International Society for the Study

and Conservation of Amphibians

(International Society of Batrachology)

SEAT

Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

BOARD FOR 1991

President: Raymond F. LAURENT (Tucumän, Argentina).

General Secretary: Alain DuBois (Paris, France).

Treasurer: Dominique PAYEN (Paris, France).

ant Secretary, Europe: Günter GOLLMANN (Wien, Austria).

ant Treasurer, Europe: Annemarie OHLER (Paris, France).

Assistant Secretary, outside Europe: David B. WakE (Berkeley, U.S.A.).

Assistant Treasurer, outside Europe: Janalee P. CALDWELL (Los Angeles, U.S.A.).

Other members of the Board: Jean-Louis FISCHER (Paris, France); David M. GREEN (Montreal,

Canada); Roy W. McDiarmiD (Washington, U.S.A.); James I. MENZIES (Boroko, Papua New

Guinea), Richard WassErsUG (Halifax, Canada).

TARIFFS 1991

Subscription to

ISSCA Circalytes Alytes Total

Individuals

ISSCA direct members 60 FF 60 FF 200 FF 320 FF

ISSCA group or section members* 25 FF 60 FF 200 FF 285 FF

Non-members r _ 220 FF 220 FF

Institutions

ISSCA direct members 120 FF 120 FF 400 FF 640 FF

ISSCA group or section members* 50 FF 120 FF 400 FF 570 FF

Non-members - _- 440 FF 440 FF

* Members through a group or section of ISSCA (Société Batrachologique de France; Société

Lémanique de Batrachologie; Working Group on Oriental Amphibians).

Tariff for the inclusive Section or Group affiliation to ISSCA: 250 FF.

Tariff for individual subscription to the ISSCA Board Circular Letters: 200 FF.

MODES OF PAYMENT

In French Francs, by cheques payable to “ISSCA”, sent to our Treasurer (address above).

— In French Francs, by direct postal transfer to our postal account: “ISSCA”, Nr. 1 398 91 L, Paris.

— In French Francs, by direct bank transfer Lo our bank account: “ISSCA”, Nr. 10207-00014-

04014048104-97, BICS Paris-Monge.

- In U.S. Dollars: please write to our General Secretary (address-above) for further information.

Source : MNHN, Paris

AIVTES

INTERNATIONAL JOURNAL OF BATRACHOLOGY

March 1991 Volume 9, N° 1

Alytes, 1991, 9 (1): 1-14. 1

Batrachology

as a distinct scientific discipline

liothèque Centrale Muséum

Laboratoire des Reptiles et Amphibiens, ULN

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

Although amphibians and reptiles have been considered as two distinct

classes of vertebrates for more than 150 years, the study of both groups is still

traditionally referred to a single scientific discipline, “herpetolog”. It is here

argued that, both for scientific reasons and in order to have a better efficiency

in the field of conservation, it is necessary to recognize two discrete scientific

disciplines: “herpetology” for the study of reptiles and “batrachology” for that

of amphibians. The first steps in the direction of this distinction already taken

by some zoologists are outlined.

A general trend in science today is towards greater specialization. This has become

more and more necessary as the amount of knowledge already accumulated and the

bibliographical corpus have been growing, as techniques have been multiplying and

becoming increasingly sophisticated, and as it has become impossible for any single

scientist to master all problems, methods and results.

Whereas this trend is unavoidable, it also implies the risk that biologists might lose

sight of the unity of biology. In order to minimize this risk, it is important that they both

reach a high specialization in their own research work (a condition for efficiency of the

latter) and keep a certain encyclopaedism of the mind and of general knowledge (a

condition for being able to put their own research back in a more general context).

Zoology is one of the domains of biology where specialization of research and of

knowledge has been growing progressively from the beginning. The specialization

movement has developed in three major directions, which correspond to three major

approaches to zoology.

(1) The first approach may be qualified of “systematic”. Following the increasing

subdivision of the animal kingdom into distinct groups, each of these became the object

Source : MNHN, Paris

2 ALYTES 9 (1)

of a particular sector of zoology, such as malacology for molluses or entomology for

insects. Each of these disciplines is centered on the animal group studied, on the peculiar

characters separating it from related groups, on its internal diversity and variability, and

on its history and evolution.

(2) In the second approach, the accent is not put on the animal groups themselves or

on their peculiarities, but rather on the study of general structures, patterns and processes

irrespective of the groups where they occur. Disciplines such as embryology (study of

development) or ethology (study of behaviour), to take only two examples, tend to use the

animals as a “material”, devoid, in itself, of any particular interest, but allowing the

discovery of (more or less) general laws. These disciplines are not alternative to those of

the first kind, but rather orfhogonal to them, since a behavioural study of a bird, for

example, both belongs to the field of ethology and to that of ornithology.

G) Finally, the important multiplication of zoological methodologies and the growing

sophistication of many techniques has made necessary the development of “disciplines”

such as histology or karyology, which do not correspond in fact to conceptual fields, but

to technical ones. Although for practical reasons of information inflation these “disci-

plines” have their own specialists, teachings, journals, they lack specific problems and are

only at the service of the disciplines of the first two categories.

Whereas disciplines of the second kind described above tend to be dominant in current

zoology, it should be clear that those of the first one are to be firmly defended. They allow

insights into the evolutionary biology of organisms and prevent zoology from becoming

purely reductionist. This need is recognized by many zoologists. However, even acknowl-

edgement of the need leaves the question of how many different ‘‘systematic” zoological

disciplines should be distinguished. In this respect, there is the question rarely addressed

until now, whether there should be a single zoological discipline, “herpetology”, for both

reptiles and amphibians, or two disciplines, “herpetology” for reptiles and “batrachology”

for amphibians. In this paper 1 will address the justification of the latter distinction.

AMPHIBIANS AND REPTILES

Amphibians and reptiles have not always been recognized as two distinct groups of

the same hierarchical rank (class) within the taxinomic system.

LINNÉ (1758) did not recognize these two groups as classes. He recognized a class

Amphibia, with three orders, each of which contained several genera: Reptiles (Testudo,

Draco, Lacerta and Rana), Serpentes (Crotalus, Boa, Coluber, Anguis, Amphisbaena and

Caecilia) and Nantes (Petromyzon, Raja, Squalus, Chimaera, Lophius and Acipenser). In

the light of our current knowledge, these orders and genera are highly heterogeneous, and

the Amphibia of LINNÉ are an assemblage of reptiles, amphibians and of various groups

of “fishes”. Later, LINNÉ (1767) recognized a fourth order of Amphibia, Meantes, for the

single species Siren lacertina (see Dugois, 1991).

LAURENTI (1768) recognized a class Reptilium, with three orders: Salientia (Pipa,

Source : MNHN, Paris

Dusois 3

Bufo, Rana, Hyla and part of Proteus!), Gradientia (part of Proteus!, Triton, Salamandra,

Caudiverbera, Gekko, Chamaeleo, Iguana, Basiliscus, Draco, Cordylus, Crocodylus, Scincus,

Stellio and Seps) and Serpentia (Chalcides, Caecilia, Amphisbaena, Anguis, Natrix,

Cerastes, Coronella, Boa, Dipsas, Naja, Caudisona, Coluber, Vipera, Cobra, Aspis, Con-

strictor and Laticauda). As can be seen from this list of genera, while his Salientia

contained only amphibians, both his Gradientia and his Serpentia were a mixture of am-

phibians and reptiles.

It is only in 1799 when Alexandre BRONGNIART (in a paper first published, in two

parts, in 1800; see Dupois, 1984: 10) presented to the French Academy of Sciences the first

classification separating, at the ordinal level, the amphibians (under the name Batraciens)

from the other “reptiles” (maintained in three distinct orders: Chéloniens, Sauriens and

Ophidiens). BRONGNIART's Batraciens contained both the anurans and the urodelans, but

not the gymnophiones (then all classified under the generic name Caecilia), which this

author considered to be insufficiently known to be properly given a place in his system.

DumékiL (1808: 317) was the first author to point to the high similarity of Caecilia

with the batrachians, and his student OPPEL (1810: 409) was the first to clearly place this

genus in the order Batracii, which he still considered part of the class Reptilia.

Another student of DUMÉRIL, [DUCROTAY] DE BLAINVILLE (1816 a-b, 1818), was the

first author to give the amphibians (including Caecilia) and the reptiles the same rank in

classification, since he considered these groups to be either two subclasses (Ornithoïdes and

Ictyoïdes) of the class Reptiles, or two independent classes (Squammifères and Nudipel-

lifères). He did not choose between these two possibilities in his 1816 a-b and 1818 works,

but he did so in his 1822 book, where he was the first author to clearly recognize a class

Reptiles and a class Amphibiens. MERREM (1820) also recognized these two classes, but

under the names Pholidota and Batrachia (Amphibia s. str.). GRAY (1825), in a famous

paper, also distinguished a class Reptilia and a class Amphibia (including Caecilia). Since

then, except for a few exceptions in the early years, all authors have adopted this

classification, which is still in use today, at least for living forms (some difficulties arise for

assigning some fossil forms, but these need not be discussed here).

It is therefore more than 150 years since the amphibians and reptiles were last

considered a single group of vertebrates. On the other hand, during the same period, the

study of both kinds of animals has been referred to a single discipline of zoology, that of

herpetology, and only recently has this been challenged. It is therefore justified to look for

the reasons of this long period of stability.

L. LAURENTT (1768) work provides an extraordinary example of non truly hierarchical classification, which

has been overlooked by most authors until now: his genus Proteus straddles his two orders Salientia and

Gradientia, with one species (P. raninus) in the former, and two (P. tritonius, P. anguinus) in the latter. The

nomenclatural status of the generic name Proteus was only fixed by STENEGERS (1936) subsequent

designation of P. anguinus as its type species, a fact which was ignorcd in Frosr's (1985) checklist (see

Dusois, 1987 b).

Source : MNHN, Paris

4 ALYTES 9 (1)

THE HISTORICAL REASONS FOR THE LONG MAINTENANCE

OF HERPETOLOGY AS A SINGLE DISCIPLINE

In my opinion, the primary reason for the long maintenance of herpetology as a single

discipline is to be found in both the relatively low number of species of amphibians and

reptiles, and in the low level of interest long given, even by scientists themselves, to these

animals. Furthermore, amphibians and reptiles are superficially very similar, which

explains that they were first placed in a single group, and once a historical tradition has

been well entrenched it is difficult to change.

In 1985, the number of recognized species was fewer than 3500 amphibians and 6000

reptiles (Dugois, 1988: 62), thus less than birds (more than 9000) and “fishes”’ (more than

21000), and only comparable to the number of known species of mammals (more than

4000). Besides, most species of amphibians and reptiles have little or no direct economic

interest or value, in contrast to fishes, birds and mammals. Amphibians and reptiles have

long been considered “inferior” vertebrates, of much less interest than mammals, which

are “closer to man”, “more evolved” and more useful for medical and other applied

reasons.

As a result, the number of scientists studying amphibians and reptiles has long been

very low, with few positions available in academic or private institutions for their study.

In many museums, a single person is in charge of the collections of both groups (and

sometimes also of others) and has to deal with scientific problems pertaining to both.

As a consequence of the few specialists working on amphibians and reptiles, the

number of scientific publications dealing with these animals has remained relatively low as

well, and the knowledge of these groups has lingered far behind that of the other groups

of vertebrates. Furthermore, the (relatively) few specialists on these animals have tended

to group themselves in learned societies devoted to the study of both groups, and which

have started, one after another, the publication of scientific journals also dealing with both

groups (and sometimes also of others). Whereas the groups amphibians and reptiles have

clearly been considered distinct since the beginning of the XIXth century, the discipline

“herpetology”, dealing with both amphibians and reptiles, has become established as a

“natural” discipline of zoology.

SOME TERMINOLOGICAL PROBLEMS

The name herpetology was coined by KLEIN (1755), under the form herpetologia, for

the branch of zoology dedicated to the study of snakes and serpentiform animals. This

term is derived from the Greek épretév, meaning, according to the authors, either reptile

or snake. It was used again by BONNATERRE (1789), under the form erpétologie, for the

study of ‘“snakes and reptiles” (the latter including the amphibians), by RAFINESQUE[-

SCHMALTZ] (1814 a: 47, 1815: 39, erpétologie; 1814 b-c-d, erpetologia) and a few other

authors of this period, but it became widely used only with the publication of DUMÉRIL &

Source : MNHN, Paris

DuBois s)

BiBRON [& DumÉRiLJ's Erpétologie Générale (1834-1854). The spelling starting with an h,

which had first been introduced by KLEIN (1755) but had then been abandoned, was

reintroduced by English-speaking authors such as HOLBROOK (1842), and has since then

won general acceptance.

Although most authors until now have referred the study of both amphibians and

reptiles to a single discipline “herpetology”, there were some exceptions, even as far back

as more than one century ago. For example, STEINDACHNER clearly distinguished between

herpetological (STEINDACHNER, 1867, 1870, 1891, 1895, 1901 a-b, 1907) and barrachological

studies (STEINDACHNER, 1864 a-b, 1882). The term batrachology is derived from the name

Batrachia, a latinization (first used by Ross & MACARTNEY, 1802: tabl. III) of the French

name Batraciens, created by BRONGNIART (1800 a: 82) for the order including all recent

amphibians. BRONGNIART (1800 a) may have based his new name either on the Latin name

Batrachi, proposed by BaTscH (1788: 437) for a taxon of the rank family, or directly on

the Greek name GBérpxyos, meaning frog, which is also the root of BATSCH’s name (see

Dusois, 1984).

Another term, that of amphibiology, has been used incidentally by a few authors, both

in the XIXth century (e.g. RAFINESQUE, 1840; BONAPARTE, 1845, 1850, 1852 a-b) and in the

XXth century (e.g. TAyLOR, 1947). This name is based on the name Amphibia, derived

from the Greek name äupi6toc (“which lives in two elements, on the ground and in

water”). The name Amphibia was the name given by LINNÉ to one of his classes of

vertebrates, and which has been retained by the subsequent authors for the class

containing the anurans, urodelans and gymnophiones.

Dusois (1984) proposed to stabilize the use of the names Amphibia and Batrachia, the

first one being retained for the class as a whole, and the second one for the subclass

including all living amphibians. For the latter, the name Lissamphibia (created by

HAECKEL, 1866) has also been used by a few authors (following ROMER, 1966), but the

name Batrachia has been used much more and is universally known, so that it should be

retained (see Dugois, 1984). If this is done, the use of the term batrachology is warranted

for all studies dealing with all amphibians belonging to the three living orders of the class

Amphibia (not only anurans). The name amphibiology, on the other hand, would be the

correct term for the field covering the study of all amphibians, including all fossil groups.

The creation of the journal Alytes in Paris in 1982 led to a wider usage of the terms

batrachology, batrachologist and batrachological. This runs counter to the direction

described above. Let us then examine the reasons for such a change.

THE NEED OF BATRACHOLOGY

The major reason for recognizing batrachology as a distinct discipline is not a simple

demand for intellectual rigour and elegance, but the need of a better efficiency in research

and conservation. Specialization permits a better communication among scientists

studying these animals and among those interested in their conservation and in that of

their environment.

Source : MNHN, Paris

6 ALYTES 9 (1)

Table I. — Number of papers dealing with the five major groups of Vertebrates listed in

the Zoological Record from 1953 to 1987.

Year Pisces Amphibia Reptilia Aves Mammalia

1953-57 5775 2717 3087 10903 8595

1958-62 8752 4719 4317 11309 11474

1963-67 10561 6493 5838 18728 25859

1968-72 22214 6720 6986 22514 30681

1973-77 24567 5092 8540 35338 27883

1978-82 32112 6110 12147 42542 36570

1983-87 38673 7397 12440 49314 46390

A first remark of some relevance is that the number of papers dealing with

amphibians is much lower than those for the other four major groups of vertebrates (see

Table I and fig. 1). However, relative progression of scientific research dealing with

amphibians has been very strong in the recent years, similar to those for the other

vertebrates (see Table II and fig. 2). The current increase in the number of species of

amphibians described yearly in the world is still very strong: its curve is still far from

reaching the plateau characteristic of such sigmoidal curves (STEYSKAL, 1965), which

means that there remains, in amphibians, a very high number of species to describe and

of systematic problems to solve (Dugois, 1977 b). The same is true in all other domains

of the biology and evolution of these animals, where our knowledge is still very

incomplete. Therefore amphibians, which have long been considered of little interest and

importance, are attracting more and more attention. As such, their study should be

highlighted by the recognition of a specific discipline devoted to it.

That living amphibians are monophyletic now seems to be admitted by all authors

(RAGE & JANVIER, 1982; MILNER, 1988). This group is relatively small and homogeneous,

which makes it possible for a single researcher or team to have a rather complete and general

approach of the knowledge accumulated on these animals in any given field of research (Du-

BOIS, 1977 b). Actually, comparative studies in various fields of research (morphology, ana-

tomy, phylogeny, genetics, biochemistry, physiology, ethology, ecology, etc.), are much

more effective, meaningful and instructive within a homogeneous group. But besides this set

of reasons, which also apply to other “systematic” disciplines of biology as defined above,

the recognition of batrachology as a discrete discipline is also strongly supported by other

arguments, which rely on the unique properties of amphibians among vertebrates.

Amphibians share indeed many strange peculiarities lacking in other vertebrates and

which make this group very special. Let us just stress some of them.

Amphibians, as their name indicates, usually go through a double life, with a larval

stage separated from a post-larval stage by a metamorphosis. This has made the

amphibians crucial for the study of development, which is true from the beginnings of

embryology as a scientific discipline. From the XVIIIth century, embryologists have

studied amphibian embryonal and larval development, and noted their peculiarities when

contrasted with other vertebrates. Physiologists have long been interested in amphibian

Source : MNHN, Paris

DuBois 7

Table II. — Relative increase in the number of papers dealing with the five major groups

of Vertebrates listed in the Zoological Record from 1953 to 1987 (measured as the

ratio to the number of papers listed for the period 1953-57, the latter being given the

value 100). (Based on the data in Table I).

Year Pisces Amphibia Reptilia Aves Mammalia

1958-62 ETES 170 140 104 133

1963-67 183 234 189 172 301

1968-72 385 242 226 206 357

1973-77 425 183 277 324 324

1978-82 556 220 393 390 425

1983-87 670 266 403 452 540

metamorphosis, which has no counterpart in other tetrapods, and in its anomalies

(neoteny, paedogenesis, etc.).

Particularly relevant to the present discussion is the observation that, of the very large

number of papers dealing with amphibian development and metamorphosis, and with va-

rious kinds of aneuchrony (Dugois, 1987 a) in these animals, published in this century, only

a very few appeared in “herpetological” journals. The reason is simple to understand: for

developmental biologists, there is little relevance in a “discipline” associating the amphi-

bians with the reptiles. If embryologists were to associate the reptiles with another animal

group, no doubt it would be the birds, and, if amphibians had by all means to be grouped

with other animals, the least bad choice would be “fishes”. This is certainly one of the rea-

sons, although not the only one, why embryologists and physiologists have tended to publish

their works on amphibian development in journals of general zoology, of experimental z00-

logy or of developmental biology, rather than in “herpetological” ones (another reason for

this is clearly the fact that developmental biologists tend to believe that they are studying

general phenomena that cross major taxa). This tendency has been disastrous with respect to

the synthetic appraisal of amphibian biology as a whole: most “herpetologists” who have

dealt with the general features of amphibian biology have ignored, or grossly under-

exploited, the literature dealing with the biology of development of these animals.

It is very striking for example to realize that the classic and historical papers of

embryologists of amphibians as important as F. BALTZER, E. BATAILLON, L. BOUNOURE,

J. BRACHET, R. BRIGGS, A. DALCQ, G. FANKHAUSER, M. FISCHBERG, L. GALLIEN, E.

Hapor\, G. & O. HERTWIG, J. HOLTFRETER, R. R. HUMPHREY, T. J. KING, J. ROSTAND,

W. Roux, H. SPEMANN or E. WITsCHI do not appear in the bibliography of an important

recent treatise on the biology of amphibians. Similarly, it cannot be denied that submission

of papers dealing with the developmental biology of amphibians to traditional “herpeto-

logical” journals often leads to very feeble reviews, including irrelevant comments or

suggestions. Many of the editors of such journals have few contacts with the world of

developmental biology and little knowledge in this discipline. Developmental biology of

amphibians is not only absent, or almost so, from “herpetological” journals, but also from

“herpetological” meetings, congresses and courses.

Source : MNHN, Paris

8 ALYTES 9 (1)

50000

Pisces

Amphibia —

Reptilia —

Aves -

Mammalia —..—

40000

30000 +

20000 -|

10000 -|

0 Year_,

TT T T FLE EE U

1953-57 58-62 63-67 68-72 73-77 78-82 83-87

Fig. 1. — Number (N) of papers dealing with the five major groups of Vertebrates listed in the

Zoological Record from 1953 to 1987. (From the data in Table I).

Source : MNHN, Paris

Dusois 9

Pisces

Amphibia ————

Reptilia

Aves

600! Mammalia

500

400+ où

300+

2004

100. —- T —- T —er,

do53Ës7 OSp62 ©6367 GH72 711 182 8387

Fig. 2. — Relative increase (RI) in the number of papers dealing with the five major groups of

Vertebrates listed in the Zoological Record from 1953 to 1987 (measured as the ratio to the

number of papers listed for the period 1953-57, the latter being given the value 100). (From the

data in Table II).

Source : MNHN, Paris

10 ALYTES 9 (1)

If the biology of amphibians is to become a unified field, this situation must change

drastically. The association, in a single journal, of papers dealing with the developmental

biology of amphibians and of the other aspects of the biology of these animals, would

certainly be extremely profitable to all amphibian biologists, whatever their specialization.

The existence of a larval stage in amphibians is not only interesting for embryologists.

Amphibian larvae have long been ignored by biologists, both as animals in themselves and

as parts of aquatic ecosystems. Recently however, more and more attention has been given

to the fact that tadpoles are full organisms that have their own morphology, anatomy,

physiology and behaviour, and that the two stages vital cycle of amphibians poses peculiar

problems to the study of their ecology. Amphibian larvae represent an important fraction

of the total animal biomass in some aquatic ecosystems, and play a significant rôle in the

filtration of these waters, possibly comparable to that of earthworms in the ground. Until

now, such factors have been underestimated by limnologists and other ecologists, possibly

because tadpoles, being “only” larvae, tend rarely to be considered as significant

components of ecosystems: in temperate countries (where many biologists live), tadpoles

of a number of species only occur seasonally in aquatic ecosystems.

The peculiarities of the life cycle and of the ecology of amphibians entail other

consequences, especially as concerns the conservation of the populations and species of

these animals. Because most species of amphibians breed in fresh water, the disappearance

of lakes, ponds, marshes, torrents, streams, or their modification (through pollution or

through other changes, usually caused by man: see e.g. DuBois, 1980), is usually a major

threat to amphibians. Some of these facts have not been given the proper attention

previously. For example, whereas the effects of the pollution of continental waters by acid

rain and other sources are being more and more studied, little interest has been devoted

to the effect of the recent important increase of the introduction of fishes into closed

continental waters, particularly of salmonids into mountain lakes, on amphibian

populations and species. Amphibians may also be threatened by other factors (collecting

for consumption of frog legs, for teaching and research; modification and pollution of

terrestrial, not only aquatic, ecosystems; introduction of alien species into ecosystems, and

even displacement of amphibians, leading to genetic or faunistic pollution) (see e.g.:

Dusois, 1976, 1977 a, 1983a-b, 1985; Dupois & MorÈRE, 1980). Altogether, the worldwide

situation of amphibians is extremely worrying. It is more than time for specialists of these

animals to assemble the information they have accumulated, and to work together to try

and find solutions to the acute problems amphibians face, to propose actions and measures

to governments and other organizations and agencies, and to see that these are carried out.

Consequently, recognition of batrachology as a discrete discipline, besides its purely

scientific reasons, has the most important justification of greatly increasing the efficiency

of their conservation. This aim is certainly worthwhile, indeed critical, if some amphibians

are to survive into the next century.

Source : MNHN, Paris

Dugois Il

FIRST STEPS TOWARDS THE RECOGNITION OF BATRACHOLOGY

The idea of the recognition of batrachology has been “in the air” for a long time, and

1 can remember having heard or supported it in conversations with colleagues more than .

20 years ago. However, the first official act was the creation (in February 1982 at the Paris

Museum) of Alytes as the first journal in the world to be specifically devoted to the biology

of amphibians. This was followed in November 1982 by the foundation of the Société

Batrachologique de France (S.B.F.), which accepted the task of publishing A/ytes, and

later, in November 1988, by the foundation of the International Society for the Study and

Conservation of Amphibians (ISSCA), to which the journal was again transferred.

The statutory aims of ISSCA are “to contribute on a world scale:

— to the study and knowledge of amphibians (batrachology);

— to the conservation of amphibians and to that of their environment;

— to the establishment, reinforcement and facilitation of bonds among batracholo-

gists;

— to the promotion of batrachology as an independent scientific discipline.” (Article

2.1 of the Statutes of ISSCA).

However the recognition of batrachology as a distinct scientific discipline cannot be

the work of a single society and a single journal. It will become a fact only if this idea is

supported by multiple specialists of amphibians, as well as of specialists of reptiles. The use

of the terms “batrachology” and “batrachologist” in the scientific literature, and,

complementarily, that of the terms “herpetology” and “herpetologist” in a restrictive

sense, referring only to the study of reptiles, will help achieve this recognition.

Organization of meetings dealing only with batrachological or herpetological topics will

also play a positive rôle in this direction.

Many biologists still are working on both groups of animals and, in all events,

specialists on these two groups will continue to collaborate for many kinds of studies

(particularly in the fields of ecology and faunistics). However, recognition of two distinct

fields of research is likely to help rather than hinder this collaboration. It is also clear that

the collaboration is likely to be fruitful between batrachologists and specialists of still other

groups, in particular ichthyologists, limnologists, or all other biologists working on animal

groups in which larvae and metamorphoses do occur: possibly, in the future, a new

biological discipline, “larvology”, could slowly emerge. But this is another story.

RÉSUMÉ

Bien que les Amphibiens et les Reptiles soient considérés comme deux classes

distinctes de Vertébrés depuis plus de 150 ans, l'étude des deux groupes est encore

traditionnellement rapportée à une seule discipline scientifique, l’“herpétologie”. Il est ici

suggéré qu'il est désormais nécessaire, à la fois pour des raisons scientifiques et pour

Source : MNHN, Paris

12 ALYTES 9 (1)

pouvoir avoir une meilleure efficacité dans le domaine de la conservation, de reconnaître

deux disciplines scientifiques distinctes: l'“‘herpétologie” pour l'étude des Reptiles et la

“batrachologie” pour celle des Amphibiens. Les premiers pas déjà effectués dans cette

direction par certains zoologistes sont soulignés.

ACKNOWLEDGEMENTS

For their comments on previous versions of this paper, I am extremely thankful to Carl Gans,

Tim HALLIDAY, Raymond F. LAURENT, Hobart M. SMirH, Richard WASsSERSUG and an anonymous

reviewer. However, their agreement with parts or all of the ideas presented above is not implied.

T'also want to express my most sincere thanks to Anne-Marie VACHOT for collecting the data in

Table I, and to Dominique PAYEN for drawing the figures.

LITERATURE CITED

Batscu, À. J. G. C., 1788. — Bersuch einer Anleitung, zur KenniniB und Geschichte der Thiere und

Mineralien. Erster Theil. Jena, Akademischen Buchhandlung: + 1-528, pl. IV

BONAPARTE, C. L., 1845. — Specchio generale dei sistemi erpetologico ed anfibiologico. /n: Atti della

sesta riunione degli scienzati italiani, tenuta in Milano in Settembre del MDCCCXLIV. Milano,

Luigi di Giacomo Pirola: 376-378.

1850. — Conspectus systematum herpetologiae et amphibiologiae. Lugduni Batavorum, Brill: 1 pl.

1852 a. — Conspectus herpetologiac et amphibiologiae. Nuovi Ann. Sci. nat. Bologna, (3), 5: 89-96.

1852 b. — Conspectus herpetologiae et amphibiologiac. (Continuazione). Nuovi Ann. Sci. nat.,

Bologna, (3), 5: 477-480.

BONNATERRE, Abbé, 1789. — Tableau encyclopédique et méthodique des trois règnes de la nature.

Erpétologie. Paris, Panckoucke: i-xxviii + 1-70, pl. 1-7 + 1-6 + 1-12.

BRONGNIART, À. 1800 a. — Essai d'une classification naturelle des Reptiles. Première partie.

Etablissement des ordres. Bull. Sci. Soc. philom., 2: 81-82.

che 1800 b. — Essai d'une classification naturelle des Reptiles. Deuxième partie. Formation et

disposition des genres. Bull. Sci. Soc. philom., 2: 89-90.

DE BLAINVILLE, H., 1816 a. — Prodrome d’une nouvelle distribution systématique du règne animal.

Bull. Sci., Soc. philom. Paris, 1816: [113]-124.

E 1816 b. — Prodrome d'une nouvelle distribution systématique du rêgne animal. J. Phys. Chim.

His. nat. Arts, 83: 244-267.

_ 1818. — Vorläufige Anzcige einer neuen systematischen Eintheilung des Thierreichs. /sis von

Oken, 1818: 1365-1384.

Dusois, À., 1976. — A propos de la protection des Amphibiens et des Reptiles. Première partie: les

problèmes. Bull. Sect. paris. Soc. herp. Fr., 1: 13-22.

ne 1977 a. — A propos de la protection des Amphibiens et Reptiles. Deuxième partie: conclusions

et suggestions. Bull. Soc. herp. Fr., 1: 18-24.

en 1977 b. — Les problèmes de l'espèce chez les Amphibiens Anoures. Mém. Soc. zool. France, 39:

161-284.

-— 1980. — L'influence de l'homme sur la répartition des Amphibiens dans l'Himalaya central et

occidental. C. R. Soc. Biogéogr., 35: 155-178.

1983 a. — A propos de cuisses de Grenouille. Protection des Amphibiens, arrêtés ministériels,

projets d'élevage, gestion des populations naturelles, enquêtes de répartition, production,

importations et consommation: une équation difficile à résoudre. Les propositions de la Société

Batrachologique de France. Alpes, 2: 69-111.

1983 b. — Renforcements de populations et pollution génétique. C. r. Soc. Biogéogr., 59: 285-294.

1984. — La nomenclature supragénérique des Amphibiens Anoures. Mém. Mus. natn. Hist. nat.

(A), 131: 1-64.

Source : MNHN, Paris

DuBois 13

- 1985. — A nouveau sur les cuisses de Grenouilles. Circalytes, 1 (8): 1-21.

- 1987 a. — Necoteny and associated terms. A/ytes, 4: 122-130.

- 1987 b. — Living amphibians of the world: a first step towards a comprehensive checklist. A/ytes,

5: 99-149.

-— 1988. — The genus in zoology: a contribution to the theory of evolutionary systematics. Mém.

Mus. natn. Hist. nat., (A), 140: 1-123.

rl 1991. — Miscellanea nomenclatorica batrachologica (XVIII). Alytes, 8: 107-120.

Dusois, À. & More, J.-J., 1980. — Pollution génétique et pollution culturelle. C. r. Soc. Biogéogr..

56: 5-22.

Ducroray DE BLAINVILLE, H. M., 1822. — De l'organisation des animaux, ou Principes d'anatomie

comparée. Tome premier, contenant la Morphologie et l'Aistésologie. Paris, Levrault: [i-üii] +

i-lix + 1-574, 10 tabl. h. 1.

DumériL, A.-M.-C. & BiBRON, G., 1834. — Erpétologie générale ou histoire naturelle complète des

Reptiles. Tome 1. Paris, Roret: i-xxiv + 1-447.

me 1835. — Erpétologie générale ou histoire naturelle complète des Reptiles. Tome 2. Paris, Roret: i-iv

+ 1-680.

1836. — Erpétologie générale ou histoire naturelle complète des Reptiles. Tome 3. Paris, Roret: i-iv

+ 1-518.

ue 1837. — Erpétologie générale ou histoire naturelle complète des Reptiles. Tome 4. Paris, Roret:

+ 1-572.

1839. — Erpétologie générale ou histoire naturelle complète des Reptiles. Tome 5. Paris, Roret:

i-vii + 1-855.

— 1841. — Erpétologie générale ou histoire naturelle complète des Reptiles. Tome 8. Paris, Roret: i-vii

+ 1-792.

_——— 1844. — Erpétologie générale ou histoire naturelle complète des Reptiles. Tome 6. Paris, Roret: i-xii

+ 1-610.

DumériL, A.-M.-C., BIBRON, G. & DUMÉRIL, A., 1854 a. — Erpétologie générale ou histoire naturelle

complète des Reptiles. Tome 7 (1). Paris, Roret: i-vii + [i-iv] + i-xvi + 1-780.

—— 1854 b. — Erpétologie générale ou histoire naturelle complète des Reptiles. Tome 7 (2). Paris,

+ 781-1536.

— 1854 c. — Erpétologie générale ou histoire naturelle complète des Reptiles. Tome 9. Paris, Roret:

i-xx + 1-440.

Dumériz, C., 1808. — Mémoire sur la division des Reptiles Batraciens en deux familles naturelles.

Mag. encycl., J. Sci. Lett. Arts, 1808 (2): 308-329.

FROST, D. R. (ed.), 1985. — Amphibian species of the world. Lawrence, Allen Press & Assoc. Syst.

Coll: [iv] + iv + 1-732.

Gray, J. E., 1825. — A synopsis of the genera of Reptiles and Amphibia, with a description of some

new species. Ann. Philos., (2), 10: 193-217.

HaëckeL, E., 1866. — Gencrelle Morphologie der Organismen. Zweïter Band. Allgemeine Entwicke-

lungsgeschichte der Organismen. Berlin, Georg Kramer: i-clx + 1-462, pl. I-VIIL.

HOLBROOK, J. E., 1842. — North American herpetology; or, a description of the reptiles inhabiting the

United States. Philadelphia, J. Dobson: vol. 1: i-xv + 17-152, pl. 1-24; vol. 2: i-vi + 9-142, pl.

1-20; vol. 3: i-ii + 3-128, pl. 1-30; vol. 4: + 7-138, pl. 1-35; vol. 5: ivi + 5-118, pl. 1-38.

KL, J. TL. 1755. — Tentamen herpetologiae. Leidae & Gottingae, Eliam Luzac: i-iv + 1-72, pl. ILII.

LAURENT, J. N., 1768. — Specimen medicum, exhibens synopsin Reptilium emendatam cum

circa venena et antidota Reptilium austriacorum. Viennae, Joan. Thom. Nob. de

Tratinern: iii + 1-215, pl. I-V.

Linné, C. A, 1758. — Systema Naturae per regna tria naturae, secundum classes, ordines, genera,

species, cum characteribus, differentiüs, synonymis, locus. Editio decima, reformata. Tomus L.

Holmiac: i-iv + 1-824.

ema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum

characteribus, differentiüs, synonymis, locis. Editio duodecima, reformata. Tomus 1, Pars II.

Holmiae, Laurentii Salvii: 533-1327 + Li-xxxvii].

MERREM, B., 1820. — Versuch eines Systems der Amphibien. Marburg, Krieger: i-vii + [viïi-xv] x 2

+ [1-188] x 2 + 189-191, 1 pl.

MILNER, À. R., 1988, — The relationships and origin of living amphibians. /n: M. J. BENTON (ed.),

Source : MNHN, Paris

14 ALYTES 9 (1)

The phylogeny and classification of the tetrapods, vol. 1, Amphibians, reptiles, birds, Oxford,

Clarendon Press, The Systematics Association Special Volume 35A, i-x + 1-377: 59-102.

OPrel, M., 1810. — Second mémoire sur la classification des Reptiles. Ann. Mus. Hist. nat., 16:

394-418.

RAFINESQUE, C. S., 1815. — Analyse de la nature ou Tableau de l'univers et des corps organisés.

Palerme, Barravecchia: 1-224, 1 pl.

me 1840. — The good book. — Number I. Amenities of nature, or annals of historical and natural

sciences. Philadelphia, Printed for the Eleutherium of Knowledge: 1-84.

RAFINESQUE-SCHMALTZ, C. S., 1814 a. — Principes fondamentaux de somiologie ou les loix [sic] de la

nomenclature et de la cl ‘ation de l'empire organique ou des animaux et des végétaux,

contenant les règles essentielles de l'art de leur imposer des noms immuables et de les classer

méthodiquement. Palerme, Abate: 1-52.

ee 1814 b. — O quadro del metodo sinottico di somiologia. Specchio Sci., Giorn. encicl. Sic

11-15.

1814 c. — Prodrono [sic] di Erpetologia Siciliana. Specchio Sci., Giorn. encicl. Sicilia, 2: 65-67.

1814 d. — Fine del Prodromo d'Erpetologia Siciliana. Specchio Sci., Giorn. encicl. Sicilia, 2:

102-104.

RAGE, J.-C. & JANVIER, P., 1982. — Le problème de la monophylie des amphibiens actuels, à la

lumière des nouvelles données sur les affinités des tétrapodes. Géobios, Mém. spéc. 6: 65-83.

ROMER, A. S., 1966. — Vertebrate paleontology. Third edition. Chicago & London, The University

of Chicago Press: i-ix + 1-468.

Ross, W. & MACARTNEY, 1802. — Lectures on comparative anatomy. Translated

G. Cuvier. Vol. I. On the organs of motion. London, Longman & Re:

ia, 1:

from the French of

-x1 + 1-542, tables

I-IX.

STEINDACHNER, F., 1864 a. — Batrachologische Mittheilungen. Verhandl. zool.-bot. Ges. Wien, 14:

239-288, pl. IX-XVII

— 1864 b. — Zusätze und Berichtigungen zu den batrachologischen Mittheilungen. Verhandl.

sool.-bot. Ges. Wien, 14: 551-552.

- 1867. — Herpetologische Notizen. Sher. kaiserl. Akad. Wiss. Wien, 55: 263-273, pl. I-IV.

1870. — Herpetologische Notizen (I). Sber. kaiserl. Akad. Wiss. Wien, 62: 326-350, pl. I-VII.

1882. — Batrachologische Bciträge. Sber. kaiserl. Akad. Wiss. Wien, 85: 188-194, pl. I-III.

1891. — Ueber neue und seltene Lacertiden aus den herpetologischen Sammlungen des k. k.

näturhistorischen Hofmuseums. Ann. k. k. naturhistorischen Hofmuseums, 6: 371-378, pl.

XI-XIT.

2, 1895. — Bericht über Dr. Sturany's herpctologische Ausbeutc in der Umgebung der Plitvicer Scen

in Croatien. Ann. k. k. naturhistorischen Hofmuseums, 10, Notizen: 78-19

— 1901 a. — Expedition S. M. Schiff “Pola” in das Rothe Meer (nôrdliche und südliche Hälfte),

1895/96 und 1897/98. Zoologische Ergebnisse. XVII. Bericht über die herpetologischen

Aufsammlungen. Denkschrift. k. Akad. Wiss. Wien, Math.-Naturwiss. CI., 69: 325-335, pl. 1-IL.

es 1901 b. — Herpetologische und ichthyologische Ergebnisse ciner Reise nach Südamerika mit

einer Einlcitung von Therese Prinzessin von Baiern. Anziger k. Akad. Wiss. Wien, 18: 194-196.

ee 1907. — Herpetologische Notizen (II). Sber. kaiserl. Akad. Wiss. Wien, 116: 1-6, 1 pl.

SrENEGER, L., 1936. — Types of the Amphibian and Reptilian genera proposed by Laurenti in 1768.

Copeia, 1936: 133-141.

SrevskaL., G. C., 1965. — Trend curves of the rate of species description in zoology. Science, 149:

880-882.

Tayuor, E. H. 1947. — A bibliography of mexican amphibiology. Univ. Kansas Sci. Bull. 31:

543-580.

Corresponding editor: Pierre JOLY.

Source : MNHN, Paris

Alytes, 1991, 9 (1): 15-22. 15

Images d’Amphibiens camerounais.

IIL. Le comportement de garde des œufs

Jean-Louis AMIET

Université de Yaoundé,

Faculté des Sciences, Laboratoire de Zoologie,

B.P. 812, Yaoundé, Cameroun

The eggs are guarded by one of the parents in six genera of Cameroonian

Amurans. Seven photos illustrate this behaviour in the genera Petropedetes,

Alexteroon, Phrynodon, Nectophryne and Hemisus. In several, if not all, cases,

the hypothesis of a protection against predators does not seem to be

acceptable.

Sur 39 genres d’Anoures représentés au Cameroun, 9 comprennent des espèces qui

accordent des soins à leur descendance. Ceux-ci se ramènent à deux types de comporte-

ments parentaux : la surveillance des œufs et la construction de nids. Fait curieux, la

batrachofaune camerounaise — et plus généralement africaine — montre un “ vide

éthologique ” en ce qui concerne deux autres types de comportements plus ou moins

fréquents ailleurs: le transport des œufs et/ou des larves et les soins accordés aux têtards.

Les photos des figures 1 à 7 montrent des exemples de la garde des œufs. Elles

fourniront l’occasion d’une mise au point sur les 6 genres d'Anoures camerounais qui

pratiquent ce comportement (le cas des constructeurs de nid sera examiné dans un autre

article).

LE GENRE PETROPEDETES

La plupart des espèces de Perropedetes, comme l'indique leur nom générique, sont

rupicoles, au moins en période de reproduction. Les œufs sont déposés en plaques,

atteignant parfois une quinzaine de centimètres de plus grande dimension, sur des rochers

recouverts d’un très mince film d’eau, plus rarement sur un support végétal, base de tronc

d'arbre ou grande feuille. Les têtards, hygropétriques, broutent le revêtement de

micro-organismes tapissant les rochers.

Chez P. newtoni (Bocage, 1895), P. parkeri Amiet, 1973, et P. cameronensis Reichenow,

1874 j'ai pu constater que, /a nuit, il y a toujours, à proximité d’une plaque d'œufs, un mâle

qui se tient immobile et qui, apparemment, en assure la garde (fig. 1: P. newtoni).

Source : MNHN, Paris

16 ALYTES 9 (1)

Fig. 2. — Cette femelle d'Alexterodn obstetricans (AI, 1931) a été trouvée à proximité de sa ponte

et ramenée au laboratoire. En {errarium, elle est restée auprès de ses œufs, ou sur eux, jusqu'à

la sortie des têtards. Ototomo, I11-73.

Source : MNHN, Paris

AMIET

PARIS

Fig. 3. —

tade, les têtards, déjà

munis de bourgeons de pattes postérieures, s à se r. Kala, X-70.

Fig. 4. — Dans une lune grosse rac ace du sol, un mâle de Nectophryne

afra Buchholz & Pete appuyé sur ses membres antérieurs et aux deux tiers immergé,

brasse l'eau où a été déposée la ponte. L'orifice du gîte était masqué par un morceau de bois

tombé qui a

Source : MNHN, Paris

ALYTES 9 (1)

Un couple d'Hemisus guincensis Cope, 1865 vient de se former après une journée de pluie

dans les savanes d’Obala, au nord de Yaoundé. On remarquera que l’accouplement est lombaire,

ce qui doit faciliter les déplacements du couple dans le sol où sera creusée la chambre de ponte.

Obala, 3-I11-72

6. — Dans la paroi meuble d’un trou d'extraction de sable, un coup de pelle vient de mettre à

jour une femelle d'A. guincensis sur sa ponte. Minkama, 6-111-78

Fig. 7. — Oeufs d'Hemisus guineensis dans la chambre de ponte, après enlèvement de la femelle.

Minkama, 6-I11-78.

Source : MNHN, Paris

AMIET 19

Un mâle d’une autre espèce, P. johnstoni (Boulenger, 1887), dont les têtards sont

probablement terrestres, a été observé près d’une ponte dans la matinée mais je ne peux

pas dire si, chez les espèces précitées, les pontes sont surveillées aussi pendant la journée,

car l’approche de l’observateur peut déclencher la fuite du mâle avant même qu'il ait pu

être repéré.

Il est cependant probable que, chez les espèces rupicoles, les pontes ne soient gardées

que la nuit.

LE GENRE PHRYNODON

Jusqu'ici, ce genre ne comprend qu’une seule espèce nommée, P. sandersoni Parker,

1935, répandue dans la plaine littorale ainsi que sur le versant et le rebord ouest du Plateau

sud-camerounais, mais il existe aussi deux espèces orophiles non nommées sur les reliefs

de la Dorsale camerounaise. P. sandersoni n’est pas rare dans les collines de la région de

Yaoundé où il se localise dans les fonds de vallon en forêt de type primaire.

Dans un travail consacré à la biologie de cette espèce (AMIET, 1981), j'ai déjà relaté

comment la femelle monte chaque soir sur sa ponte, déposée sur une feuille à quelques

dizaines de centimètres (parfois jusqu’à 2 m) au-dessus de zones mouilleuses, et la “couve”

jusqu’au lendemain matin.

Ce comportement se manifeste pendant toute la période de développement des œufs,

soit une douzaine de jours. Les œufs, au nombre de 12 à 17, mesurent 2,3 mm de diamètre

et sont disposés sur une seule couche. La gangue gélatineuse qui entoure chaque œuf est

d’abord peu épaisse mais elle va gonfler progressivement, en même temps que s’élargit la

cavité occupée par le jeune têtard. De ce fait, la ponte finit par ressembler à une masse

mamelonnée atteignant 3 cm de plus grande dimension. L’attitude de la femelle est alors

assez comique car, même en écartant largement ses membres, elle n’arrive plus à recouvrir

complètement son encombrante progéniture (fig. 3).

Une femelle ne paraît pas capable de reconnaître sa propre ponte : elle peut, en

captivité, accepter une autre ponte que la sienne, même si elle n’est pas au même stade de

développement. En revanche, si on présente une ponte à une femelle non ‘“‘incubante”, elle

s’en désintéresse.

LE GENRE ALEXTEROON

Chez 4. obstetricans (Ah, 1931), espèce placée auparavant dans le genre Hyperolius,

les œufs sont pondus, au nombre d’une cinquantaine, sur une feuille surplombant le cours

d’une petite rivière, à au moins 1,5 m de la surface de l'eau (AMIET, 1974).

Il est assez paradoxal que chez cette espèce peu commune, difficile à capturer, le

Source : MNHN, Paris

20 ALYTES 9 (1)

comportement de garde des œufs soit connu depuis près de 90 ans. BRANDES &

SCHOENICHEN ont en effet publié en 1901 une photo (reproduite dans AHL, 1931) qui

représente une femelle surveillant ses œufs déposés sur une feuille.

J'ai pu refaire cette observation et constater que la femelle se tient pendant la journée

à proximité de sa ponte, sur laquelle elle se place durant la nuit. Je l’ai vue “ratisser” à

l'aide de ses membres postérieurs la surface de la ponte: il est possible, mais non certain,

que ce comportement ait pour effet d’aider les têtards à sortir de la gelée qui les entoure!.

LE GENRE NECTOPHRYNE

La ponte et le développement de N. afra Buchholz & Peters, 1875 ont été décrits par

SCHEEL (1970). J'ai pu refaire les mêmes observations sur cette espèce ainsi que sur N.

batesi Boulenger, 1913 et découvrir de plus quels sites de ponte elles utilisent dans la

nature.

Les Nectophryne pondent leurs œufs, réunis en cordon comme chez les autres

Bufonidae mais entièrement blancs et assez volumineux, dans des petites cavités d’arbre

contenant de l’eau. Ces cavités n’occupent pas toujours une situation élevée : celle qui a

permis de réaliser la photo de la figure 4 se trouvait dans une racine affleurant à la surface

du sol. Il est possible que d’autres “aquariums naturels” soient utilisés, en particulier les

bases de pétiole de Macabo, où peut s’accumuler une quantité d’eau suffisante pour le

développement des minuscules têtards de cette espèce, mais j'ai prospecté en vain ce type

de biotope. En revanche, on m'a apporté une ponte trouvée dans une boîte de conserve

abandonnée sur le sol où s'était accumulé un peu d’eau!

La surveillance des œufs est assurée par le mâle. Comme l’a montré SCHEEL (1970),

la garde est permanente et, de plus, le mâle, en s'appuyant sur ses mains aux parois de la

cavité, “pédale” dans l’eau avec ses pattes postérieures, ce qui a probablement pour effet

d’aérer la ponte.

LE GENRE TRICHOBATRACHUS

PERRET (1966) rapporte qu'il a trouvé un mâle de grenouille poilue, T. robustus

Boulenger, 1900, “immergé à environ 60 cm de profondeur sous un fort courant d’eau dans

un trou rocheux alors qu'il semblait protéger une masse d'œufs avec des embryons déjà

bien avancés”.

1. Le nom donné par Au (1931) à cette espèce doit s'inspirer d'une observation similaire rapportée par

Branpts & SCHOENICHEN (1901), dont je n'ai pas eu l'article en mains et que je cie ici d'après PERRET (1988).

e d'autre part que le nom générique Alexteroon, dont il est l'auteur, est dérivé du grec

” = œuf. Peu d'espèces doivent avoir un nom générique et un nom spécifique

Source : MNHN, Paris

AMIET 21

LE GENRE HEMISUS

Remarquablement adapté à la vie fouisseuse, ce genre est représenté au Cameroun par

deux espèces savanicoles, H. marmoratus sudanensis Steindachner, 1863 et H. guineensis

Cope, 1865. Grâce à WAGER (1965), on sait que les Hemisus pondent leurs œufs dans une

chambre creusée dans la berge d’une mare et que la femelle reste sur sa ponte. Les têtards,

après l’éclosion, rejoignent le milieu aquatique grâce à une galerie de sortie forée par leur

mère.

J'ai eu la chance de pouvoir observer une femelle d’H. guineensis juchée sur ses œufs,

dans une cavité de la grosseur du poing à peu près, creusée dans la paroi d’un trou d’extrac-

tion de sable. Les clichés des figures 5 à 7 montrent cette femelle photographiée juste après

que la chambre de ponte eût été éventrée (fig. 6), ainsi que la masse d'œufs sur laquelle elle

était installée (fig. 7).

Il est difficile de se représenter comment la chambre de ponte est creusée car la femelle,

nécessairement, n’y arrive pas seule mais portant un mâle solidement agrippé à son arrière-

train (l’accouplement se fait en effet à la surface du sol, comme le montre la photo 5). D'au-

tres points restent à élucider : le mâle, après l’accouplement, quitte-t-il la chambre de ponte

par la galerie empruntée auparavant par le couple ou en fore-t-il une autre? La voie suivie par

les têtards n'est-elle pas la galerie de sortie du mâle plutôt qu'une galerie creusée par la

femelle?

DISCUSSION

Dans ce qui précède, les termes de “garde” ou de “surveillance” des œufs ont été

utilisés faute de mieux. Ils ont l'inconvénient de laisser supposer que le parent concerné

puisse exercer une protection de la ponte contre d'éventuels prédateurs ou agresseurs. Cette

fonction, en fait, est très improbable pour les raisons suivantes.

— Bien que DUELLMAN & TRUEB (1986) mentionnent divers animaux susceptibles de se

nourrir de pontes d’Anoures (Opilions, Araignées, Crabes et Serpents) je n'ai pu déceler jus-

qu'ici de cas semblables au Cameroun, où j'ai pourtant effectué près de 1 200 sorties de nuit? .

— Même si de tels prédateurs existent, on ne voit pas comment de petits Batraciens,

eux-mêmes sans défense, pourraient protéger efficacement leur ponte contre un agresseur.

Chez Phrynodon, et probablement Petropedetes, les pontes restent d’ailleurs abandonnées à

elles-mêmes pendant la journée.

— Beaucoup d’autres Anoures de la faune camerounaise ont des pontes aériennes très

apparentes (“pontes suspendues”) auxquelles ils n’assurent aucune protection.

— La femelle, en se plaçant sur ses œufs comme elle le fait chez Phrynodon et Alexte-

roon, doit attirer l'attention des prédateurs mieux que ne le fait la ponte seule.

En fait, la présence d’un des parents sur ou à proximité de la ponte peut avoir une fonc-

tion tout autre que la protection.

2. Au Gabon, les pontes aériennes de Chiromantis rufescens, déposées dans une masse d'écume battue,

peuvent être pillées par un petit oiseau, Nigrita bicolor, ainsi que l'a montré Brosser (1976). Cela se produit

aussi au Cameroun, mais très rarement.

Source : MNHN, Paris

22 ALYTES 9 (1)

— Chez Phrynodon, le rôle de la femelle semble être d’humecter les œufs, non pas pour

les protéger de la dessiccation, peu à craindre là où vit l'espèce, mais pour entraîner le gonfle-

ment de la gangue gélatineuse qui les entoure. J’ai constaté en effet (AMIET, 1981) que, en

l'absence de femelle, les pontes fraîchement déposées montrent un début de développement

des œufs mais que leur gangue reste mince: les embryons, comprimés dans leur enveloppe,

dépérissent alors. Les pontes récoltées dans la nature à un stade plus avancé de gonflement

ont une évolution normale, même en l'absence de femelle.

— En brassant l’eau avec ses pattes postérieures, le mâle de Nectophryne contribue pro-

bablement à aérer la ponte. Même si ce comportement a en réalité une autre fonction, il est

difficile d'imaginer qu'il soit destiné à éloigner d'éventuels agresseurs.

— La présence de la femelle Hemisus dans la chambre de ponte (en elle-même protec-

tion efficace contre les prédateurs) peut s'expliquer par la nécessité de forer une voie d’accès à

l’eau pour les têtards.

Ainsi, l'humectation des œufs chez Phrynodon, leur aération chez Nectophryne, le forage

de la galerie de sortie chez Hemisus suffisent à justifier la présence d’un des parents. La ques-

tion reste posée pour les Petropedetes, Alexteroon et Trichobatrachus mais, là encore, une

fonction de protection contre les prédateurs semble peu plausible.

RÉSUMÉ

La ponte est gardée par l’un des parents dans six genres d’Anoures camerounais. Sept

photos illustrent ce comportement chez les genres Petropedetes, Alexteroon, Phrynodon,

Nectophryne et Hemisus. Dans plusieurs cas, sinon dans tous, l'hypothèse d’une protection

contre les prédateurs ne paraît pas pouvoir être retenue.

RÉFÉRENCES BIBLIOGRAPHIQUES

Au, E. 1931. — Amphibia, Anura, III, Polypedatidae. Das Tierreich, 55: 1-45.

Amiër, J.-L. 1974. — Le têtard d'Hyperolius obstetricans Ahl (Amphibien Anourc). Bull. L.F.A.N.,

(A), 36 (4): 973-981.

— 1981. — Écologie, éthologie et développement de Phrynodon sandersoni Parker, 1939 (Amphibia,

Anura, Ranidae). Amphibia-Reptilia, 2: 1-13.

BRANDES, G. & SCHOENICHEN, W., 1901. — Die Brutplege der schwanzlosen Batrachier. 4bh.

h. Ges. Halle, 22: 394-461 (in PERRET, 1988).

sr, A, 1976. — La vie dans la forêt équatoriale. Paris, Nathan: 1-26.

DUELLMAN, W.E. & TRuën, L., 1986. — Biology of Amphibians. New York, McGraw Hill: ixix +

1-670.

PERRET, J.-L. 1966. — Les Amphibiens du Cameroun. Zoo!. Jb. Syst., 8: 289-464.

1988. — Sur quelques genres d'Hyperoliidae (Anura) restés en question. Bull. Soc. neuchat. Sc.

nat., LU: 35-48.

Senerz, J.J., 1970. — Notes on the biology of the African trce-toad, Nectophryne afra Buchholz &

Peters, 1875 (Bufonidae, Anura) from Fernando Po. Rev. Zool. Bot. afr., 91 (3-4): 225-236.

WaGER, VA. 1965. — The Frogs of South Africa. Cape Tow & Johannesburg, Purnell & Sons: 1242.

Corresponding editor: Alain DuBois.

Source : MNHN, Paris

Alytes, 1991, 9 (1): 23-32. 23

Faunal deficit of anurans

in tropical farmland of Amazonian Peru

Manfred AICHINGER

Institut für Zoologie, Universität Wien, AlthanstraBe 14, 1090 Wien, Austria

Fifty-eight (= 89.2 %) of 65 identified anuran species at Panguana, a study

site in the upper Amazon Basin in east-central Peru, occur in primary

rainforest, 35 (= 53.9 %) in secondary forest, 21 (— 32.3 %) at the forest edge,

8 (= 12.3 %) at the margin of the Rio Llullapichis, and 9 species (= 13.8 %)

in farmland. The diversity of anuran reproduction in forest habitats was 2.3

times higher than in areas under cultivation. The faunal impoverishment of

84.5 % in relation to the number of species living in primary forest is caused

by unfavorable environmental conditions, by the absence of appropriate

habitats, and by changes in diet supply.

INTRODUCTION

In the humid environments of tropical rainforests, anurans have developed a great

diversity of reproductive modes (DUELLMAN, 1988; HôDL, 1990). Because of the different

environmental conditions it seems reasonable to compare forest and non-forest habitats

with respect to species composition and reproduction.

A study area at the lower Rio Llullapichis in the upper Amazon Basin in east-central

Peru disposed the required characteristics: (1) an area of about 2 km? mainly covered with

unexploited primary forest and (2) adjacent areas under cultivation. The Panguana site

(fig. 1b) was founded in 1968 by H.-W. KoEPCKE (Hamburg). Between 1959 and 1984 the

areas under cultivation rapidly increased (compare fig. la, b). Within a few years the

rainforest plot of Panguana will be surrounded by farmland if rainforest destruction

continues at the present rate.

Seventy-three anuran species occur at Panguana (DUELLMAN & TorT, 1979;

SCHLÜTER, 1979, 1981, 1984; HôDL 1990; personal observations). Eight species of the

genus Eleutherodactylus are still unidentified. According to the classification system of

DUELLMAN & TRUEB (1986), the 65 identified species show 8 of 29 described reproductive

modes: (mode 1) eggs deposited in open water with feeding tadpoles (25 species); (mode

3) eggs and early larval stages in constructed basins, subsequent to flooding feeding

tadpoles in streams (Hyla boans); (mode 4) eggs and feeding tadpoles in water in tree holes

(Phrynohyas resinifictrix, supposed for Osteocephalus leprieurii); (mode 14) eggs on ground,

feeding tadpoles are carried by adult to water (7 species); (mode 17) eggs deposited out of

water with direct development (13 species, 8 species unidentified, supposed for /schno-

Source : MNHN, Paris

24 ALYTES 9 (1)

Le) 1000 2000m

Fig. 1. — Distribution of farmland at the mouth of the Rio Llullapichis in 1958 (a) and 1984 and

location of Panguana (b). Stippled areas: under cultivation. The circle in b marks the investigated

farmland pond site. (Drawn after photographs taken by the Peruvian air-force in June 1958 and

August 1984). The course of the river and the situation of islands changed between 1958 and

1984.

Source : MNHN, Paris

AICHINGER 25

cnema quixensis); (mode 18) eggs on vegetation above water, eggs hatching into tadpoles

that drop into water (8 species, supposed for Hyla rossalleni); (mode 21) eggs in foam nests

in burrows, subsequent to flooding, feeding tadpoles in ponds or streams (6 species); (mode

22) eggs in foam nests on land, non feeding tadpoles complete development in nest (2

species).

Because many anuran species with a variety of reproductive modes occur in such a

small area in tropical rainforest it seems especially interesting to know: (1) how many and

which species inhabit the different habitats; (2) if species do exist that are restricted to open

areas; and (3) which reproductive modes are favoured by the environmental conditions in

farmland.

DESCRIPTION OF STUDY SITE

Panguana is located on the south bank of the lower Rio Llullapichis, a tributary of

the Rio Pachitea, in the upper Amazon Basin, 9°37' S, 74°56° W, at an elevation of 260 m

(fig. 1b). From November 1981 through October 1982, the total annual precipitation was

2635 mm. The rainfall is seasonal, with a marked dry period lasting from mid-March to

mid-September (AICHINGER, 1987). On sunny days, daily maximum temperatures were

about 6°C higher in farmland than in primary forest. Minimum relative humidity was

between 35-40 % in farmland whereas it was seldom below 90% in the forest

(CHANAGARTH, 1981).

The investigation area of Panguana is mainly covered with unexploited primary

forest. Vegetation at sites of abandoned cultivated areas (compare fig. la, b), along a

temporary stream, at the border of the Rio Llullapichis, and at the forest edge are assigned

to secondary forest. The edge of the forest is characterized by dense vegetation near

ground level. The settlers around Panguana gain farmland by clearing primary forest.

After burning the areas are planted with maize and subsequently with grass for

cattle-breeding.

The two investigated ponds are situated about 2.8 km northwest of Panguana near the

Rio Pachitea in farmland belonging to Elvio MODENA (fig. 1b). The rainforest in this area

was cleared in the late fifties; since then it has been used for cattle-breeding. The two ponds

are surrounded by a dense vegetation of high grass and low “Niejilla” palm trees (Bactris

sp.). They are about 300 m away from primary forest. Their maximum diameter was

reached in January and measured about 8 m.

METHODS

The study on annual activity patterns of anurans at Panguana, lasting from

November 1, 1981 until October 31, 1982 (AICHINGER, 1987), included observations at two

farmland ponds. I compare the anuran fauna of 4 different forest habitats (primary forest,

secondary forest, forest edge, border of the Rio Llullapichis) with that of the farmland.

Source : MNHN, Paris

26 ALYTES 9 (1)

In primary and secondary rainforest I investigated the anuran fauna in six plots of 100

m° each which were established at different aquatic sites (AICHINGER, 1987). The temporary

pond at the forest edge had about the same size as the two farmland ponds (maximum

diameter about 8 m). The border of the Rio Llullapichis was observed along the limit of

the study area. The course of the river and subsequently the situation of islands changed

between 1958 and 1984 (compare fig. la, b).

Species are treated according to their reproductive modes. According to the number

of ovarian eggs, coloration, and egg diameter (AICHINGER, 1985), the reproductive modes

of {schnocnema quixensis and Hyla rossalleni are preliminarily assigned. No observations

on the egg-deposition site of O. leprieurii could be made. This large hylid frog calls

dispersed in primary forest and did not aggregate at aquatic sites. I assume that this

species, like Phrynohyas resinifictrix, deposits its eggs in water-filled tree holes (HÔDL, in

press; ZIMMERMANN & HÔDL, 1983). Dendrophryniscus minutus, which lays strings of eggs

on low vegetation up to 50 cm above water level, is assigned to mode 14.

Data for uncommon species observed only a few times or not at all during the course

of this study are supplemented by CRUMP (1974), DUELLMAN (1978), ToFr & DUELLMAN

(1979), and SCHLÜTER (1984).

RESULTS

The habitat distribution of 65 anuran species is given in Table I. Fifty-eight species (=

89.2 %) inhabited the primary forest. Twenty-seven species (= 41.5 %) were restricted toit.

Twenty-nine species living in primary forest also occurred in secondary forest, which pro-

vided a habitat for 35 (= 53. 8 %) species. At the forest edge I found 21 species (= 32.3 %)

at a temporary pond (AICHINGER, 1985, 1987): all also occurred in secondary forest, 17 in

primary forest. Eight species (= 10.8 %) were found at the Rio Llullapichis. Bufo marinus,

B. typhonius, and Hyla boans called during the dry season along the Rio Llullapichis. In

mid-September (early in the 6 month rainy season), Leptodactylus pentadactylus called at the

river bank where no other anurans were found throughout the whole rainy season.

Nine species (= 13.8 %) occur at the two farmland ponds. They belong to the families

Hylidae (8 species) and Leptodactylidae (1 species). I observed frogs only in the dense

vegetation surrounding these ponds. No anurans were found in the pasture. Eight of the

farmland species were also common in secondary forest. Phyllomedusa tarsius was

observed in primary forest too. Ololygon rubra also occurred in the palm-leave roofs of

native huts. Bufo marinus, most common near houses, hunting after insects attracted to

artificial light, called at the river’s edge and was not observed in farmland.

The distribution of the reproductive modes of 65 anuran species at 5 habitats is shown

in Table Il. I included only species which were actually breeding (calling males and/or

gravid females). Thus 3 species observed at the river edge (Adenomera hylaedactyla, Bufo

glaberrimus, Leptodactylus wagneri) were excluded. Species occurring in primary forest

used 7 modes of reproduction. Poison-dart frogs (mode 14) and species of the genus

Eleutherodactylus (mode 17) occurred predominantly in primary forest. The number of

Source : MNHN, Paris

AICHINGER 27

species with oviposition in water increased in secondary forest. Fourteen (= 66.7 %) out

of 21 species found at the forest edge laid their eggs in water (mode 1). At the border of

the Rio Liullapichis, 60 % of the species underwent complete aquatic development.

The 9 species found at the two farmland ponds showed 3 reproductive modes. Five

species (= 55.6 %) represent mode 1. Two species laid their eggs on vegetation above

water (Hyla leucophyllata, Phyllomedusa tarsius). Arboreal oviposition is also supposed for

H. rossalleni. The foam-nest producing Leptodactylus wagneri (mode 21) was heard calling

at these aquatic sites. In farmland no species of poison-dart frogs (mode 14) or of the

genus Eleutherodactylus (mode 17) occurred.

DISCUSSION

Of 65 anuran species known from Panguana, only 9 species (13.8 %) were found at

the two farmland ponds. Only habitat generalists breed at these aquatic sites. No anuran

species occurs exclusively in areas under cultivation. This faunal impoverishment seems to

be due to changes in microclimate, habitat diversity, and food supply.

Microclimatic conditions in farmland favor only few amphibians. In open areas, high

maximum temperatures correlated with low humidity restrict the occurrence of amphib-

ians. Due to intensive sun irradiation, maximum temperatures are higher and humidity is

lower in farmland than in forest habitats (HANAGARTH, 1981). The danger of desiccation

forces the anurans living in farmland to hide during daytime in shady shelter sites. During

night hours, when temperatures and humidity of rainforest and farmland are nearly equal

(HANAGARTH, 1981), anurans become active. All species breeding in open areas are

nocturnal (HEYER, 1976). The closed canopy of primary rainforest reduces sun irradiation

and maintains high humidity; this is important for all anurans, especially those with

terrestrial development.

Species with egg deposition directly in water (with subsequent development occurring

there as well) represent the primitive mode of reproduction (DUELLMAN & TRUEB, 1986).

These species are less dependent on the high humidity provided by a forest than those

depositing eggs in terrestrial environments where all needed water is extracted from the air

(Lyc, 1979). Environmental conditions in farmland favor species depositing many eggs

in open water. Reproductive generalists with high fecundity are better adapted for

unpredictable environments (DUELLMAN, 1978). The mean egg number of the 5 aquatic

breeding farmland species examined by AICHINGER (1985) ranged between 394 (Hyla

riveroi) and 1170 (Ololygon rubra) (n = 10 — 15).

Four species at the farmland ponds do not deposit their eggs directly in the water. The

species were less abundant than aquatic breeders. Two species lay their eggs on vegetation

above water. This mode is also assumed for Hyla rossalleni. Development is endangered

by a possible desiccation of eggs. Phyllomedusa tarsius encloses the egg mass with leaves.

The top and the bottom of the clutch consist mainly of eggless capsules providing

protection from sun irradiation (CRUMP, 1974). Hyla leucophyllata lays its eggs on leaves

on low vegetation (up to a height of 50 cm), where humidity from the evaporating water

surface may prevent desiccation. Leptodactylus wagneri reflects the primitive Leptodactylus

Source : MNHN, Paris

ALYTES 9 (1)

Table I. — Distribution of 65 anuran species in 5 different habitats. PF = primary forest,

SF = secondary forest, FE = forest edge, RE = river edge, FL = farmland. Categories corre-

spond to reproductive mode numbers according to DUELLMAN & TRUEB (1986), * indicates

species where data are supplemented by CRUMP (1974), DUELLMAN (1978), TOFT & DUELLMAN

(1979), and SCHLÜTER (1984).

SPECIES

LEPTODACTYLIDAE

1. Adenomera andreae

2. À. hylaedactyla

3. Ceratophrys cornuta

4. Edalorhina perezi

5. Eleutherodactylus acuminatus

6. E. altamazonicus

7. E. carvalhoi

E. diadematus

. E. imitatrix

10. E. lacrimosus

11. E. mendax

12. E. ockendeni

13. E. peruvianus

14. E. sulcatus

15. E. toftae

16. E. ventrimarmoratus

17. Ischnocnema quixensis

18. Leptodactylus pentadactylus

19. L. rhodomystax

20. L. wagneri

21. Lithodytes lineatus

22. Physalaemus petersi

BUFONIDAE

23. Bufo glaberrimus

24. B. marinus

25. B. typhonius

26. Dendrophryniscus minutus

DENDROBATIDAE

27. Colostethus marchesianus

28. C. peruvianus

29. Dendrobates quinquevittatus

30. Epipedobates femoralis

31. E. petersi

32. E. pictus

33. E. trivittatus

only

RE ponds

21 21

Source : MNHN, Paris

AICHINGER

HYLIDAE

34. Hyla boans

35. H. brevifrons

* 36. H. calcarata

37. H. fasciata

38. H. geographica

39. H. granosa

40. H. leucophyllata

41. H. marmorata

42. H. minuta

43. H. parviceps

44. H. rhodopepla

45. H. riveroi

46. H. rossalleni

47. H. sarayacuensis

48. Ololygon cruentomma

49. O. funerea

50. O. garbei

51. O. rubra

52. Osteocephalus leprieurit

53. O. taurinus

54. Phrynohyas coriacea

* 55. P. resinifictrix

56. P. venulosa

* 57. Phyllomedusa palliata

58. P. tarsius

59. P. tomopterna

60. P. vaillanti

CENTROLENIDAE

61. Centrolenella midas

62. C. munozorum

MICROHYLIDAE

63. Chiasmocleis ventrimaculata

64. Ctenophryne geayi

65. Hamptophryne boliviana

RENE ©

Sé gets

ll il ll

18 18

1 ||

[l 1

ll l

ll ll

Source : MNHN, Paris

30 ALYTES 9 (1)

Table II. — Distribution of reproductive modes (after DUELLMAN & TRUEB, 1986) in 5 different

habitats.

Primary

Reproductive Primar: Secondar: Forest- River- Farmland

mode Que font Pet font edge cd ponds

= 1 23 354% 21362% 4154% 19500% 1466.7 % 3 600% 5 55.6 %

1 15% 1 26% 1 200%

4 ER 234% 1 38% 1 26%

14 T 108% 7121% 7269%

17 13 200% 13224% 9346% 3 79%

18 11 169% 8138% 3115% 6158% 4190% 3 333%

À 6 92% 6103% 2 77% 410.5% 2 95% 1 200% 1 111%

22 2 11% 1 17% 1 38% 1 26% 1 48%

Total 65 1000% 5889.2% 2741.5% 3553.8% 21323% 5 77% 9 138%

pattern (HEYER, 1969) in that a foam nest is placed on top of the water. Eight gravid females

of L. wagneriexamined by AICHINGER (1985) contained a mean number of 1726 eggs.

Tropical rainforests provide a great diversity of habitats. Anurans are well adapted to

live in leaf litter, bushes, or in the forest canopy. At Panguana no anuran species with

terrestrial breeding (Dendrobatidae, Eleutherodactylus spp., Adenomera hylaedactyla)

occurs in farmland. These species require leaf litter as oviposition and shelter sites. AIl are

restricted to forest habitats where leaf litter is abundant and humidity high throughout the

day. Many anurans, especially hylid frogs, are adapted to the dense vegetation of bushes.

In farmland this microhabitat was found only around the two investigated ponds. In forest

habitats, 40 anuran species were found in bushes, whereas in farmland only 8 species

occurred in this microhabitat. Even the species diversity of birds was remarkably higher

in undisturbed primary forest than in cacao or coffee plantations (TERBORGH & WESKE,

1969). In California, logging of redwood forest opened the forest canopy and apparently

favored only a few amphibian species to the detriment of the majority — the

forest-dependent species (BURY, 1983).

Farmland habitats favor only broad-niched species. Leptodactylus wagneri is well

adapted to changing environments and probably an excellent colonizer species (HEYER &

BELIN, 1973). This seems also to be true for Ololygon rubra, which was found especially

at ephemeral watersites.

Diet supply for food specialists, such as certain leaf-litter frogs (TorT, 1980), may limit

the existence in farmland. There, compared with primary forest, abundances of ants and

termites are remarkably reduced; beetles, grasshoppers and crickets, however, are more

abundant (HANAGARTH, 1981). The ant fauna is nearly completely destroyed by the cutting

and burning of forests (VERHAAGH, in press).

Rainforest destruction leads to a reduction in the numbers of individuals of those

species bound to primary rainforest (JACOBS, 1988). At Panguana environmental

Source : MNHN, Paris

AICHINGER 31

conditions, shortage of food supply, and a remarkable deficit of habitat types in farmland

result in a faunal impoverishment of 84.5 % compared with the anuran species living in

primary forest.

RESUMEN

58 especies de batracios (= 89,2 %) de 65 especies identificadas de Panguana, una

estacion de estudios en la parte alta de la cuenca amazénica central, ocurren en el bosque

primario, 35 (= 53,9 %) en el bosque secundario, 21 (= 32,3 %) en el borde del bosque,

8 (= 12,3%) en la orilla del Rio Llullapichis y 9 especies (= 13,8 %) en los campos

cultivados. La diversidad de reproduccién de los batracios en los habitates selväticos fue

2,3 veces mäs alto que en los campos cultivados. El empobrecimiento faunistico de 84,5 %

en relaciôn al nümero de especies que viven en el bosque primario es posiblemente causado

por las condiciones desfavorables del ambiente, la ausencia de habitates apropriados y los

cambios de la provisién alimenticia.

ACKNOWLEDGEMENTS

Thanks are due to Prof. H.-W. KorPCKE (Univ. Hamburg) for permission to carry out the study

at Panguana and to W. HôpL and W. MORAWETZ for reviewing the manuscript. Financial support

was provided by “ ôsterreichischer Fonds zur Fôrderung der wissenschaftlichen Forschung ” (Project

P6399B) and by the Austrian ministry for science and research.

LITERATURE CITED

AicHinGer, M., 1985.— Niederschlagsbedingte Aktivitätsmuster von Anuren des tropischen Regenwal-

des: eine quantitative Studie durchgeführt im Forschungsgebiet von Panguana (Peru). Wien,

Univ. Diss.: 1-68.

ee 1987. — Annual activity patterns of anurans in a seasonal ncotropical environment. Oecologia,

71: 583-592.

BURY, R. B., 1983. — Differences in amphibian populations in logged and old growth redwood forest.

Northwest Sci., 57: 167-178.

Crume, M. L., 1974. — Reproductive strategies in a tropical anuran community. Misc. Publ. Mus.

Nat. His. Univ. Kansas, 61: 1-67.

DuëLiMAN, W. E. 1978. — The biology of an equatorial herpetofauna in Amazonian Ecuador. Misc.

Publ. Mus. Nat. Hist. Univ. Kansas, 65: 1-352.

= 1988. — Patterns of species diversity in anuran amphibians in the American tropics. Ann.

Missouri Bot. Gard. 75: 79-104.

DUELLMAN, W. E. & TRUEB, L., 1986. — Biology of Amphibians. New York, MeGraw Hill: 1-670.

HANAGARTH, W., 1981. — Vergleichend-ükologische Untersuchungen an epigäischen Arthropoden aus

Naturbiotopen und Kulturland im tropischen Regemwald Perus. - Ein Beitrag zur Agrarëkologie

der Tropen. Hamburg, Univ. Diss. Nr. 125: 1-240.

Hever, W.R., 1969. — The adaptive ecology of the s

(Amphibia, Leptodactylidac). Evolution, 23: 421-428.

pu 1976. — Notes on the frog fauna of the Amazon Basin, Acta Amazonica, 6: 369-378.

ecies groups of the genus Leprodactylus

Source : MNHN, Paris

3 ALYTES 9 (1)

Hever, W. R. & BeuiN, M.S., 1973. — Ecological notes of five sympatric Leptodactylus (Amphibia,

Leptodactylidae) from Ecuador. Herpetologica, 29: 66-72.

Hôp, W., 1990. — Reproductive diversity in Amazonian lowland frogs. Fortschritte d. Zoologie, 38:

41-60.

ass in press. — Phrynohyas resinifictrix (Hylidae). Calling behaviour. Wiss. Film.

Jacoss, M., 1988. — The tropical rainforest. À first encounter. R. Kruk, Berlin, Heidelberg, New

York, London, Paris, Tokyo, Springer Verlag: 1-295.

LYNCH, J. D., 1979. — The amphibians of the lowland tropical forests. /n: W. E. DUELLMAN (ed.), The

South American herpetofauna: its origin, evolution, and dispersal, Monog. Mus. Nat. Hist.

Univ. Kansas, T: 189-215.

SCHLÜTER, A., 1979. — Bio-akustische Untersuchungen an Hyliden in einem begrenzten. Gebiet des

tropischen Regenwaldes von Peru (Amphibia: Salientia: Hylidae). Salamandra, 15: 211-236.

en 1981. — Erstnachweis von Bufo glaberrimus Günther, 1868 für Peru. Stud. Neotrop. Fauna and

Environment, 16: 221-223.

Le 1984. — Ükologische Untersuchungen an einem Stillgewässer im tropischen Regenwald von Peru

unter besonderer Berücksichtigung der Amphibien. Hamburg, Univ. Diss.: 1-300.

TERBORGH, J. & WESskE, J.S., 1969. — Colonization of secondary habitats by peruvian birds.

Ecology, 50: 765-782.

Torr, C. A., 1980. — Feeding ecology of thirteen syntopic species of anurans in a seasonal tropical

environment. Oecologia, 45: 131-141.

Torr, C. A. & DUELLMAN, W.E., 1979. — Anurans of the lower Rio Llullapichis, Amazonian Peru:

a preliminary analysis of community structure. Herpetologica, 35: 71-77.

VERHAAGH, M. in press. — Clearing a tropical rainforest — effects on the ant fauna. /n: W. ERDELEN,

N. IsHWARAN & P. MÜLLER (eds), Tropical Ecosystems, Saarbrücken, Proc. Symp.,

15-18.6.1989, in press.

ZIMMERMANN, B. & HôDL, W., 1983. — Distinction of Phrynohyas resinifictrix (Goeldi, 1907) from

Phrynohyas venulosa (Laurenti, 1768) based on acoustical and behavioral parameters. Zoo!.

Anz., 211: 341-352.

Corresponding editors: Walter Hôp & Alain DuBois.

Source : MNHN, Paris

AINTTES

International Journal of Batrachology

published by ISSCA

EDITORIAL BOARD FOR 1991

Chief Editor: Alain Dumois (Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire

naturelle, 25 rue Cuvier, 75005 Paris, France).

Deputy Editor: Günter GOLLMANN (Institut für Zoologie, Universität Wien, Althanstr. 14, 1090 Wien,

Austria).

Other members of the Editorial Board: Jean-Louis AMIET (Yaoundé, Cameroun); Stephen D. BUSACK

(Ashland, U.S.A.); Tim HarriDAY (Milton Keynes, United Kingdom); William R. HEYER

(Washington, U.S.A.); Walter Hôp (Wien, Austria); Pierre JOLY (Lyon, France); Milos

KaALEzIC (Beograd, Yugoslavia); Raymond F. LAURENT (Tucumän, Argentina}; Petr ROTH

(Libechov, Czechoslovakia), Borja SANCHIZ (Madrid, Spain); Dianne B. SEALE (Milwaukee,

U.S.A.); Ulrich SINsCH (Bonn, Germany).

Index Editor: Annemarie OHLER (Paris, France).

GUIDE FOR AUTHORS

Alytes publishes original papers in English, French or Spanish, dealing with amphibians. Beside

papers reporting results of original research, consideration will be given for publication to review

articles, comments and replies, and to papers based upon original high quality color photos of

amphibians, showing beautiful or rare species, interesting behaviors, etc.

The title should be followed by the name(s) and address(es) of the author(s). The text should be

organized as follows: English abstract, introduction, method, results, discussion, conclusion, French

or Spanish abstract, acknowledgements, literature cited.

Figures and tables should be mentioned in the text as follows: fig. 4 or Table IV. Figures

should not exceed 16 x 24 cm. The size of the lettering should ensure its legibility after reduction.

The legends of figures and tables should be assembled on a separate sheet. Each figure should be

numbered using a pencil.

References in the text are to be written in capital letters (SOMEONE, 1989; EVERYBODY et al.,

1980; So & So, 1987). References in the literature cited section should be presented as follows:

= when in a periodical:

INGER, R. F., Voris, H. K. & Voris, H. H., 1974. - Genetic variation and population ecology of some

Southeast Asian frogs of the genera Bufo and Rana. Biochem. Genet, 12: 121-145.

— when in a multi-authors book:

GRAF, J.-D. & PoLLs PELAZ, M., 1989. — Evolutionary genetics of the Rana esculenta complex. In:

R. M. DAWLEY & J. P. BOGART (eds.), Evolution and ecology of unisexual vertebrates, Albany, The

New York State Museum: 289-302.

— when a book:

BOURRET, R., 1942. - Les Batraciens de l'Indochine. Hanoï, Institut Océanographique de l’Indochine:

i-x+1-547, pl. I-IV.

Manuscripts should be submitted in triplicate to Alain DUBois (address above) if dealing with

amphibian morphology, systematics, biogeography, evolution, genetics or developmental biology, or

to Günter GOLLMANN (address above) if dealing with amphibian population genetics, ecology,

ethology or life history.

Âcceptance for publication will be decided by the editors following review by at least two

referees.

No page charges are requested from author(s), but the publication of color photographs is

charged. For each published paper, 25 free reprints are offered by A/ytes to the author(s). Additional

reprints may be purchased.

Published with the support of

the Muséum national d'Histoire naturelle (Paris, France).

Directeur de la Publication: Alain Dumois.

Numéro de Commission Paritaire: 64851.