biospeology is the study of the cavernicolous organizations.


History of the biospeology

Precursors and others

biological is the study of the world living inside the terrestrial cavities (or hypogean). This scientific discipline, connects related to speleology, bore the name of biospeleology first of all (Armand Viré 1895), then was renamed biospeology (Armand Viré 1904, Emil Racovita 1907): it is this term which will be finally retained.

The first certificate of an observation by the man of cavernicolous animals dates from average Magdalénien (approximately 15000 years): Count Bégouen indeed discovered in the cave of the "Three-Brothers" in Ariège, a bone of engraved bison on which one can recognize Troglophilus (species of grasshopper troglobie). Historically, the first cavernicolous alive being which was the subject of a written description was protée, vertebrate (batrachian urodele) a nearest relative to the salamanders. Discovered in a cave of Yugoslavia, the area of Carniole by the baron Johan Weichard von Valsavor , it will be quoted in a publication published in 1689 in Laibach (Lubjana). Then terrestrial cavernicoles will be discovered and studied in the cave of Postumia (Adelsberg) like Leptodirus in 1831 by Franz Von HohenWart. The first cavernicoles discovered in France were it in 1857 in the Pyrenees Ariégeoises by CH. Lespes. Later, thanks to and with its many explorations, the vocation of biospeologist came to his companion Armand Viré who installed even the first underground laboratory dedicated to the biospeology in the catacombs under Botanical garden in Paris between 1896 and 1910.

Other great names marked since this discipline (alphabetically): Pierre Alfred Chapppuis, Claude Delamare-Debouteville, Carl Eigenmann, Louis Fage, Rene Jeannel, Curt Kosswig, Albert Vandel ...

When it is known that celebrates it itself, studied the cavernicolous animals and published articles on their subject (Thermosbanea mirabilis, 1927, in "Fauna of the French colonies"), one will understand easily that the bibliography quoted at the end of the article is only one infinitesimal piece of what was published to date in the field.

First test of classification of the cavernicoles

The first attempt at classification of the hypogean animals (alive under ground) was published in 1849 per J. C. SCHIÖDTE in a work entitled "Specimen Faunae Subterraneae". This first attempt at classification was based on the zone occupied by the animals in the caves: shade, total or partial darkness, concretions. A significant modification will be made by J. R. SCHINER in 1854 in an article on the caves of Adelsberg. It will exclusively introduce the denomination of "troglobie" for the animals living into the caves, it will call "troglophiles" those which are not completely dependent there and finally "occasional hosts" all the others.

Since one preserved the terms given by Emil Racovita in : troglobies, troglophiles and trogloxenes. It is important to note that this classification does not have anything systematic nor phylogenetic: it gathers in each three category of the very diverse sets of animals. This sorting, only based on ecological considerations, is however sufficiently practical so that it is always used.

With share of the speleologists, that there of alive under ground is

The underground world: place of life

One cannot give general and universal description characters particular to the underground life, because the caves and their "climate" are quite different according to the latitudes or the altitude to which they open. These differences are all the more sensitive as one is close to the entry or surface. However as one is inserted under ground and in the moderate areas of the sphere common characteristics are found.

Darkness : only point common to all the caves of the sphere, last a few tens of meters of the entry (variable distance according to the topography of the cavity) the absence of light is total there. Nothing to see (it is the case to say it) with one night without the moon. All spéléo made some one day the experiment, amusing if it is voluntary or unpleasant in the event of unexpected incident, and can testify some. When the light dies out, all that you see results from your imagination or occurred directly on your optic nerve without passing by the box departure, i.e. the retina! Black it is black...

Hygroscopy : the percentage of moisture of the cavities is generally very important in the moderate zone and in any event higher than outside. There are of course "dry" caves rather, and the percentage of moisture varies according to the season, the zones, the depth, the circulation air, meteorology, the presence of hypogean rivers etc. In the cavities of our areas, the water content of the air is often higher than 90 % and can sometimes come very close to 100 %, which makes it possible certain aquatic animals to leave water to change place by the terrestrial way (Niphargus). One can also meet terrestrial animals as the isopode Scotoniscus macromelos which can remain drowned during several days without dying for as much! Actually, of very many cavernicoles troglobies are amphibious.

Temperature : it differs notably from that recorded outside. Let us tell it good once, in the world there are caves where it freezes and others where it makes more than 20 °C. For as much, the variations in temperature are increasingly weaker than outside and the climate is more lenient there. If it makes 0 °C in an Alpin swallow-hole of altitude, there can be 10 to 20 degrees less at the same time outside, if in New Mexico one raises more than 20 °C in a gallery of Lechuguilla, mercury can go up to 50 °C in the shade of the external cacti. In the moderate areas of Europe, the temperature of the cavities of average altitude turns around 12 or 13 °C, the variation in temperature is weak, even negligible there. One can summarize all that as follows: under ground the temperature is much more stable than outside and the seasonal or daily variations definitely reduced beyond a certain depth or distance of the entry. It is a well-known phenomenon of the Languedocien speleologists, in summer one is well under ground because it made fresh there and in winter one because it is just as easily made soft there there: two good reasons to practise the spéléo all the year!

On the other hand, the current living conditions can be quite different from what they were in the past. One should not forget only if cavities which seem today favorable to the development of the cavernicolous animals of are completely deprived, the origin of this phenomenon goes up sometimes well far. The caves are not populated beyond a certain latitude which corresponds appreciably to the extension of the glaciers during two last glaciations of the Christian era (Mindel and Würm). The ices coming from North destroyed their settlement on the whole of the sphere. A long time, it was believed that there was not troglobies in the tropical caves but recent research shows that many species are present.

To finish, let us add that if no house plant can push far from the entry of the caves (consequently no herbivore saw there), food for the cavernicoles does not miss. First of all the density of population is not enormous (put aside in the heaps of guano where that grouille really), then water involves there sufficient organic matter for repaître the troglobies. I do not even speak to you about the myriads of bacteria contained in clay! Useless thus to let trail your snacks hardly started to nourish Aphaenops. Never forget that the cavernicolous animals nourish themselves under ground without you since glosses with what comes to them naturally from outside and, that in the absence of another thing, they can make belt (some troglobies have particular faculties for that) or devour themselves between them since they are often carnivorous! Consequently, less we will influence their diet, better that will apply to them...

Order please

As soon as it was noted that the underground world was populated, to make a sorting among all these alive beings had become essential. Several possibilities are offered to that which attacks this task:

  • to classify according to anatomical characteristics, it is what make the systematicians and who allows to associate such or such species an order and to a given family,
  • to classify according to the preferred places of life of the cavernicoles: water, guano, clay, the concretion formation... Tests in this direction took place but the result is not convainquant: the adult animals can, for example, food in a zone of the cave completely different from the larval stage of the same species. However there remains interesting to study the various underground ecosystems: input area - slightly enlightened of the cavities, heaps of crawling guanos of life, mixtures of stone, of clay and walls stalagmitic, free subsoil water and finally drowned interstitial medium of the ground water. In each one of these parts one will find preferentially such or such types of animals, such as for example in the input area whose biotope was studied in details and to which the name "of parietal association" per R was given. JEANNEL.
  • to classify according to faculty with living exclusively or not in the hypogean world.

This last method was refined with the passing of years to be currently recognized like most practical in biospeology. As we said it above to the 1.2), three categories of cavernicoles are defined by the narrowness of their bond in the deep underground world: trogloxenes, troglophiles and troglobies. The speleologists, following the example clamping plate or gun dogs fallen by inadvertency in swallow-holes, do not form part of any of these categories. We are only accidental or voluntary visitors underground world, not of the cavernicoles!

Three categories of cavernicoles


In fact animals use, when that is possible, the underground world during a part of their existence for reasons particular to each species. The physical characteristics of the cavities theirs are temporarily favorable, for example, to hibernate (bear and bat), for estiver (batrachians of the hot countries) or quite simply to shelter (snakes, rodents). These animals can find elsewhere conditions similar (semi-darkness, temperature stable...) and to use other places in the absence of cavities. Moreover these animals do not carry out their complete cycle of reproduction under ground: even the species of bats which use the caves as nursery couple outside at another period of the year.


These animals, although far from different morphologiquement from the épigées forms, are particularly well adapted to the underground life. During the history of their line appeared, following genetic variability, of the characters which made them more suited than of others to the life under the specific conditions of the underground world. It is what is called "preadaptation".

Certain animals use these predispositions to exploit the hypogean world in their favour, they can then see their behavior differing appreciably from that of the épigés members of the same species: the épigés Oxychilus snails hibernate whereas cavernicolous Oxychilus troglophile has a constant activity throughout the year. More extremely still, if one analyzes the gastric juices of a species of épigé Oxychilus one realizes that they contain ten times more enzyme able to digest the carapaces of the insects than those of other snails, and yet épigé Oxychilus is détrititivore (animal died, dead sheets)! And well, guess what, under ground the species troglophile of Oxychilus is carnivorous: it eats waste of carcasses of insects and drives out even butterflies! Beautiful example of preadaptation: without this preliminary physiological specificity, never Oxychilus could not have given a line of viable cavernicolous animals.

For as much, and even if their cycle of life proceeds entirely in the cavities, the morphology of the troglophiles or did not evolve/move very little: perhaps they are not hypogean since a sufficiently raised number of generations, because as S underlines it. J. GOULD: "the organizations are not a modelling clay that the environment works with its liking, nor of the billiard balls on the green carpet of the natural selection. The morphology of these organizations and the behavior which they inherited the past exert a constraint and slow down their evolution: it is impossible for them to acquire new optimal characteristics quickly each time their environment changes. ». Perhaps only the bacteria, still badly known, seem to make exception to this rule and to have an adaptive capacity above the average of the other alive beings but, in spite of an important bacterial presence in clays of the caves, this is another history...


In fact the true cavernicoles have surprised the first observers by their physical aspect, different from that of the épigés animals. Although in a remote way resulting from animals of surface, they are since so distant physiologically and morphologiquement that they cannot survive any more well a long time outside. Their development depends completely on the caves, swallow-holes, ground water which they populate and to which one says that they are pledged. For all these reasons, they form new species with whole share, cousins distant from those which live outside.

There are not thus herbivores troglobies since there is no chlorophyllian vegetation in the total darkness, not of birds nor of mammals (Guacharo and the bats are trogloxenes), some rare vertebrate (fish, batrachians) and an immense crowd of invertebrates (insects, shellfish, molluscs, towards, unicellular).

Particular characters of Troglobies

The hypogean species troglobies true present compared to their épigés cousins of the same family, of the distinctive features of which most frequent and most known are as follows:


Many species do not present any more eyes (at least at the adulthood), others have eyes extremely or not apparent tiny rooms (hidden by skin). But attention, anophtalme and blind man are two different things: certain cavernicoles blind although are equipped with eyes. Moreover, heaps of épigés animals are blind (with or without eyes). The absence of eyes is thus not an immutable rule in the underground world of the troglobies, but at most a tendency much more frequent than at the épigées species or troglophiles.


The animal fabrics are more or less coloured and these colors have various origins: optical phenomena (reflections of the feathers of the peacock), coloured pigments (ornaments of the skin of the salamanders, red colour of haemoglobin), mélanine (bronzing of the plagists homo sapiens). The biospeologists could note that many species troglobies had the rather pale dye or were almost transparent (niphargus, protée). Others however still have dark colors (Staphyllin). It would seem that this provision to lose certain pigments is not always irreversible besides, at protée for example which becomes brownish when he is exposed a long time with the artificial light. At other species, the exposure to sunlight is straightforwardly mortal (over-sensitiveness with UV) within a few going second (Planar) or a few minutes (Sphodrides) with a few tens of hours (Niphargus).

The absence of wings (whereas the family is ptérygote)

All hypogean the troglobies whose épigés cousins are winged, is deprived of complete wings. Although their élytres is still present, the wings are always atrophied, it remains about it often only traces, stubs. Once again, this character also meets at certain épigées species (Trechus which live in the humus).

Cut and forms body

Although one has, in the past, often written that the troglobies saw their size increasing (like that their antennas for example) compared to their cousins of the same épigés groups, any general rule does not seem to arise from the systematic examination of the species of the underground world. It seems simply that the evolution accentuated certain characters already present on the épigées animal lines once that they were found insulated under ground. The opilionidés cavernicolous ones have for example legs even longer than those their cousins. If the cavernicolous crayfish Cambarus tenebrosus is larger than crayfish of the brooks, the isopodes, them, became tiny.

Afflicted if I still have just demolished one of your certainty, but the idea according to which the cavernicoles had seen their sensors increasing in the face and a number to compensate for the loss of the sight, although tempting for certain species, cannot be generalized. Moreover, this interpretation of the evolution is a tantinet obsolete.

Life and evolution of the cavernicolous animals.

From which do they come?

The totality of the cavernicolous animals result from épigées animal lines from which they diverged slowly. The majority of the watery fresh water cavernicoles comes curiously from marine forms. But now let us try the machine to go up time...

Let us climb on the seat and insert the button corresponding to the tertiary era. We have just made a good behind some 50 million years. The position of our area (Languedoc Roussillon) is several hundred kilometers more in the south that today, and the Atlantic Ocean, in the west, is ridiculously narrow. An enormous mountainous solid mass emerges towards the south and extends gradually from the Iberian peninsula to current Italy. But neither Corsica, nor Sardinia are yet places from there, just starts one to guess them in the west. Jurassic and cretaceous limestones, compressed by this push of south-west towards the North-East, are folded, fractured and with all these slits the slow process of karstification starts. It should be said that the climate is rather hot and wet, one would be believed under the equator. Precipitations are strong and the luxuriant vegetation: a vast forest covers the areas with average altitude, their ground is made of a thick layer of humus, fresh, wet and obscure where our future cavernicoles grouillent; in the zones less touffues, the small ancestors of the horses graze close to animals resembling tapirs: lophodions, beavers and raccoons play about near. In the lakes among fish, one finds limnées, crocodiles and tortoises, and the hot sea is populated enormous oysters, sea urchins and sharks.

Towards -25 My, a basin of collapse opened in the south, it was invaded by the sea whose limits move once again, Corsica and Sardinia separates by taking their current site. Already, water in withdrawal one left behind them the ancestors of Caecosphaeroma and Monolistra (shellfish isopodes) which begin their long training of the fresh water life in this immense network of slits drowned of more and more fractured limestone. This sea will end up being drained (-14 My) and at the time of its return towards -5 My, the climate softened already a little. The remainders of the great tropical foliations are neighbourly with forests of birches, maples and oaks. And terrestrial cavernicolous fauna in all that? After a great mixing of several animal migrations coming from the south and the east, a part perhaps begins its hiding. Large mammals play about in the plains: rhinoceros, mastodons (small elephants), antelopes, hyenas, lions and tigers with teeth of sabre. Soon the climate will become seriously fresher, then increasingly hard on the whole of the sphere: for many species the end is close...

We are in Pleistocène (-1,65 My), from Africa where it began its development since approximately 10 the My man, the last born of the primates, arrived to Europe. In a few millenia, the climate became very unstable: three long ice ages will alternate with others more lenient but dry. The polar caps extend, the alpine and Pyrenean glaciers advance and the general level of the seas drops of more than 100 meters. Limestones are eroded always because they are extremely sensitive to the gélifraction which breaks them and widens them diaclases. Under the crust of ice which covers all north with Europe, the animal forms, resulting from the tropical climate of the tertiary era and the end of the secondary, irremediably and are definitively eliminated. In our areas with the variation of the polar ices, the forest was replaced by a grassy steppe strewn with woodland pines often swept by cold winds and violent one. Small the bébêtes frileuses does not have any chances of survival outside, only the nivicoles species resist in the moraines the face of the glaciers. Certain species migrated certainly benefitting from the draining of the Straits of Gibraltar. Those which succeeded in hiding and surviving benefit from an underground space that the karstification in full rise does not cease extending. Nothing will come any more to disturb them in their quiet retirement until the intrusion of the men.

How do they live?

To eat, reproduce, die... What a life! Fortunately as we saw to the 2.1), food generally does not miss. A part comes from outside brought by water or the air: it is made up of refuse and various remains of plants or animals. Another important source of food is in the cavities themselves within a complex food chain. This one could be summarized as follows: BACTERIA of the ground - > PROTISTES bacteriophages (which nourish bacteria) - > ANIMAL nourishing organic contents of the silt and clay (Oligochètes, Nématodes, Mollusques, larvae of Protée, young Niphargus) - > CARNIVOROUS truths which consume others troglobies, as well as troglophiles and trogloxenes. Between two meals, the life of the troglobies seems to proceed with the idle, compared to the épigées close forms. Experiments showed that it is a constant characteristic of the physiology of the troglobies: they breathe slowly, consume little oxygen, lay?ufs larger but fewer. The larvae of Coleopters less often moult and remain little with the free air before isolating themselves in clay from long months. Some are not even nourished before being transformed into adults. On the other hand Myriapodes pass, them, by a number of higher larval stages. At all events, in all the cases, the lengthening of the duration of the larval stage will lead to a total larva/adult longevity larger than that of the species living outside.

Which evolution?

With. VANDEL expressed the difficulty in very well interpreting the evolution of the cavernicoles while writing: "the idea of adaptation became at such a point obsessing which one could write that the depigmentation and the anophtalmy represented adaptations to the cavernicolous life. As much to say that the catarrhe, rheumatisms and presbyopia are adaptations to old age ". Is this with saying that the biospeologists are annoyed with Darwin and joined the clan of the creationnists? That nenni, far from us the idea to deny the role of the Darwinienne evolution while being based on the appearance of the characters specific to the troglobies, but it is necessary well to seize the significance of it.

Admittedly, in the caves, there can there be competition for food, but the evolution is generally the consequence of a selective pressure of the medium. Here, as one saw, the atmospheric conditions are stable, large the predatory non-existent ones, the density of population low. This medium thus forgives many "errors" and of the heaps of decays, which outside would be fatal, can persist. Morphological characters still unstable at certain species, but supported by the conditions of the external medium, for example will disappear. The absence of a real adaptation to the underground medium thus does not mean that there is no evolution. This evolution known as "regressive" resembles rather a bottom of evolutionary bag, with the multiple causes: épigée preadaptation followed of insulation in a not very selective medium since many generations: this design is called ORGANICISM (A. VANDEL 1964).

What preadaptation?

Let us leave, for some lines, the serious references and see how one could clear up this concept in a smiling way...

Imagine that a club misses members and that by forming part, you decide to convert with speleology two of your friends. The first, Édouard, are reprocessed merchant navy, impassioned model making, Breton, large, corpulent, short-sighted, claustrophobic and prone to the giddiness. The second, Alfred, of size average, slender, sporting, nyctalope, is originating in Ariège, it collects minerals and followed a time the occupation of minor.

With your opinion, with which of your two buddies, Édouard or Alfred, are likely you the most to succeed? Preadaptation it is a little that...

In the animal population, a species has hundreds of genes which code thousands of proteins and the whole is expressed in the form of a deal even more great of physical natures, physiological or behavioral distinct. The variation of some of these genes inevitably does not have particular adaptive interest for the épigée life: it is said that it is neutral. But among this genetic inheritance "neutral", certain genes can bring an adaptive advantage for the hypogean medium: it will appear like a predisposition when, by chance, the animal species which carries this character is carried under ground and succeeds better than another to adapt to it.

Not easy to sort all these small beasts


Here above the list (extremely incomplete) of the drawers in which are arranged the alive beings!

And yes, there is much of it... To arrange the room of a 14 year old teenager it is nothing beside the mission which the zoologists systematicians gave each other and called taxonomy. The more so as by discovering every year of new species or even of the odd animals that one does not know too much where to arrange, it is sometimes necessary to create new drawers. Moreover, each one goes there from its preference and some alternatives exist according to the researchers. And like there was a life before you, it is necessary also to classify all the fossils if one wants to include/understand the evolution!

All these enthralling characteristics, has been just added for a few years, the study of the genetic inheritance (ADN or ARN) of the alive beings in order to compare them, with beautiful surprises with the key. Only for the clade (drawer) largest, that of the "Reign", a very clear evolution took place during fifty last years. One passed from the elementary opposition "Animal kingdom"/"Vegetable kingdom", with the trilogy "Animal kingdom"/"Protistes unicellular"/"Vegetable", then with a group of 5 reigns: "Plants"/"Animal"/"Mushrooms"/"Protistes (unicellular complexes)"/"Monères (bacteria and cyanophycées). All this was recently corrected and a new classification is born, living it there is shared originally in three categories: Eubactéries/Archées (other types of bacteria)/Eucaryotes (all other living organisms which they are mono or pluricellular). In this phylogenetic classification (classification in groups containing a common ancestor and the whole of its descendants), the Mushrooms for example are gathered with the animals or métazoaires (cow, worm of ground, sponges...) in a taxon (contained concrete of a category) called "Opisthochontes" because they have common points currently well defined. For more details you defer to the excellent work of G. LECOINTRE and H. The GUYADER quoted in bibliography.

In spite of this apparent difficulty of taxonomy, it is the only means of finding and of including/understanding the complexity of the alive one there. I will give you an example. For the author of these lines, impassioned speleologist of his state, which is?

INDIVIDUAL: Doc. Carbur, 1 m 86, 87 kg (Homo sapiens) SPECIES: Hominiens KIND: Homo FAMILY: Hominides (Man, Gorilla, Chimpanzee) UNDER GROUP: Hominoïdes (Man and "large monkeys") GROUP: Simiiformes (Spider monkey, Ouistiti, Macaque, Gibbon, Orang-outang, Gorilla, Chimpanzee...) INFRA ORDER: Haplorrhiniens (Tarsiers, Monkeys) ORDER: Primates (Monkeys, Tarsiers, Lemurs, Lorisoïdes) CLASSIFY: Mammals [ Cetacea, Carnivores, Chiroptères, Artiodactyles (cow), Périssodactyles (horse) ] UNDER PHYLUM (UNDER JUNCTION): Vertebrate PHYLUM (JUNCTION): Chordés REIGN: Animal

Good, in the series "Nothing is simple, to say it immediately as much to you! ", we now will present to you, a modern classification good more detailed where you will find in fat some" drawers "of preceding classification, much more traditional. You hang, it left!


And well here, one smells oneself immediately better when one knows from where one comes not? If you want to know of it still more, it does not remain you any more that to launch you in the genealogy, but that it is another history...

A simplified sorting (systematic almost summary) of the hypogean alive forms.

To present a systematic classification simplified in a field of zoology as this one is a challenge!

All divisions will not appear, but only those which are relatively well represented in the underground world (except exception: the mention "not of hypogean forms" is then added). Moreover I will as often as possible mention a small general description and very synopsis of the type of animal concerned with such or such group. When that is possible, I will add to it one or more examples of relatively known épigés animals and cousins far away from the hypogean forms of the same group. Sometimes also the line will be supplemented by the name of a famous cavernicole. Generally the lines will be incomplete and will stop with the families, or even with the kinds or the sub-orders.

Which is the goal of such an incomplete presentation? It is double: initially to show the extent and the extraordinary variety of the hypogean animal forms and then to give some bench marks on troglobies about which one often intends to speak in spéléo without never knowing too much, neither what they resemble, nor of which kind of animal it acts.

CAUTION: here the sorting are carried out on anatomical criteria, biological or phylogenetic but do not hold account of separation between Troglophiles and Troglobies!

NB: in the tables, beside the names of animals quoted in example, the mention (EAR) will mean épigé and (HYPO) hypogean.

TABLE 1: the vegetable kingdom

TABLE 2: INVERTEBRATES PROTOSTOMIENS (except arthropods classified in table 3)


ARTHROPODS or EUARTHROPODES: they are animals with the members articulated with an external chitinous skeleton. Classification indicated hereafter date of about thirty years. Considerable evolutions took place since, placing for example Myriapodes, the Shellfish (renamed Pancrustacés) with Hexapodes (Insects, Protoures and Collemboles) in a single group called: MANDIBULATES or ANTENNATES. These two last names have the advantage of being rather clear!


TABLE 5: HEXAPODES (having 6 legs) INSECTS PTÉRYGOTES (winged)


TABLE 7: the TÉLÉOSTÉENS (FISH whose upper jaw is mobile and the symmetrical caudal fin outside)

For reasons practise these tables were not reproduced here, they are consultable on the heading of origin of this article:

[ 1 ]


Currently the systematicians described approximately 1 750 000 living organisms on the ground and the scientists consider their number probable in a fork from 10 to 40 million: crowned uncertainty!

At all events, the biodiversity varied during geological times, developed much thanks to the genetic variability, controlled by the natural selection, reduced violently by interplanetary cataclysms (falls of comets or meteorites) or ecological, it is currently in the course of fast reduction (about several thousands of species each year) under the essential action of the man (voluntary destruction, pollution, modification of the ecological niches). As us points out it the American paleontologist S. J. GOULD, incidentally destroyer of the creationnists, "extinctions [ of the alive species ], (...) are final. Each time that an experimental prototype complexes disappears, it is for always ". Admittedly within the genomes, of the modifications always occur and of new species appear, but the balance is negative currently by far. Moreover the biodiversity was decreasing to the wire of geological times: never it was also strong only in Cambrien 530 Million Years ago. Since, in fact the species diversified, but not the other taxinomic categories more general (phylum, class, order, family): the branches of the tree of the life divide at their end but the trunk and the main branches seem fixed.

Voir aussi

Mini bibliography

The list of the publications concerning the biospeology is impressive (thousands of references). Here thus a small selection of works headlights (they is least things to see there a little more clearly in the caves). The list thus concise, but is spread out chronologically and enough varied in order to find your happiness there and more if affinities...

General works on the evolution

DARWIN (Charles), "One the Origin of Species", London, John MURRAY, November 1859 (first French edition: "Of the origin of the species by the means of the natural selection" translation J-J. MOULINIE, REINWALD and Co, Paris, 1873) GOULD (Stephen Jay), "the life is beautiful. Surprises of the evolution ", THRESHOLD, Paris, 1991 WRONG (Patrick) to dir., "Dictionary of the Darwinism and the evolution", PUF, 1996 LECOINTRE (Guillaume) and GUYADER (Herve), "phylogenetic Classification of alive", BELIN, Paris, 2001

Works on the insects

CHINERY (Michel), "Insects of Western France and Europe", ARTHAUD, 1998 NEL (Andre), "insects: a success of the evolution "in" For Science "N° 293, Mars 2002

Works on the cavernicolous animals

TRANSFERED (Armand), "the underground fauna of France", Paris, 1900 RACOVITZA (Emile George), "Test on the problems biospeologic" in BIOSPEOLOGICA I, 1907 MARTEL (Édouard Alfred), "underground evolution" pp 242 to 289, FLAMMARION, Paris, 1908 * JEANNEL (Rene), "cavernicolous Fauna of France", LECHEVALIER, Paris, 1926 JEANNEL (Rene), "alive fossils of the caves", GALLIMARD, Paris, 1943 VANDEL (Albert), "Biospéologie: the biology of the cavernicolous animals ", GAUTHIER-VILLARS, Paris, 1964 GEZE (Bernard), "scientific speleology", pp 137 to 167, THRESHOLD, Paris, 1965 DELAMARE-DEBOUTEVILLE C., "life in the caves", P.U.F., What do I know?, Paris, 1971 THINES G. and TERCAFS R. "Atlas of the underground Life - cavernicolous Animals". Boubée (Paris) and De Visscher (Brussels), 1972. GINET R. and DECOU V., "Initiation with biology and ecology underground", Editions university DELARGE, 1977 SIFFRE (Michel), "animals of the pits and the caves", HATCHET, 1979 Scientific COLLIGNON (Bernard), "Speleology, approaches. ", EDISUD, 1988 SLICE (Fabien), "Approach of the biospeology" File of instruction EFS N° 116, 1st edition, 1997. SIFFRE (Michel), "France of the caves and the caves" (pp 136 to 153), ED. PRIVAT, 1999

  • It is interesting to very compare the position Lamarckienne of MARTEL and its acceptance of the heredity of the acquired features, with the modern positions and much more Darwiniennes of its successors.

Biographical works

WRONG (Patrick), "Charles DARWIN", 1999

Group Franco-héllénique de Recherches Biological (Toulouse) and Nature and Man (Athens), "Albert VANDEL (1894-1980)" Published by the Company of natural history of Toulouse, Université Paul Sabatier, 118 road of Narbonne, Bât.: 4R3, 31062 TOULOUSE cedex, France., 2002

Scientific periodic publications

· memories of BIOSPEOLOGICA: publication of 1907 to 1955: more than one hundred volumes published by the "Files of Experimental and General Zoology".

Ref. BNF 1: Type: text printed, periodic Title: Biospeologica. Studies on the natural history of the underground field [ printed Text ]. 1907/09 1950/55 (I-VIII) Publication: Paris: [ S.N.?] material Description: , 8 ° Note: Constitute certain booklets of: "Files of experimental and general zoology"; also published in flight. independent Others the shape of the title: Files of experimental and general zoology - Note n°: FRBNF32712801.

Ref. BNF 2: Type: text printed, periodic Title: Files of experimental and general zoology. Natural history, morphology, histology, evolution of the animals... [ Printed Text ] 1872-1882 (I-X). 1883-1892 (2e S. I-X). 1893-1902 (3e S. I-X). 1903-1909 (4e S. I-X). 1909-1912/13 (5e S. I-X). 1912/13 (LI) - Publication: Paris: [ S.N.?] material Description: , 8 ° Note: Tables all 10 flight. : 1872-1912/13 (I-5e S. X) in X, 2e S. X, etc. - Contains: ?Files of experimental and general zoology... Notes and re-examined? who becomes since 1898 (3e S. VI) a suppl. independent; starting from 1907 certain booklets are consisted: "Biospeologica... "and since 1925 by: "Prostistologica... "which was published also independently Other the shape of the title: Files of experimental and general zoology... Notes and re-examined; Biospeologica... ; Prostistologica... ; Note n°: FRBNF32701334.

Ref. BNF 3:Type: text printed, periodic Title: Files of experimental and general zoology... Notes and re-examined [ printed Text ]. 1898-1902 (3e S. VI-X). 1903-1909 (4e S. I-X) . 1909-1912/13 (5e S. I-X). 1912/13 (LI) - Publication: Paris: [ S.N.?] material Description: , 8 ° Note: Before 1898 contents in: "Files of experimental and general zoology... "it preserves the volume number while becoming suppl. Other the shape of the title: Files of experimental and general zoology... - Note n°: FRBNF32701335.

· Memories of the International Company of Biospéologie: 30 annual bulletins published to date (available at the SIBIOS 09200 MOULIS/ST-GIRONS, France or in Spelunca Bookshop FFS)

· ENCYCLOPAEDIA BIOSPEOLOGICA: two volumes (flight 1, 1994) published since 1997 by the SIBIOS 09200 MOULIS/ST-GIRONS, France (see the synopsis)

Ref. BNF 1: Type: printed text, monograph Title: Encyclopaedia biospeologica. T. I [ printed Text ]/Christian Juberthie and Disappointed Vasile, éd. ; [ publ. by the Company of biospeology with the collab. of underground Laboratory of CNRS with Moulis and Institute of speology Emil Racovitza ] Bond with the overall title: Encyclopaedia biospeologica. Publication: Moulis: Underground laboratory of CNRS; Bucharest: Institute of speology Emile Racovitza, 1994 Printer/Manufacturer: 09-Saint-bosoms: Impr. Fabbro Material Description: XII-834 p.-[17 ] F. of pl. in black and coul. : ill., voub. ill. in coul. ; 31 cm Note: French or English texts. - Notes bibliogr. Other author: Juberthie, Christian. Disappointed scientific editor, Vasile. Scientific editor Company of biospeology. Scientific editor underground and edaphic Research center (Moulis, Ariège). Scientific editor Institute of speleology Emil Racovitza (Cluj-Napoca, Romania). Scientific editor Subject: Cavernicolous animals -- cavernicolous Encyclopaedias Plants -- Encyclopaedias Ecology of the caves -- Research -- History -- Encyclopaedias Speleology -- History -- Encyclopaedias (Rel.) : 550 F Note n°: FRBNF35730550.

Ref. BNF 2: Type: printed text, monograph Title: Encyclopaedia biospeologica [ printed Text ]. Volume II/Christian Juberthie and Vasile Decu, éd. ; [ publ. by the underground Laboratory of CNRS with Moulis and the Institute of speology Emile Racovitza in Bucharest ] Bond with the overall title: Encyclopaedia biospeologica. Publication: Moulis (09200): Company of biospeleology, 1998 Printer/Manufacturer: 09-Saint-bosoms: Impr. Fabbro Material Description: VIII p. and p. 836-1373: ill. in black and coul., couv. ill. in coul. ; 31 cm Note: English or French texts. - Notes bibliogr.. - the structure supports by error: ISSN 0398-7973 Other author: Juberthie, Christian. Disappointed scientific editor, Vasile. Scientific editor underground and edaphic Research center (Moulis, Ariège). □diteur scientifique Institut de sp □l □ologie Emil Racovitza (Cluj-Napoca, Roumanie). Scientific editor Subject: Cavernicolous animals cavernicolous Plants Ecology of the caves Dewey Index: 591.758 4 (Rel.) : 490 F Note n°: FRBNF37622750.

Internal bonds

External bonds


The phylogeny of the bacteria: [ 2 ]

A bank of data of unicellular and their classification: [ 3 ]


Cavernicolous animals: [ 4 ] Cavernicolous animals. Biospeology by Raymond Tercafs, Doctor of Zoological Sciences.

Run of Biospéologie: [ 5 ] Patent of initiator, the small course of biospeology of the school of Belgian speleology available on line: cheer!

Myriapodes: [ 6 ] Of Jean-Jacques GEOFFROY, and Didier Geffard.

Underground laboratory of Moulis: [ 7 ] The site of the underground laboratory of Moulis.

Hétéroptères: [ 8 ] Hétéroptères and biospeology.

Bernard LEBRETON: HTTP:// Site perso of one impassioned, with very many bonds.

Rene JEANNEL: [ 9 ] Who was Rene JEANNEL?

Biospeologica bibliographia: [ 10 ] Our minis biblio being more than brief, you have there what to inform you!!!

Bulgarian biospeleology bibliography: [ 11 ] Bulgarian biospeologic bibliography (in English).

Underground Hydrobiology and ecology: [ 12 ] A field of research of CNRS in Lyon (hydrogeology, underground ecology, biospeology).

Bonds of the FFS: [ 13 ]


  > French to English > (Machine translated into English)