|Bosna i Hercegovina||
PROTEUS PROJECT IN
THE TREBIŠNJICA RIVER BASIN"
AND ITS NATURAL KARST HABITAT IN THE TREBIŠNJICA RIVER BASIN."
Eastern Hercegovina and the Hercegovinian Littoral, Bosnia & Hercegovina; Dalmatian Littoral and Konavle, Croatia; Old Hercegovina, Montenegro.
Project Title [BiH-Hrv Lat] :.."Zajednička Strategija za Zaštitu Ugroženog Podzemnog Endem Čovječje Ribice (Proteus anguinus) i Njezinog Prirodnog Krškog Staništa u Slivu Rijeke Trebišnjice."
Istočna Hercegovina, Bosna i Hercegovina.
Project Title [BiH-Srpski Cyr] :.."Заједничка Стратегијаза Заштите Угроженог Подземног Ендема Човјечије Рибице(Proteus anguinus) и Његовог Природног Кршког Станишта у Сливу Ријеке Требишњице."
Источна Херцеговина, Босна и Херцеговина.
Project Title [Magyar] :.."Együttműködési Stratégia a Veszélyeztetett Barlangi Vakgőte (Proteus anguinus) Természetes Karsztos Élőhelyének Védelmére a Trebišnjica Folyó Völgyében."
Kelet- Hercegovina, Bosznia - Hercegovina.
"Proteus Project" Partner,
Updated.01 April 2018.
....."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. "Proteus Project" Webpage [4.].
.....Link to. "Proteus Project" Webpage [5.] - The "Proteus Project in Other River Basins in Bosnia & Hercegovina." (not yet active)
.....Link to Webpage on.."Proteus Project" Monograph Publication Series.
.....Link to Webpage on.."Proteus Project" "Observations in Speleology" Publication Series.
.....Link to. "Trebinje Proteus Observatorium" Webpage
anguinus anguinus (Laur 1768.)
1. Listed in App. II of the Convention of European Wildlife and Natural Habitats of 1979.
2. Appendix 2 of the Bern Convention.
3. EUDirective 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."
4. IUCN (Red List) Status 2013.2 = Vulnerable (VU).
5. CITES = No CITES Listing.
6. Present National Status :-
R. Italia = (no information);
R. Slovenia = "protected" since 1949;
(Formerly before the modern Republic, since 1922.)
R. Croatia = protected;
R. Bosnia & Hercegovina = Not yet protected.
(During the period of Austro-Hungarian Administration,
it was protected from 23 February 1914 - 1918 under the Decree № 35.380 V.)
.....Images..(above),...A photo-montage of a specimen of Proteus anguinus anguinus (Laurenti 1768) in its natural underwater hypogean habitat (phreatic conditions) in Eastern Hercegovina, RS-Bosnia & Hercegovina..
.....(Photos : The "Proteus Project's" Hungarian Cave Diving and Technical Support Team "Caudata Hungarian Cave Research" 07 September 2009.)
(This Column, Above and Below)
"Proteus Project" Partners,
Co-ordinator Groups, Sponsors,
its Title indicates, the Society's BiH Project No. 3. is primarily
concerned with the protection of the endangered population of the endemic
Proteus anguinus through the protection and conservation
of its natural karst habitats.
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 numbers of viable Habitat Locations and of the Proteus itself.
We cannot be a partner to encouraging the further demise of this unique and highly endangered species. In this regard, we draw your attention to relevant details, which expand on this problem, in Section 6.0.
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 Co-ordinator Group 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" Partner,
RS-BiH Local Co-ordinator Group,
Trebinje, Bosnia & Hercegovina.
Introductory Webpage...Short General Introduction to the "Proteus Project" and to Speleobiology.
"Proteus Project" Page 1...[Information Section on this Page] :
Section 1...Short Scientific Introduction to Proteus anguinus anguinus (Laur. 1768)..
"Proteus Project" Page 2...for the following Information Section on another Page :
Section 2...Bibliography and Further Reading..
"Proteus Project" Page 3...for the following Information Sections on another Page :
Section 3...Broad Aims and Objectives of the "Proteus Project" :
Section 4...Theoretical and Practical Fieldwork Support-Documentation for the "Proteus Project" :
Section 5...Glossary of Biological Terms & Developmental-Biology Terminology, together with General Speleobiological Terms
.................and Terms used in the Taxonomic Classification System of Fauna :
"Proteus Project" Page 4...for the following Information Sections on another Page :
Section 6...Synopsis of the Conservation Problems Associated with the Survival of Proteus anguinus and its Natural Karst Habitats in the Trebišnjica River Basin :
Section 7...Incidental Non-Project-related Data Acquisition - Exokarst Flora and Fauna :
"Proteus Project" Page 5...for the following Information Section on another Page :
Section 8...Incidental Non-Project-related Data Acquisition - Exokarst Flora and Fauna :
"Proteus Project" Originator
UK Co-ordinator Group,
Plymouth, England, UK.
Suppliers of Electronic Water-quality Testing Equipment to the
"Proteus Project", since 2000.
Palintest House, Kingsway, Team Valley,
Gateshead, Tyne & Wear, NE11 0NS,
All Publication Abstracts are cited fully in the BIBLIOGRAPHY Section on Proteus Project Webpage [2.] A GLOSSARY is now in Section 5. on Proteus Project Webpage .
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 our 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.
Scientific name :..Proteus anguinus anguinus (Laurenti 1768).
......................................[There may be other, as yet undifferentiated, sub-species of the White Proteus.]
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 or Trivial Names : "The White Proteus"; "Human Fish"; "Cave Olm"; "Cave Salamander";
..................................also :-..Srpski = "Човјечија рибица";..Bosanski = "Čovječija ribica";...Hrvatski = "Čovječja ribica";...Slovene = "Človeška ribica" and "Močeril";..Magyar = "Barlangi Vakgőte";
.............................................Polish = "Odmieniec Jaskiniowy";...French = "Protée anguillard"; ..German = "Grottenolm";
.....other related sub-Species : Proteus anguinus parkelj, (Sket et Arntzen,.1994)..[.=The Black Proteus], currently found only in Slovenia.
Vlada Republike Srpske-BiH.
Ministarstvo za prostorno uređenje, građevinarstvo i ekologiju.
Government of RS-BiH
Ministry for Spatial Planning,
Construction and Ecology.
Biodiversity specialists in the
Administrative Support-Department for
Republički Zavod za Zaštitu Kulturno,
Istorijskog i Prirodnog Nasleđa.
Republic Institute for the
are the extant descendants of the first vertebrate class to successfully
colonize terrestrial environments; hence they occupy a unique position
between fish and reptiles.
In general, amphibian skin provides essentially no resistance to evaporative water-loss and consequently, daily water turnover rates are an order of magnitude greater than in other terrestrial vertebrate groups. This has led to a suite of physiological, morphological and behavioural adaptations that have allowed a successful terrestrial existence in spite of this apparently spendthrift water retention strategy. Proteus anguinus, however, is a predominantly aquatic amphibian .......
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 as the eggs are laid outside in the aquatic habitat.
Discovery of Proteus anguinus anguinus and its accepted Type Locality :
Proteus anguinus was first recorded in the literature 327 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.
Short Note about Josephus Nicolaus Laurenti :
Josephus Nicolaus Laurenti (born 4 December, 1735 - died 17 February, 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.
Short Note about the work of Dr Mauro Rusconi and Pietro Configliachi with Proteus :
The 19th century Italian naturalists Dr. Mauro Rusconi (1776-1849) and Sen. Pietro Configliachi.(1779-1844) published their "Monografia del Proteo anguino di Laurenti" in the year 1819.
Dr. Rusconi , was the dentist in Pavia and he performed much anatomical research on live and dead Proteus specimens. The results of their work was based on the deaths of many specimens of Proteus and included dissection and
experimentation on live as well as dead individuals!
Shortly after the publication of their Monograph, it was translated into English and reproduced in the Edinburgh Philosophical Journal, Vol. 5. (1821)*.
As protectors of Proteus anguinus, to us involved in the "Proteus Project", the translated text, in places, reads like a gothic horror story as it references and details the dissection and experimentation on the live animals, often yielding no conclusive results!
For our readers' enlightenment, we shall include further extracts from the translated work of M. Rusconi and P. Configliachi in the relevant Sections of text below and we especially reproduce their anatomical drawings in Section(s) below.
If you wish to aquaint yourselves with the full text of the English Translation of their work, please refer to "Proteus Project Webpage ." (Bibliography) and download the relevant document.
Notes on abandoned 19th Century Taxonomic Descriptions of Proteus :
Fitzinger (1850) described a series of supposed "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).
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 during 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.
Another current line of thought proposes that its origin was as an epigean species living in freshwater surface lakes, from where it was forced underground by the increasingly cold climate of the last glacial period and where it then had to adapt to hypogean conditions.
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.
Speleobiological 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.
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 & Hercegovina.
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 Hercegovina, to the Trebišnjica River in Eastern Hercegovina. 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 & Hercegovina 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 Hercegovina 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.
Environment Protection Order,
RS-Official Gazette No.
for the Karst Biosphere Reserve
GENERAL PHYSICAL CHARACTERISTICS & BEHAVIOURS - supported by our own observations of Proteus anguinus anguinus in its natural habitat :.
....a)...GENERAL APPEARANCE :
This mainly aquatic animal is well adapted to its underground environment. Proteus anguinus anguinus is a true troglobiont often referred to as a stygobiont.
Its elongated snake-like cylindrical body consists of a Head, Trunk, and a laterally-flattened Tail, which is markedly shorter than the trunk. The Trunk has 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 its mouth contains tiny teeth. Males and females differ only slightly in their appearance, with the male of the species usually being 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."
It is interesting to note that Proteus exhibits a marked attachment to particular "favoured residence sites" within a habitat location. Our direct observations in Eastern Hercegovina 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 - 31 metres, (0 - 98 feet). At the greater depths, Proteus has only been observed in very small numbers.
At the shallower depths occur the greater population numbers.
..........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 fast flow of flood-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 obligate cave-dwelling animal in Europe. Proteus is the only cave amphibian in Europe. In 2007., we 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 other useful biometric 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.":-
"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 PHOTO-RECEPTORS :
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 optically 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 possible for Proteus as with 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 nearby low-frequency water displacements (Bulog & Schlegel 2000) and (Schlegel et. al. 2006).
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).
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).
(Schlegel 1996) reports on the observations of some experiments with Proteus and Helmholtz coils, whereby it was shown that Proteus aligns itself to natural and artificially modified magnetic fields. The animal's movements were observed by an infrared video camera.
The "Proteus Project" is continuing to analyse cave sediments from many Proteus anguinus habitat locations and locations where Proteus cannot possibly exist. Thus far, we have determined that there is a significant proportion of paramagetic ferrous / ferric material in many of these locations.
Although we do not yet have sufficient data to come to any particular conclusion in the matter, early indications would seem to suggest some interesting correlations.
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 major 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.
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 immediately behind and 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, only one of which remains fully formed and functional, whilst the other would seem to have degenerated and may not be able to function at all.
It has been claimed that it can also breathe through its skin, (ie. three respiratory systems in total), although claims for the skin-breathing method are questionable.
However, the idea that the internal rudimentary lungs are merely "secondary" respiratory organs has now been dispelled because groups of Proteus have been observed leaving the cave aquatic environment and feeding in the cave's terrestrial environment and
sometimes even at the cave entrance area!!
The "Proteus Project" has also recorded Proteus "gulping" atmospheric air through its mouth in a cave under various circumstances.
..........Adaptation to survive Anoxic (=hypoxic) conditions :
(NB. The term "anoxic" [or hypoxic] refers to such a condition relative to a named subject species. For example, a given availability of dissolved oxygen may be oxic for some species whilst being anoxic [or hypoxic] for others.
Anoxia [or hypoxia] does not necessarily imply a zero oxygen situation in absolute terms, but is a condition relative to the requirements of the named subject species.)
The ability of amphibians to survive extremes of oxygen deficiency 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 in such adverse conditions. The low metabolic rate of Proteus would seem to assist in its survival in low-oxygen conditions. For long-term anoxic survival, enhanced storage of glycogen in the liver is also necessary.
However, in the "Proteus Project", we have observed an interesting repeated pattern of survival behaviour in conditions of anoxia [or hypoxia] in certain populations of Proteus.
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.
The longevity in the lifespan of Proteus has again been the subject of renewed and recent investigations and has called into question the currently perceived theories about the factors which were thought to have contributed to the natural ageing process and the limitation
on maximum age. Several physiological traits are normally associated with long-lived animals: larger size, low metabolic rates, and high protection against oxidative stress. Examples include giant tortoises and elephants - animals that have large body masses and low
basal metabolic rates. However, Proteus does not show any of these traits. So why does it live so long?
The "free-radical / oxidative damage" theory, formulated in the early 1950's and which rose to prominence in the 1990's, was widely accepted as being the responsible mechanism for the restriction in the age or lifespan of many fauna.
However, evidence resulting from the study of Proteus would strongly suggest that this theory does not explain the controlling factor in the longevity of a species. In a useful overview, Speakman and Selman (2011) provide detailed implications of recent research carried out by Voituron, Y. et al (2010) on Proteus, which revealed that Proteus appears to have unexceptional defences against oxidative damage. This research is also reported upon by PhysOrg.com. and completely throws open to speculation the whole question of ageing.
With access to the captive breeding colony in the Moulis Underground Laboratory, together with relevant data on 400 specimens over many decades of data-collection, the researcher Yann Voituron estimates that the maximum age that Proteus can achieve is 103 years.
The problem for members of the "Proteus Project" is that the report by Yann Voituron is about a captive colony and is not about observations of a colony living in any natural habitat. Whilst we do not disagree with his general conclusions about longevity, basing a proposed maximum age figure on data from a captive colony in a totally unnatural living environment is already subject to much doubt. We also do not agree with several other behavioural traits reported in his Paper and which seem to be contradicted by observations in the field.
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.
....q)...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.
..........-.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, under certain conditions, can give rise to fully-formed live young. 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 stygobitic 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 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.
.....-.Egg-laying :-..The optimum water temperature for this species is 7º -10º C. Females normally lay up to 80 eggs, but curiously enough if the water is warm enough (about 15º C) they can give birth to two larvae instead.
The female has been observed undertaking egg-guarding and defending behaviour. Vandel (1966b) again reports that under laboratory conditions at a temperature of 11.9º C, eggs were not all laid at once but over a period of 1 to 3 weeks. He noted that under his artificial conditions, he obtained 10 egg-laying events producing batches varying from 20 to 60 eggs. The eggs were always laid on the undersides of large flat stones. The parents remained close by thereafter, maintaining a circulation of water around the eggs by the regular waving of their tails to prevent the settling of mud on the sticky mucilage which envelops the eggs. He reports that these eggs were from 8mm to 9mm in diameter, each surrounded by a layer of mucilage, by which the egg adhered to the stone on which it was laid.
..........-..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.
.....-..Hatching and Post-embryonic Development :-..At hatching, the young Proteus retains an important reserve of yolk, which appears as a whitish mass on the underside of the body. This is progressively resorbed.
The yolk and its resorption enabled Vandel et al to distinguish two distinct stages in the post-embryonic development of Proteus anguinus,.(Vandel, Durand and Bouillon, 1966.).
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.
Amphibians do not have a fixed number of vertebrae. Proteus anguinus anguinus has 29–32 vertebrae.
We have now appended images of old diagrams of the anatomical structure of the Proteus anguinus anguinus, drawn by the Italian explorers M. Rusconi and P. Configliachi.and published in "Monografia del Proteo anguino di Laurenti", (1819).
To make the information more widely accessible to our readers, we have drawn the Explanations of the Plates from the English Language Translation as published in the Edinburgh Philosophical Journal, Vol. 5. (1821).
Image PA7. (left) as PLATE VI.
We quote the following explanatory text of PLATE VI., with some modernizing of Old English words.
[NB. All sizes quoted herein relate to the scaled drawings in the original publication.]
Skeleton (Figs 1. and 2.) :
"Fig.1..View of the Head from below, 8x greater than natural.
..a-a,..the two branches of the Lower Jaw;
..b-b,..processes of the Temporal Bones to which they unite;
..c,..the roof of the Palate;
..d,..the os hyoides;
..e,e,e,..the three Branchial Arches of the right side;
..f,g, the intermediate bones of the 1st and 2nd arches;
..h,h,h, the 1st three Vertebrae;
..n,n, the branches of the os hyoides.
"Fig.2...The Skeleton of the Proteus of its natural size.
..a,..the three cartilages forming the Shoulder;
..b, the Pelvis."
The Digestive, Excretory and Reproductive Systems (Fig 3.) :
"Fig.3..The lower half of the trunk of a Female Proteus laid open.
..a,..the Alimentary Canal shortened and straightened from the action of ardent spirits:-- at its termination, it is slit up to show the common focus of the Ureters and Oviducts;
into which two pins are inserted, the opening from the Urinary-Bladder is indicated by a single pin.
..b, the left ovarium, containing minute ova, and drawn back to one side to display the Kidney underneath;
..c, a portion of the Oviduct straightened by the action of the spirits;
..d, the left Kidney;
..e, the Ureter running along its margin and terminating with the Oviduct in the rectum;
..f, a portion of the left Air-bladder, in this instance remarkably enlarged; the corresponding one of the right side was very small."
The Central Nervous System (Fig 4.) :
"Fig.4..The Cranium laid open to show the cerebral mass and certain nerves springing from it.
..a-a, the two hemispheres of the Cerebrum;
..b, the Cerebellum;
..c, the medulla oblongata;
..d, the right olfactory nerve;
..e, the origin of the fifth pair of nerves;
..f, the vestibule of the organ of hearing, laid open, in which the little sac is seen, and the origin of the acoustic nerves;
..g, the facial nerve;
..h, the entrance of the carotid [artery] into the Cranium, from which springs the opthalmic going to the eye (n);
..m, the origin of the par vagum."
The Head and Trunk (Fig 5.) :
"Fig.5..Profile of the Head and Part of the Trunk.
..a,..the external aperture of the Nostrils surrounded by Pores;
..b, the doubling of the Inferior Lip, which is in part covered by the Superior;
..c, the swelling or protuberance produced by the Heart."
Image PA8. (left) as PLATE VII.
We quote the following explanatory text of PLATE VII., with some modernizing of Old English words.
The Digestive, Excretory and Reproductive Systems (Fig 1.) :
"Fig.1..A Male Proteus laid open, to exhibit the relative size and position of the viscera.
..a,,.the Heart, with its pericardium, opened and turned back;
..b,b,b, the Liver drawn aside, to show the viscera beneath it;
..c, the Stomach;
..d, the Alimentary Canal;
..e, the Spleen;
..f, the Pancreas;
..g, the Testicle of the left side;
..h, a part of the left Kidney;
..i, the Urinary-bladder;
..k, the left Air-bladder, with its tube, opening into the conical cavity above;
..l, the Anus or Cloaca;
..m, the sinus of the vena cava."
The Heart and Air-bladder (Fig 2.) :
..a, the Heart reversed and turned upward;
..b, the short conical canal cut longitudinally, which communicates anteriorly with the Glottis and posteriorly with the cavity from which the two tubes, (c,c) terminating in the Air-bladder, proceed."
The Head (Fig 3.) :
"Fig.3..Head of the Proteus viewed from below, 8x greater than natural, displaying the circulating and respiratory systems.
..b,.the arterious trunk springing from it;
..c, its bulb;
..d-d, the two primary trunks arising from the bulb, and again subdividing;
..e, the first branch of the primary trunk, or artery corresponding to the common carotid [artery], and which subdivides into two, one branch (f) being continued to the first or exterior Gill, and the other (g) proceeding to the muscles of the os hyoides;
..h, the vein which carries back the florid blood from the Gill.
The second branch of the primary trunk (d) soon also subdivides, sending off the branch (i) to the third Gill, and another (l) to the middle Gill. To these two branchial arteries, the two veins (m-m) which carry back florid blood, correspond.
The principal trunk of this second branch, after receiving this florid blood, sends off the artery (n), which, descending along the air-tube, supplies the Air-bladder and generative organs in each sex; it then curves upward, and from its curvature gives off the vertebral artery (o), which, after sending some twigs to the Occiput, enters the vertebral canal, and descends along it; it also gives off another branch (p) to the Temporal bone, and then making another curvature downwards, it becomes a branch (q) of the Aorta, which by uniting with its fellow of the opposite side, it contributes to form. The Aorta (r) gives off the branchial arteries (s-s), the mammary (u) and the vessel (t) going to the Stomach; the letter x denotes a portion of the vena cava cut off."
The Gills (Fig 4.) :
"Fig.4..A Leaflet of the Gill highly magnified,
..exhibiting the branchial artery (a), conveying dark blood to the Gill, and the branchial vein (b), returning florid blood to the Aorta."
....t)...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.
....u)...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.
..........Official Information in the Public Domain from various sources :
The Olm is listed as "Vulnerable" in the IUCN Red List of Threatened Species, 2013.2., 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.
From our own perspective in Bosnia & Hercegovina, we would suggest that the status of Proteus anguinus anguinus should be set at because of the very low number of extant viable hypogean ecosystems that are able to support its existence in the
wild and because of the increasing threats to those that are still able to support this animal and associated species.
....w)...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 & Hercegovina is the only known project that is fully underway (and has been for 16 years) which has been specifically designed to protect Proteus anguinus by understanding its behaviour and the problems within
its remaining natural habitats ... 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 2. continues the overwhelming success already achieved in Phase 1., producing a great wealth of information about the behaviour of Proteus and the characteristics and problems of its natural habitats across a whole region within its geographical range.
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 Hercegovina 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 throughout 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 subject to continuous updating.
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 of restocking habitat locations by translocating some specimens from one location to another, not only because of the natural variations in conditions that exist between individual habitats but also to protect the gene-base of existing populations.
Proteus anguinus is indeed a critically 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 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.
....x)...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."
.....Proteus anguinus is not listed on CITES.
.........IUCN Red List, 2013.2 :
.....the Olm is listed as being "Vulnerable" because of its fragmented and limited distribution and ever-decreasing population numbers.
.........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, Slovene and collectors from other countries such as Italy, France, the Czech Republic and Slovakia continue to remove specimens for private collections or for illegal trading for onward use in Asia or for "scientific study".
.........Global Biodiversity Information Facility (GBIF) :
.....Entries for the Proteus anguinus species in this database are either mostly useless or misleading.
....y)...PROTECTION AND CONSERVATION OF THE ENVIRONMENT & BIODIVERSITY IN THE REPUBLIC OF BOSNIA & HERCEGOVINA.
.........(i) Summary of what R. BiH has agreed do :
UNITED NATIONS DEVELOPMENT ASSISTANCE - FRAMEWORK FOR BOSNIA AND HERCEGOVINA 2010 – 2014
A UNDAF has been prepared by the United Nations Country Team in Bosnia and Hercegovina in consultation with the Government of Bosnia and Hercegovina 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 Hercegovina 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."
Whilst both the Bošnjak-Croat Federation-BiH and the Bosnian-Serb RS-BiH Entity-Governments have their own separate Cultural, Historical and Nature Protection Institutions, none exists at a Central State level for the entire Republic.
In the case of F-BiH., they have a Culture and Sports Ministry, whilst in RS-BiH., the responsibility for Cultural and Natural Heritage now operates within the Ministry for Ecology.
........(ii) Summary of what the RS-BiH Government is doing :
During November 2011., we are very pleased to be able to announce that the "Proteus Project" has at last been able to engage with the RS-BiH Government's Nature Protection Institute within the Ministry for Ecology. Furthermore, as of September 2013.,
we have engaged directly with the RS-BiH Ministry of Ecology in Banja Luka, to progress as urgently as possible the Legal Protection of a defined cave part-Ecosystem containing a large breeding population of Proteus anguinus.
This has been a long time coming, but we are nearly at a successful conclusion and more details about this exciting progress will be published on these webpages later in 2016.
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