ARSOLOGICA Tanja PiPan •E Pikars T – a Promising habiTaT C CoPEPod fauna, iTs divErsiTy and ecology: a CasE sTudy from slovEnia (EuroPE) A-Carsologica_Pipan.indd 1 7.3.2007 9:43:14 Carsologica 5 Urednik zbirke / Series Editor Franci Gabrovšek Tanja Pipan Epikarst – A Promising Habitat Copepod Fauna, its Diversity and Ecology: A Case Study from Slovenia (Europe) © 2005, Založba ZRC, Inštitut za raziskovanje krasa ZRC SAZU ZRC Publishing, Karst Research Institute at ZRC SAZU Recenzenta / Reviewed by David C. Culver in/and Horton H. Hobbs III Jezikovni pregled / Language editing David C. Culver, Trevor R. Shaw Oblikovanje / Graphic art and design Milojka Žalik Huzjan Izdala in založila / Published by Inštitut za raziskovanje krasa ZRC SAZU, Založba ZRC Karst Research Institute at ZRC SAZU, ZRC Publishing Zanj / Represented by Tadej Slabe, Oto Luthar Glavni urednik / Editor-in-Chief Vojislav Likar Tisk / Printed by Collegium graphicum, d. o. o., Ljubljana Izdajo je finančno podprla / The publication was financially supported by Agencija za raziskovalno dejavnost Republike Slovenije Slovenian Research Agency Fotografiji na naslovnici / Front cover photos by Marko Pršina CIP - Kataložni zapis o publikaciji Narodna in Univerzitetna knjižnica, Ljubljana 595.34(497.4) PIPAN, Tanja, 1970- Epikarst, a promising habitat : copepod fauna, its diversity and ecology : a case study from Slovenia (Europe) / Tanja Pipan. - Postojna : Inštitut za raziskovanje krasa ZRC SAZU = Karst Research Institute at ZRC SAZU ; Ljubljana : Založba ZRC = ZRC Publishing, 2005. - (Carsologica ; 5) ISBN 961-6500-90-2 (Založba ZRC) 220477696 A-Carsologica_Pipan.indd 2 7.3.2007 9:43:14 EPIKARST – A PROMISING HABITAT COPEPOD FAUNA, ITS DIVERSITY AND ECOLOGY: A CASE STUDY FROM SLOVENIA (EUROPE) TANJA PIPAN POStOjnA – LjubLjAnA 2005 A-Carsologica_Pipan.indd 3 7.3.2007 9:43:14 A-Carsologica_Pipan.indd 4 7.3.2007 9:43:14 This book is dedicated to my daughter Laura with all my love. A-Carsologica_Pipan.indd 5 7.3.2007 9:43:14 A-Carsologica_Pipan.indd 6 7.3.2007 9:43:14 Contents FOREWORD ........................................................................................................................9 PREFACE ...........................................................................................................................11 INTRODUCTION ..............................................................................................................13 Introduction to the Epikarst ...........................................................................................14 Definition of Project Goals and Research Questions/Hypotheses ................................15 Introduction to the Copepods ........................................................................................16 Copepods in Subterranean Habitats and their Adaptations ...........................................17 Diversity of Hypogean Fauna in the Dinaric Region and Distribution Patterns of Copepods...................................................................................................................20 Survey of Researches of Copepods in Slovenia up to the Present ................................21 GEOGRAPHICAL LOCATION OF THE RESEARCH CAVES AND DESCRIPTION OF RESEARCH METHODS ..................................................................30 Precipitation ..................................................................................................................32 Field Work .....................................................................................................................33 Laboratory Work ...........................................................................................................34 Physical and Chemical Analysis ...................................................................................35 Biological Analysis and Statistical Treatment ..............................................................35 EPIKARST fauna FROM SIx caves ..........................................................................38 Composition of Epikarst Copepods in Six Karst Caves ................................................39 Similarity of Copepod Communities.............................................................................45 Spatial and Temporal Distrtibution of Copepods in Trickles and Pools Filled with Percolation Water ..................................................................................................46 Postojnska jama .......................................................................................................46 Pivka jama ...............................................................................................................47 Črna jama .................................................................................................................49 Škocjanske jame ......................................................................................................50 Dimnice ...................................................................................................................52 Županova jama ........................................................................................................53 Canonical Correspondence Analyses (CCA) ................................................................55 Abundance in Different Types of Pools ........................................................................59 A-Carsologica_Pipan.indd 7 7.3.2007 9:43:15 Diversity of Copepod Communities ..............................................................................59 Estimating Epikarst Copepods’ Richness ......................................................................59 PATTERNS OF EPIKARST COPEPOD BIODIVERSITY ...............................................61 Differences in Faunal Composition between Drips and Pools ......................................66 Population Parameters ...................................................................................................68 Ecosystem Structure and Dynamics of Epikarst ...........................................................69 Conclusions ...................................................................................................................71 SUMMARY ........................................................................................................................73 REFERENCES ...................................................................................................................77 APPENDIx .........................................................................................................................87 A-Carsologica_Pipan.indd 8 7.3.2007 9:43:15 FOREWORD lovenia is situated in the very northwestern end of the SlDinaric area, the site of the richest subterranean fauna in the world. The subterranean fauna increases only slightly to global biodiversity, but is of a high scientific importance in the study of mechanisms of colonisation of extreme environments as well as the nature of adaptations to these environments. I started my scientific career with both the taxonomic and the ecological investigations of copepods in hypogean environments of Slovenia. Copepod crustaceans may be the most abundant metazoans on Earth and can be found in virtually every kind of aquatic habitat from the deepest ocean trenches to damp leaf litter at high altitude in the Himalayas and from hot springs to polar seas. They are frequently found in all kinds of hypogean habitats. My research was aimed at unravelling the spatial structure of copepod fauna in the epikarst. In particular, I was interested in understanding the distribution and the ecology of copepods in the epikarst zone. For understanding of the structure and physico-chemistry of epikarst, I also studied the ecological and microgeographic distribution of epikarst copepods. Research presented in this monograph was conducted at the Karst Research Institute ZRC SAZU (Postojna) and National Institute of Biology (Ljubljana) within the framework of the Ministry of Education, Science and Sport’s programme under the mentorship of Dr. Anton Brancelj and Dr. Andrej Blejec. The hypogean fauna from percolation water and from pools, filled with percolation water from six cave systems in Slovenia, was studied in the period 2000–2003. Besides collecting fauna, parallel measurements of physical properties of the percolation water, performing the chemical analyses of water in the laboratory and the determining of the specimens were conducted. The methodologies used to asses epikarst habitats and fauna living there were also useful for the evaluation of human impacts on the environment, especially karst waters. Research represents the pioneer work on selected ecological aspects of the epikarst fauna. Field observations and studies are continuing. After concluding my doctoral studies I advanced with my research of the epikarst fauna within the framework of the research programme of the Karst Research Institute ZRC SAZU. I enlarged my studies by par- ticipating in the bilateral project USA-Slovenia Cooperation in Science and Technology in years 2003 and 2004 which was supported by the Ministry of Education, Science and Sport (Slovenia) and Karst Waters Institute (USA). The book has benefited from comments by two referees and discussion with several specialists: Dr. Anton Brancelj, Dr. Andrej Blejec, Prof. Dr. Tone Novak and Dr. Peter A-Carsologica_Pipan.indd 9 7.3.2007 9:43:15 Trontelj who guided me during my PhD work. Dr. Anton Brancelj trained me in tax- onomy