This course is part of the programme
Karstology (Third Cycle)

Objectives and competences

The primary goal of this course is to give students the theoretical and practical knowledge for understanding the biology of subterranean habitats, to present them types of subterranean habitats, their ecological characteristics, biotic characteristics of subterranean organisms and troglomorphoses, as well as knowledge on sampling methods for ecological research work in underground habitats.




• Historical review of speleobiology and modern trends
• Survey of subterranean habitats and physical characteristics in different types of subterranean habitats
• Shallow subterranean habitats: ecology, evolution, conservation
• Survey of general characteristics of subterranean organisms
• Adaptations to subterranean life
• General ecological conditions in different types of subterranean habitats
• Research methods for biological investigation and collecting of organisms in subterranean habitats
• Conservation and protection of subterranean habitats and organisms

Intended learning outcomes

Knowledge and understanding:
Students get skills on ecological characteristics of subterranean habitats, biotic characteristics of subterranean organisms, history of speleobiology and trends in modern scientific researches, they get practical knowledge for sampling in subterranean habitats, for ecological research work in subterranean habitats and identification of troglomorphic organisms.


• Chapman P. 1993: Caves and cave life. Harper Collins, London.
• Culver D. C., Kane T. C., Fong D. W. 1995: Adaptation and Natural Selection in Caves. The Evolution of Gammarus minus. Harvard Univ. Press, Cambridge.
• Culver D. C., Pipan T. 2009: The Biology of Caves and Other Subterranean Habitats. Oxford University Press, Oxford.
• Culver D. C., Pipan T. 2014: Shallow Subterranean Habitats. Ecology, Evolution, and Conservation. Oxford University Press, Oxford.
• Gunn J. 2004: Encyclopedia of Caves and Karst Science. Fitzroy-Dearborn, New York.
• Pipan T. 2005: Epikarst – a promising habitat. Copepod fauna, its diversity and ecology: a case study from Slovenia (Europe). Založba ZRC, Ljubljana.
• White W. B., Culver D. C. 2012: Encyclopedia of Caves. Elsevier, Amsterdam.
• Wilkens H., Culver D. C., Humphreys W. 2000: Ecosystems of the World. Subterranean Ecosystems. Elsevier, Amsterdam.
• Selected articles from scientific journals.


Short written seminar, its presentation and exam. 50/50

Lecturer's references

Adjunct Professor of Karstology



CULVER, David C., DEHARVENG, Louis, PIPAN, Tanja, BEDOS, Anne. An overview of subterranean biodiversity hotspots. Diversity, ISSN 1424-2818, 2021, vol. 13, iss. 10, 8 str. https://www.mdpi.com/1424-2818/13/10/487, doi: 10.3390/d13100487. [COBISS.SI-ID 81063171]

PIPAN, Tanja, CHRISTMAN, Mary, CULVER, David C. Abiotic community constraints in extreme environments : epikarst copepods as a model system. Diversity, ISSN 1424-2818, 2020, vol. 12, iss. 7, 16 str. https://www.mdpi.com/1424-2818/12/7/269, doi: 10.3390/d12070269. [COBISS.SI-ID 22691331]

CULVER, David C., PIPAN, Tanja. The biology of caves and other subterranean habitats, (Biology of habitats). 2nd ed. New York: Oxford University Press, 2019. XX, 301 str., ilustr. ISBN 978-0-19-882076-5. ISBN 978-0-19-882077-2. [COBISS.SI-ID 44574765]

PIPAN, Tanja, CULVER, David C. Shallow subterranean habitats. V: WHITE, William Blaine (ur.), CULVER, David C. (ur.), PIPAN, Tanja (ur.). Encyclopedia of caves. 3rd ed. London [etc.]: Academic Press, an imprint of Elsevier. cop. 2019, str. 896-908, ilustr. [COBISS.SI-ID 44674093]

PIPAN, Tanja, CULVER, David C. The unity and diversity of the subterranean realm with respect to invertebrate body size. Journal of caves and karst studies : a publication of the National Speleological Society, ISSN 1090-6924, 2017, vol. 79, no. 1, str. 1-9, ilustr. https://caves.org/pub/journal/PDF/V79/cave-79-01-full.pdf. [COBISS.SI-ID 41823533]