as a constantly working diapausing stage in many freshwater cyclopids micro-evolutionary machine aimed at selecting (Alekseev, 1991). So, population structure of this and creating species for inhabiting in species in wells and streams even in summer underground environment. From this point of months had abnormally high number of possibly view, this Goby desert area seems like a very diapausing stages (85-100% of total population convenient polygon for understanding the density). mechanism of penetration of surface species At the same time, in open water, their populations into underground waters. had normal population structure and adult female and male were dominated. M. afganicus found in 2. MATERIAL springs were morphologically identical to those In 1995, 1999 and 2000, along with the Joint collected in lakes, so the only difference in the Russian-Mongolian Biological Expedition, I species collected in the two types of ecosystems worked in different karstic areas in Mongolia and was a different percentage of resting stage 4 in collected crustaceans in lakes, rivers and springs their populations. To my mind, this species could

SWSB December 2004 143 POSTERSPOSTERS: EXTENDED ABSTRACTS use diapause for a preliminary adaptation to life copepodids 4 dominated. It was close to the in underground biotope. Diapause as an same in M. afganicus from Mongolian wells. Few adaptation to low food ability and low adults of the species found together with their temperature could help this surface species in resting stages did not show any morphological adjusting its life cycle to its conditions in differences from the same species from surface underground environment. waters. They had a red pigment in their eye, Additional profit for this cyclopid, which recently appendages of normal sizes with invaded in springs and wells, could come from homogeneously setated and normal size seta. smaller size and lower metabolism in diapausing This result seems to confirm my hypothesis and stages than in actively growing stages. enables to use a disproportion in population structure (accumulation of diapausing stages) as Eucyclops dumonti Alekseev was another an indicator of recently invaded to caves species, species dominated in a spring in Central at least in Cyclopids. Mongolia. This also small size cyclopid looked much better adapted to life in springs than other surface species of this genus. E. dumonti poses several signs of typical interstitial cyclopid Ð 4. CONCLUSIONS small size; shorten furca, spiniform seta P1-P4, ¥ In the cyclopid species, recently penetrated into small amount of large size eggs in females. underground environment, the main adaptive Interesting is that this form living in cold water change was the transformation of population springs had normal population structure with structure resulting in accumulation of equal proportion of adults to other age stages. As diapausing copepodid stages. Such recent it was shown, this species was well separated invaders had no clear seen morphological both genetically and morphologically from related adaptation to life in this environment. surface species (Alekseev 2000). ¥ Species with longer history of inhabiting Another example of good adaptation to its underground environment (that was indicated environment is an endemic cyclopid Diacyclops both by DNA and morphological differences eulittoralis Alekseev et Arov, described from from surface relatives) have normal population similar to spring biotope - interstitial pores in structure and clear seen morphological and sand beach near Baikal. This species is also very biological adaptations to life in such conditions. small and well separated from surface congeners ¥ Disproportion in population structure and it has equal proportion in density of different (accumulation of diapausing stages) if found in stages. This species was morphologically very a species inhabiting underground environment well adapted to life in interstitial pores of the sand can possibly be used as an indicator of recent beach (Alekseev, Arov 1986). invasion to caves of this species, at least in From results of these observations, I proposed Cyclopids. the hypothesis that : ¥ both E. dumonti and D. eulitoralis were under longer micro-evolutionary selection by their AKNOWLEDGEMENTS underground environment than Microcyclops This study was partly supported by a research afganicus in Goby desert wells, grant N 04-04-49121-a and a travel grant ¥ duration of life in underground environment is N 04-04-58834-3 from The Russian Foundation correlated with the above mentioned difference for Basic Researches. I thank Dr. Peter Krylov for in population structure found in species sampling copepods for me in the Sablinskije inhabiting springs, wells or other close to Caves. surface or recently isolated from surface underground environments. To test this hypothesis, I analyzed population BIBLIOGRAPHY structure in several cyclopid species from Alekseev, V.R. 1991. ‘Diapause in Crustaceans: samples collected in Sablinskije Caves. These ecological and physiological aspects’. Academy caves created by human have not existed for Scientific Publishers “Nauka”: Moscow, (In Russian, more than 300 years, so cyclopid fauna in their English summary). lakes must be of the same age or about it. Alekseev, V.R. 1996. The role of copepods in Goby desert Duration of isolation of species in the lakes from lake ecosystems (Mongolia). Russian Hydrobiological surface relatives could be regarded as a control Journal, special issue: 81-94. in this natural experiment. Alekseev, V.R. 2000. Eucyclops dumonti sp. nov. from Central Mongolia. Hydrobiologia, Vol. 441:63-71 In Sablinskije caves, all studied cyclopid species (Acanthocyclops vernalis, Megacyclops viridis, Alekseev, V.R., and Arov, I.V. 1986. A New species from the Genus Diacyclops (Crustacea, Copepoda) from Mesocyclops leuckarti, Eucyclops serrulatus) the Eulittoral of Lake Baikal. Zoologicheskij Journal, had very simple population structure with Vol. 65:1084-1088 (In Russian, English summary).

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INTERSTITIAL HARPACTICOIDS FROM VOLCANIC LAKES OF LATIUM (ITALY) : STATE OF THE ART.

R. BERERA, M.C. BRUNO, L. PARICIANI and V. COTTARELLI

Università degli Studi della Tuscia, Dipartimento di Scienze Ambientali, Largo dell’Università s.n.c Blocco D, 01100, Viterbo, Italy. ([email protected]), ([email protected]), ([email protected]), ([email protected]).

ABSTRACT We present faunistic and ecological data on interstitial harpacticoids we studied for thirty years. Our long-term investigation focused on microcrustaceans communities of Vico, Bracciano and Bolsena lakes. In the last five years we also studied the small Martignano and Mezzano lakes, even if sporadically. All the investigated lakes provided interesting faunistic, taxonomic, and biogeographical data. We discussed the endemic and rare taxa. Particularly interesting is the recent finding of Schizopera sp. nov. in the psammon of Bolsena and Mezzano. Lakes. This taxon was unknown in Italian lakes until now; it is a thalassic taxon and its origin may be related to marine transgressions phenomena.

KEYWORDS : HARPACTICOIDS, INTERSTITIAL, LATIUM, VOLCANIC LAKE.

1. INTRODUCTION be discussed briefly for each of the five lakes. Bolsena Lake. The first studies on the interstitial The study of some the volcanic lakes of Latium harpacticoids of this lakes date the early 70s and (Bracciano, Martignano; Vico, Bolsena, and 80s (Cottarelli, 1972; Cottarelli and Maiolini, Mezzano Lakes) lead to the description of 1980); a long-term study on the interstitial several new species of harpacticoids (Cottarelli, community was developed in the late 80s and 1972, Cottarelli and Maiolini, 1980; Bruno and 90s (Sozio, 1985; Bruno, 1997; Bruno and Cottarelli, 1998), and long-term studies allowed Cottarelli, 1998, Bruno et al., 1996, 1997). This to describe spatial and temporal changes of lake has the highest diversity among the five copepod communities, in relation to biotic and lakes, probably due to the good water quality of abiotic factors (Bruno, 1997; Bruno et al., 1996, the lake. Twelve taxa were recorded from 1997; Pariciani, 2000, in prep.). The unpublished interstitial samples collected along the lake results of several more collections allowed us to sandy shores: 6 species were stygophiles and 6 complete an updated list of taxa for each of the species were stygobites. Among these latter taxa lakes. In this paper, we will shortly describe the Parapseudoleptomesochra italica Pesce e result for each lake, thus presenting the state-of- Petkovski, 1980 and Schizopera sp. are, the-art of the faunistic knowledge of these according to Pesce and Petkovski (1980) and volcanic lakes. Cottarelli et al. (1995), stygobites and thalassoids, i.e., originated from marine 2. METHOD ancestors which, after colonising the littoral The researches discussed here were conducted sands, entered continental groundwater with a from 1968 to 2000. The samples have been passive horizontal transition (Coineau and collected using the Karaman-Chappuis method Boutin, 1993). Schizopera sp. is a new species. (Delamare Deboutteville, 1960); all the material The remaining stygobites belong to the genus has been fixed with 5% buffered formaldehyde Parastenocaris Kessler, which groups and sorted and identified in laboratory. harpacticoids of small dimensions with Permanent slides have been prepared morphological and biological features suitable for immersing specimens in Faure liquid. the “subterranean life”, which has a limnicoid origin (Noodt, 1955, Bruno et al., 1998), i.e. the ancestor first colonised the surface freshwater, 3. RESULTS AND DISCUSSION then with a vertical transition entered In table 1 we present an up-to-date list of the groundwater (Coineau and Boutin, 1993). known subterranean harpacticoids for the above- Parastenocaris amalasuntae Cottarelli and mentioned volcanic lakes of Latium. Result will Bruno, 1998 is a taxon with a limited distribution

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Table 1 : List of the collected Harpacticoids area because, until now, it was collected only in Mezzano Lake. This small lake, located 10 Km Bolsena Lake and in the hyporheos of Fiora from Bolsena Lake, was investigated only in (Viterbo province, Northern Latium) and Cecina 1995 and in 1999 by Pariciani, 2005. The shores (Grosseto province, Southern Tuscany) rivers, of this basin are not suitable for a stygobitic and in a well near Manciano (Grosseto province, community, given the high content of silt, thus Southern Tuscany). In Bolsena Lake copepods were collected by rinsing the Parastenocaris amalasuntae seems to replace submerged vegetation along the shore. A rich Parastenocaris proserpina Chappuis, 1938 population of Canthocamptus staphylinus was which was known in the past for this lake, but present, together with one specimen of the same was never collected in the latest researches. Schizopera sp. present in Bolsena Lake. In addition to the interstitial samples collected Bracciano Lake. This lake was studied from 1968 along the sandy shores, divers at 30 and 42 m to 1972 (Cottarelli and Drigo, 1978), in 1987 depth collected benthic samples. Taxa collected (Desiderio, 1988) in 1988 (Calvitti, 1989) and a in those deep samples were: Canthocamptus long-term study of the interstitial community has staphylinus (Jurine, 1820), Bryocamptus been completed recently (Pariciani, in prep.) it (Rheocamptus) zschokkei (Schmeil, 1893), has investigation. The taxa collected in this lake Bryocamptus (Bryocamptus) pygmaeus (G.O. were: Parastenocaris proserpina, Parastenocaris Sars, 1863), Attheyella (Attheyella) crassa (G.O. orcina Chappuis, 1938, Parastenocaris italica Sars, 1862), Moraria (Moraria) poppei (Mrázek, Chappuis, 1953, Parastenocaris pasquinii 1893) and Paracamptus schmeili (Mrázek, Cottarelli, 1972, Phyllognathopus viguieri 1893). (Maupas, 1892), Canthocamptus staphylinus,

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Bryocamptus (B.) pygmaeus, Paracamptus specimens of Attheyella (A.) crassa and schmeili, Parapseudoleptomesochra italica, Canthocamptus staphylinus were collected. The Epactophanes richardi Mrázek,1893, Nitokra sp., actual status of this lake suggests deterioration Schizopera sp. and Onychocamptus mohammed of phreatic water quality probably induced by (Blanchard and Richard, 1891). All taxa were human impacts during the last decade (Leone abundant, but the most recent data denote and Ripa, 2003). qualitative and quantitative impoverishment of the taxocoenosis. For example, Parastenocaris orcina, which is an important Italian endemism, BIBLIOGRAPHY was never collected after 1987; Parastenocaris Barbanti, L., G. Bonomi, A. Carollo, G. Chiaudani, I. italica, another endemic taxon, was never Ferrari, M. Gerletti, A.M. Nocentini, D. Ruggiu, and L. collected between 1987 and 2002 (Cottarelli, Tonelli. 1971. Limnologia ed ecologia dei laghi di Bruno, Berara, unpubl. data, Pariciani in prep.) it Bolsena, Bracciano, Trasimeno e vico: situazione reappeared in 2003-2004 samples (Pariciani, in attuale e prevedibili conseguenze derivanti da una loro utilizzazione multipla. Istituto Italiano di prep.). Some other species, such as Idrobiologia, 263 pp. Parastenocaris proserpina, Parastenocaris Bazzanti, M. 1975. I Chironomidi (Diptera) dei sedimenti pasquinii, Parapseudoleptomesochra italica and del Lago di Martignano (Lazio). Bollettino Pesca Paracamptus schmeili, had reduced abundance. Piscicoltura e Idobiologia 30(1):139-142. Martignano Lake. This small lake is located about Bazzanti, M., O. Ferrara, L. Mastrantuono, and M. 3 km from Bracciano Lake. Meiofauna from this Seminara. 1994. Water quality monitoring of Italian lake was studied with sporadic collections in the lakes using zooplankton and zoobenthos: the case of Lake Vico (Central Italy). Contributions to Animal 70s (Cottarelli, unpubl. data) and then regularly Biology 79-85. from 1999 to 2000 (Pariciani, 2000). A Benfatti, D., M. Marisa, and I. Morselli. 1992. preliminary study of the shore interstitial habitat Halacaroidea (Acari, Actinedida) from four lakes of developed by Pariciani in 1999 did not provide volcanic origin in Lazio (Central Italy). Bollettino di sufficient data for qualitative and quantitative Zoologia 59:105-111. analysis, because harpacticoids were not Bruno, M.C. 1997. Ricerche su Copepodi Arpacticoidi di collected in any of the sampling sites, although acque sotterranee italiane: osservazioni sulla struttura few years earlier Cottarelli (1975, unpubl. data) di comunità meiobentoniche interstiziali del Lago di collected the stygobite Parastenocaris Bolsena; osservazioni sulla geonemia, ecologia e sistematica di alcune specie di Parastenocaris proserpina, from the interstitial habitat on the Kessler, 1913. Ph. D. thesis. University “della Tuscia”, shores of this lake. Therefore we switched to Viterbo, Italy. studying the benthic meiofauna from shallow Bruno, M.C., and Cottarelli, V. 1998. Description of shores recording a simplified biocoenosis. This Parastenocaris amalasuntae n. sp. And new data on “scarcity” had been already detected by Stella et Parastenocaris proserpina and Parastenocaris al. (1972) who reported only 4 species: Nitokra pasquinii from subterranean waters of central Italy lacustris (Schmankevitch, 1875), (Copepoda, Harpacticoida). Italian Journal of Zoology Onychocamptus mohammed, Attheyella (A.) 65:121-136. crassa and Canthocamptus (C ) stapylinus. We Bruno, M.C., P. Frantoi, and V. Cottarelli, 1996. Prime osservazioni sulla struttura della comunità di collected the same species and also Nannopus Copepodi Arpacticoidi interstiziali del Lago di Bolsena palustris Brady, 1880 that represents a new (Italia Centrale). Atti VII Congresso Nazionale della faunal element for central and southern Italy. The Società Italiana di Ecologia, Napoli 11-14 Settembre simplified community can be due to altered 1996: 519-522. quality of the sediments. Bruno, M.C., V. Cottarelli, and P. Franzoi. 1997. Ruolo dei Vico Lake. Meiofauna of this lake was fattori ambientali nella struttura della comunità di Copepodi Arpacticoidi interstiziali del Lago di Bolsena investigated during a long time-span and with (Italia Centrale). Atti VIII Congresso Nazionale della particular reference to stygobite elements Società Italiana di Ecologia, Parma 11-12 Settembre (Cottarelli, 1972; Cottarelli and Maiolini, 1980). 1997. 151-154. From 1972 to 1989 four species new for the lake Bruno, M.C., V. Cottarelli, and R. Berera. 1998. fauna were collected: Parastenocaris proserpina, Preliminary remarks on the cladistic systematics in Parastenocaris italica, Simplicaris veneris and some taxa of Leptopontidae and Parastenocarididae Parapseudoleptomesochra italica (Cottarelli (Copepoda, Harpacticoida). Memorie del Museo Civico di Storia naturale di Verona, 2 ser. 13:69-79. unpubl. data). Further sampling along the sandy shores of Vico Lake from 1993 (Cottarelli, Calvitti, M. 1989. Osservazioni sulla tassonomia e geonemia di alcune specie di Harpacticoida unpublish data) lead to the collection of only (Crustacea, Copepoda) con particolare riferimento a specimens of Parastenocaris italica, a species forme del meiobenthos del Lago di Bracciano. Tesi di with wide ecological tolerance. The apparent Laurea in Scienze Biologiche, Università degli Studi disappearance of Simplicaris veneris indicates “La Sapienza”, Roma. the impoverishment of the harpacticoid Coineau, N. and Boutin, C. 1993. Biological processes in communities in this lake. During the latest space and time. Colonization, evolution and collections by Bruno in 1996, only same speciation in interstitial stygobionts. In: A.I. Camacho

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[Eds.], The Natural History of Biospeleology. Margaritora, F., O. Ferrara, and D. Vagaggini. 1999. Le Monographias, 7:423-451. comunità zooplanctoniche del Lago di Martignano Cottarelli, V. 1972. Parastenocaris (Copepoda, (Lazio Italia). Atti Associazione Italiana Oceanografia Harpacticoida) di alcuni laghi vulcanici del Lazio. e Limnologia 13(1):281-289. Rdiconti dell’Istituto Lombardo di Scienze e Lettere, B, Mastrantuono, L. and Natale, A. 1998. Struttura dello 106:137-155. zoobenthos dei sedimenti sabbiosi del Lago di Cottarelli, V., and Maiolini, B. 1980. Parastenocaris Martignano (Italia Centrale) e valutazione delle attuali veneris n. sp., nuovo arpacticoide interstiziale del condizioni ambientali del litorale. XIII Congresso Lago di Vico (Crustacea, Copepoda). Fragmenta A.I.O.L. Ancona. Entomologica 15 (2):243-251. Mayer, A.S., A.J. Rabideau, R.J. Mitchell, P.T. Imhoff, Cottarelli, V., and Drigo, E. 1972. Sulla presenza di M.I.Lowry, and C.T. Miller 1993. Groundwater quality. Parastenocaris orcina Chappuis (Copepoda, Water Environmental Research 65, 4:486-534. Harpacticoida) in acque interstiziali del Lago di Noodt, W. 1955. Die Verbreitung des Genus Bracciano. Notizie del Circolo Speleologico Romano Parastenocaris, ein Beispel einer subterranean 1-2:138-155. Crustaceen-Gruppe. Verhandlungen der Deutschen Cottarelli, V., M.C. Bruno, and F. Venanzetti. 1995. Two Zoologischen Gesellschaft 429-435. new species of Parastenocaris from the interstitial Notenboom, J., S. Plénet, and M.-J. Turquin. 1994. waters of rivermouths in Latium and Sardinia Groundwater contamination and its impact on (Crustacea, Copepoda, Harpacticoida). Fragmenta groundwater animals and ecosystems. In: J. Gibert, Entomologica 26(2):229-247. D.L. Danielopol, and J. Stanford [Eds.]. Ground water Danielopol, D.L., and Rouch, R. 1991. L’adaptation des ecology. Academic Press, San Diego. organismes au milieu aquatique souterrain. Pariciani, L. 2000. La comunità di arpacticoidi bentonici di Réflexions sur l’apport des recherches écologiques costa del Lago di Martignano. Graduation thesis, récentes. Stygologia 6(3):129-142. Biological Sciences Faculty, University “della Tuscia”, Delamare Deboutteville, C. 1960. Biologie des eaux Viterbo, Italy. souterraines littorales et continentales. Hermann, Pariciani, L. 2005. Caratterizzazione della comunità Paris, 740 pp. maiobentonica ad arpacticoidi del lago di Bracciano. Desiderio, A. 1988. Osservazioni sulle Parastenocaris Ph. D. thesis. University “della Tuscia”, Viterbo, Italy. (Crustacea, Copepoda, Harpacticoida) della fauna (in prep.). interstiziale italiana, con particolare riferimento alle Pesce, G.L. and Petkovski, T.K. 1980. specie del Lago di Bracciano. Tesi di Laurea in Parapseudoleptomesocra italica n.sp., a new Scienze Biologiche, Università degli Studi “La harpacticoid from subterranean waters of Italy Sapienza”, Roma. (Crustacea, Copepoda, Ameridae). Fragmenta Dyer, M. 1995. The water quality at Lago di Vico during Balcanica 5 (247):33-42. 1992-93. Riserva Naturale del Lago di Vico, 11 pp. Piver, W.T. 1993. Contamination and restoration of Forneris, G., M. Pascale, and G.C. Perosino. 1996. groundwater aquifers. Environmental Health Idrobiologia. Consorzio Regionale per la tutela, Perspectives 100:237-247. l’incremento e l’esercizio della pesca – Valle d’Aosta. Provini, A., R. Marchetti, and G. Tartari. 1992. The Italian 372 pp. lakes: trophic status and remedial measures. Leone, A. and Marini, R. 1993. Assessment and mitigation Memorie dell’Istituto Italiano di Idrobiologia 147-169. of the effects of land use in a Lake Basin (Lake Vico Sozio, C. 1985. Osservazioni sullo psammon del Lago di in Central Italy). Journal of Environmental Bolsena, con particolare riferimento agli Arpacticoidi. Management 39-50. Graduation thesis, Biological Sciences Faculty, Leone, A. and Ripa, M.N. 2003. Land use, pollutant University “La Sapienza”, Rome, Italy. nonpoint sources and related modelling for lakes Stella, E., F.G. Margaritora, G.B. Palmegiano, and M. management. The Lake Vico experience. Bazzanti. 1972. Il lago di Martignano: prime International Conference on: Residence Times In osservazioni sulla struttura e distribuzioni delle Lakes: Science, Management, Education Bolsena biocenosi. Rendiconti dell’Accademia Nazionale dei (Viterbo - Italy) September 29th - October 3rd Lincei, XL, 22:1-17. 2002:148-155. Maggioni, M. 1994. Nematofauna del Lago di Bolsena. Tesi di Laurea in Scienze Biologiche, Università degli Studi di Milano.

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DISTRIBUTION ATLAS OF THE STYGOBIONT MOLLUSCS OF FRANCE

A. BERTRAND

Etude et Conservation des Mollusques Continentaux, Abéla, 09320 BOUSSENAC ([email protected]).

In some regions, especially karst ones, ground ¥ the number of species which has been waters have a particularly rich and diversified described during the last few years is very fauna (Culver et al. 2001, Gibert & Deharveng important (fig.1) and the rhythm of new 2002). discoveries is sustained ; This is also the case for the stygobiont mollusc ¥ several genera which count numerous taxa fauna in western Europe and especially in today (e.g. Bythiospeum) need to be re- France, even if the number of listed taxa may not evaluated. reflect reality. In fact :

Figure 1 : Percent of stygobionts species described before and after 1960.

The nomenclature, systematics and phylogeny of This is also true for many other species (a total of the taxonomic groups and taxa of the French 33, which represent 45% of the stygobious fauna which are entirely or partially stygobiont, molluscs) which, to my knowledge, have never are still very confuse and their morphological, been found alive. anatomical and genetic approach put forward The available data on the species biology and results which sometimes differ considerably (see ecology are almost nonexistent and this makes e.g. Boeters 1998 and Wilke et al. 2001). their status difficult to assess from the point of In the present state of our knowledge (Bertrand view of groundwater environment. 2004), the stygobiont molluscs of France number Nearly all stygobiont species are from karstic today 73 published species and sub-species ; aquifers; the few exceptions being from species 96% are endemic from France or micro-endemic which are known only from specimen found in from a little area (fig.2). To include species of the flood deposits (Bythiospeum michaudi, B. terveri, genus Bythinella has been a problem as it counts Istriana falkneri, Moitessieria bourguignati). The many poorly defined crenobiont taxa ; the few presence of stygobious species in the aquifer “stygobiont” populations of Bythinella have not and in the hyporheic environment is poorly either been studied or their study is not yet documented because of the evident under- finished. And two of them, B. bouloti and B. gale- exploration of these two habitats. rae, have never been found alive.

Figure 2 : Percent of endemic stygobiont species in France.

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The number of species/site varies (karstic basin illustrations posterior to the description, has been sampled directly or at exurgences); the distribution and eventually some comments. You “record” comes from the south-east edge of will note that the real confusion within some Massif Central, from the Ardèche département to genera does not allow an exhaustive list of the Hérault département where it often exceeds synonyms to be established. 5 species and reaches sometimes 8 species in It is still difficult to propose a detailed analysis of some very rich sites. the main patterns of distribution of species in The proposed mapping of taxa comes from the France. The effort of inventory in the different compilation of references and personal regions is still very uneven. However, we can unpublished data. They are part of a data base reasonably conclude that the general outlines of which counts 600 bibliographic data and 1500 the distribution of genera and the main centres of unpublished data. The squares of the grid are 0.2 endemism are today clearly identified (fig.3). x 0.2 grad. The superimposing of all data on the map of karst The atlas part is preceded by notes on each (fig.3) shows that some regions with well- genus and every retained taxon. For the taxa, we developed groundwater habitats, especially in indicate reference of the original description, the Alps, have revealed no species. The possible synonyms, locus typicus, localisation of inventory in progress on the south-west and the typical material, possible references to south edge of the Alps, from Vaucluse to Alpes-

Figure 3 : Stygobiont molluscs in France : geographic repartition for all species and karstic areas.

150 SWSB December 2004 POSTERS: EXTENDED ABSTRACTS

Figure 4 : Number of species, endemic species, genus and endemic genus for French administrative regions.

Maritimes where relatively few species are to my knowledge no specific conservation known, looks promising. The number of known measure of stygobiont molluscs has been taxa per region fluctuates a lot and, the south and enforced. However, the presence of some south-east edge of Massif Central, from the species in Protected Areas, National Parks or Ardèche département to the Hérault Natural Reserves is known and some important département, appears especially rich and inventory work is being carried out. original. The gradient showing the decrease in Fig. 4 is a synthesis of the current knowledge and the number of species towards the north-east underlines the regional responsibilities regarding and the west is really outstanding. From the east the stygobiont molluscs. Note that 46 species or of Hérault to the south of Ardèche, the number of sub-species are endemics in just one known taxa per square varies from 5 to 9. administrative region. Today, 21 taxa are protected (Ministerial Decree Within the scope of the updating of the ZNIEFF, of 07/10/1992). Eighteen are listed as Rare in the these species have been taken into account in national Red Book. Eleven species are qualified several regions (Midi-Pyrénées, Rhône-Alpes, Vulnerable in the IUCN Red List, three are Provence-Alpes-Côte d’Azur and Languedoc- thought to be at low risk and data are insufficient Roussillon) and this is important progress for three others. towards conservation. These two Red Lists are characterized by For every species, conservation implies good noteworthy inconsistencies and a lack of serious knowledge of every facet of its Natural History. work on the species status, although the data But the knowledge of the stygobiont molluscs of now available have considerably increased. It is France is still very fragmentary. essential to review all the stygobious taxa of France, especially in comparison with the IUCN BIBLIOGRAPHY criteria. Bertrand, A. (2004). Atlas préliminaire de répartition Although these species are listed in these texts, géographique des mollusques stygobies de la faune

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de France (Mollusca, Rissoidea, Caenogastropoda). - Gibert, J. and Deharveng, L. (2002). Subterranean Documents Malacologiques, Hors Série, 2, 81 p. ecosystems : a truncated functional biodiversity. Boeters, H. D. (1998). Mollusca : Gastropoda : BioScience, 52, 6, 473-481. Superfamilie Rissooidea. In : Schwoerbel, J. et Wilke, T., Davis, G. M., Falniowski, A., Giusti, F., Bodon, Zwick, P. (Hrsg.), Süßwasserfauna von Mitteleuropa M., Szarowska, M. (2001). Molecular systematics of (Begründet von A. Brauer), 5 (1, 2) : IX + 76 pp. Hydrobiidae (Mollusca : Gastropoda : Rissoidea) : Stuttgart (G. Fischer). Testing monophyly and phylogenetic relationships. - Culver, D. C., Deharveng, L., Gibert, J. and Sasowsky, I. Proceeding of the Academy of Natural Sciences of (Eds.) (2001). Mapping subterranean biodiversity. Philadelphia, 151 : 1-21. Cartographie de la biodiversité souterraine. Karst Waters Institute, Special Publication, 6, 82 pp.

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HYPORHEIC ZONE AS REFUGIUM FOR THE MACROINVERTEBRATE FAUNA IN THE PO RIVER (NW ITALY) : FIRST DATA

S. FENOGLIO, T. BO, M. CUCCO and G. MALACARNE

Di.S.A.V., Università del Piemonte Orientale, Via Bellini 25 - 15100 Alessandria, Italia. ([email protected]), ([email protected]), ([email protected]), ([email protected])

ABSTRACT In this study we analysed the role of hyporheic zone as refugium for the benthic macroinvertebrates in the high reach of the Po river (NW Italy). Aim of our study was to investigate the vertical distribution of macroinvertebrates within the substratum. We positioned 12 hyporheic traps in the streambed: each trap consisted in an outer structure containing three inside bags.The structures were buried in the streambed in June 2004 with a mini excavator Kubota and still remain in place. The inside traps were filled with clean sterile substrate, similar to the streambed composition, and placed at different depths: the top trap was positioned from 0 to 30 cm, the medium one from 30 to 60 cm and the bottom one from 60 to 90 cm. In this preliminary work we present our first data about distribution, stratification and abundance of stream benthos within the substratum during the dry season.

KEYWORDS : DROUGHTS, HYPORHEIC ZONE, MACROINVERTEBRATES, PO RIVER.

1. INTRODUCTION 2. METHODS There is a growing interest in the vertical To investigate the presence and the colonisation dimension of stream systems: traditionally, process of stream macroinvertebrates in the sub- aquatic ecologists perceived streams and rivers substratum, we constructed and positioned 12 as bounded systems, consisting in riverbed and hyporheic traps in the streambed: each trap overlying water, and only in the last years, the consisted in an outer structure containing three exchanges of water, detritus, nutrients and inside bags (Fig. 1). organisms between groundwater and stream The structures were buried in the streambed in channels have become a central element in June 2004 in two sites (Martiniana Po and freshwater ecology (Vallett et al. 1993). The Revello) with a mini excavator Kubota (Fig. 2) importance of the hyporheic zone as refuge for and still remain in place. The inside traps were stream macroinvertebrates is not widely filled with clean sterile substrate, similar to the accepted (Dole-Olivier and Marmonier, 1997): streambed composition, and placed at different some studies underlined that in unfavourable depths: the top trap was positioned from 0 to 30 hydrological conditions, stream invertebrates cm, the medium one from 30 to 60 cm and the move into the depth zone, escaping from dry bottom one from 60 to 90 cm. periods or from floods (Cooling and Boulton, At the same time, we conducted quantitative 1993), while some others found little evidence for samplings, using a Surber net, to evaluate the use of hyporheic zone as refuge (Palmer et structure, density and composition of benthic al., 1992). macroinvertebrate communities. Aim of this study was to investigate the importance of the sub-substratum for macroinvertebrates inhabiting a zone 3. RESULTS characterised by numerous droughts. We also Unfortunately, an unusual and prolonged investigated the vertical distribution of shortage affected the study area and the entire macroinvertebrates within the substratum. Piemonte during the study period. After This study was realised in the subalpine reach of displacing the traps, we removed the inside the river Po. This area belongs to the Parco del baskets only two times, because the water level Po Cuneese (near Saluzzo, CN) and it is in the hyporheic zone was always deeper than 90 characterised by high streambed permeability cm (i.e. our traps were above the water level in and summer droughts. The length of the the sub-substratum). We removed the traps in droughts increase along a longitudinal gradient, September and in November 2004, when no from Sanfront (water in the streambed for 12 water was present in the surface. Because of this months/year) to Saluzzo (water in the streambed problem, we can provide only partial and for 2-3 months/year). incomplete data. In the baskets removed, mean

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Figure 1 : Hyporheic traps scheme

Figure 2 : Traps displacement macroinvertebrate abundance was 843.3 to assess patterns and mechanisms on the basis organisms/m3, varying from a minimum of 33.3 of the resilience of stream communities to organisms/m3 to a maximum of 2,733.4 droughts. In this work, we present in particular organisms/m3. We also provide a preliminary our experimental design, because environmental taxonomic list of macroinvertebrates found in the problems have not allowed us to conduct more sub-substratum: Plecoptera (Leuctra sp.), complete studies until now. Quantitative data Ephemeroptera (Serratella ignita, Baetis sp.), collected in the period January - December 2004 Trichoptera (Odontocerum albicorne, using Surber samplers in four stations along a Limnephilidae, Hydropsyche sp.), Diptera gradient of water permanence indicated that (Chironomidae, Ceratopogonidae, Psychodidae, drought strongly influence structure and Limoniidae, Tabanidae, Dolichopodidae), composition of macrobenthos: taxonomic Coleoptera (Helicus substriatus, Dytiscidae, richness decrease from the Station 1 (values Hydraena sp., Elminthidae), Irudinea (Dina sp.), similar to data reported in previous studies Oligochaeta (Eiseniella tetraedra, Lumbricidae, realised in southern Piedmont, Fenoglio et al., Lumbriculidae), Hydracarina. 2002, 2004) to the Station 4 (very poor communities). These findings indicate the 4. CONCLUSIONS existence of an evident longitudinal gradient. Our preliminary results support the hypothesis This work is a part of a greater project, founded that interstitial zone represents an important by the Natural Park of the Po River, with the aim

154 SWSB December 2004 POSTERS: EXTENDED ABSTRACTS refuge for stream macrobenthos, although it may stream macroinvertebrates. J. Fresh. Ecol. 19:485- not be used by all taxa. 492. Fenoglio, S., P. Agosta, T. Bo, and M. Cucco. 2002. Field experiments on colonization and movements of BIBLIOGRAPHY stream invertebrates in an Apennine river (Visone, Cooling, M.P. and Boulton, A.J. 1993. Aspects of the NW Italy). Hydrobiologia 474:125-130. hyporheic zone below the terminus of a South Palmer, M. A., A.E. Bely, and K.E. Berg. 1992. Response Australian arid-zone stream. Austr. J. Mar. Fresh. of invertebrates to lotic disturbance: a test of the Res. 44:411-426. hyporheic refuge hypothesis. Oecologia 89:182Ð194. Dole-Olivier, M.-J., P. Marmonier, and Beffy, J.L. 1997. Vallett, H.M., C.C. Hakenkamp, and A.J. Boulton. 1993. Response of invertebrates to lotic disturbance: is the Perspectives on the hyporheic zone: integrating hyporheic zone a patchy refugium? Freshwat. Biol. hydrology and biology. Introduction. J. N. Am. Benthol. 37:257-276. Soc. 12:40-43. Fenoglio, S., T. Bo, G. Gallina, and M. Cucco. 2004. Vertical distribution in the water column of drifting

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MICROINVERTEBRATES OCCURRENCE IN GROUND-WATERS

1L. MANCINI, 1D. VENANZI, 1L. VOLTERRA, 2B. PENNELLI and 1P. FORMICHETTI

1. Istituto Superiore di Sanità, Laboratorio di Igiene Ambientale, V.le Regina Elena 299, 00161 - Rome, Italy ([email protected]), ([email protected]), (paolo formichetti [[email protected]]). 2. Istituto Sperimentale per la Nutrizione delle Piante, Via della Navicella 2-4, 00184 - Rome, Italy ([email protected]).

ABSTRACT Drinking waters in Italy are mostly taken from ground- or spring-waters (84%) and only in minor part come from surface waters (15%) or other sources (1%). In this study, we investigated for the presence of microinvertebrates, and Nematodes in particular, in drinking water samples collected from pipelines in nine Italian regions. For each sample, ten litres of water were filtered through cellulose membranes and the residues were fixed and then investigated for the presence of microinvertebrates. Among them nematodes were the most frequently collected. A relationship between biological risk and nematodes occurrence in drinking water was not straightforward, rather the presence of such fauna might be a reliable indicator of the vulnerability of water networks.

KEYWORDS : DRINKING WATERS, GROUND WATERS, NEMATODA

1. INTRODUCTION Lazio, Molise) were collected and analysed. Drinking waters in Italy are mostly taken from Each ten-litres sample was screened through ground- or spring-waters (84%) and only in minor ester cellulose and nitrate cellulose filters from part come from surface waters (15%) or other 500 to 0.45 µm, sequentially. The material caught sources (1%). The presence of metazoan in in the filters was fixed, screened at the drinking waters (WHO, 1993) is known in many stereoscope or, depending on the size of Italian areas (Domenichini and Molinari, 1984; Di organisms, under an inverted microscope, and Girolamo et al., 1995; Lupi et al., 1994) and some taxonomically detected. inspections revealed plenty of identifiable nematodes in piped waters (Volterra et al., 1993) 3. RESULTS AND DISCUSSION as well as reported in other countries (Cobb, Table I shows the location and amount of water 1918; Chang et al., 1959; Chang et al., 1960; samplings, sampling season, typology of water Mackenthun and Keup, 1970; Thombes et al., supplied, water treatment carried out in 1979; Gerardi and Grimm, 1982; Mott et al., aqueducts, taxa and specimen collected. 1983; Levy et al., 1984; Levy et al., 1986; Gray, Nematoda were found to be the majority of 1994). A relationship between biological risk and findings in 6 out of 9 regions and in 91 out of 408 nematodes occurrence in drinking water has not water samples (22%). If the case of Umbria been reported in literature; rather the presence of (3,564 Cladocera found) is not considered, such fauna might be a reliable indicator of the Nematoda rise up to 85.2% of the whole catch. vulnerability of water networks. In any case, it However, none of the water treatments (including cannot be excluded that microinvertebrates could disinfections) looks successful against such be of potential health hazard, as vectors of contamination. Furthermore, our study revealed pathogens and risk cannot be completely no seasonal variation of Nematoda occurrence, excluded especially for unhealthy population. In in contrast with other studies (Volterra and this study, we investigated for the presence of Santamaria, 1993; Chang et al., 1959; Chang et microinvertebrates, and Nematodes in particular, al., 1960; Mackenthun and Keup, 1970; in drinking water samples collected from Thombes et al., 1979; Mott et al., 1983). pipelines in nine Italian regions. The abundance of Nematoda in tap water is likely to be correlated with the nourishment of a biofilm 2. METHODS inside the pipelines. These in fact, are a relatively 408 water samples coming from 92 water stable environment in terms of physical, chemical supplies of nine Italian regions (Piemonte, Friuli, and biological properties and consequently are Emilia Romagna, Toscana, Marche, Umbria, suitable to Nematoda colonization. Differently,

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Table 1 : The location (region), sampling season, type of water, water treatment, and taxa found are listed. In parenthesis is reported the number of specimen collected for each taxon.

Cladocera in tap water are clue of external genera and species of Nematoda can be contaminations, usually due to breaks and spills indicators of trophic level and contamination of aqueducts. (Zullini, 1982). The examination of Nematoda in network waters In the past, the percentage of Nematoda can be used to justify organoleptic alterations, to removal, in drinking water, was used as an reveal biological colonization of the network, to indicator of potabilisation efficiency (Thombes et correlate this population with that of ecotones, al., 1979; Mott et al., 1983), where: >90% springs and aquifers. Moreover, Nematoda might excellent; 75-90% good; 50-74% poor; 50% bad be used, as well as other invertebrates, as (APHA et al., 1995). To improve the potabilisation indicators of environment modifications. processes, the routine monitoring of Nematoda According to some researchers, in fact, specific in filtration beds and rinsing waters may be a

158 SWSB December 2004 POSTERS: EXTENDED ABSTRACTS reliable and easy-to-run tool. After all, Nematoda acque potabili”, Atti del I Convegno Accademia are not sensitive to disinfections. Italiana di Entomologia, 165-185. Although the role of metazoan, and in particular, Gallo, L., P. Polli, and Sartori, R. 1993. “Rapporto sulla presenza di elminti in acque potabilizzate”, Ambiente of Nematoda, as active and passive vectors of Risorse e Salute 18:38-40. microorganisms, is not well known, it is likely that Gerardi, M.H. and Grimm, K.J. 1982. “Aquatic invaders”, Nematoda, in some larval stages, could be Water/Engineering & Management 10:22-23. responsible for allergies in consumers. Such an Gray, N.F. 1994. Drinking water quality. Problems and event might be enhanced by the short life cycle solutions. J. Wiley and Son, New York. of these organisms: Rhabditidae, for instance, Haney, P.D. 1978. “Evaluational microbiological standards have a complete generation cycle from egg to for drinking water”, Water Sewage Works 125:126- egg, lasting 3 days. In European fresh waters, 134. 500 Nematoda species have been identified, King, C.H., E.B.J., Hote, R.E. Wooley, and Porter, K.G. each having its own environmental and trophic 1988. “Survival of coliforms and bacteriological requirements (Zullini, 1982). Furthermore, the list pathogens within protozoa during chlorination”, Appl. of species of a given site can be considered an Envion. Microbiol. 54:3023-3033. indicator of the environmental conditions. These Italy (I). 1988. Gazzetta Ufficiale della Repubblica Italiana, organisms might therefore be used as serie generale No. 152. bioindicators for drinking water quality (Zullini, Italy (I). 2001. Gazzetta Ufficiale della Repubblica Italiana, 1982). serie generale No. 52. Levy, R.V., R.D. Cheetam, J. Davis, D. Winer. and Hart, F.L. 1984. “Novel method for studying the public 4. CONCLUSIONS health significance of macroinvertebrates occurring in potable water”, Appl. Environ. Microbiol. 47:889-894. A huge monitoring performed on drinking waters revealed a remarkable nematode contamination Levy, R.V., F.L. Hart, and R.D. Cheetam. 1986. “Occurrence and public health significance of but without any relationship between biological invertebrates in drinking water systems”, J. A. W. W. risk and nematode’s occurrence. The presence A. 78:105-110. of metazoan in drinking waters has been widely Lupi, E., V. Ricci, and Burrini, D. 1994. “Occurrence of documented in the past; however it was never Nematodes in Surface water used in a drinking water included into the set of compulsory assays for plant”, J. Water STR Ð Acqua 43:107-112. human consumption. Lupi, E., V. Ricci, and Burrini, D. 1995. “Recovery of Despite the fact that in temperate areas non bacteria in nematodes isolated from a drinking water supply”, J. Water STR Ð Aqua 44:212-218. parasitic metazoan which host human parasites are not recorded, it cannot be excluded that Mackenthun, K.M. and Keup, L.E. 1970. “Biological problems encountered in in water supplies”, J. A. W. microinvertebrates could be of potential health W. A. 62:520-526. hazard, as vectors of pathogens. Mott, J.B., G. Mulamoottil, and A.D. Harrison. 1983. “A 13- On the other hand, the presence of such fauna month survey of nematodes at three water treatment might be a viable indicator of the vulnerability of plants in Southern Ontario, Canada”, Wat. Res. water networks. 15:729-738. Nelsson, J.R. 1987. “Structural aspect of digestion of Escherichia coli in Tetrahymena”, J. Protozool. 34:1-6. BIBLIOGRAPHY Smerda, S.M., H.J. Jensen, and Anderson, A.W. 1971. APHA, AWWA, WPCF. 1995. Standard methods for the “Escape of Salmonellae from chlorination during examination of Water and Wastewater, Washington ingestion by Pristionchulus lheritieri (Nematoda D.C.: American Public Health Association, New York. Diplogasterinae)”, J. Nematol. 3:201-204. Chang, S.L., J.H. Austin, H.W. Poston, and Woodwart, Thombes, A.S., A.R. Abernathy, D.M. Welch, and Lewis, R.L. 1959. “Occurrence of a nematode worm in a city S.A. 1979. “The relationship between rainfall and water supply”, J. A. W. W. A. 51:671-676. nematode density in drinking water” . Wat. Res. Chang, S.L., G. Berg, N.A. Clark, and P.W. Kaber. 1960. 13:619-622. “Survival and protection against chlorination of human Volterra, L. 1995. “I materiali usati per venire in contatto enteric pathogens in free-living nematodes isolated con l’acqua potabile”, Ambiente Risorse e Salute from water supplies”, Am. J. Trop. Med. Hyg. 9:136- 14:17-21. 142. Volterra, L., M. De Mattia, R. Marini, , L. Mancini, T. Cobb, N.A. 1918. “Filter-bed nemas: nematodes of the Famiglietti, M. Migliola, N. Focchi Ricci, I. Trotta, M. low filter-beds of American cities”, Contr. Sci. Ercolessi, M. Ricci, M.R. Aliquò, G. Bottinelli, D. Nematol. 7:189-212. Viglione, M. Molina, L. Spadafino, A. Lolini, and Curds, C.R. 1977. “Microbial interations involving Sansoni, G. 1990. “Microbiologia dell’acqua potabile. protozoa”, Soc. Appl. Bacteriol. 6:69-105. I parametri C 4 - D.P.R. 25/5/88”, Biol. It. 20:42-47. Di Girolamo, I., G. Bezziccheri, L. Bonadonna, M. Volterra, L., A. Bertolotti, and Gallo, L. 1993. “Ecosistema Ercolessi, I.Trotta, and Ottaviani, M. 1995. “Nematoda rete”, Ing. Sanit. 41:29-34. in acque potabili: analisi di un caso italiano”, Ann. Ig. Volterra, L. and Santamaria, F. 1993. “Nematoda delle 7:291-299. acque con particolare riguardo a quelle potabili”, Ing. Domenichini, G. and Molinari, F. 1984. “Artropodi delle Amb. 22:200-202.

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WHO. 1993. Guidelines of drinking water quality (2nd Zullini, A. 1982. Nematoda.Guida per il riconoscimento ed.), World health Organization, Geneva. delle specie animali delle acque interne italiane. CNR AQ/1/190.

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DIVERSITY OF SUBTERRANEAN FISHES IN BRAZIL

E. TRAJANO and M.E. BICHUETTE

Departamento de Zoologia, Instituto de Biociências da USP, C.P. 11461, 05422-970, São Paulo, Brazil. ([email protected]), ([email protected]).

ABSTRACT Brazil distinguishes at a worldwide scale by its diversity of troglomorphic fishes, with nearly 20 subterranean species recorded so far, most catfishes belonging to the families Heptapteridae, Trichomycteridae and Loricariidae. These species differ in their degree of reduction of eyes, pigmentation and other troglomorphisms, without a taxonomic correlation, but with a geographic correlation indicating common vicariance events affecting subterranean populations living in the same areas. These species also differ in habitat, population sizes and densities.

KEY WORDS : SUBTERRANEAN FISHES, BIODIVERSITY, BRAZIL, SUBTERRANEAN EVOLUTION, POPULATION ECOLOGY.

Brazil has a remarkable subterranean Trichomycterus itacarambiensis Trajano & Pinna, ichthyofauna, similar to few other countries or 1996, from Itacarambi, eastern Brazil, regions with comparable geographical extension Trichomycterus sp. 1 (as in Trajano, 1997), from of karst areas, such as Mexico, China and south- Serra da Bodoquena, Trichomycterus sp. 2 (ditto) eastern Asia. To date, nearly 20 species (several from Serra do Ramalho, north-western Brazil, awaiting formal description) of Brazilian Trichomycterus sp. 3 from Mambaí region, subterranean fishes are known to present central Brazil; Heptapteridae: Phreatobius reduction of eyes and pigmentation at least at cisternarum Goeldi, 1904, from the Amazon some degree beyond that observed in their delta, northern Brazil, Pimelodella kronei epigean congeners, suggesting a troglobitic (Ribeiro, 1907), from the Upper Ribeira Valley, (exclusively subterranean) status for these south-eastern Brazil, Pimelodella spelaea fishes, and this number is increasing every year. Trajano, Reis & Bichuette, 2004, from São The majority (all but two) are siluriformes Domingos, Rhamdia sp. from Serra do Ramalho, belonging to three out of 12 catfish families found Rhamdiopsis sp. from Cordisburgo region, in Brazil. Catfish species are also predominant in eastern Brazil, Taunayia sp. from Campo groundwater of Mexico (although the proportion Formoso region, north-eastern Brazil, and a new in relation to non-siluriforms is lower than in genus from Chapada Diamantina, north-eastern Brazil) and other Neotropical countries, whereas Brazil; Gymnoti-formes: Sternopygidae: cypriniforms largely outnumber catfishes in Eigenmannia vicentespelaea Triques, 1996, from China, Thailand and other Asian countries. São Domingos region. The subterranean troglobitic Brazilian fishes In addition, at least 13 troglomorphic species recorded so far are: Characiformes: Stygichthys have been found in epigean water bodies in the typhlops Brittan & Böhlke, 1965 (although Amazon basin, including the bottom of large previously thought to be a tetra characin, recent rivers, litter banks at river margins, and rocky studies indicated that it cannot be assigned to pools (Trajano, 1997). These fishes also belong any of the known families), from the Jaíba region, to typically nocturnal and chemo-oriented eastern Brazil; Siluriformes Loricariidae Siluriformes families (Trichomycteridae, (armoured catfishes): Ancistrus cryptophthalmus Heptapteridae, Pimelodidae) or to the Reis, 1987, from the São Domingos region, Gymnotiformes (Sternopygidae, Apteronotidae), Central Brazil, A. formoso Sabino & Trajano, which are also adapted to poorly illuminated 1998, from Serra da Bodoquena, north-western environments. Brazil; Trichomycteridae: Ituglanis passensis Brazilian subterranean fishes greatly differ in Fernández & Bichuette, 2002, I. bambui their degree of troglomorphism, from totally Bichuette & Trajano, 2004, I. epikarsticus anophthalmic and depigmented species to those Bichuette & Trajano, 2004, I. ramiroi Bichuette & showing population variability in eye and Trajano, 2004, all from São Domingos, pigmentation development, with no taxonomic

SWSB December 2004 161 POSTERSPOSTERS: EXTENDED ABSTRACTS correlation, i.e., the same range of differences towards the breakdown of the hydrological may be observed within and among families and continuity, due to the lowering of the phreatic genera. This is well illustrated in the cases of level may be hypothesized. In fact, significant Ancistrus (A. cryptophthalmus, with different Quaternary climatic fluctuations have been well populations showing a mosaic distribution in eye documented for karst areas such as those in the and pigmentation development, versus the highly Upper Ribeira Valley, south-eastern Brazil (home specialized, homogeneously depigmented and to P. kronei) and in Central (the new heptapterid anophthalmic A. formoso), Trichomycterus (from genus) and Northern Bahia (Taunayia sp.), and Trichomycterus sp. 3, with only slightly reduced seem to have been widespread, although dry eyes and pigmentation, to Trichomycterus sp. 1, and wet peaks were not synchronous throughout anophthalmic and mostly depigmented, going the country. through T. itacarambiensis, with highly variable The São Domingos karst area, Central Brazil, is eyes and one third of the only known population distinguished worldwide as a hotspot of diversity, being truly albinic while the remaining two thirds with seven troglobitic species, including the only present variable melanic pigmentation Ð (Trajano confirmed case of syntopy in Brazil (A. & Pinna, 1996) and Pimelodella (from P. spelaea, cryptophthamus and I. passensis, in Passa Três with slightly reduced eyes and pigmentation, to cave). The occurrence of significant the moderately troglomorphic P. kronei, mostly paleoclimatic fluctuations in this particular area anophthalmic and with highly variable remains to be demonstrated. In spite of intensive pigmentation). Intrafamilial variation is collecting efforts, no epigean Pimelodella and exemplified by the heptapterids Rhamdiopsis sp., Ituglanis catfishes were found in São Domingos with only slightly reduced eyes, on one hand, and region, hence extinction of epigean relatives due the highly specialized Taunayia sp. and the new to unknown factors may be the cause of genus, totally anophthalmic and depigmented, on differentiation resulting in exclusively the other. subterranean species. On the other hand, These species occur in different karst areas, topographic isolation as a consequence of situated in distinct phytogeographic domains, alluvial downcutting was also proposed, as a including the humid Atlantic Forest (Upper subsidiary process, to account for the origin of Ribeira Valley), the savannah-like Cerrado (São troglobitic species living in upper vadose Domingos, Mambaí), the semiarid Caatinga tributaries, such as I. bambui and P. spelaea; (Campo Formoso, Serra do Ramalho) and other similar processes affecting the epikarst may be at more complex vegetational types (Serra da the origin of I. epikarsticus and I. ramiroi Bodoquena, Jaíba, Itacarambi, Chapada (Bichuette & Trajano, 2004). Topographic Diamantina). They occupy different kinds of isolation may also explain the significant habitat, from open channel streams including morphological differences described for distinct base-level streams (A. cryptophthalmus, T. populations of A. cryptophthalmus separated by itacarambiensis, Trichomycterus sp. 2, P. kronei, high and very strong waterfalls inside the caves. Rhamdia sp., Rhamdiopsis sp., E. It can be concluded that the high diversity of the vicentespelaea) and vadose tributaries and pools subterranean ichthyofauna in Brazil results from fed by epikarst waters (I. bambui, I. epikarsticus, a combination of epigean megadiversity I. ramiroi, P. spelaea) to the upper phreatic zone (potentially colonizing species) and opportunities in caves (A. formoso, Trichomycterus sp. 1, for isolation. Taunayia sp., the new heptapterid genus) and in Several Brazilian subterranean fishes have been non-cave areas (S. typhlops); P. cisternarum is investigated in detail with regards to their found in shallow wells in the alluvial fan around population ecology, representing the majority of the Amazon delta. Although no taxonomic studies with this focus for cavefish around the correlation is observed with habitat types, there world. Five species (P. kronei, T. itacarambiensis, seems to be a geographic correlation with A. cryptophthalmus, I. bambui and I. passensis) degree of troglomorphism, i.e., even distantly have been studied using mark-recapture (MR) related species co-occurring in the same karst techniques. Population densities estimated on areas tend to present similar degrees of basis of MR data and/or visual censuses greatly troglomorphism, suggesting vicariant events varied, from very low (0.01 ind. m-2 in Taunayia affecting the whole subterranean fauna. In fact, sp.) to relatively high (average of 1.0 ind. m-2, areas harbouring highly to moderately maximum up to 10 ind. m-2 in A. specialized troglobitic fishes, such as the cryptophthalmus and Rhamdia sp.), without a currently semiarid karst areas in Bahia and the clear taxonomic correlation. Apparently, the main forested Upper Ribeira, are also hotspots of factor conditioning population densities is food biodiversity for troglobitic invertebrates. availability. Population sizes, as a result of the For several species, allopatric differentiation combination of population densities and related to paleoclimatic fluctuations leading extension of geographic range, also vary from

162 SWSB December 2004 POSTERS: EXTENDED ABSTRACTS few thousands (P. kronei, A. cryptophthalmus in Ramalho), and disturbance due to human Passa Três Cave) to several dozens of visitation (A. formoso, P. kronei). Few species thousands of individuals (A. cryptophthalmus in live within the boundaries of Conservation Units, Angélica Cave). It is noteworthy that populations and even this is not enough to guarantee their with relatively high densities may be quite low as protection because of the poor control of a result of restricted geographic distribution, as is potentially harmful activities by the authorities. the case of T. itacarambiensis, with a population density of 0.15 ind. m-2 and a total population of BiBLIOGRAPHY approximately 1,000 individuals (Trajano, 2001). Bichuette, M.E. and Trajano, E. 2004. Three new However, the knowledge gathered is still beneath subterranean species of Ituglanis from Central Brazil the necessary in view of the serious (Siluriformes: Trichomycteridae). Ichthyological environmental problems threatening several of Explorations of Freshwaters 15(3):243-256. these species. Such threats include lowering of Trajano, E. 1997. Synopsis of Brazilian troglomorphic the water table (an immediate threat for S. fishes. Mémoires de Biospéologie 24:119-126. typhlops and a potential one for Taunayia sp., Trajano, E. 2001. Ecology of subterranean fishes: an both among the most specialized and interesting overview. Environmental Biology of Fishes 62(1- Brazilian cave fishes), water pollution (the new 3):133-160. heptapterid genus), deforestation of headwaters Trajano, E. and Pinna, M.C.C. 1996. A new cave species (all the cave ichthyofauna in São Domingos of Trichomycterus from eastern Brazil (Siluriformes, Trichomycteridae). Revue française d’Aquariologie region), substitution of the natural vegetation by 23(3-4):85-90. soyabean and cotton cultures (fishes in Serra do

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164 SWSB December 2004 POSTERS: ABSTRACTS

A NEW SUBTERRANEAN CATFISH, GENUS RHAMDIA, FROM BRAZIL (SILURIFORMES : HEPTAPTERIDAE) WITH NOTES ON ECOLOGY

M.E. BICHUETTE and E. TRAJANO

Departamento de Zoologia, Instituto de Biociências da USP, C.P. 11461, 05422-970. São Paulo, SP. Brasil. ([email protected]), ([email protected]).

A new species of Rhamdia was recorded from subterranean waters in South-western Bahia, North-eastern Brazil. This species represents the first record of the genus for the subterranean realm in Brazil and one of the largest subterranean fishes recorded until now (225.0 mm SL). From about 20 species of cave fishes registered in Brazil, five are heptapterids. Troglomorphic characters are present in this species, such as reduction of eyes and melanic pigmentation, showing high variability. The population of Rhamdia from Enfurnado Cave comprises thousands of individuals distributed along 3,000 km of the subterranean stream. We observed 12 individuals.m-2 in a stream reach. High amount of organic matter is present in the cave, which shows that the food is not scarce and dissolved oxygen is really low (1.43 mg.l-1). Adult individuals are distributed in the bottom, midwater and most in the surface. Juveniles were observed hiding under boulders. Type-locality of Rhamdia is not legally protected and the region is under treat of deforestation. This can lead this population to extinction, once the food came entirely from the surface (floods between October and March). Protection actions are urgent and for this we are describing this species. Besides, we are going to start a population study of two years, using mark-recapture methods, in the next year.

HABITAT DETERMINATION AND SUBTERRANEAN SINGULARITY

A.I. CAMACHO, A.G. VALDECASAS and J. RODRIGUEZ

Museo Nacional de Ciencias Naturales, C/ Jose Gutierrez Abascal, 2, 28006-MADRID, Spain. ([email protected]), ([email protected]).

Historically, the aquatic subterranean environment has been clearly associated with caves. The main diagnostic characteristic being the absence of light. More recently, other habitats like the interstitial, ankialine and profound benthos of sea have been added. The biological counterpart to a physical definition of the subterranean environment is to look for shared characteristics of organisms living in this realm and not presented anywhere else. These characteristics could be used to define the stygobiont (strictly subterranean) organism. Both sides, the physical and the biological, have their bias and controversies. In this work we look at another side of the “subterranean singularity”: the sampling gear reliance. We studied the chemistry and biology of subterranean samples taken from the interstitial environment, wells, caves and springs with different methods: simple hand-nets, Karaman-Chappuis digging method and Bou-Rouch pipe. Data analysis included multidimensional scaling ordination, and permutation test of similarity among samples. It can be concluded that some subterranean habitats are more “subterranean” than others, and that a “subterraneanbility” gradient could be established among the sampling gear/habitat sampled combinations.

SWSB December 2004 165 POSTERSPOSTERS: ABSTRACTS

EFFECTS OF BASE FLOW REDUCTION ON AQUATIC ECOSYSTEMS

G. CECCHI, P. FORMICHETTI and L. MANCINI

Laboratorio di Igiene Ambientale, Istituto Superiore di Sanità, Roma Viale Regina Elena, 299 00161 Roma, Italy. ([email protected]), ([email protected]), ([email protected]).

Italian National Institute of Health (ISS) strongly supports the integrated study of the surface water ecosystems. Following this principle, one major subject of investigation is the interaction between surface water ecosystems and all surrounding systems, both physical and ecological ones. In order to apply this strategy, ISS and other institutions developed the “Fluvial Functioning Index” (FFI) which accounts for vegetation conditions of the banks and the territory, physical and morphological structure of the banks, structure of the wet river bed, biological habitat composition and biological characteristics of the periphyton, macrophytes and macrobenthos. In many studies carried out by ISS, the impact of groundwater status on surface ecosystems turned out to be a major issue of concern. In the Almone River, in the “Appia Antica” Regional Park, the lowering of the water table and the infiltration of poor quality water from surface (mostly collected waters) determined poor groundwater quality (assessed by microbiological indicators). In the “Doganella wetland”, located in the “Castelli Romani“ Regional Park, heavy water abstraction for drinking water supply determined the progressive drying of the area which caused a negative ecosystem impact measured in terms of biodiversity reduction (ex. absence of Simulidae Diptera Nematocera species). Last, in the ”Topino” River, monitored by the Environmental Regional Authority of Umbria (ARPA), the pressure on surface aquatic ecosystem took the form of hydromorfological alterations such as riverbed straightening, artificialization and riparian vegetation removal, these alterations are likely to affect surface and sub-surface waters connection; in this site, a heavy impact was measured in terms of biological quality (low values for EBI) and ecological functionality (low values for FFI) despite good physiochemical quality and unaltered flow regime.

INTERSTITIAL FAUNA DYNAMIC IN AN ISLAND-BRAIDED FLOOD PLAIN (TAGLIAMENTO RIVER, ITALY)

1C. CLARET and 2K. TOCKNER

1. Institut Méditerranéen d’Ecologie et de Paléoécologie (UMR CNRS 6116), Université Paul-Cézanne Aix-Marseille 3, Case 31, 13397 Marseille cedex 20, France. ([email protected]). 2. Department of Limnology, EAWAG/ETH, Überlandstrasse 133, 8600 Dübendorf, Switzerland. ([email protected]).

Surface-subsurface exchanges, hydrological flow paths, and sediment characteristics (size and distribution) lead to a mosaic of habitat conditions in the interstitial compartment that influence density, diversity, and distribution of faunal assemblages. In riffle-pool sequences or gravel bars, floods that modify the extent and configuration of surface-subsurface exchange patches, resulted in variations in the composition and distribution of invertebrate within sediments at fine scale. However, at large scale interstitial fauna dynamics in response to flow and flood pulse are less documented. The Tagliamento River (Italy) experienced rapid and frequent changes of water level. In this highly dynamic river, the combination of small scale vertical exchanges, and large scale disturbances should result in complex dynamics of interstitial animals at the floodplain scale. Monthly sampling of faunal assemblages beneath lentic and lotic water bodies distributed across an island-braided flood plain was used to investigate, at large scale, the response of invertebrate assemblages to changes in water level.

166 SWSB December 2004 POSTERS: ABSTRACTS

DIVERSITY AND ORIGIN OF THE INTERSTITIAL ISOPOD MICROCHARON (CRUSTACEA, MICROPARASELLIDAE) FROM THE ROUSSILLON AND WESTERN LANGUEDOC REGIONS (FRANCE)

1N. COINEAU, 2M. ARTHEAU, 3A. BEDOS, 4M. BOULANOUAR, 2C. BOUTIN, 5F. BREHIER, 3L. DEHARVENG and 6N. GIANI

1. Université P et M Curie-Paris 6, Observatoire Océanologique de Banyuls, Laboratoire Arago, 66650 Banyuls/Mer, France. ([email protected]). 2. Université de Toulouse, LaDyBio-UMR CNRS 5172, Bât 4R3, 118 route de Narbonne, 31062 Toulouse cedex 04, France. ([email protected]), ([email protected]). 3. Museum National d’Histoire Naturelle, ESA CNRS 8043, Laboratoire d’entomologie, 45 rue Buffon, 75005 Paris, France; ([email protected]). 4. Université de Marrakech, Faculté des Sciences Semlalia, UFR Sciences de l’eau, Laboratoire d’Hydrobiologie, BP 2390, Marrakech, Maroc. 5. Alas, 09800 Balaguères, France. 6. Université de Toulouse, EDH-Ecologie des Hydrosystèmes, Bat 4R3, 118 route de Narbonne, 31062 Toulouse Cedex 04, France. ([email protected])

The interstitial genus Microcharon (Isopoda Microparasellidae) is highly diversified within the Mediterranean basin. In the framework of the PASCALIS project, several species new for science were discovered, whereas the distribution pattern of other species was extended. In the Roussillon and western Languedoc regions, endemic species inhabit the four study basins of the Tech, the Têt, the Agly-Verdouble and the Aude rivers. They belong to the western Mediterranean phylogenetic group of species. These stygobites are derived from marine ancestors which formerly lived in the littoral interstitial of the Tethys. The two-step model of colonization and evolution, the second phase of which explains both the establishment in fresh groundwater and vicariance during Tethys regressions, provides an understanding of their evolutionary history. The species from the Aude basin evolved from an ancestor which might have settled in fresh groundwater at the end of the Middle Ilerdian regression of the Pyreneo-Provençal gulf (=lower Eocene, 48 MY BP). In the Tech and Têt basins, the ancestors of the two-three extant representatives of the genus might have been left by the regression of the Pliocene marine embayment which covered the southern Roussillon. Other species might result from a much more older Tethys regression in the late Senonian (end of the Cretaceous) together with further vicariance evolution due to the Pyrenees orogenesis and passive drift dowstream. The palaeobiogeographic history in the Verdouble basin within the Corbières is much more complicated since a large allochtonous part of this mountain sliped northwards from the Pyrenees. Nevertheless, a late Senonian gulf flooded the region. Finally, regressions of more or less old marine embayments may have played a major role in the evolution through vicariance in the genus Microcharon.

SWSB December 2004 167 POSTERSPOSTERS: ABSTRACTS

OBLIGATE GROUNDWATER FAUNA OF FRANCE : SPECIES DIVERSITY PATTERNS AND CONSERVATION IMPLICATIONS

1D. FERREIRA, 2F. MALARD, 2M.-J. DOLE-OLIVIER and 1J. GIBERT

1. University Claude Bernard of Lyon 1, UMR CNRS 5023, Ecologie des Hydrosystèmes Fluviaux, Equipe Hydrobiologie et Ecologie Souterraines, 43, Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France. ([email protected]), ([email protected]). 2. CNRS - University Claude Bernard of Lyon 1, UMR CNRS 5023, Ecologie des Hydrosystèmes Fluviaux, Equipe Hydrobiologie et Ecologie Souterraines, 43, Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France. ([email protected]), ([email protected]).

A data base of species occurrences was established on obligate groundwater fauna (stygobites) and used to examine taxonomic and geographic patterns at the scale of France Studies carried out at the community level were selected to estimate the spatial distribution of sampling effort and revealed that biological exploration of ground waters was mainly restricted to karst areas. Large unconsolidated aquifers situated in northern and western zones, the centre area, and the main fluvial corridors, remained quasi-unexplored. The total number of stygobite species and subspecies (381) is distributed within a high number of genera (100) and families (39). Conversely to the surface water biodiversity, obligate groundwater fauna is strongly dominated by the group of Crustacea. The species accumulation curve over time (sampling effort) did not reach a plateau, indicating that groundwater biodiversity was still largely underestimated. The contribution of stygobite richness to the biodiversity of continental freshwater was higher than previously expected and in some cases exceeded the number of obligate surface water species. Two measurements of the geographic range size (area of occupancy and latitudinal extent of occurrences) demonstrated, that despite the accumulation of data over time, the geographic range size of the majority of obligate groundwater species remained spatially limited supporting their high level of endemism. An optimal selection of sites has been processed by using complementary software to obtain a proposition combining the highest biodiversity within the lowest set of sites. This selection should be considered as a trial, a first element of discussion towards the integration of fauna into groundwater protection processes.

168 SWSB December 2004 POSTERS: ABSTRACTS

DIVERSITY PATTERNS OF COPEPOD ASSEMBLAGES (CRUSTACEA, COPEPODA) OF DIFFERENT CAVES FROM SOUTHERN AND CENTRAL ITALY

E. GATTONE, T. DI LORENZO, B. FIASCA, P. DE LAURENTIIS and D.M.P. GALASSI

Dipartimento di Scienze Ambientali, University of L’Aquila, Via Vetoio, Coppito, 67100 L’Aquila, Italy. ([email protected]), ([email protected]), ([email protected]), ([email protected]).

This contribution presents the first results of an analysis on a biological data set collected in six distinct active caves, four of which located within carbonatic massifs, the remaining two in gypsum formations. Faunal samples have been collected following different methodologies for each habitat typology encountered. The taxonomic group of Crustacea Copepoda is often found to be the most relevant component, in both terms of abundance and species richness, in the communities of most groundwater habitats, and it seems quite likely that the taxonomic diversity of copepod assemblages closely matches the overall diversity of the entire communities under consideration, as they may represent a focal group sensu Hammond (1995), possibly allowing an extension of the results of this study to the groundwater biodiversity of the areas investigated as a whole for the purpose of the establishment of future conservation strategies. The biological data have been processed by PRIMER v5 software in order to measure the taxonomic diversity based on phylogenetic relatedness of the species within samples, starting by a matrix of presence/absence of species. Information about geology, palaeogeography, microhabitat heterogeneity, hydrological regime has been used to evaluate the major determinants affecting the different values observed in the taxonomic distinctness for each cave. Paleogeographical factors, habitat heterogeneity and the degree of isolation of the aquifer seem to be the most important parameters affecting the measured diversity patterns. Moreover, the taxonomic dispersion of the diversity might adequately integrate the information resulting from diversity indexes based only on species richness, evenness and abundance.

THE STYGOBIONTIC FAUNA OF TUNISIA : PRELIMINARY RESULTS

1A. GHLALA, 2D. DELLA VALE, 1F. CHARFI and 2G. MESSANA

1. Faculty of Sciences of Tunis, University campus, El Manar II, 2092 Tunis, Tunisia. ([email protected]). 2. Istituto per lo Studio degli Ecosistemi, CNR, Sezione di Firenze, Via Madonna del Piano, 10 - 50019 Sesto Fiorentino (FI), Italy ([email protected]).

A research work on biodiversity of Tunisian subterranean fauna of carcinological stygobies was begun since October 2003. In this presentation, a list of the subterranean aquatic isopods revealed in the North of Tunisia is detailed. We indicate, for the first time, in the Ichkeul’s National Park, the presence of the genus Typhlocirolana. Specimens of Typhlocirolana sp. population, from a well of surface in the park, are described and their specific morphological characteristics are clarified. Finally, a brief apercu on the geographical distribution of the stations of Proasellus in the northwest of Tunisia is given.

SWSB December 2004 169 POSTERSPOSTERS: ABSTRACTS

RELATIONSHIP BETWEEN MORPHOLOGICAL TAXONOMY AND MOLECULAR DIVERGENCES : EVIDENCES FROM CRUSTACEANS

1T. LEFÉBURE, 1C.J. DOUADY, 2M. GOUY and 1J. GIBERT

1. University Claude Bernard of Lyon 1, Laboratoire d’Ecologie des Hydrosystèmes fluviaux, UMR-CNRS 5023, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne Cedex, France. ([email protected]), ([email protected]), ([email protected]). 2. Laboratoire de Biométrie et Biologie Evolutive, UMR CNRS 5558, Université Claude Bernard Lyon 1, 43, bd. du 11 Novembre 1918, 69622 Villeurbanne Cedex, France. ([email protected]).

Species level is usually recognized as the major unit of biodiversity. Thus biodiversity assessment is directly dependent on taxonomy. However, morphological traits that represent the base for current taxonomy are non-neutral, complex and potentially highly biased makers. As noticed by the supporter of the DNA taxonomy, molecular markers seem of promising help. Nevertheless, to be employ conjointly morphological taxonomy and molecular markers should first show an overall correlation. This analysis tests this hypothetical relationship for two mitochondrial genes (COI and 16S) within the diversified group of Crustacea. Our analysis revealed that, when appropriately measured, there is a general but poor correlation between molecular divergence and taxonomy. At low taxonomic level morphology and molecular divergence tend to corroborate each other. Therefore our analyses tend to suggest that comparable amount of molecular and morphological evolution accompanies the phenomenon of speciation. In this way molecular divergences may be used to delimit what is usually called species by taxonomists. At upper level (i.e., genus, family), we find almost no correlation between morphology and molecule. If corroborated in other groups this would strongly compromise the possibility of using the COI as a universal marker for identification of any taxonomic level.

170 SWSB December 2004 PARTICIPANTS

SYMPOSIUM PARTICIPANTS

NAME ADDRESS E-MAIL

Zoological Institute of the Russian ALEXEEV Viktor Academy of Sciences, University emb. [email protected] 1, 199034 St Petersburg, Russia UMR-CNRS 5023, Ecology of Fluvial Hydrosystems, University Claude AMOROS Claude Bernard Lyon 1, Bât Darwin C, [email protected] 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France CNRS - Délégation Rhône-Alpes Site Vallée du Rhône ANDRAL Bruno [email protected] 2, Avenue Albert Einstein - BP 1335 69609 Villeurbanne cedex - France Université Paul Sabatier - Laboratoire Dynamique de la Biodiversité ARTHEAU Malvina UMR-CNRS 5172 [email protected] 118 Route de Narbonne - 31062 Toulouse cedex 04 - France Università della Tuscia Viterbo, Dipartimento di Scienze Ambientali, BERERA Raffaella [email protected] Largo dell`Università s.n.c. Blocco D, 01100 Viterbo - Italy ECMC - Abéla- 09320 Boussenac BERTRAND Alain [email protected] France Muséum national d’Histoire naturelle - Dpt Systématique et BICHAIN Jean-Michel Evolution - USM 602 - Taxonomie et [email protected] Collections - Eq Malacologie - CP 051 - 55, Rue Buffon - 75005 Paris - France Departamento de Zoologia, IBUSP, Rua BICHUETTE Maria Elina do Matao, travessa 14, n 101, 05508- [email protected] 900 SAO PAUL Brazil Università del Piemonte Orientale “A. BO Tiziano Avogadro”, Via Bellini 25 [email protected] 15100 Alessandria - Italy University Claude Bernard Lyon 1, UMR- CNRS 5023 - Laboratory of Fluvial Hydrosystems Ecology, Subterranean BOUGER Guillaume [email protected] Hydrobiology and Ecology, Bât. Forel, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France Université de Toulouse - LaDyBio UMR 5176 - Bât 4R3 BOUTIN Claude [email protected] 118 Route de Narbonne 31062 Toulouse cedex 04 - France National Institute of Biology, Vecna pot BRANCELJ Anton [email protected] 111, 1000 Ljubljana - Slovenia Università della Tuscia, Viterbo Dipartimento di Scienze Ambientali, BRUNO Maria Christina [email protected] Largo dell\’Università s.n.c. - Blocco D, 01100 Viterbo - Italy

SWSB December 2004 171 PARTICIPANTS

NAME ADDRESS E-MAIL

Agence de l’ Eau Rhône-Méditerranée- Corse - Délégation Rhône-Alpes CADILHAC Laurent [email protected] 14, rue Jonas Salk 69363 LYON Cedex 07 - France Museo Nacional de Ciencias Naturales, Dpto. de Biodiversidad y Biología CAMACHO Ana Isabel [email protected] Evolutiva,C/ José Gutiérrez Abascal 2, 28006 MADRID, Spain University of Rome “La Sapienza”, Dpt CAMPANARO Alessandro of Animal and Human Biology, Viale [email protected] dell’Università 32, 00185 ROME , Italy University Claude Bernard Lyon 1, UMR- CNRS 5023 - Laboratory of Fluvial Hydrosystems Ecology, Subterranean CASTELLARINI Fabiana [email protected] Hydrobiology and Ecology, Bât. Forel, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France CECCHI Giulano Via Tiburtina 720 - 00159 Rome - Italy [email protected] Ecologie des Eaux Continentales Méditerranéennes, IMEP-UMR CNRS 6116 - Faculté des Sciences et CLARET Cécile [email protected] Techniques de Saint-Jérôme (case 31), Université Aix-Marseille III, 13397 MARSEILLE Cedex 20 - France Observatoire Océanologique de COINEAU Nicole Banyuls, Laboratoire Arago, [email protected] 66650 BANYULS-SUR-MER, France The American University, Department of CULVER David Biology, 4400 Massachussetts Avenue - [email protected] NW 20016 WASHINGTON, DC - USA IRBI - UMR-CNRS 6035, Université de DANGLES Olivier Tours, Parc Grandmont, 37200 TOURS - [email protected] France Austrian Academy of Sciences, Institute of Limnology, Department Mondsee - DANIELOPOL Dan L. [email protected] Mondseestrasse 9, 5310 MONDSEE, Austria Institut Royal des Sciences Naturelles DE BROYER Claude de Belgique - Rue Vautier 29 [email protected] 1000 Bruxelles - Belgique Museum National d’ Histoire Naturelle de Paris, USM 601 - CP 50 DEHARVENG Louis [email protected] FR 2695 du CNRS - 45 rue Buffon 75005 Paris - France Dipartimento di Scienze Ambientali, DI LORENZO Tiziana University of L’Aquila, Via Vetoio, [email protected] Coppito, 67100 L’Aquila, Italy University Claude Bernard Lyon 1, UMR- CNRS 5023 - Laboratory of Fluvial Hydrosystems Ecology, Subterranean DOLE-OLIVIER Marie-José [email protected] Hydrobiology and Ecology, Bât. Forel, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France

172 SWSB December 2004 PARTICIPANTS

NAME ADDRESS E-MAIL

University Claude Bernard Lyon 1, UMR- CNRS 5023 - Laboratory of Fluvial Hydrosystems Ecology, Subterranean DOUADY Christophe [email protected] Hydrobiology and Ecology, Bât. Darwin C, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France Department of Conservation and Land Management - Woodvale Research EBERHARD Stefan [email protected] Center - PO Box 51 - Wanneroo - WA 6946 - Australia Raiffeisenstrasse 5, 85457 Woerth- FALKNER Gerhard [email protected] Hoerlkofen, Germany Raiffeisenstrasse 5, 85457 Woerth- FALKNER Margrit [email protected] Hoerlkofen, Germany Università del Piemonte Orientale - C.so FENOGLIO Stefano [email protected] Borsalino 54 -15100 Alessandria - Italy University Claude Bernard Lyon 1, UMR- CNRS 5023 - Laboratory of Fluvial Hydrosystems Ecology, Ecology and FERREIRA David [email protected] Subterranean Hydrobiology, Bât. Forel, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France Dipartimento di Scienze Ambientali, FIASCA Barbara University of L’ Aquila, Via Vetoio, [email protected] Coppito, 67100 L’Aquila, Italy University of Western Australia - School FINSTON Terrie of animal biology (MO92) - 35 Stirling [email protected] Hwy - 6009 CRAWLEY - Australia Facultés Universitaires, Notre Dame de FLAMEN Caroline la Paix, Département Géologie, Rue de [email protected] Bruxelles, 61, 5800 Namur, Belgium University Claude Bernard Lyon 1, UMR-CNRS 5023 - Laboratory of Fluvial Hydrosystems Ecology, Subterranean FOULQUIER Arnaud [email protected] Hydrobiology and Ecology, Bât. Forel, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France University of Western Australia - School FRANCIS Cara of animal biology (MO92) - 35 Stirling [email protected] Hwy - 6014 PERTH - Australia University of Landau - Department of FUCHS Andreas Biology, Im Fort 7, 76829 Landau, [email protected] Germany Dipartimento di Scienze Ambientali, GALASSI Diana University of L’ Aquila, Via Vetoio, [email protected] Coppito, 67100 L’Aquila, Italy Dipartimento di Scienze Ambientali, GATTONE Enrico University of L’ Aquila, Via Vetoio, [email protected] Coppito, 67100 L’Aquila, Italy Faculté des Sciences de Tunis - GHLALA Adnène Campus universitaire El Manar II - 1060 [email protected] TUNIS - Tunisia

SWSB December 2004 173 PARTICIPANTS

NAME ADDRESS E-MAIL

University Claude Bernard Lyon 1, UMR-CNRS 5023 - Laboratory of Fluvial Hydrosystems Ecology, Subterranean GIBERT Janine [email protected] Hydrobiology and Ecology, Bât. Forel, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France University Claude Bernard Lyon 1, UMR-CNRS 5023 - Laboratory of Fluvial Hydrosystems Ecology, Ecology and GINET René - Subterranean Hydrobiology, Bât. Forel, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France University of Landau - Department of HAHN Hans Jürgen Biology, Im Fort 7, 76829 Landau, [email protected] Germany University Claude Bernard Lyon 1, UMR-CNRS 5023 - Laboratory of Fluvial Hydrosystems Ecology, Subterranean HERVANT Frédéric [email protected] Hydrobiology and Ecology, Bât. Forel, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France Western Australian Museum, Locked HUMPHREYS Williams Bag 49, Welshpool DC, WA 6986, [email protected] Australia The Natural History Museum, Dpt of HUYS Rony Zoology, Cromwell Road, London, [email protected] SW7 5BD, UK. University Claude Bernard Lyon 1, UMR-CNRS 5023 - Laboratory of Fluvial Hydrosystems Ecology, Subterranean ISSARTEL Julien [email protected] Hydrobiology and Ecology, Bât. Forel, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France University Claude Bernard Lyon 1, UMR- CNRS 5023 - Laboratory of Fluvial JOLY Pierre Hydrosystems Ecology, Bât. Darwin C, [email protected] 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France Karst Waters Institute - PO Box 537 - JONES Williams K. [email protected] Charles Town - WV - USA University Claude Bernard Lyon 1, UMR-CNRS 5023 - Laboratory of Fluvial JUGET Jacques Hydrosystems Ecology, Bât. Darwin C, - 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France Cemagref - 3 bis quai Chauveau - CP LAFONT Michel [email protected] 220 - 69336 Lyon cedex 09 - France University Claude Bernard Lyon 1, UFR Chimie Biochimie, Laboratoire LANTERI Pierre Chimiométrie, ERT 11, Bât CPE, [email protected] 43 Bd du 11 Novembre 1918 69622 VILLEURBANNE cedex - France

174 SWSB December 2004 PARTICIPANTS

NAME ADDRESS E-MAIL

University Claude Bernard Lyon 1, UMR- CNRS 5023 - Laboratory of Fluvial Hydrosystems Ecology, Subterranean LEFEBURE Tristan [email protected] Hydrobiology and Ecology, Bât. Darwin C, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France 4, Avenue Allende - 69100 Villleurbanne LIPS Josiane [email protected] France University of Bourgogne - Life Sciences- MAGNIEZ Guy Bât. Gabriel - 6, bd Gabriel [email protected] 21000 DIJON - France University Claude Bernard Lyon 1, UMR-CNRS 5023 - Laboratory of Fluvial Hydrosystems Ecology, Subterranean MALARD Christine - Hydrobiology and Ecology, Bât. Forel, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France University Claude Bernard Lyon 1, UMR-CNRS 5023 - Laboratory of Fluvial Hydrosystems Ecology, Subterranean MALARD Florian [email protected] Hydrobiology and Ecology, Bât. Forel, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France UMR-CNRS 5117 - LEH - Université Paul Sabatier - Bât 4R3 b2 - 118 Route MANGIN Alain [email protected] de Narbonne - 31062 TOULOUSE cedex 4 - France University Claude Bernard Lyon 1, UMR-CNRS 5023 - Laboratory of Fluvial Hydrosystems Ecology, Subterranean MARTIN Dominique [email protected] Hydrobiology and Ecology, Bât. Forel, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France Institut Royal des Sciences Naturelles MARTIN Patrick de Belgique, Biologie des Eaux Douces, [email protected] 29 rue Vautier, 1000 Bruxelles, Belgium University Claude Bernard Lyon 1, UMR-CNRS 5023 - Laboratory of Fluvial Hydrosystems Ecology, Subterranean MATHIEU Jacques [email protected] Hydrobiology and Ecology, Bât. Forel, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France University Claude Bernard Lyon 1, UMR-CNRS 5023 - Laboratory of Fluvial MERMILLOD-BLONDIN Hydrosystems Ecology, Subterranean [email protected] Florian Hydrobiology and Ecology, Bât. Forel, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France Ecole Française de Spéléologie - comite.speleo.rhone- MEYSSONIER Marcel 28, rue SÏur Janin - 6905 Lyon - [email protected] France

SWSB December 2004 175 PARTICIPANTS

NAME ADDRESS E-MAIL

Commission Wallone d’Etude et de Protection des Sites Souterrains, MICHEL Georges [email protected] CWEPSS, 21, Avenue Rodin, 1050 Bruxelles, Belgium CBGP - UMR INRA-IRD-CIRAD-Agro.M. Campus Int. De Baillarguet MORAND Serge [email protected] CS 300016 - 34988 Montferrier sur Lez Cedex - France National Institute of Biology, Vecna pot MORI Natasa [email protected] 111, 1000 Ljubljana, Slovenia MULOT José 4 rue du Village - 80160 Thoix - France [email protected] University Claude Bernard Lyon 1, UMR-CNRS 5023 - Laboratory of Fluvial Hydrosystems Ecology, Subterranean NOGARO Géraldine [email protected] Hydrobiology and Ecology, Bât. Forel, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France Irlands Brygge 63C 5tv - 2300 NOTENBOOM Jos [email protected] Copenhagen - Denmark University Claude Bernard Lyon 1, UMR-CNRS 5023 - Laboratory of Fluvial Hydrosystems Ecology, Subterranean NOUNE Valérie [email protected] Hydrobiology and Ecology, Bât. Forel, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France Centre SITE - Ecole nat. Sup. des Mines PARAN Frédéric de St-Etienne - 158 cours Fauriel - [email protected] 42023 Saint-Etienne cedex 2 - France University Claude Bernard Lyon 1, UMR-CNRS 5023 - Laboratory of Fluvial PATTEE Eric Hydrosystems Ecology, Bât. Darwin C, [email protected] 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France Karst Research Institute ZRC SAZU, PIPAN Tanja [email protected] Titov trg 2 - 6230 POSTOJNA - Slovenia University Claude Bernard Lyon 1, UMR-CNRS 5023 - Laboratory of Fluvial PONT Didier Hydrosystems Ecology, Bât.Forel, [email protected] 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France La Clastre PRIE Vincent [email protected] 34520 St Maurice Navacelles - France British Cave Research Association Biological Recorder Department of PROUDLOVE Graham Zoology, The Manchester Museum, [email protected] University of Manchester, MANCHESTER M13 9PL, UK Australian National University REEVES Jessica Dpt of Earth and Marine Sciences [email protected] 0200 ACT - Australia

176 SWSB December 2004 PARTICIPANTS

NAME ADDRESS E-MAIL

Environmental Agency of REJEC-BRANCELJ Irena the Republic of Slovenia, Vojkova 1b, [email protected] 1000 Ljubljana, Slovenia SNSD Mus. Für Tierkunde SCHMIDT Susanne Königsbrücker Str. 159 [email protected] 01099 Dresden - Germany University of Maryland SCHNEIDER Katie 1204 Biology Psychology Building [email protected] 20742 College Park MD - USA Department of Biology, Biotechnical Faculty, University of Ljubljana, SKET Boris [email protected] Vecna pot 111, P.P. 2995, 1001 Ljubljana, Slovenia Museum of Natural History, Viale le XXV STOCH Fabio [email protected] Aprile 24, 34015 - MUGGIA (TS), Italy University of Sao Paulo, Instituto Biociências da USP, Departamento de TRAJANO Eleonora [email protected] Zoologia CEP -05422-970 SAO PAULO, SP - Brazil Department of Biology, Biotechnical Faculty, University of Ljubljana, TRONTELJ Peter [email protected] Vecna pot 111, P.P. 2995, 1001 Ljubljana, Slovenia University Claude Bernard Lyon 1, UMR-CNRS 5023 - Laboratory of Fluvial Hydrosystems Ecology, Subterranean TURQUIN Marie-José [email protected] Hydrobiology and Ecology, Bât. Forel, 43 Bd du 11 Novembre 1918, 69622 Villeurbanne cedex, France Museo Nacional de Ciencias Naturales, Dpto. de Biodiversidad y Biología VALDECASAS G. Antonio [email protected] Evolutiva, C/ José Gutiérrez Abasca 2, 28006 MADRID, Spain

SWSB December 2004 177

AUTHORS

AUTHOR INDEX

Alexeev Viktor R...... 143 Giani Narcisse...... 167 Argano Roberto...... 131 Gibert Janine ... 13, 39, 73, 75, 79, 83, 107, 168, 170 Artheau Malvina...... 129, 167 Gouy Manolo...... 73, 170 Barron Harley J...... 61 Hahn Hans Jürgen...... 89 Bedos Anne...... 167 Halse Stuart A...... 61, 69 Berera Raffaella...... 145 Hobbs III Horton H...... 138 Bertrand Alain...... 149 Humphreys Williams F...... 57, 69 Bichuette Maria Elina...... 161, 165 Huys Rony...... 134 Bo Tiziano...... 153 Johnson Mike S...... 69, 133 Boulanouar Mohamed...... 167 Joly Pierre...... 25 Boutin Claude...... 167 Jones William K...... 134 Bradbury John H...... 57 Ketmaier Valerio...... 131 Brancelj Anton...... 39, 55,129 Lafont Michel...... 99, 135 Brehier Franck...... 167 Lefébure Tristan...... 73, 132, 170 Briolay Jérôme...... 73 Madden Molly...... 132 Bruno Maria Christina...... 130, 145 Malacarne Giorgio...... 153 Camacho Ana I...... 39, 165 Malard Florian...... 39, 75, 79, 83, 99, 107, 135, 168 Campanaro Alessandro...... 131 Mancini Laura...... 157, 166 Castellarini Fabiana...... 39, 83 Marmonier Pierre...... 99 Cecchi Giuliano...... 166 Martin Patrick...... 39, 95 Charfi F...... 169 Mathieu Jacques...... 99 Christman M.C...... 132 Messana Giuseppe...... 169 Claret Cécile...... 166 Michel Georges...... 39, 95 Cocking Jim S...... 61 Morand Serge...... 21 Coineau Nicole...... 167 Mori Natasa...... 129 Cottarelli V...... 130, 145 Paran Frédéric...... 99 Cucco Marco...... 153 Pariciani L...... 145 Culver David C...... 27, 55, 132, 136, 138 Pennelli B...... 157 Danielopol Dan L...... 29 Pipan Tanja...... 136 Datry Thibault...... 107 Pospisil Peter...... 29 De Broyer Claude...... 39, 95 Proudlove Graham S...... 139 De Laurentiis Paola...... 115, 133, 134, 169 Ranalli F...... 115 Deharveng Louis...... 39, 55, 167 Reeves Jessica...... 137 Della Vale D...... 169 Rodriguez Jaime...... 165 Di Lorenzo Tiziana...... 115, 133, 169 Sablon Rose...... 95 Dole-Olivier Marie-José...... 39, 75, 79, 83, 168 Scanlon Mike D...... 61 Douady Christophe J...... 39, 73, 132, 170 Schneider Katie...... 138 Eberhard Stefan M...... 61, 69 Sket Boris...... 39, 55, 138 Fenoglio Stefano...... 153 Stoch Fabio...... 39, 115 Ferreira David...... 75, 168 Tockner Klement...... 166 Fiasca Barbara...... 115, 133, 169 Trajano Eleonora...... 161, 165 Fiers Frank...... 95 Trontelj Peter...... 39, 73 Finston Terry...... 69, 133 Valdecasas Antonio G...... 39, 165 Fong Daniel...... 138 Venanzi D...... 157 Formichetti Paolo...... 157, 166 Volterra Laura...... 157 Francis Cara. J...... 133 Watts Chris H.S...... 57 Fuchs Andreas...... 89 White D...... 132 Galassi Diana M.P...... 39, 99, 115, 133, 134, 169 Wood Paul J...... 139 Gattone Enrico...... 115, 169 Wouters Karel...... 95 Ghlala Adnen...... 169

SWSB December 2004 179

SYMPOSIUM - PICTURES

SWSB December 2004 181 SYMPOSIUM - PICTURES

182 SWSB December 2004