Shallow subterranean habitats

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, ecology, and evolution of shallow subterranean habitats.




• Shallow subterranean habitats (SSHs):
• Seepage springs and the hypotelminorheic habitats
• Epikarst: the soil-rock interface in karst
• Intermediate-sized terrestrial shallow subterranean habitats
• Calcrete aquifers
• Interstitial habitats along rivers and streams
• Soil
• Lava tubes
• The role of light in SSHs
• Environmental fluctuations and stresses in SSHs
• Organic carbon and nutrients in SSHs
• Evolution of morphology in SSHs
• Colonization and dispersal in SSHs
• Phylogeny in SSHs
• Conservation and protection in SSHs

Intended learning outcomes

Students get knowledge about types of shallow subterranean habitats (hypotelminorheic, epikarst, MSS, calcrete aquifers, aquatic interstitial—hyporheic, soil, lava tubes), general features of SSHs like absence of light, surface-subsurface connections, availability of organic carbon and nutrients, geographic distribution patterns of SSHs, habitat size, features of SSHs of general ecological and evolutionary interest and conservation and protection of shallow subterranean habitats.


• Culver D. C., Pipan T. 2014: Shallow Subterranean Habitats. Ecology, Evolution, and Conservation. Oxford University Press, Oxford.
• Culver D. C., Pipan T. 2009: The Biology of Caves and Other Subterranean Habitats. Oxford University Press, Oxford.
• 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., 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., 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. [COBISS.SI-ID 41823533]