....."PROTEUS PROJECT" WEBPAGE [1.] Contents :
.....INDEX..to All Sections.
.....Section 1....Short Scientific Introduction to Proteus anguinus anguinus (Laurenti 1768.)
.
.....Click on the underlined Links.below to navigate to the other "Proteus Project" Webpages more quickly.
.....Link to. "Proteus Project" Introductory Webpage :
.....Link to. "Proteus Project" Webpage [2.].
.....Link to. "Proteus Project" Webpage [3.].
.....Link to Webpage on.."Proteus Project" Monograph Publication Series.
 

Status :
1.  Listed in App. II of the Convention of European Wildlife and Natural Habitats of 1979.
2. EU Directive 92/43/EEC, Annexes II, IV and V.
The species is a "priority species" of Annex II - 
"Species of Community Interest whose
Conservation Requires the Designation of
Special Areas of Conservation."
3.  IUCN (Red List) Status = Vulnerable (VU).
4. CITES =  No CITES Listing.
5. Present National Status :- 
R. Italia = (no information);
R. Slovenia = "protected" since 1949;
R. Croatia = (no information);
R. Bosnia & Hercegovina = (not protected).

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Members, Project Research Assistants, 
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should note that SAR will automatically 
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level of Systems Access, which 
is reviewed on a regular basis.

Introductory Webpage..Short General Introduction to the "Proteus Project" and to Speleo-biology. (Public Access)

Section 1...Short Scientific Introduction to Proteus anguinus (Laur. 1768)...[This Page]..(Public Access)

Section 2...Bibliography and Further Reading..(Public Access)

Section 3...Background Administrative Data to the Project. (Public Access)
Section 4...Project Participants and their Roles :
.............4.1...Who it is necessary to involve?  (Public Access)
.............4.2...Voluntary Sector. (Not Available On-line)
.............4.3...Industry Sector. (Not Available On-line)
.............4.4...Academic Sector. (Not Available On-line)
.............4.5...Local and Regional Government Sector. (Not Available On-line)
.............4.6...Education Sector. (Not available On-line)
.............4.7...The Press and Media Sector. (Not Available On-line)
.............4.8...Future Participants. (Not Available On-line)
.............4.9...Conclusion. (Summary is Available On-line)
Section 5...Synopsis of the Problems with Proteus anguinus and its Natural Karst Habitat in Eastern Herzegovina : (Public Access)
.............5.1...Proteus Habitat Threat No. 1. -  ERS / HET Co. Hydro-electric Scheme.
.............5.2...Proteus Habitat Threat No. 2. -  Introduction of Foreign Fauna.
.............5.3...Proteus Habitat Threat No. 3. -  Industrial Pollution.
.............5.4...Proteus Habitat Threat No. 4. -  Ubiquitous Dumping of General Domestic and Farm Waste.
.............5.5...Proteus Habitat Threat No. 5. -  Illicit Trading in Proteus Specimens and Uncontrolled Publicity of Proteus anguinus Habitat Locations.
.............5.6...Proteus Habitat Threat No. 6. -  The Apathy of Society in General and of the Competent Authorities in Particular :-
....................(Illustrated by an Interesting, Typical Case History - Vrelo "Oko", Zasad, Trebinje.)
Section 6...Aims and Objectives of the Project :
.............6.1...Proposed Aims and Objectives. (Public Access)
.............6.2...Progress to Date. (Summary is Available On-line)
Section 7...Geographical Scope of the Project Area as Defined by Proteus anguinus Habitat Locations and Associated Catchments : (Not available On-line)
Section 8...Financial Costs and Funding Sources : (Not Available On-line)

Information about the Slovene Black Proteus
is not yet applicable to Bosnia & Herzegovina.
 
 
 
 

IUCN RED LIST STATUS :
"Vulnerable"
 
 
 
 
 
 
 
 

Proteus anguinus
features in the Zoological Society of London,
EDGE AMPHIBIAN'S Programme
as one of the 10 most endangered amphibian species on the planet.
 
 
 
 
 
 
 
 
 
 
 
 

Link to another webpage on this Website

Protecting the
ENDANGERED HYPOGEAN BIODIVERSITY
of Eastern Herzegovina,
Bosnia & Herzegovina.

As is indicated by the token number of entries in the Bibliography Section 2., the amount of research that has been conducted on the animal is both extensive and on-going. 
-..Where it has been reasonable to do so, the following descriptive text has been augmented with Abstracts of the results obtained from scientific research by many specialists, almost always under artificial conditions.
To support readers in locating the full Articles to which the Abstracts refer, all Papers and the technical journals in which they are published, are cited within the text and further referenced in the extensive Bibliography Section 2. in the next "Proteus Project" Webpage.
-..Additionally, where it has been possible for us to do so, without compromising any Proteus anguinus habitat locations, we have now included some of our own observations about Proteus anguinus in its natural habitat.
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Scientific name :..Proteus anguinus anguinus (Laurenti 1768).
.
Systematic Classification :
.....Kingdom / Carstvo / Царство :  Animalia.
.........sub-Kingdom :  Eumetazoa.
.....Phylum :  Chordata. (Possesses a notochord).
.........sub-Phylum :  Vertebrata. (Possesses a back-bone or spinal column).
.........Infra-Phylum :  Gnathostomata.
.....Class / Razred / Разред :  Amphibia. (Linnaeus,.1758). (A Class of Vertebrates typically gill-breathing in the larval state and lung / skin breathing as adults).
.........sub-class / Podrazred / Подразред :  Lissamphibia.
.....Order / Red / Ред :  Caudata (Scopoli,.1777) / Urodela. (Amphibia with permanent tails).
.....Super-Family :  Salamandroidea.
.........Family / Porodica / Породица :  Proteidae.
.....Genus / Rod / Род :  Proteus.
.....Species / Vrsta / Врста :  Proteus anguinus.
.........sub-Species :  Proteus anguinus anguinus (Laurenti,.1768).
.....English Common / Trivial Names :  "The White Proteus"; "Human Fish"; "Cave Olm"; "Cave Salamander"; 
..................................also :-..Srpski = "Човјечија рибица";..Hrvatski / Bosanski = "Čovječja ribica";...Slovene = "Človeška ribica" and "Močeril";..Magyar = "Barlangi Vak Gőte";..German = "Grottenolm".
.....
.....other related sub-Species :  Proteus anguinus parkelj, (Sket et Arntzen,.1994)..[=The Black Proteus], currently found only in Slovenia.
..............................................[There may be other, as yet unknown sub-species.]
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Proteidae is an ancient family of salamanders comprising just six species, commonly referred to as the olm, mudpuppies and waterdogs. The lineage diverged from their closest relatives 190 million years ago in the early Jurassic period in the era of the dinosaurs.
This pre-dates the first fossil bird by 40 million years! The family Proteidae is part of a sub-order of salamanders that contains all of the internally-fertilising species (known as the “Salamandroidea”). Most salamanders reproduce via external ferlilsation,
but in the proteids, females take the spermatophore (or sperm packet), deposited by the male, into their bodies, allowing fertilization of her eggs to occur within the body cavity. 
.
Discovery of Proteus anguinus anguinus and its accepted Type Locality :
Proteus anguinus was first recorded in the literature 320 years ago by the Slovene, Baron Janez Vajkard Valvasor.(also Johann Weikhard Freiherr von Valvasor), in 1689. 
It was subsequently mentioned again by Steinberg in 1761 and also by Jeršinović von Loewengreif.in 1797.
The Type Locality was originally set as Cerkniško Jezero (=Cerknica Lake) in Slovenia, because it was mentioned in the original taxonomic description by Laurenti in 1768. 
Due to subsequent uncertainty and confusion about the validity of the location, the Type Locality was re-set by the accomplished Hungarian zoologist, Baron G.J. de Fejérváry (1926) and accepted as being Magdalene Grotto in Črna Jama,
part of the Postojna-Planina Cave System, Slovenia.
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Short Note about Josephus Nicolaus Laurenti :
Josephus Nicolaus Laurenti (born December 4, 1735 - died February 17, 1805) was an Austrian naturalist.
Laurenti was the author of "Specimen medicum, Exhibens synopsis reptilium emendatum, cum experimentis circa venena et antidota reptilium austriacorum" (1768),.[="Medical Treatise, Exhibiting an Emended Synopsis of Reptiles, with Experiments Concerning Venoms and Antidotes for Austrian Reptiles"] about the poisonous function of reptiles and amphibians and their antidotes. This was an important book in herpetology, defining 30 genera of reptiles...Carolus Linnaeus's "Systema Naturae" (1758) defined only 10 genera.
More relevant to us, in 1768, Laurenti also published an important manuscript titled "Il Dragone" (="The Dragon"), describing the blind salamander (amphibian): Proteus anguinus, purportedly collected from cave waters in Slovenia (or possibly western Croatia);
this description represented the first published account of a cave animal in the western world.
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Notes on abandoned 19th Century Taxonomic Descriptions of Proteus  :
Fitzinger (1850) described a series of "new species" under the Genus "Hypochthon". Although Proteus exhibits a natural variation in appearance and genetic composition across its geographical range, Fitzinger's taxa were never accepted.
He did provide a list of all Proteus habitats known at that time and even falsified other entries !  We include details of his taxa here, merely for historical interest.
For various "species" he described in Slovenia, he ascribed the following taxa:..Hypochthon zoisii (Fitzinger); Hypochthon schreibersii (Fitzinger); Hypochthon freyeri (Fitzinger); Hypochthon haidingeri (Fitzinger); Hypochthon laurentii (Fitzinger);
Hypochthon xanthostictus (Fitzinger) and for those he described in Croatia, he added Hypochthon carrarae (Fitzinger). In 1880, Proteus croaticus (Brusina, 1880).
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Origin of the Species :
Unexplained; archaic species.  Several theories for its origin have been suggested by various "specialists". 
In determining its true origins, a more thorough knowledge is required of its actual geographical range and more work has to be undertaken in regard to DNA profiling.
The lineage supposedly diverged from its closest relatives 190 million years ago in the early Jurassic period in the era of the dinosaurs.
One theory is that it is a suspected Tertiary fauna relict, supposedly originating as a marine animal progressing firstly into brackish then adapting to fresh water conditions inland as a result of the Messinian Salinity Crisis in the Meditteranean Sea
over the period 7.2 to 5.33 million years ago. However, because the species retains the ability to darken its skin colour in light conditions and because it has atrophied eyes rather than no eyes, its previously assumed ancient lineage is now a matter for fresh debate.
Current research in progress at the Biotechnical Faculty of Ljubljana University is seeking to develop a gene database of Proteus anguinus to highlight any differences in the genetic make-up of specimens across the range of its geographical distribution.
Speleo-biological specialists at the Faculty are also engaged upon the task of trying to develop a plausible schema for the evolution of the species and an explanation for its geographical distribution. More information about this can be obtained by referring to the relevant entries in the Bibliography in.Section 2. of the "Proteus Project" Webpages.
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Geographical Range of its Natural Habitats (1), as recorded in the published literature :
Subterranean karst water environment from Italy, (east of the River Isonzo in the areas around Gradisca, Fogliano, Redipuglia and Monfalcone) and Slovenia, through Croatia to Bosnia & Herzegovina.
This is partly illustrated by an Abstract of Biogeography by Sket, B..(May 1997),*.in his Paper "Distribution of Proteus (Amphibia: Urodela: Proteidae) and its possible explanation." :-
.
"Approximately 250 localities of the nominal species Proteus anguinus (Laurenti 1768) have been evaluated and listed. The species is limited to the Dinaric Karst; it ranges from the Tsonzo-Soca River in southeastern Venezia Giulia, Italy, through the southern half of Slovenia, southern Croatia, and parts of Bosnia and Herzegovina, to the Trebišnjica River in Eastern Herzegovina. In some regions, Proteus anguinus populations have been extinguished or endangered by pollution or human-induced hydrographical changes.
The distribution of Proteus is comparable with that of some cave Crustacea: Troglocaris (Crustacea: Decapoda), Monolistra, and Titanethes (Crustacea: Isopoda). The similarity of distribution patterns within this ecologically diverse assemblage supports their paleogeographic rather than ecological foundation. The paleogeographical and paleoclimatological data, in combination with the physiological requirements of Proteus, strongly suggest that these animals invaded the caves, at least in the NW parts of their ranges,
only after the last glaciations, within the last 10,000 years. It is suggested that the high heterozygosity of populations can best be explained by fusion of some locally restricted immigration waves."

[Ed. :  This list is grossly inaccurate in its listings for Bosnia & Herzegovina and serves no useful purpose, either actual or comparative.]
.
Geographical Range of its Natural Habitats (2), as evidenced by direct observation by Project Workers and Research Assistants within the "Proteus Project" :
The generally accepted and published detailed geographical range of Proteus anguinus in Herzegovina is substantially not in agreement with our direct observations. Due to current in-Project restrictions with the dissemination of this type of data,
we are presently unable to clarify this statement.
.
Additional unnatural Occurrences :
Due to ill-conceived human intervention, the animal also occurs unnaturally in the Harz in Germany, in the CRNS underground cave laboratory at Moulis in France and in the Grotte Oliero in Italy.
Other ridiculous and ill-advised attempts have also been made to establish it in artificial environments in Hungary (Baradla barlang, Aggtelek) and in Devon, UK (in the 1970's) without success.
.
GENERAL PHYSICAL CHARACTERISTICS & BEHAVIOURS - supported by our own observations of Proteus anguinus anguinus in its natural habitat :

....a)...GENERAL APPEARANCE :
This aquatic animal is well adapted to its underground environment. Proteus anguinus anguinus is a true troglobiont often referred to as a stygobiont. 
Its body consists of a Head, Trunk, Tail, two slender but fully functional Forelimbs and two Rear Limbs. The front legs bear 3 toes and the rear legs have 2 toes each.
Its relatively large triangular head ends bluntly in a flattened snout and in its mouth contains tiny teeth. The male of the species is usually smaller than the female of equivalent age.
....b)...ENVIRONMENTAL ADAPTATION :
..........i)..Repeated attachment to particular Residence Sites within a general Habitat Location :
During the progress of the Project, we have observed that Proteus anguinus.is reliably found, most often in groups, in specific locations year after year. It seems to favour certain "residence sites" within a given habitat location, not only year after year but also during the year. Where it is found in groups, this is often in cave lakes with a water surface, ie. vadose conditions, or in shallow phreatic and/or epiphreatic cave passages. Where the species is found in regularly used "residence sites" deep under water in phreatic conditions, they are usually found as single representatives.
We can perhaps enlarge on this by considering a recent contribution in the field of Experimental Zoology by Guillaume, O. (2002), in his Paper."The Importance of Chemical Communication in the Social Behaviour of Cave Salamanders. Comparison Between a
Strict (Proteus anguinus L., Proteidae) and a Facultative (Euproctus asper D., Salamandridae) Cave Dweller." :-
.
"Chemical communication is known to be a great importance in animals. It is generally expected that chemical cues will dominate in cave forms, which cannot use vision to regulate their behaviour. To obtain evidence that might support or refute this hypothesis, we have studied the importance of chemical communication in the social behaviour of a stygo-bitic salamander, Proteus anguinus, and in a facultative cave dweller Euproctus asper.
The first part of this work deals with social interactions between conspecifics. Ethological two-choice tests and observations during rearing have shown that adults of P. anguinus exhibit an attachment to a residence site, and mark it using a substrate-adherent substance and faecal pellets. This marking is used as directional information, allowing individuals to relocate their residence site. Furthermore, this marking attracts conspecifics that then display a gregarious behaviour. Similar tests with E. asper, together with a field study on four populations representative of the different types found in the Pyrenees (epigean populations from low, middle and high attitudes and a cave population), show that E. asper do not exhibit site attachment and do not mark the site they use. In fact, individuals travel continuously through their environment and shelter on the way. They thus tolerate the presence of conspecifics. The results of this study therefore indicate that P. anguinus establishes stable open groups, while E. asper forms facultative and unstable aggregations."
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It is interesting to note that Proteus exhibits a marked attachment to particular residence sites within a habitat location. Our observations in Eastern Herzegovina have begun to show that the type of such residence sites can often be predicted, in that they have at least two major common characteristics, unspecified here in this text for reasons of habitat-location protection.
..........ii)...Adaptation to Water Depth and Pressure :
We have observed and have recorded still and moving images of Proteus anguinusat depths of water varying from 0 - 31 metres, (0 - 98 feet).
..........iii)..Adaptation to survive high water flow conditions :
Again, it is interesting to note that Proteus seems to be able to maintain its specifically-favoured residence sites year after year, often after the most tremendous flows of water through the karst conduits that it occupies. Therefore, it must be able to select refuge in small crevices where it can safely shelter out of the main flow of water in the karst conduits during the winter months.
.
....c)...LENGTH and WEIGHT  :
At 20cm to 35+ cm (8 to 13.75+ inches) long, Proteus anguinus is the largest permanently cave-dwelling animal in Europe. Proteus is the only cave amphibian in Europe. In 2007., we have successfully developed a methodology for directly obtaining biometric data of the animal in its natural habitat with the result that we are now accumulating body-mass and dimensional information.
....d)...SKIN and SKIN COLOURING :
..........i)....In common with many other amphibians, including salamanders, the skin of Proteus.contains essential mucus and waxy layers in the epidermis that protect it from dehydration, viruses, bacteria and fungi.
..........ii)...According to Vandel (1966b)., the young larvae after hatching from the egg have a greyish appearance, which, if observed under a microscope, is actually a sprinkling of black chromatophores (pigmentation cells) covering the entire body with the exception of the underside. He reports that in laboratory conditions, this pigmentation develops in total darkness and persists for several months before progressively diminishing. After one year, the appearance becomes greyish-white and it is only near the age of 18 months that the young Proteus becomes completely white. Vandel also reports that if young individuals are reared in laboratory conditions in daylight, the pigmentation persists and the Proteus keeps the black colour it had at the very beginning of the larval development and becomes even darker. He reports that one particular larva, which had always been kept in the light in the Moulis Underground Laboratory in France, had become completely black after 4 years.
Proteus has lost its dark skin-colouring when usually observed under normal conditions underground, although the young do retain more of the skin pigmentation. All Protei darken when exposed to light, the whole body being dermatopic. The skin without dark pigmentation is slightly pink due to blood circulation visible through the skin and in some places is yellowish as a result of riboflavin. Some of the internal physiological arrangement of the Proteus can often be observed through its skin. The overall colouration represents the normal skin colour of white European Humans, which is why the species is often referred to in the South Slavic Languages as Човјечија рибица (Srpski);..Čovječja ribica..(Hrvatski and Bosanski); Človeška ribica (Slovene), which, in the English Language, all mean "Human Fish". It should be noted that pterin, a pigmentation typical for amphibians, is not present in the Proteus physiology.
....e)...METABOLIC RATE :
The metabolic rate of Proteus is remarkable in being unusually low, a factor that also results in a very low growth rate. It saves energy in every way possible by streamlining its movements and by adopting highly efficient foraging and reproductive strategies. These are obvious specializations to cope with a low food supply. It has been described in the technical journals as being a "good example of a low-energy-system vertebrate", which, from our perspective of studying the animal in its natural environment is both glaringly obvious and a gross understatement!
We can perhaps enlarge on this by considering a recent contribution in the field of Experimental Zoology by Hervant, F.; Mathieu, J. and.Durand, J. (Jan 2001) in their Paper "Behavioural, physiological and metabolic responses to long-term starvation and re-feeding in a blind cave-dwelling (Proteus anguinus) and a surface-dwelling (Euproctus asper) salamander.":-
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"The effects of long-term starvation and subsequent refeeding on haematological variables, behaviour, rates of oxygen consumption and intermediary and energy metabolism were studied in morphologically similar surface- and cave-dwelling salamanders. To provide a hypothetical general model representing the responses of amphibians to food stress, a sequential energy strategy has been proposed, suggesting that four successive phases (termed stress, transition, adaptation and recovery) can be distinguished. The metabolic response to prolonged food deprivation was monophasic in the epigean Euproctus asper (Salamandridae), showing an immediate, linear and large decrease in all the energy reserves. In contrast, the hypogean Proteus anguinus (Proteidae) displayed successive periods of glucidic, lipidic and finally lipido-proteic-dominant catabolism during the course of food deprivation. The remarkable resistance to long-term fasting and the very quick recovery from nutritional stress of this cave organism may be explained partly by its ability to remain in an extremely prolonged state of protein sparing and temporary torpor. Proteus anguinus had reduced metabolic and activity rates (considerably lower than those of most surface-dwelling amphibians). These results are interpreted as adaptations to a subterranean existence in which poor and discontinuous food supplies and/or intermittent hypoxia may occur for long periods. Therefore, P. anguinus appears to be a good example of a low-energy-system vertebrate."
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Further information is given by the same Authors Hervant, F.; Mathieu, J. and.Durand, J. (Aug 2000)in their Paper "Metabolism and circadian rhythms of the European blind cave salamander Proteus anguinus and a facultative cave dweller, the Pyrenean newt (Euproctus asper)." :-
.
"Comparisons of circadian rhythmicity, behavior, and metabolism between surface- and cave-dwelling salamanders allow evolutionary trends in these processes to be elucidated. The proteid Proteus anguinus, an obligate cave-dweller, showed no apparent daily rhythm of activity or resting metabolic rate. In contrast, the salamandrid Euproctus asper, a surface-dweller/facultative cave-dweller, had a circadian resting metabolic rate and activity cycle. These circadian rhythms had an endogenous component. The lives of both studied salamanders were characterized by long periods of inactivity punctuated by bouts of foraging or exploratory/predatory behavior. Proteus anguinus had reduced resting metabolic and spontaneous activity rates (considerably lower than those of most surface-dwelling amphibians), and therefore appears to be a good example of a vertebrate as a low-energy system. The low metabolic and activity rates of P. anguinus are interpreted as adaptations to a subterranean environment, where a poor and discontinuous food supply and (or) intermittent hypoxia may be present for long period."
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....f)...VISUAL SYSTEM AND SENSORY ORGANS  :
The eyes of Proteus appear as two tiny dots in the young and are atrophied and covered with skin in the mature animal. In the adult, it is sunk into the skin. Effectively, Proteus anguinus is visually blind. On the other hand, the eyes are very conspicuous in the larvae and very young individuals, in whom it persists for a comparatively long time.  Between hatching and the age of 2 months the eyes reach their maximum size. The development of the eyes then stops before they have acquired a normal functional structure and therefore, never develop beyond the embryonic state. The cornea fails to appear and a thin and fibrous sclerotic membrane covers the eyes. The eyes are still discernable as a microscopic dot in individuals of 2½ to 3 years of age.
The retina is very thick and has a simple structure and a very small number (ca. 2000) of degenerated sensors, which are either very poorly or not differentiated. The lens never has any fibrous differentiation. The optic nerve is very thin, being 5-8µ in diameter, although it is complete and reaches the brain. The cessation of growth of the eyes is followed by a phase of occular regression, which particularly affects the lens. The latter usually disappears completely during the animal's Juvenile Stage of development. In an immature Proteus of about 100 - 220 mm. in length, the diameter of the eye ranges from 0.3mm to 0.4mm.
Vandel (1966b) reports that adult Proteus can tolerate low-level light for extended periods of time, unlike its eggs and embryos, which are very sensitive to illumination and are killed by the light of a 100 watt bulb.
..........Other Photo-receptors :
Other light-sensitive areas of the olm’s body include a “pineal body” in the centre of the head, and even the skin registers light through the proposed presence of the photosensitive pigment melanopsin inside specialised cells called melanophores.
However, since no light penetrates into the hypogean system, this light sensitivity is seldom, if ever, used. 
....g)...AUDIO SENSORY ORGANS :
For hearing, the inner ear has specially-developed crystal masses performing as electro-receptors. Little is known about the hearing of Proteus, but occasionally observed reactions to sounds have indicated the possibility of a hearing capability under water (Prof. Bulog, pers. observation).
....h)...TASTE & OLFACTORY SENSORY ORGANS :
For the senses of taste and smell, there are well developed chemo-receptors on its tongue. The Olm is capable of sensing very low concentrations of organic compounds in the water. They are better at sensing both the quantity and quality of prey by smell than related amphibians (Guillaume 2000). The nasal epithelium, located on the inner surface of the nasal cavity and in the Jacobson's organ, is thicker than in other amphibians (Dumas & Chris 1998). The taste buds are in the mucous epithelium of the mouth, most of them on the upper side of the tongue and on the entrance to the gill cavities. Those in the oral cavity are used for tasting food, where those near the gills probably sense the chemical composition of water.(Istenič & Bulog, 1979).
....i)...MECHANO-RECEPTORS :...The sensory epithelia of the inner ear are very specifically differentiated and enable the Olm to receive sound waves in the water, as well as vibrations from the ground. The complex functional-morphological orientation of the sensory cells enables the animal to register the sound sources (Bulog 1989). 
As this animal stays neotenic throughout its long life span, it is only occasionally exposed to normal adult hearing in air which is probably also possible for Proteus as in most salamanders. Hence, it would be of adaptive value in caves, with no vision available, to profit from underwater hearing by recognizing of particular sounds and eventual localization of prey or other sound sources, i.e. acoustical orientation in general. Experiments indicate that the best hearing sensitivity of Proteus is from 10 Hz and up to 15.000 Hz. The lateral line supplements inner ear sensitivity by registering low-frequency nearby water displacements (Bulog & Schlegel 2000) and (Schlegel et. al. 2006).
....j)...ELECTRO-RECEPTORS :
A new type of sensory organs have been analyzed by light and electron microscopy on the head of Proteus and described as ampullary organs.(Istenič & Bulog 1984). Proteus has the ability to register weak electric fields (Schegel & Bulog, 1997).
Ampullary electroreceptors are responsible for this ability in Urodelans. Proteus senses electrical current fields and their polarity. It reacts to current density of 100 nA/cm² and the lowest threshold of its ampullary organs is 3mV cm­¹ at best frequencies of 30Hz.
Prey capture is obviously performed by a combination of mechano-, chemo- and eventually by electro-perception (Schlegel et. al. 2006).
....k)...CHEMO-RECEPTORS :
Proteus is capable of sensing very low concentrations of organic compounds in the water. They are better at sensing both the quantity and quality of prey by smell than related amphibians (Guillaume 2000). The nasal epithelium, located on the inner surface of the nasal cavity and in the Jacobson's organ, is thicker than in other amphibians (Dumas & Chris 1998). The taste buds are in the mucous epithelium of the mouth, most of them on the upper side of the tongue and on the entrance to the gill cavities. Those in the oral cavity are used for tasting food, where those near the gills probably sense the chemical composition of water (Istenič & Bulog, 1979). 
.
....l)...LOCOMOTION :
For locomotion out of water, two pairs of weak legs, spaced wide apart, with three digits on the front and two on the rear pair, play a minor role. It walks with a snake-like movement. When swimming, it uses its flat tail which is surrounded by a fin of thin skin, for propulsion, whilst simultaneously holding its limbs in tight against its body. It is a highly efficient swimmer and can move extremely fast.

....m)...RESPIRATION :
For respiration, it retains its external larval gills in the adult stage and is thus able to live permanently underwater. It breathes through these gills, which branch out in three bunches on either side of the head. These are coloured light-pink to bright-red by the blood visible through the gill structures. The external gills can sometimes appear larger in low-oxygen conditions, when a greater surface-area of the gill tissue is required for oxygen take-up.
In what were thought to be secondary respiratory organs, it also has two rudimentary internal lungs and can also breathe through its skin, (ie. three systems in total). However, the idea that these are "secondary" respiratory organs has now been dispelled because groups of Proteus have been observed leaving the cave aquatic environment and feeding at the cave entrance area!!
..........Adaptation to survive Anoxic (=hypoxic) conditions :
The ability of amphibians to survive extremes of oxygen availability derives from a core triad of adaptations: profound metabolic suppression, tolerance of ionic and pH disturbances, and mechanisms for avoiding free-radical injury during reoxygenation.
From our own observations and results of physico-chemical determinations in many hypogean habitat locations, the swing from oxic to anoxic and then back to oxic conditions, can be relatively sudden. We have observed various survival strategies at work.
For long-term anoxic survival, enhanced storage of glycogen in critical tissues is also necessary.
....n)...LIFESPAN :
The lifespan of Proteus can be in excess of 100 years, although the norm is often less. Post-mortem autopsies undertaken on a deceased Proteus body can approximate its age of its life-span by determining the number of growth rings on the vertebrae.
....o)...NUTRITION :
For nutrition, it eats a variety of food. It can consume large amounts of food at once, and store nutrients as large deposits of lipids and glycogen in the liver...Niphargus and other cave crustacea such as Troglocaris can also occasionally feature in the natural diet of Proteus, as must phyto-plankton carried in from the surface. We have photographic evidence that it eats cave snails.
It can apparently undergo years without eating. In 1926, an experiment by Gadeau de Kerville confirmed that the animal could not only live in captivity but could survive without being fed. He kept a captive specimen alive for 14.5 years, during the last 8 of which he did not feed it..However, some more recent speleo-biologists have noticed that captive Olms regularly "slough" and then eat the shed bacterial mucus layer, which, like an extra skin, covers and protects their whole body. This mucus is sticky and microscopic examination has shown that in captive amphibians, it becomes encrusted with bacteria, algae and protozoa.
Vandel (1966b) reports that in order to support reproduction, correct feeding was one of the essential conditions for rearing this species in the Moulis Underground Laboratory in France. Here, the adults were fed with crustacea, notably Gammarus, together with aquatic insect larvae. They also ate earthworms, small fish and tadpoles! He notes that the Proteus larvae were fed with micro-organisms contained in the mud of wells or drinking troughs.
....p)...TISSUE REGENERATION :
In the matter of tissue regeneration, the animal has remarkable limb-regeneration capabilities. A substantial amount of experimental biological investigations have been undertaken on live Proteus specimens over a great many years. We do not support such activities being carried out on an already endangered species.
....q)...REPRODUCTION :
..........-.System :-..For reproduction, the female has three systems of gestation according to the most appropriate for the prevailing circumstances. It is most commonly oviparous (egg-laying). It can lay eggs, larvae or live young. Males and females differ only slightly in their appearance. In captivity, a female Proteus has laid 70 eggs of about 12mm diameter.
..........-.Mating and Fertilization :-..To enlarge on the reproductive behaviour, we can provide the following Abstract from Guillaume, O. (2002), in his Article "The Importance of Chemical Communication in the Social Behaviour of Cave Salamanders. Comparison Between a Strict (Proteus anguinus L., Proteidae) and a Facultative (Euproctus asper D., Salamandridae) Cave Dweller."
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"Chemical communication is known to be a great importance in animals. It is generally expected that chemical cues will dominate in cave forms, which cannot use vision to regulate their behaviour. To obtain evidence that might support or refute this hypothesis, we have studied the importance of chemical communication in the social behaviour of a stygo-bitic salamander, Proteus anguinus, and in a facultative cave dweller Euproctus asper. ......
...........In P. anguinus, contrary to the non-sexually active conspecifics, sexually active males shelter apart. The coupling between site-residence marking and aggressive behaviour towards intruders (except sexually active females), allows sexually-active males to establish territories for reproduction. Mate identification requires a close contact, during which pheromones may be released from cloacal glands. Mates may then exhibit courtship, but differences between the descriptions of various authors do not allow the establishment of an unambiguous pattern. After courtship, the male releases on a stone a spermatophore composed of a peduncle topped by a spermatic cap. The cap is composed of a coat that ensheathes the spematozoa packed in a matrix full of mitochondria. The matrix may serve as an energetic substrate for the sperm survey while the female introduces ther spermatophore into her cloacae. The female then demarcates a territory for laying, and stays near the eggs until they hatch. The eggs release a chemical signal that repels young and adult conspecifics.
In E. asper, the male initiates amplexus. He seems not to identify his partner beforehand and often tries to mate with other males, juveniles and even individuals of other species. However, mate identification may occur during amplexus via behavioural interactions and/or pheromones. Then, the male releases several spermatophores consisting of bundles of a great number of spermatozoa, embedded in a viscous substance. Spermatophores are apparently not introduced into the female cloacal ducts, but handled by the male near the cloacal orifice. This manipulation may facilitate the release of the spermatozoa from the gangue that pack them. The spermatozoa may subsequently swim to the female cloacal orifice, using their undulating membrane.
The second part of this study deals with research on the nature and localization of the production sources of chemical signals. We have examined the cytological structure and the development of the cloacal glands, which are good candidates as the production sources of communication signals between conspecifics.
P. anguinus possesses additional tegumental cloacal glands, which are not homologous with other glands described in salamanders. These glands occur in both sexes, but show sexual dimorphism, being better developed in males.
Female E. asper possesses tegumental glands that, on the basis of the present knowledge, must also be considered as forming an additional type. However, there is no evidence that these glands are homologous with those of P. anguinus.
The data clearly show a preponderant influence of chemical communication on social behaviour in P. anguinus, while this cue is less developed in E. asper. Despite this difference, gregariousness is a common character of both species. This characteristic is also found in some cave fishes, which supports the hypothesis that gregariousness may be propitious to cave life. According to some authors, attraction towards conspecifics increases the chances of finding a mate and helps in the search of food."
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By observing the behaviour of captive specimens, Parzefall (1976).and.Briegleb (1962) have attempted to demonstrate that males only show aggressive behaviour and territoriality for a very short reproductive period. Normally the animals rest under stones in groups of both sexes without any aggressive reaction. Also, during breeding activity, males have a larger and more elongated swollen cloaca (urinogenital opening) than the females. When a male becomes sexually active it starts to control its conspecifics by contacts with the snout and allows only females in the reproductive state to remain in the hiding place. All the females had well-developed eggs visible in their oviducts. Intruders were attacked by tail-beating, ramming and biting. It was also observed that males patrol along territorial borders with oscillating tail movements. Such territories may exist for only a few days and animals being attacked in such a territory avoid it thereafter on the basis of a substrate-specific chemical signal. This avoidance reaction also persists for only a few days.
Others report that adult Protei congregate in suitable areas such as cracks and under rocks. The males establish a territory when breeding, which is furiously defended against competing males. When a female enters such a territory, the courtship commences. The male fans with his tail in the direction of the female's head (possibly even secreting a female-attracting pheromone) and touches the female's cloaca (the combined reproductive and urinogenital opening) with his snout. The female then touches the male's cloaca with her snout and then follows the male who walks 50-100 mm forward and deposits a spermatophore (a small packet of sperm). The pair then moves forward again until the female can take up the spermatophore with her cloaca. Courtship may be repeated several times within a few hours. After leaving the male's territory, the female establishes her own egg-laying territory.

Image PA5. (left)
Proteus anguinus anguinus (Laurenti 1768) in its egg development stage in a vivarium.

..........-..Embryonic Development :-..Vandel (1966b) reports that just as the egg-laying process in artificial conditions at a temperature of 11.9º C was gradual, the hatching was also spread over several weeks.
He determined an estimated embryonic period of 111 days.

 
 

Image PA6. (left)
Proteus anguinus anguinus (Laurenti 1768) in its Larval Stage of development in a vivarium.

The Juvenile Stage is the second, during which it extracts its food from its surroundings. It is from the Juvenile Stage onwards that Proteus begins to lead an active life. The increase in size is very slow during this Stage with the growth curve showing a strong deflection after the age of 2 to 3 years. Vandel again reports that the largest of the specimens reared in the Moulis Underground Laboratory was only 200mm long and therefore, as Proteus does not become sexually mature until it reaches a length of 240mm., it seems likely that these cave salamanders do not reproduce until they are at least 10 years old.
....r)...ANATOMY :
Above, we append an image of an old diagram of the anatomy of the Proteus anguinus, drawn by the Italian explorers M. Rusconi and P. Configliachi.and published in "Monografia del Proteo anguino di Laurenti" in the year 1819.
Amphibians do not have a fixed number of vertebrae. Proteus anguinus anguinus has 29–32 vertebrae.
....s)...DISEASE AND INFECTION :
As well as being susceptible to various infections, the adults also suffer from parasites. 
We are aware that populations of certain European amphibians have succumbed to a fungal virus. We are currently trying to obtain more information about this and will publish the results of our search when completed. In August 2005., we have extracted the stomach contents of 2 animals, together with body tissue from the tip of the tail of one animal for laboratory tests for the presence of fungal viruses. Further information will be given in due course.
....t)...MISCELLANEOUS PHYSIOLOGY :
............i)..Thyroxine - Proteus retains a highly developed and endocrinologically active Thyroid Gland system, producing thyroxine. It has been proven under laboratory conditions that in Proteus and other cave salamanders, this is not to help control the process of metamorphosis, unlike in other types of salamanders. Therefore in Proteus, the thyroxine must have another use.
....u)...VULNERABILITY :
..........Official Information in the Public Domain from various sources :
...................The olm is listed as "Vulnerable" in the IUCN Red List of Threatened Species because its area of occupancy is less than 2,000 km². Its distribution is severely fragmented and there is a continuing decline in the extent and quality of its habitat and in the number of mature individuals. The olm is thought to be in decline by the IUCN Red List of Threatened Species.
There is currently little information available on the abundance of this species. However the IUCN Red List of Threatened species states that the olm is apparently most common in Slovenia and Croatia, although a decline has been observed in the populations of Goriza (Italy) and Postojna (Slovenia). Specifically, the number of individuals of the subspecies Proteus anguinus parkelj is thought to be very low.
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....v)...PROTECTION AND CONSERVATION OF THE SPECIES,
.........Summary of what the Society is doing :
.....The long-term protection and conservation of the species can only be achieved through the protection and conservation of its natural karst habitats.
The species is greatly endangered within its natural habitats from over-collection by so-called "cave scientists" and by the industrial chemical pollution of karst groundwaters, together with anthropogenic changes in natural hydrographic conditions.
The Society's Joint International Programme in Bosnia & Herzegovina is the only known project that is fully underway (and has been for 9 years) and which has been specifically designed to protect Proteus anguinus by understanding its behaviour and the problems in its natural habitat and then addressing those problems. The 30-year Programme, (about which this is the second of four dedicated Webpages), is in 3 Phases. With only minimal voluntary-sector funding but enormous quantities of voluntary-sector effort, the current Phase 1. has been overwhelmingly successful and has produced a great wealth of information about the behaviour of Proteus and the characteristics and problems of its natural habitats across a whole region.
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We have designed and implemented a successful strategy for identifying previously unknown Proteus anguinus Habitat Locations across a whole region. As such, we are able to preliminarily identify such locations before we actually enter them for the first time to eventually confirm their status. The same strategy also allows us to predict the location of once-viable habitats that have now become degraded and non-viable. 
Our work includes regular water-quality monitoring of the hypogean habitats and of all those locations where it is no longer found. Population monitoring and a species biometric programme are included. Vulnerability assessments are made about each habitat location and Habitat Viability studies are also undertaken.
Habitat Restoration and Habitat Protection schemes have been successfully designed and implemented at various locations, restoring some to their former "viable habitat" condition, whilst others are the subject of on-going pro-active protection.
The habitat locations of Proteus anguinus in Eastern Herzegovina vary greatly in their hydrological functionality and other prevailing conditions. We have found that Conservation Action Plans have to be designed to be site-specific and that the concept of a "universal conservation model" is inappropriate. Our Project has designed and implemented site-specific Conservation Action Plans with great success.
As a necessary adjunct to all of our practical work with Proteus anguinus, we have identified critical issues that must be observed by all Project Workers undertaking work in all parts of the Project. These have been realised through the implementation of a Code of Ethics, Operational Protocols & Procedures and Methodology, detailing what work can be undertaken with Proteus anguinus and how the work must be conducted. A Prime Directive has been included in this document.
Our Project recognizes the need to work with many sectors of the local population and with its political representatives. As such, we are fully engaged with all of them in support of our objectives. Public Education Programmes at various times throuhgout the year support our connection with the local population and "stakeholders".
We are well placed to succeed in our final objectives of protecting this incredible animal through the protection and conservation of its natural habitats and in knowing how to restore the viability of many of its previous habitats, which are currently degraded.
Further details about the Society's work on this Project can be found in the Proteus Project Webpage 3., which is currently being updated.
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Finally, from studying the species in its natural habitat, we do not think it is possible to use the system of "captive breeding" of the Proteus as a viable contribution towards ensuring the survival of the species. This may be appropriate for some amphibians but we are of the opinion that it is not necessarily so for Proteus. We have learned that the natural variations in the characteristics of some Proteus habitats differ greatly from others. We are not even contemplating the idea yet of restocking habitat locations by transplanting some specimens from one location to another, because of the natural variations that exist between individual habitats!
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Proteus anguinus is indeed a highly endangered species. However, due care must be undertaken by anyone or any organization who thinks that it is a wise and practical idea to just "dive in" and to do "something or anything" to help save this species.
It really is not that easy. More harm than good will arise as a result of such ill-considered "knee jerk" decisions. This is not an animal like any other. This is not an amphibian like any other.
This is a unique and highly adapted hypogean species, endemic to a comparatively tiny area of our planet and which seems to have further adapted to prevailing localised conditions within and across its geographical range.
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....w)...PROTECTION AND CONSERVATION OF THE SPECIES (continued ....),
.........Summary of the status of Proteus in the European Union's Habitats Directive :
.....Proteus anguinus is mentioned in the List of animal species covered in EU Directive 92/43/EEC, Annexes II, IV and V.
.....The species is a "priority species" of Annex II - "Species of Community Interest whose Conservation Requires the Designation of Special Areas of Conservation."
.........CITES :
.....Proteus anguinus is not listed on CITES.
.........IUCN Red List :
.....the Olm is listed as vulnerable because of its fragmented and limited distribution and ever-decreasing population.
.........In Slovenia :
.....Protected in Slovenia since 1949. 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, which continues to exist. In 1982 it was placed on a list of rare and endangered species in Slovenia. This list also had the effect of theoretically 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. Although the Olm is included in a Slovenian red list of endangered species and notwithstanding the aforementioned protection mechanisms, non-Slovene collectors continue to visit the country from Italy, France, the Czech Republic and Slovakia and remove specimens for private collections or for illegal trading for onward use in Asia.
.........Global Biodiversity Information Facility (GBIF) :
.....Entries for the Proteus anguinus species in this database are either mostly useless or misleading.
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....x)...PROTECTION AND CONSERVATION OF THE ENVIRONMENT & BIODIVERSITY,
.........Summary of what BiH has agreed do :
UNITED NATIONS DEVELOPMENT ASSISTANCE - FRAMEWORK FOR BOSNIA AND HERZEGOVINA 2010 – 2014
A UNDAF has been prepared by the United Nations Country Team in Bosnia and Herzegovina in consultation with the Government of Bosnia and Herzegovina and other partners, with the aim of
improving the lives of the people of BiH, and particularly the most vulnerable. This was signed by Mr. Nikola Spirić in Sarajevo, Bosnia and Herzegovina in March 2009 on behalf of the Council of Ministers of Bosnia & Hercegovina.
Four main goals have been identified that will set the direction and scope of action of UN system development assistance in the next 5 years. One of these is:
"By the end of 2014, BiH Government meets requirements of EU accession process and multi-lateral environment agreements (MEA), adopts environment as a cross-cutting issue for participatory
development planning in all sectors and at all levels, strengthens environmental management to protect natural and cultural resources and mitigate environmental threats."
Within the document on p.13., we note the statement : "While environmental protection is emerging as a priority for the Government, there is a lack of sufficient capacities, strong policy and a legal framework at the state-level."