THE PROJECTS IN BOSNIA & HERZEGOVINA

....."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" Webpage [2.].
.....Link to. "Proteus Project" Webpage [3.].

..........PROTEUS ANGUINUS IS YOUR RAREST AND GREATEST NATURE RESOURCE

..........ČOVEČIJA RIBICA JE VAŠA RIJETKOST I VELIKO PRIRODNO BOGATSTVO.
..........Ona nema prirodnog neprijatelja i samo mi možemo ugroziti njezin opstanak. 
..........Zato, upoznajmo i čuvajmo prirodu Istočne Hercegovine.

..........ЧОВЕЧИЈА РИБИЦА JE ВАША РИЈЕТКОСТ И ВЕЛИКО ПРИРОДНО БОГАТСТВО.
..........Она нема природног непријатеља и само ми можемо угрозити њезин опстанак. 
..........Зато, упознајмо и чувајмо природу Источне Херцеговине.

..........
.........
.....Image PA1. (above-left),...2 specimens of Proteus anguinus anguinus (Laurenti 1768) in their natural habitat (vadose conditions) in Eastern Herzegovina, RS-Bosnia & Herzegovina..(206 kB.)
.....(Photo :  Hungarian Cave Diving Team, 07 August 2005.)
.....Image PA2. (above-right),...Portrait of a living miracle - Proteus anguinus anguinus (Laurenti 1768) in its natural habitat (vadose conditions) in Eastern Herzegovina, RS-Bosnia & Herzegovina, undergoing examination..(211 kB.)
.....(Photo :  Bogdan Prelević, [SDZB], 07 August 2005.)
.
.......
.....Image PA3. (above-left),..Proteus anguinus anguinus (Laurenti 1768) in its natural habitat (phreatic conditions) in Eastern Herzegovina, RS-Bosnia & Herzegovina..(164 kB.)
.....(Photo :  Hungarian Cave Diving Team, 18 August 2005.)
.....Image PA4. (above-right),..Proteus anguinus anguinus (Laurenti 1768) in its natural habitat (phreatic conditions) in Eastern Herzegovina, RS-Bosnia & Herzegovina.
.....Is this how it maintains its presence in the Habitat during periods of high water-flow in the phreatic conduits?.(200 kB.)
.....(Photo :  Hungarian Cave Diving Team, 18 August 2005.)

Members, Project Workers and 
other Authorized Users should 
note that SAR will automatically 
allocate you with the  appropriate 
level of Systems Access, which 
is reviewed on a regular basis.

If the Registry has failed to do so, 
or if you have problems accessing 
specific pages, please contact 
the DKRS Webmaster.

Accessibility to the published pages of this Project has had to be restricted in the past for many reasons, some of which are explained below. Most of these reasons are unstated but relate specifically to the prevailing geo-political situation in the Project area in Bosnia & Herzegovina. It is hoped that in due course, as the Project proceeds into its second stage, all participants can agree on publishing more information.

One of our main problems is that of disclosing Proteus anguinus Habitat Locations. We are not at liberty to freely disclose this information, either directly or indirectly under the present situation that exists in BiH. As with our partners in this Project, we are acutely aware of the prevailing minimum critical level of both the viable Habitat Locations and of the Proteus itself. We cannot be a partner to encouraging the further demise of this highly endangered species. In this regard, we draw your attention to relevant details, which expand on this problem, in Section 4.5. below.

We do welcome any serious offers of help and involvement that non-BiH individuals or groups may wish to make to the Project. All offers are closely scrutinised in the UK and locally approved in BiH. As of 2004, letters of invitation for approved non-BiH individuals or groups to visit BiH as a Project participant(s) can only be issued by the Speleološko Društvo "Zelena Brda" in Trebinje.  In the first instance, please contact the UK Project Control Administrator  via e-mail or by normal post, using the contact details on the main Homepage of this Website.
Serious offers of help and involvement from BiH Nationals should be sent direct to Speleološko Društvo "Zelena Brda" in Trebinje.

"Proteus Project" Page 1...[Information Section on this Page] :

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. (Restricted Access)
.............4.3...Industry Sector. (Restricted Access)
.............4.4...Academic Sector. (Restricted Access)
.............4.5...Local and Regional Government Sector. (Restricted Access)
.............4.6...Education Sector. (Restricted Access)
.............4.7...The Press and Media Sector. (Restricted Access)
.............4.8...Future Participants. (Restricted Access)
.............4.9...Conclusion. (Restricted Access)
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. (Restricted Access)
Section 7...Geographical Scope of the Project Area as Defined by Proteus anguinus Habitat Locations and Associated Catchments : (Restricted Access)
Section 8...Financial Costs and Funding Sources : (Restricted Access)

Information about the Slovene Black Proteus
is not 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.

To read more about the ZSL EDGE PROGRAMME,
click on the Link below ..

ZSL EDGE AMPHIBIANS PROGRAMME
 

As is indicated by the token number of entries in the Bibliography Section 2. below, the amount of research that has been conducted on the animal is both extensive 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 at the bottom of this page.
-..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.

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).
.....Family / Porodica / Породица :  Proteidae.
.........super-Family :  Salamandroidea.
.....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, (Parkelj, 1768). [The Black Proteus].

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 suborder 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 spermatophone (or sperm packet) deposited by the male into their bodies, allowing fertisation of her eggs to occur within the body cavity. 

Origin of Species :  Unexplained; archaic species.  The lineage diverged from their closest relatives 190 million years ago in the early Jurassic period in the era of the dinosaurs. This predates the first fossil bird by 40 million years. 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 makeup 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 Bibliography in Section 2. of the "Proteus Project" Webpages.

Geographical Range of its Natural Habitats 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 best 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. The close morphological similarity of nearly all populations of Proteus is probably due to the convergent evolution of previously differentiated populations (or even species) after their withdrawal underground."

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.

[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 in its natural habitat :

....a)...GENERAL APPEARANCE :
This aquatic animal is well adapted to its underground environment. Proteus anguinus is a true troglobiont. Its body consists of a Head, Trunk, Tail, two 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 it has 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. 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."

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 anguinus at depths of water varying from 0 - 30 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 25 cm to 30 cm (10-12 inches) long and sometimes even longer, Proteus anguinus is the largest permanently cave-dwelling animal in Europe. Proteus is the only cave amphibian in Europe. Until recently, we were of the opinion that the variety living further south in Herzegovina was in general, markedly shorter than its Slovene or Croatian counterparts. However, our direct observation of a specimen well in excess of 25 cm was observed in September 2006 at a habitat location in Trebinjsko Polje. We have no data relating to the body-weight of the animal. The female of the species tends to be bigger.
In 2007., the Project aims to begin recording body-measurements and mass of the Herzegovinian Proteus. 
....d)...SKIN and SKIN COLOURING :
..........i)....In common with many other amphibians, including salamanders, the skin of Proteuscontains 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 yellowish as a result of riboflavin. Some of the internal physiological arrangement of the Proteus can be observed 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." :-

"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."

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."

....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.
....g)...AUDIO SENSORY ORGANS :
For hearing, the inner ear has developed electro-receptors.
....h)...OLFACTORY SENSORY ORGANS :
For a sense of smell there are well developed chemo-receptors. Along the length of its body on either side are mechano-receptors.
....i)...TASTE SENSORY ORGANS :
For a sense of taste there are well developed chemo-receptors on its tongue.
....j)...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.

....k)...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 bright red by the blood visible through the gill structures. 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!!
....l)....LIFESPAN :
The lifespan of Proteus can be in excess of 100 years, although the norm is often less.
...m)...NUTRITION :
For nutrition, it eats a variety of food. 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. 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.
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.
....n)...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.
....o)...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."

"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."

By observing the behaviour of captive specimens, Parzefall (1976) and Briegleb (1962) have shown 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 adults 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 (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 (Laurenti 1768) in its Larval Stage of development in a vivarium.

The Larval Stage is the first, when the animal lives on the reserve of yolk. Vandel reports that under his artificial conditions, this lasted for about an average of 36 days. He reports that total duration of both the Embryonic and Larval stages was on average about 5 months and that in the other group of Proteidae, Necturus, which is a non-cavernicolous form, it is only 2½ months. 
Proteus is a neotenic or “paedomorphic” salamander. It does not undergo metamorphosis but permanently retains the physical characteristics of its larval stage into adulthood.
The retention of the external gills and its long finned-tail into adulthood are evidence of neoteny
 
 

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.
....p)...ANATOMY :
Above, we append an image of an old diagram of the anatomy of the Proteus anguinus, drawn by the Italian explorers M. Rusconi P. Configliachi and published in "Monografia del Proteo anguino di Laurenti" in the year 1819.
....q)...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.
....r)...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.
....s)...VULNERABILITY :
..........(i) 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.
..........(ii) Our Direct Observations :
As is the usual problem with general statements given by large international organizations, the actual situation often needs a more detailed and considered statement.
....................In Italy, the occurrence of Proteus anguinus is in those areas contiguous with or otherwise hydrographically connected with the Slovene Karst. The situation in Italy is partly within the control of the Italian Authorities and is substantially affected by the environmental and political situations that currently exist in Slovenia. As a Member State of the European Union, Italy has sufficient understanding and funding to take control of the situation.
....................In Slovenia, now a Member State of the European Union, the Government continues to ignore the advice of its own scientists regarding the effects of inherited environmental pollution on the future of the country's amazing hypogean bio-diversity. In addition to not addressing the environmental problems inherited from the period of industialization under the previous Yugoslav regime, it is currently embarked upon further industrialization which will only add to these environmental problems.
The Slovene Government clearly has the future of its Proteus anguinus population in its own hands and has no-one else to blame but itself for the decline in numbers of this species. While it continues to choose to ignore its own specialists, we certainly have no sympathy for the Slovene situation. The country is now politically mature enough and financially able to solve these critical problems for itself. In the meantime, its Proteus anguinus anguinus population continues to decline and the future of the "Black Proteus", Proteus anguinus parkelj, which are only found in a very tiny geographical range in Slovenia, is put at a critical status.
....................In Croatia, a candidate country for European Union membership, the situation is quite different. Here the Proteus is more widely distributed. It faces the same general type of threats as exist in Slovenia, but not in such a concentrated form. Additionally, in the Dalmatian Littoral Karst, a more gentle decline in Proteus populations took place under the previous Yugoslav regime than in Slovenia but, more specifically it can be stated that right now in Dalmatia, within the area of hydrographic influence of the Trebišnjica River Basin, hypogean habitats are under threat as never before. This threat is principally caused by industrial pollution of the Trebišnjica River within the Repulika Srpska political entity of Bosnia & Herzegovina.
....................In Bosnia & Herzegovina, where the present geographical range of Proteus anguinus within Western and Eastern Herzegovina remains unclear, it is evident that more than anywhere else in any other part of its geographical range, there is the greatest need to protect Proteus anguinus and its hypogean habitats. Here the occurrence of Proteus in Eastern Herzegovina was once prolific. Due to the growth of the hydro-electric industry under the former Yugoslav regime, most notably in the Hidroelektrane na Trebišnjici Project, serious disruption to the natural karst hydrographic conditions has caused the serious demise of many hypogean species and has badly affected Proteus. However, it is as a result of post-Communist-era industrial degradation of the karst environment within the Trebišnjica River Basin that we are now witnessing the final and rapid demise of Proteus anguinus in its natural habitat.
Due to the "difficult" politics of Bosnia & Herzegovina, it is not yet even a candidate country for membership of the European Union. It does not have the actual or potential access to great wealth which Slovenia and Croatia enjoy. Yet here in Bosnia & Herzegovina are remarkable "hotspots" of Proteus anguinus Habitats. These habitat "hotspots" are the remnants of a once much bigger and contiguous "hotspot". This is where the work must be done and the funds provided to recover the situation!
....................As for Montenegro, we have no comments to make about the prevailing situation, for reasons of diplomacy!

....t)...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 Society's Joint International Programme in Bosnia & Herzegovina is the only known project that is fully underway (and has been for 8 years) and which has been specifically designed to protect Proteus anguinus by understanding its behaviour and problems in its natural habitat and then addressing those problems. The 30-year Programme, (about which this is the first of three dedicated Webpages), is in three 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.

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 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 idea of a universal 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 Conduct, 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.

.........Summary of what the Zoological Society of London is doing :
The Zoological Society of London (ZSL) has initiated an EDGE Project. The mission of the EDGE of Existence programme is to prevent the extinction of the world's most Evolutionarily Distinct and Globally Endangered (EDGE) species. Its first step towards achieving this mission is to raise awareness of these species. In the case of the EDGE Amphibians, the ZSL have highlighted the ten most endangered species of amphibians, with Proteus anguinus being on their list. 
The ZSL's "Proposed Actions" of the EDGE Amphibians Programme for Proteus anguinus are :-
"EDGE would like to fund an EDGE Fellow to carry out surveys on the distribution of the species in Slovenia, which is currently very poorly known.
EDGE hopes to fund research into the distribution of the olm in Slovenia so that the range of this species is more accurately known; currently it is suspected that the olm may be found beyond the existing confirmed range, however this needs verification by field studies. This research is required to feed into a Conservation Action Plan for the olm and may identify additional threats to this species that need to be addressed."
In our opinion, this action-proposal is highly insufficient to be able to do anything to address the wider problems of this species across its full geographical range in the Balkans. Slovenia has all the expertise and knowledge it needs to be able to deal with the problems of the Olm within its territory. It has its own specialists who have been working with the Slovene Proteus for decades. The only aspect that is missing in Slovenia is a political willingness to listen to its own scientific experts and then to act accordingly on their advice. In the absence of this, work should be dedicated elsewhere.
Clearly, the more critical problems for Proteus anguinus lay elsewhere in the geographical range of the species; namely in Bosnia & Herzegovina.
More about the ZSL EDGE Amphibians Programme can be found at :-
.........http://www.edgeofexistence.org/amphibians/default.php

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 habitats!

Proteus anguinus is indeed a highly endangered species. However, due care must be undertaken by anyone who thinks that it is a wise and practical idea to just "dive in" and 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 across its geographical range.

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