Olm, Blind cave salamander, Grottenolm, hulepadde, koopaolm, Proteo, Protée anguillard, hulesalamander, White Olm, Black Olm, odmieniec jaskiniowy, olmi, covecija ribica, cloveska ribica
© 2015 Dr. Joachim Nerz (1 of 22)
The average total length lies between 23-25 cm. They may grow up to 30 cm and rarely more than 30 cm. Black Proteus can grow up to 40 cm or more. Males are somewhat smaller than females. Other sexually dimorphic characteristics include the shape and size of the cloaca during breeding activity, with the males having a larger and more elongated swollen cloaca than the females.
Distribution and Habitat
Country distribution from AmphibiaWeb's database: Bosnia and Herzegovina, Croatia, Italy, Slovenia. Introduced: France.
This species prefers underground water systems in karst formations, with calm, usually well-oxygenated water and a constant low water temperature between 8ºC (winter) and 11ºC (in one locality rarely up to 14ºC in summer) (Sket 1997; Honnegger 1981). It occurs in caves and may also be found in abandoned mines, from close to the surface to depths as much as 300 m underground, depending on how thick the karst formation is (Stuart et al. 2008). During periods of high rainfall and flooding it may be found in cave entrances (Stuart et al. 2008). The variant known as the black olm can be found in warmer surface waters (Stuart et al. 2008).
Life History, Abundance, Activity, and Special Behaviors
Although adults aggregate in suitable spots as in cracks and under rocks, males establish a territory when breeding, which is furiously protected from competing males. When a female enters such a territory, the courtship begins. The male fans with his tail in the direction of the female's head. The male touches the female's cloaca with his snout. The female then touches the male's cloaca with her snout and then follows the male who walks 5-10 cm forward after which the male deposits a spermatophore. The pair then moves forward again until the female can take up the spermatophore with her cloaca. Courtship can be repeated several times within a few hours. After leaving the male's territory, the female establishes an egg-laying territory. After 2-3 days the female starts to lay eggs and can continue doing so for up to 25 days, laying a total of up to 70 eggs under rocks. Eggs are guarded by the female. The diameter of the eggs directly after laying is 4-5 mm and can increase through water uptake to 8-9 mm. Unconfirmed historical observations of vivipary exist; it was long thought that female Proteus gave birth to only two well-developed young at lower temperatures and laid eggs at higher temperatures, but this has not been confirmed by rigorous observations. The eggs develop in 182 days at 8ºC, 140 days at 10ºC, 123 days at 11ºC, and in 86 days at 15ºC. Development of larvae is highly temperature-dependent. At 10ºC it takes another 14 years to reach sexual maturity. There is no clear metamorphosis; P. anguinus is a neotenic salamander, maintaining external gills, tail fin and other juvenile characteristics throughout its life.
Proteus anguinus is thought to be the longest-lived amphibian species. Using data spanning more than 50 years from a 400-animal captive breeding colony at the CNRS in Moulis, France, the predicted maximum lifespan is over a century, and the average adult olm lifespan is 68.5 years (Voituron et al. 2010). If the predicted maximum lifespan is accurate, it is more than double that of the next longest-lived species, the Japanese giant salamander (Andrias japonicus, at 55 years. Individual specimens have been kept under semi-natural conditions in concrete basins for up to 70 years (Prof. B. Bulog, personal communication). This species reaches sexual maturity at 15.6 years and lays 35 eggs every 12 years, on average (Voituron et al. 2010).
The diet consists of insect larvae, mostly Trichoptera, Ephemeroptera, Plecoptera and Diptera larvae, molluscs (Belgrandiella), and amphipods (Niphargus, Asellus, Synurella) (Bizjak-Mali 1995; Bizjak-Mali and Bulog 2004). In captivity worms are also readily eaten (Boehme et al. 1999).
Sensory adaptations to cave dwelling
Visual pigments in the regressed eye and the pineal of the depigmented and pigmented subspecies were studied by immunocytochemistry (Kos et al. 2001). The presence of visual pigments indicates retained light sensitivity in both subspecies. In the retina of the black Proteus are principal rods, red-sensitive cones and a third photoreceptor type, which might represent a blue- or UV-sensitive cone. The majority of these outer segments of the regressed eye of unpigmented Proteus showed immunolabelling for the red-sensitive cone.
The pineal organ influences skin pigmentation, metamorphosis and gonadal development, and controls circadian rhythms through secretion of pineal hormones. The pineal structure is very similar in all Proteus individuals analyzed. In Proteus the pineal organ is reduced in size; it has degenerated photosensitive cells and can be found only by serial sectioning of the brain. The pineal organ probably possesses some control over the physiological processes also in Proteus, taking into account the presence of visual pigments (Kos et al. 2001). Behavioral experiments revealed that the skin itself is also sensitive to light, and immunocytochemical analysis also supported the existence of photosensitive pigment in Proteus' integument. Photosensitivity of the integument is due to the pigment melanopsin inside pigment cells called melanophores. (Kos et al. 2001).
Trends and Threats
The Olm is extremely vulnerable to changes in its environment due to its adaptation to the specific conditions in caves. Water resources in the karst are extremely sensitive to all kinds of pollution (Bulog et al. 2002) . The contamination of the karst underground waters is due to the large number of waste disposal sites leached by rainwater, as well as to the accidental overflow of various liquids. The reflection of such pollution in the karst underground waters depends on the type and quantity of pollutants, and on the rock structure through which the waters penetrate. Self-purification processes in the underground waters are not completely understood, but they are quite different from those in surface waters. Among the most serious chemical pollutants are chlorinated hydrocarbon pesticides, fertilizers, polychlorinated biphenyls (PCBs), which are (or were) used in a variety of industrial processes and in the manufacture of many kinds of materials; and metals such as mercury, lead, cadmium, and arsenic. All of these substances persist in the environment, being slowly, if at all, degraded by natural processes. In addition, all are toxic to life if they accumulate in any appreciable quantity.
Slovenian caves became famous for the animals they contained and which could not be found elsewhere. Due to its rarity the Olm is also popular among collectors, leading to possible overcollection. Honnegger (1981) also lists overcollecting, for scientific use or as pig-food by farmers, as a threat to this species.
The Olm is included in Appendices II and IV of the EU Habitats Directive (92/43/EEC). Appendix II seeks to preserve favorable conservation status in animal and plant species along with their habitats by protecting the species or defining special areas of conservation. These areas of conservation form the Natura 2000 network. Appendix IV further defines "animal and plant species of community interest in need of strict protection." Hunting or keeping a limited number of Olms is allowed only under strictly controlled circumstances, determined by local authorities.
The Olm was first protected in Slovenia in 1922 along with all cave fauna, but the protection was not effective and a substantial black market came into existence. In 1982 it was placed on a list of rare and endangered species in Slovenia. This list also had the effect of prohibiting trade of the species. After joining the European Union, Slovenia had to establish mechanisms for protection of the species included in the EU Habitats Directive. The Olm is included in the Slovenian Red List of endangered species. The Postojna cave and other caves inhabited by the Olm were also included in the Slovenian part of the Natura 2000 network.
On the IUCN Red List, the Olm is listed as Vulnerable because of its fragmented and limited distribution and ever-decreasing population.
In Slovenia much of the range lies within proposed national or international protected areas (Kocevski Regional Park; Kraski Regional Park; NATURA 2000 sites). In Italy it occurs within the Riserva Naturale Regionale dei Laghi di Doberdò e Pietrarossa (Stuart et al. 2008).
Relation to Humans
Nearly 300 years after the first written mention of the unpigmented Proteus in Janez Vajkard Valvasor's “The Glory of the Duchy of Carniola (1689)”, a pigmented form (described as a subspecies, Proteus anguinus parkelj, the black olm) was discovered at Jelsevniscica in Jelsevnik near Crnomelj in Slovenia. This limited area is the unique locality of the black olm. A presentation on the unique black population has been prepared by the research group for functional-morphological studies of vertebrates, under the guidance of Prof. dr. Bulog. This presentation is located in the house of the family Zupancic in Jelsevnik and is prepared on ten color panels with texts and photo documentation. The presentation is intended for tourists and also for teaching and research purposes, for better recognition of the unique part of Slovenian natural heritage.
A popular scientific film dealing with our studies of biological peculiarities of endemic cave salamander supports this presentation and was recently created in collaboration with National TV of Slovenia (Bulog et al. 2003).
Possible reasons for amphibian decline
General habitat alteration and loss
Proteus is the only cave-adapted vertebrate in Europe. Current genetic research under the direction of Dr. Boris Sket of the University of Ljubljana suggests that Proteus anguinus is actually a complex of several species, with phylogenetic analysis revealing six cryptic lineages (see Trontelj et al. 2007).
Functional-morphological and environmental studies of Proteus have been performed at the Department of Biology, Biotechnical Faculty (BF), University of Ljubljana, Slovenia for more than thirty years, with the most recent twenty years under the guidance of Prof. dr. Boris Bulog.
Arntzen, J. W., and Sket, B. (1997). ''Morphometric analysis of black and white European cave salamanders, Proteus anguinus.'' Journal of Zoology (London), 241(4), 699-707.
Bizjak-Mali, L. (1995). Histological, histochemical and ultrastructural analysis of the digestive tract of Proteus anguinus (Amphibia, Caudata), Master of Science Thesis. University of Ljubljana, Biotechnical Faculty, Department of Biology, Slovenia.
Bizjak-Mali, L. and Bulog, B. (2004). ''Histology and ultrastructure of the gut epithelium of the neotenic cave salamander, Proteus anguinus (Amphibia, Caudata).'' Journal of Mophology, 259, 82-89.
Boehme, W., Grossenbacher, K., and Thiesmeier, B. (1999). Handbuch der Reptilien und Amphibien Europas, band 4/I:Schwanzlurche (Urodela). Aula-Verlag, Wiesbaden.
Bons, J. and Beniez, P. (1996). Amphibiens et Reptiles du Maroc (Sahara occidental compris). Asociacion Herpetologica Española, Barcelona.
Bulog B. (1989). ''Differentiation of the inner ear sensory epithelia of Proteus anguinus (Urodela, Amphibia).'' Journal of Morphology, 202, 325-338.
Bulog B. et al. (2003). Black Proteus: mysterious dweller of the Karst in Bela krajina . Ljubljana: TV Slovenia, Video tape
Bulog B., and Schlegel, P. (2000). ''Functional morphology of the inner ear and underwater audiograms of Proteus anguinus (Amphibia, Urodela).'' Pflügers Archive, 439(3), 165-167.
Bulog, B., Mihajl, K., Jeran, Z., and Toman, M. (2002). ''Trace element concentrations in the tissues of Proteus anguinus (Amphibia, Caudata) and the surrounding environment.'' Water, Air, and Soil Pollution, 136(1-4), 147-163.
Dumas, P. and Chris, B. (1998). ''The olfaction in Proteus anguinus.'' Behavioural Processes, 43, 107-113.
Durand, J.P. (1976). ''Ocular development and involution in the European Cave Salamander, Proteus anguinus Laurenti.'' The Biologial Bulletin, 151(3), 450-466.
Gasc, J.-P. (1997). Atlas of Amphibians and Reptiles in Europe. Societas Europaea Herpetologica, Bonn, Germany.
Gorički, S., and Trontelj, P. (2006). ''Structure and evolution of the mitochondrial control region and flanking sequences in the European cave salamander Proteus anguinus.'' Gene, 378, 31-41.
Griffiths, R.A. (1996). Newts and Salamanders of Europe. T. and A. D. Poyser, London.
Guillaume, O. (2000). ''Role of chemical communication and behavioral interactions among conspecifics in the choice of shelters by the cave-dwelling salamander Proteus anguinus (Caudata, Proteidae).'' Canadian Journal of Zoology, 78(2), 167-173.
Honegger, R. E. (1981). Threatened Amphibians and Reptiles in Europe. Akademische Verlagsgesellschaft, Wiesbaden.
Istenič, L. and Bulog, B. (1979). ''The structural differentiations of the buccal and pharyngeal mucous membrane of the Proteus anguinus Laur.'' Biološki Vestnik, 27, 1-12.
Istenič, L. and Bulog, B. (1984). ''Some evidence for the ampullary organs in the European cave salamander Proteus anguinus (Urodela, Amphibia).'' Cell and Tissue Research, 235, 393-402.
Istenič, L. and Sojar, A. (1974). ''Oxygen consumption of Proteus anguinus.'' Acta Carsologica, 6, 299-305.
Istenic, L. and Ziegler, I. (1974). ''Riboflavin as ''pigment'' in the skin of Proteus anguinus L.'' Naturwissenschaften, 12, 686-687.
Kalezic, M., and Dzukic, G. (2001). ''Amphibian status in Serbia and Montenegro (FR Yugoslavia).'' FROGLOG, 45.
Kos, M. (1992). ''Fine structure of of the skin of Proteus anguinus Laurenti (Urodela, Amphibia) and comparison of the skin of the pigmentless and pigmented specimen. Unpublished dissertation.''
Kos, M., Bulog, B., Szél, A., and Röhlich P. (2001). ''Immunocytochemical demonstration of visualpigments in the degenerate retinal and pineal photoreceptors of the blind cavesalamander (Proteus anguinus).'' Cell and Tissue Research, 303(1), 15-25.
Nöllert, A. and Nöllert, C. (1992). Die Amphibien Europas. Franckh-Kosmos Verlags-GmbH and Company, Stuttgart.
Schegel, P., and Bulog, B. (1997). ''Population-specific behavioral electrosensitivity of the European blind cave salamander, Proteus anguinus.'' Journal of Physiology (Paris), 91, 75-79.
Schlegel P. (1996). ''Behavioral evidence and possible physical and physiological mechanisms for earth magnetic orientation in the European Blind Cave Salamander, Proteus anguinus.'' Mémoires de Biospéologie, 23, 5-16.
Schlegel P.A., Briegleb W., Bulog B., Steinfartz S. (2006). ''Revue et nouvellesdonnées sur la sensitivité a la lumiere et orientation non-visuelle chez Proteus anguinus, Calotriton asper et Desmognathus ochrophaeus (Amphibiens urodeles hypogés).'' Bulletin de la Société herpétologique de France, 118, 1-31.
Sket, B. (1997). ''Distribution of Proteus (Amphibia: Urodela: Proteidae) and its possible explanation.'' Journal of Biogeography, 24, 263-280.
Stet, B., and Arntzen, J. W. (1994). ''A black, non-troglomorphic amphibian from the karst of Slovenia: Proteus anguinus parkelj n. ssp. (Urodela: Proteidae).'' Bijdragen tot de Dierkunde, 64(1), 33-53.
Stuart, S., Hoffmann, M., Chanson, J., Cox, N., Berridge, R., Ramani, P., and Young, B. (eds) (2008). Threatened Amphibians of the World. Lynx Edicions, IUCN, and Conservation International, Barcelona, Spain; Gland, Switzerland; and Arlington, Virginia, USA.
Stumpel-Rieks, S. E. (1992). Nomina Herpetofaunae Europaeae. AULA-Verlag, Wiesbaden.
Trontelj, P., Douady, C., Fišer, C., Gibert, J., Gorički, S., Lefébure, T., Sket, B., and Zakšek, V. (2007). ''A molecular test for cryptic diversity in ground water: how large are the ranges of macro-stygobionts?'' Freshwater Biology, 54, 727-744.
Uiblein, F., Durand, J. P., Juberthie, C., and Parzefall, J. (1992). ''Predation in caves: the effects of prey immobility and darkness on the foraging behaviour of two salamanders, Euproctus asper and Proteus anguis.'' Bahavioural Processes, 28, 33-40.
Voituron, Y., de Fraipont, M., Issartel, J., Guillaume, O., and Clobert, J. (2010). ''Extreme lifespan of the human fish (Proteus anguinus): a challenge for ageing mechanisms .'' Biology Letters, Published online before print July 21, 2010, doi: 10.1098/rsbl.2010.0539 .
Written by Boris Bulog and Arie van der Meijden (boris.bulog AT bf.uni-lj.si), University of Ljubljana
First submitted 1999-12-26
Edited by Kellie Whittaker (2014-08-05)
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