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 1989

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

General Secretary : Alain DuBoIs (Paris, France).

Treasurer : Dominique PAYEN (Paris, France).

Assistant Secretary for Europe : Olivier BEHRA (Paris, France).

Assistant Treasurer for Europe : Annemarie OHLER (Paris, France).

Other members of the Board : James P. BoGaRT (Guelph, Canada) ; Jean-Louis FISCHER (Paris, France) ;

David M. GREEN (Montreal, Canada) ; Pierre JoLY (Lyon, France) ; James I. MENZ1ES (Boroko, Papua

New Guinea) ; Manuel POLLS PELAZ (Barcelona, Spain) ; Richard WassrsuG (Halifax, Canada).

TARIFFS 1989

Subscription to

ISSCA Circabts Abytes Total

Individuals

ISSCA direct members 50 FF 50 FF 130 FF 230 FF

ISSCA group or section members* 15 FF 50 FF 130 FF 195 FF

Non-members “ - 150 FF 150 FF

Institutions

ISSCA direct members 100 FF 100 FF 260 FF 460 FF

ISSCA group or section members* 30 FF 100 FF 260 FF 390 FF

Non-members ee = 300 FF 300 FF

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

de Batrachologie ; Working Group on Oriental Amphibians).

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 to 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:

AIR7TES

INTERNATIONAL JOURNAL OF BATRACHOLOGY

January-April 1989 Volume 8, N° 1

Alytes, 1989-1990, 8 (1): 1-2. 1

Editorial

Alain Dugois* & Pierre JoLy**

*Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France Bibliothèque Centrale Muséum

3 58

3001 00111585 5

The journal Alytes was founded at the Paris National Museum of Natural History in

February 1982. From November 1982 to November 1989, it was published by the Société

Batrachologique de France (S.B.F.), although since March 1984 it has been subtitled

“International Journal of Batrachology”, and opened, besides those in French, to papers

in English.

On 11 November 1988, the General Assembly of the S.B.F. decided the foundation

of the ISSCA (International Society for the Study and Conservation of Amphibians), the

transfer to this new international society of the journal Alytes, and the opening of the latter

also to papers in Spanish. Since, however, publication of volume 7 of Alytes, due for the year

1988, was late, it was decided to wait until completion of this publication to transfer the

journal to the ISSCA. This was done only in November 1989, and volume 8 of the journal

will now be published by the ISSCA.

In order to suppress the one-year delay in the publication schedule of Alytes inherited

from the S.B.F., the ISSCA Board decided that volume 8 of Alytes will formally cover both

years 1989 and 1990 : therefore, at the end of this volume, the journal should have recovered

normal dates of publication.

All subscribers who have paid in 1989 for their subscription to volume 8 of Alytes have

therefore no additional subscription to pay for 1990, and will receive all issues of this

volume.

**Laboratoire de Biologie animale et Ecologie, |

Université Claude Bernard Lyon I,

69622 Villeurbanne Cedex, France

Source : MNHN, Paris

2 ALYTES 8 (1)

Alytes is therefore starting again on new, truly international, bases. We invite warmly

all batrachologists worldwide to submit to us for publication papers in English, French or

Spanish dealing with all aspects of the scientific study and conservation of amphibians.

Although the journal has until now mostly published papers in the fields of amphibian

systematics and biogeography, we want to make it quite clear that its scope is not limited to

these topics, but is much wider, including all fields of amphibian biology as such (e.g. am-

phibian ecology, ethology, physiology, karyology, genetics, evolution, developmental biol-

ogy and embryology, larval biology, conservation biology, etc.), and excluding only exper-

imental works where amphibians are used merely as a “material” but without paying attention

to these organisms as such.

The Editorial Board of Alytes has been expanded and now includes 13 members from

8 countries. Any paper submitted for publication in Alytes will be entrusted to one of these

members, who will act as Corresponding Editor for this paper, and send it out for review to

two referees at least before making the final decision.

The delay for publication of an accepted paper in Alytes is currently of 3 to 6 months

after submission, which is much less than in many other international journals of zoology or

herpetology. No page charges are requested from authors, but we regret to have to charge

the latter for the publication of color photographs.

We invite you all to join the ISSCA and to participate in the life and development of

its journal Alytes, the first journal of batrachology in the world.

Source : MNHN, Paris

Alytes, 1989-1990, 8 (1): 3-10. 3

Images d’Amphibiens camerounais.

I. Sacs vocaux et postures de chant

Jean-Louis AMIET

Université de Yaoundé,

Faculté des Sciences, Laboratoire de Zoologie,

B.P. 812, Yaoundé, Cameroun

Eight photos of anurans from Cameroon, taken during the emission of nuptial

calls, are commented upon. These photos illustrate various aspects of the vocal sac and

of the body during the call, as well as the postures taken by the frogs in relation to the

shape of the vocal sac.

INTRODUCTION GÉNÉRALE

Sous le titre “Images d’Amphibiens camerounais” se succéderont de courts articles

centrés sur des planches de photos en couleurs illustrant divers aspects de la biologie des

Amphibiens Anoures du Cameroun, et plus spécialement leur reproduction.

Dans ces articles, conçus plutôt comme des “essais”, le texte sera subordonné aux

images et se limitera aux espèces de la batrachofaune camerounaise. Je laisse à d’autres le

soin d'intégrer les informations ainsi apportées dans des travaux plus synthétiques. Dans la

même perspective, seules seront citées les publications touchant directement aux thèmes

traités.

Sur le plan technique, les documents photographiques présentés ne sont pas tous

irréprochables. La plupart ont en effet été réalisés sur le terrain, dans des conditions parfois

difficiles. Certains ont pâti de ma précipitation à fixer une scène rare ou fugace. J'espère que,

tels quels, ces clichés parviendront quand même à faire partager aux lecteurs l’émotion que

j'ai souvent ressentie en les réalisant.

REMERCIEMENTS

Ces articles n’auraient probablement jamais vu le jour sans les amicales pressions qu’ont exercées

sur moi À. Dugois et J.-J. MORÈRE. Ils doivent aussi beaucoup aux suggestions et corrections de deux

lecteurs à qui ils avaient été confiés : j'espère qu’ils se reconnaîtront dans les améliorations apportées

au texte initial. tout comme je les ai reconnus à travers leurs critiques. A tous, j’exprime ma vive

reconnaissance.

Source : MNHN, Paris

4 ALYTES 8 (1)

ASPECTS DES SACS VOCAUX

Une des particularités remarquables de la phonation chez les Anoures est que les vo-

calisations (sauf les cris de détresse) sont produites la bouche fermée. Chez la plupart des

espèces, il y a cependant une manifestation visible des émissions sonores, correspondant au

gonflement d’un ou de deux sacs vocaux. Si l’on se réfère à l’une des synthèses récentes sur

la question, celle de DUELLMAN & TRUEB (1986)!, trois types de sacs vocaux peuvent être

reconnus : subgulaire médian, subgulaire double et latéraux. A l’intérieur de chacune de ces

catégories, en particulier la première, la forme, le volume, la coloration et la texture du té-

gument varient beaucoup suivant les genres et même les espèces considérés.

Chez les Anoures à sac médian subgulaire, le tégument de la gorge, dans le cas le plus

simple, ne montre pas de différenciation particulière et a le même aspect dans les deux sexes.

C'est ce qui s’observe, dans la faune camerounaise, chez les rainettes du genre Leptopelis ainsi

que chez divers Bufonidae, Astylosterninae et Arthroleptinae.

Dans d’autres cas, le tégument gulaire a, chez le mâle, une coloration particulière, en

général plus foncée. C’est ainsi que Nyctibates corrugatus (Astylosterninae) ne montre pas

d’autre caractère sexuel secondaire que la coloration de la gorge, d’un noir d’encre chez le

mâle (AMIET, 1973). Chez certaines espèces, le tégument gulaire des mâles peut aussi être

plus ou moins spinuleux (Petropedetes, Leptodactylodon).

Ces diverses particularités de texture et de coloration n’ont pas de rapport évident avec

l'émission des sons, contrairement aux plis gulaires, plus ou moins serrés et profonds, qui

témoignent d’une plus grande extensibilité de la gorge lors du gonflement du sac vocal. De

tels plis peuvent s’observer chez des Leptodactylodon (albiventris et le groupe d’ovatus) ou chez

certains Phrynobatrachus (où ils sont, suivant les espèces, longitudinaux ou transverses).

Dans plusieurs genres de petites rainettes (Hyperolius, Afrixalus, Kassina…), cette dif-

férenciation du tégument gulaire s'accompagne de l’existence d’une zone glandulaire épais-

sie, de forme et de taille variables suivant les espèces. Cette structure a reçu diverses dé-

nominations : “disque gulaire”, “opercule”, “protective flap”, etc., auxquelles le terme de

“glande gulaire” (DREWES, 1984) paraît préférable. La fonction de cette glande reste énig-

matique (productions de substances stimulantes lors de l’accouplement, durant lequel le mâle

appuie étroitement sa gorge sur la région occipitale de la femelle?) mais son rôle protecteur

ne semble pas négligeable. Chez la plupart des espèces appartenant aux genres précités, la

partie fine et vulnérable du tégument gulaire se replie en effet plus ou moins, au repos, sous

le pourtour de la glande gulaire. Celle-ci, lorsque le sac vocal est en pleine extension, n’en

représente qu’une faible surface, comme il est facile de le constater par les photos des fig. 1

à 3 (fig. 1 : Hyperolius viridiflavus pallidus ; fig. 2 : Hyperolius ocellatus purpurescens ; fig. 3 :

Kassina senegalensis).

Un perfectionnement remarquable de l’appareil vocal s’observe chez les grenouilles des

genres Ptychadena et Hildebrandtia (voir fig. 5-6, Hildebrandtia ornata). Ici, comme chez

1. On pourra aussi consulter avec profit, sur ce même sujet, la mise au point de PAILLETTE (1986).

2. On trouvera dans DREWES (1984) une analyse détaillée de la structure et de l’évolution du sac vocal chez les Hy-

peroliidae.

Source : MNHN, Paris

AMIET o

ig. L. — Hyperolius viridiflavus pallidus gonflant son sac vocal. Garoua, juillet 1975. is

ig. 2. — Hyperolius ocellatus purpurescens chantant sur une feuille. Yaoundé, novembre 1970. N

Fig. 3 et 4. — Kassina senegalensis, même individu photographié sous deux angles différents, en phase

e gonflement du sac vocal (3) et du corps (4). Minkama, octobre 1977.

de gonflement du sac vocal (3) et du corps (4). Minkama, octobre 1977 DRÉRICRRES

6 ALYTES 8 (1)

d’autres Raninae, il y a deux sacs latéraux, mais qui peuvent s’invaginer ou faire saillie à

l'extérieur en passant par deux fentes obliques situées sous les commissures buccales.

Dans la plupart des cas, l'émission d’un cri est précédée par le gonflement des pou-

mons au-delà de leur capacité respiratoire normale. L’air ainsi accumulé est propulsé vers la

cavité buccale et le sac vocal, faisant au passage vibrer les cordes vocales. L’air peut effectuer

de la sorte plusieurs allers et retours successifs entre les poumons et le sac vocal.

Lors du remplissage des poumons, tout le corps du Batracien s’enfle de façon plus ou

moins marquée. Ceci est rendu possible par la grande extensibilité du tégument : sur les

photos des fig. 5 à 8, qui illustrent bien le gonflement alternatif du corps et du sac vocal, on

remarquera que chez Leptopelis brevirostris (fig. 7-8), comme chez Hildebrandtia ornata (fig.

5-6), le tégument forme des plis sur les flancs lors de la phase de dégonflement.

ATTITUDES LORS DE LA PHONATION

La possession d’un sac vocal, ou de deux, est un avantage pour la reproduction puis-

qu’elle permet l’amplification des appels nuptiaux produits par les mâles. Elle entraîne ce-

pendant — surtout chez les espèces à sac volumineux — quelques contraintes relatives au choix

des postes de chant et au positionnement de l’animal lors de l’émission des appels.

En effet, chez les espèces à sac vocal subgulaire, le gonflement de celui-ci peut être

entravé par son contact avec le substrat. Les fig. 7 et 8 montrent la curieuse posture qu’a dû

adopter un mâle de Leptopelis brevirostris chantant sur une large feuille de Marantacée. Pour

éviter que son sac vocal n’appuie sur le support, l’animal se soulève sur ses quatre membres,

les postérieurs étant à demi fléchis, dans une attitude tout à fait inhabituelle chez un Anoure.

Chez la Kassina senegalensis et l'Hyperolius ocellatus des fig. 2 et 3, on voit que le sac vocal a

tendance à faire saillie vers l’avant plutôt que vers le bas et il suffit alors au chanteur de tendre

ses seuls membres antérieurs.

Souvent, pour permettre le libre jeu du sac vocal, l’animal choisira de se tenir sur le

bord d’une large feuille ou en travers d’une brindille : c’est le cas de l’'Hyperolius viridiflavus

de la fig. 1, dont le sac vocal ovoïde, à grand axe dorso-ventral, est presque aussi volumineux

que l’animal lui-même.

Pour les espèces chantant sur le sol, les sacs latéraux paraissent représenter une solu-

tion pratique car l’animal n’est pas obligé de se camper sur ses membres antérieurs. Cette

facilité ne joue guère pour Hildebrandtia ornata (fig. 5-6) car ses sacs vocaux sont si encom-

brants qu’elle ne peut pas chanter tapie sur le sol comme le font les Prychadena.

INFORMATIONS COMPLÉMENTAIRES SUR LES ESPÈCES FIGURÉES

LEPTOPELIS BREVIROSTRIS (WERNER, 1898) (fig. 7-8)

Ce Leptopelis est largement répandu dans toute la zone forestière camerounaise, jus-

qu’à environ 1100 m d’altitude. C’est une espèce sylvicole qui, dans les régions à forte plu-

viosité, peut vivre aussi dans les formations secondaires denses. En revanche, il est peu fré-

quent en forêt mésophile et manque dans les forêts galeries les plus avancées.

Source : MNHN, Paris

AMIET 7

L. brevirostris se distingue de ses congénères par deux particularités biologiques im-

portantes :

— son régime alimentaire est essentiellement hélicophage, comme l’a montré PERRET

(1966) ;

— plusieurs indices montrent que son développement doit être direct : grande taille des

ovules (5,2 mm de diamètre : PERRET, 1958), sites de chant des mâles souvent très éloignés

de tout point d’eau (AMIET & ScHigTz, 1974), absence de Protozoaires endocommensaux dans

le rectum (AMIET & AFFA’A, 1985).

HYPEROLIUS VIRIDIFLAVUS PALLIDUS MERTENS, 1940 (fig. 1)

Le nom de cette sous-espèce est bien choisi car la livrée dorsale peut devenir d’un blanc

de craie. Au Cameroun, elle ne se rencontre que dans les plaines septentrionales, depuis le

pied de l’Adamaoua jusqu’au Lac Tchad. A partir de la latitude de Poli, c’est le seul repré-

sentant du genre Hyperolius dans le Nord Cameroun.

HYPEROLIUS OCELLATUS PURPURESCENS LAURENT, 1943 (fig. 2)

C’est une espèce banale des endroits marécageux en forêt ou dans les formations se-

condaires denses juxta-forestières.

Le mâle figuré ici est en livrée nocturne, d’un roux ferrugineux, parfois rougeâtre, avec

deux bandes jaune d’or latéro-dorsales et un triangle de même couleur sur le museau. En

livrée diurne, l’animal est d’un vert translucide avec des bandes d’un jaune très pâle. La fe-

melle a une livrée bien différente, ornée de petites taches claires circulaires pupillées de bleu.

KASSINA SENEGALENSIS (DUMÉRIL & BIBRON, 1841) (fig. 3-4)

Largement répandue dans les savanes et les steppes de l'Afrique sub-saharienne, K.

senegalensis est peut-être en fait un amalgame de plusieurs espèces sub-jumelles.

Les mâles chantent souvent à plusieurs mètres de l’eau, dissimulés au pied des touffes

d'herbe. Chez les Kassina (et les Phlyctimantis, qui en sont très proches), la glande gulaire

est très développée et forme plus ou moins un “pont” reliant le bord antérieur de l’arc man-

dibulaire à la région pectorale. Une partie du tégument gulaire, très fine et pigmentée de

noir, s’invagine de part et d’autre de la glande gulaire. Cette disposition pourrait laisser croire

que, lors des émissions vocales, deux protubérances font saillie de chaque côté de la zone

glandulaire. La fig. 3 montre que chez K. senegalensis il n’y a en fait qu’un volumineux sac

vocal à contour régulièrement arrondi et orné de deux aires pigmentées de noir, ce qui avait

déjà été observé par WAGER (1965).

3. Les informations données par PERRET sont ignorées par DREWES & ROTH (1981) dans leur travail sur deux autres

Anoures hélicophages, Tomnierella kounhiensis et T. obscura, mais ont été relevées par DuBoIs (1987 : 38).

4. En revanche, d’après le même auteur, chez K. maculata (autrefois appelée Hylambates maculatus) “two smaller

ballons” apparaissent à la surface du sac vocal.

Source : MNHN, Paris

8 ALYTES 8 (1)

Fig. 5 et 6. - Hildebrandtia ornata en train de chanter, avec 1és sacs vocaux gonflés (6) et dégonflés (5).

Mora, juillet 1973

Source : MNHN, Paris

AMIET

. 7 et 8. - Deux pl le l'émission du chant chez Leptopelis brevirostris. Zamakoé, avril 1977.

Source : MNHN, Paris

10 ALYTES 8 (1)

HILDEBRANDTIA ORNATA (PETERS, 1878) (fig. 5-6)

Les Hildebrandtia sont une version fouisseuse des Ptychadena, avec lesquelles elles par-

tagent de nombreux caractères, en particulier la possession de fentes sous-commissurales

permettant la dévagination des sacs vocaux.

Localisées dans les savanes sèches et les steppes, ces grenouilles n’ont qu’une très brève

période d’activité vocale en début de saison des pluies (AMIET, 1974) et les photos de mâles

en train de chanter doivent être assez exceptionnelles.

RÉSUMÉ

Huit photos d’Anoures camerounais, prises pendant l’émission des appels nuptiaux,

sont commentées. Ces photos illustrent divers aspects du sac vocal et du corps lors du chant,

ainsi que les postures adoptées en fonction de la forme du sac vocal.

RÉFÉRENCES BIBLIOGRAPHIQUES

AMIET, J.-L., 1973. - Notes faunistiques, éthologiques et écologiques sur quelques Amphibiens Anoures

du Cameroun (2ème série). Ann. Fac. Sc. Cameroun, 13 : 135-161.

_—— 1974. — Voix d’Amphibiens camerounais. IV. Raninae : genres Ptychadena, Hildebrandtia et Di-

croglossus. Ann. Fac. Sc. Cameroun, 18 : 109-128.

AMIET, J.-L. & ArF4’A, F.-M., 1985. — À propos des stratégies d’infestation chez les Protozoaires pa-

rasites ou endocommensaux des Amphibiens Anoures du Cameroun. Rev. Ecol. (Terre Vie), 40 :

389-398.

AMIET, J.-L. & ScHigtz, A., 1974. — Voix d’Amphibiens camerounais. III. Hyperoliinae : genre Lep-

topelis. Ann. Fac. Sc. Cameroun, 17 : 131-163.

DREWES, R.C., 1984. — A phylogenetic analysis of the Hyperoliidae (Anura) : treefrogs of Africa, Ma-

dagascar and the Seychelles Islands. Occ. Pap. California Acad. Sci. 139 : 1-70.

DREWES, R.C. & ROTH, B., 1981. — Snail eating frogs from Ethiopian highlands : a new anuran spe-

cialization. Zool. 7. Linnean Soc., 72 (3) : 267-287.

Dusois, A., 1987. - Miscellanea taxinomica batrachologica (1). Alytes, 5 : 7-95.

DuELLMAN, W.E. & TRUEB, L., 1986. — Biology of Amphibians. New York, McGraw Hill : i-xix + 1-

PAILLETTE, M., 1986. — La communication acoustique chez les Amphibiens. Jn : GRASSÉ, P.P. & DEL-

soL, M. (réd.), Traité de Zoologie, t. XIV, fasc. 1b, Batraciens, Paris, Masson, 1-828 : 389-416.

PERRET, J.L., 1958. - Observations sur les rainettes africaines du genre Leptopelis Günther. Rev. suisse

Zool., 65 : 259-275.

1966. - Les Amphibiens du Cameroun. Zool. Fb. Syst., 8 : 289-464.

WAGER, V.A., 1965. — The Frogs of South Africa. Cape Town, Purnell & Sons : 1-242.

Corresponding editor : Alain DuBois.

Source : MNHN, Paris

Alytes, 1989-1990, 8 (1): 11-16. ll

Tattooing as an individual marking technique

in urodeles

Pierre JoLy & Claude MIAUD

URA CNRS 367 Ecologie des Eaux Douces,

Université Claude Bernard,

Lyon 1, 69622 Villeurbanne Cédex, France

A review of the literature shows that no marking method of urodeles is entirely

satisfactory. The choice of a method has to take into account the number of animals to

be marked, the duration of the observations and the goals to be reached. Skin staining

by tattooing is not often used even though such a technique gives interesting results.

Tattooing is a quick procedure which makes it possible to mark a great number of

animals individually. The remanence of the mark reaches at least three ears. No harmful

effects have been observed.

INTRODUCTION

Marking has always been a problem in studies of population dynamics in urodeles. No

technique is entirely satisfactory and most often the solution chosen is a compromise.

Using rings, like bird bands, is made difficult because of the small size of urodeles

limbs. Such rings involve a loss of mobility. We tried caudal rings derived from those ap-

plied to anuran limbs (DELY, 1954 ; NACE & MANDERS, 1982). Such rings, on which beads

or symbols can be threaded, made it possible to study individual behaviour in the laboratory

(MARTIN et al., 1989). But we sometimes observed newts trapped in a clump of vegetation,

a blade of Myriophyllum wedged in the ring. After some time, necrosis may occur around

the ring and may involve the loss of both a piece of the tail and the tag. The animal mobility

would then be reduced until the lost part was regenerated.

Jaw tags used in anurans (RANEY, 1940) are too large for most urodeles and the risks

of necrosis are the same as for caudal rings (RANEY, 1941).

Marking by branding avoids the need of carrying a foreign object and seems to be less

traumatizing. Heat-branding using red-hot metal wire may make it possible to recognize a

toad at least one year after marking (CLARK, 1971). But such a method has never been ap-

plied to urodeles. Freeze branding, using wires immerged in dry ice gives interesting results

in fish (RICKER, 1956). In the anuran Ascaphus truei, scars remain readable for at least two

years (DAUGHERTY, 1976). In urodeles however, freeze-branding scars don’t persist longer

than a few weeks (GEIGER et al., 1982, NACE & MANDERS, 1982, and personal observations)

and may be traumatizing (KLEWEN, 1982). It is probable that such a regeneration capacity

of the derm also rules out the heat branding technique.

Source : MNHN, Paris

12 ALYTES 8 (1)

Marking by toe-clipping is the technique which has been most frequently used for in-

dividual marking. Formalized by MARTOF (1953) in the study of a green frog population (Rana

clamitans), such a method has been used in urodeles in the studies of TWITTY and his co-

workers about the homing capacities of Taricha rivularis (synthesis in Twrrry, 1966). If it

makes it possible to recognize a great number of individuals, it nevertheless has three major

disadvantages :

(1) Urodele toes regenerate quickly : sometimes total regeneration requires less than

one year. HEATWOLE (1961) suggested applying beryllium nitrate to inhibit regeneration. But,

besides the use of such a highly toxic chemical is dangerous, its efficiency seems to be

limited to temperatures below 18°C (EFFORD & MATTHIAS, 1969).

(2) The amputation of more than two digits involves a loss of mobility in anurans. The

probability of recapturing a marked individual is inversely related to the number of clipped

toes (CLARKE, 1972). The same observation is to be expected in urodeles.

G) News often show natural digit amputation in the field ; this may be due to pred-

atory insects like dragonfly larvae, particularly benthic Libellula larvae, or to small bivalves,

like Sphaerium, which are often found gripping a toe.

Recognition of an individual thanks to its coloration pattern means organizing a col-

lection of photographs. That efficient method may however only be used in species in which

a pattern of spots varies from one individual to another, as on the backs of Salamandra sa-

lamandra (JoLY, 1968) and of Notophthalmus viridescens (HEALY, 1974) or on the belly of Tri-

turus cristatus, T. vulgaris (HAGSTROM, 1973) and T. boscai (DIAZ-PANIAGUA, 1986).

In other cases, RAFINSKI (1974, 1977) recommended marking by autotransplantation

of a piece of ventral skin on the back and vice-versa. An Alpine newt (Triturus alpestris) may

support ten such grafts. Despite the advantage of giving permanent marks, that method has

two major drawbacks :

(1) it concerns only species with contrasted body colours ;

(2) the time needed to perform several grafts and the limited number of combinations

are handicaps for the study of numerous populations.

Lastly there are the skin staining techniques. For marking toads, WISNIEWSKI et al.

(1980) and GrTrrins et al. (1980) suggested the injection of alcyan blue into the limbs by use

of a high speed anaesthetic injector, usually used by odontologists (Panjet or Dermojet). Using

such an injector we tried to mark the belly skin of Alpine newts. The speed of projection

was so strong that the jet of dye crossed the animal’s body and came out on the other side,

but without involving the animal’s death. We estimated however the risk of organ injury to

be too high to accept that technique for urodele marking.

The intradermic injection of a dye may be made possible by scarification or tattooing,

as applied in anurans by KAPLAN (1958, 1959) or by using a spray gun (NISHIKAWA & SER-

VICE, 1988). Marks remain readable for at least two years and the entire marking procedure

takes only a few minutes. The aim of our paper is to describe testing of such a technique in

the newts Triturus alpestris and T. helvericus.

Source : MNHN, Paris

Jouy & MIauD 13

METHOD

TECHNIQUE

Its principle is to draw a pattern of spots on the animals belly. Each spot is drawn by

tattooing using an electric tattooer ; such a device is usually used by veterinarians for mark-

ing pets or farm animals. Alcyan blue and Indian ink provided colouring.

The newt is first anaesthetized by phenoxyethanol. Drawing a spot involves applying

the tattooer for about 20 s. In order to obtain a 2 mm spot, the needles must be shifted slightly

several times.

After rinsing the surplus dye (a paint brush is useful), the spot must be uniformly col-

oured and stand out well. The tracks of the tattooer needles have to be invisible. If the de-

sign is not correct, the procedure has to be repeated.

CHOSEN PATTERNS

Marking according to a site

That coding procedure makes it possible to estimate the newt flow between several sites.

One spot indicates the site where the first capture occurred. If the newt is caught again in

another site, a second spot is drawn according to a marking code. In the study of a Triturus

helveticus population inhabiting four neighbouring ponds, each pond corresponded to a spot

situated near a limb. Two spots meant that the animal had been caught at least twice in two

different ponds, which were identified. Newt flows were thus estimated by a quick marking

procedure and the organization of a complex file is unnecessary. But it does not provide in-

formation about individual behaviour.

Individual marking

It requires drawing a higher number of spots. In the case of a population of Triturus

alpestris, we used a maximum pattern of 11 points. They were placed according to three lon-

gitudinal rows, which are situated on the right side, the middle and the left side of the belly.

Each row can contain 5 spots. The middle row indicates the site of the first capture. It had

only one spot. Lateral rows are used for individual recognition. If the newt moved from one

site to another, no new spot is necessary, the animal retaining the individual code. In the

data file, the individual code consists of three numbers :

- the first corresponds to the site of first capture, and ranges from 0 to 5 ;

- the second and the third correspond to right and left rows respectively ; they are

composed of 0 to 5 figures, each ranging from 0 to 5.

Fig. 1 gives an example of the reading of such a code. Assuming the observer is able

to recognize sex, such a coding system makes it possible to identify 2048 individuals. With

only four spots per row, 512 newts may be recognized.

Source : MNHN, Paris

14 ALYTES 8 (1)

Fig. 1. - Marking patterns of newts. Left : overall view of the possible marking sites on the newt’s

belly : in the study of an Alpine newt population which was shared among 5 ponds, the two

lateral rows were used for the individual marking whereas the middle row corresponded to the

site of first capture. Right : example of a code : male newt n° 2/3/14 (a point at the second place

on the middle row, a point at the third place on the right row, two points at the first and the

fourth places on the left row).

RESULTS

PERMANENCE OF THE MARKS

In the laboratory, the suppression of hibernation, the high level of temperature (rang-

ing from 16 to 25°C) and the abundance of food constitute conditions of frequent skin re-

newal. After two years under such conditions, marked Alpine newts could still be identified,

despite a considerable decrease in the spot contrast. Under natural conditions, we suppose

that the spot remains readable for at least 3 years.

INNOCUOUSNESS

In the laboratory, none of the 4 marked newts died during the first year following the

marking procedure. One of them died after one year. We can attribute such a death to nat-

ural mortality because the animals were relatively old when marked.

Source : MNHN, Paris

Jouy & MrauD 15

Table I. - Relation between number of spots and recapture probability.

Spots number N newts marked in 1987 % recapture in 1988

2 68 19

3 140 12

4 41 22

Estimating the impact of marking on a wild population is more difficult. One possible

way may be to compare the probabilities of recapturing a newt according to the number of

spots drawn on its belly. If the method is harmful, the probability of recapturing should be

related to the number of drawn spots. But no clear relation appears between percentages of

recapture between two successive years and the number of spots (Table I).

Discussion

Electric tattooing exhibits some interesting advantages :

- it makes the recognition of a great number of individuals possible;

- the marking lasts longer than that provided by toe-clipping;

- numerous animals may be treated quickly;

- the method seems innocuous.

However it does not provide the life-long permanence of the autotransplantation

method. But the time needed for individual marking restricts the use of the latter method

to small populations. The choice of one or the other (when recording of body pattern is not

possible) depends on the kind of problem being dealt with and on the chosen experimental

design.

Such a technique may be used in the field with a portable generator to provide elec-

tricity.

Because of their ventral position, the spots drawn on the belly are not apparent when

the animal shows a normal posture. One may suppose that they do not interfere with the

visual signals acting during courtship behaviour.

The use of such a marking technique is restricted to adult animals, and the problem

of marking young news in order to estimate juvenile dispersion is still unsolved.

The tattooing technique could be improved by trying new dyes which would last lon-

ger or would increase the number of combinations by diversifying the colours used.

Useful address : we used an electric tattooing device, which may be found at Veto-

équipement , 188 avenue Roger Salengro, 69120, Vaulx-en-Velin, France.

ACKNOWLEDGEMENTS

Gly THOIRON corrected the English manuscript.

Source : MNHN, Paris

16 ALYTES 8 (1)

LITERATURE CITED

CLARK, D.R., 1971. - Branding as a marking technique for Amphibians and Reptiles. Copeia, 1971 :

148-151.

CLARKE, R.D., 1972. - The effect of toe-clipping on survival in Fowler’s toad (Bufo woodhousei fow-

leri). Copeia, 1972 : 182-185.

DAUGHERTY, C.H., 1976. - Freeze-branding as a technique for marking Anurans. Copeia, 1976 : 836-

838.

DELY, O.G., 1954. - Markierungsversuche an Frôschen. Annales Historico-Naturales Musei Nationalis

Hungarici, 1954 : 457-463.

Diaz-PANIAGUA, C., 1986. — La variaciôn del diseneno natural como método de reconocimiento indi-

vidual en Triturus boscai. Doñana, Acta Vertebrata, 13 : 175-179.

Error, LE. & MATHIAS, J.A., 1969. — À comparison of two salamander populations in Marion Lake,

British Columbia. Copeia, 1969 : 723-736.

GEIGER, A., KLEWEN, R., & NIEKIsCH, M., 1982. — Beitrag zur Tiefentemperatur-Markierung von

Amphibien im Freiland. Salamandra, 18 : 41-48.

GITTINS, S.P., PARKER, A.G., & SLATER, F.M., 1980. - Population characteristics of the common toad

(Bufo bufo) visiting a breeding site in Mid-Wales. Ÿ. anim. Ecol., 49 : 161-173.

HAGsTROM, T., 1973. — Identification of newt specimens (Urodela Triturus) by recording the belly

pattern and a description of photographic equipment for such registration. Brit. J. Herpetol., 4 :

321-326.

Hay, W.R., 1974. - Population consequences of alternative life histories in Notophthalmus v. viri-

descens. Copeia, 1974 : 221-229.

HEATWOLE, H., 1961. - Inhibition of digital regeneration in salamanders and its use in marking indi-

viduals for field studies. Ecology, 42 : 593-594.

Jouy, J. 1968. - Données écologiques sur la Salamandre tachetée Salamandra salamandra (L.). Ann.

Sc. Nat. Zool. Paris, 10 : 301-366.

KAPLAN, H.M., 1958. - Marking and banding frogs and turtles. Herpetologica, 14 : 131-132.

a 1959, — Electric tattooing for permanent identification of frogs. Herpetologica, 15 : 26.

KLEWEN, R., 1982. - Beitrag zur Tiefentemperatur-Markierung von Amphibien im Freiland. Sala-

mandra, 18 : 342-347.

MarTIN, E., JoLy, P. & BOVET, P., 1989. — Diel pattern of activity in the Alpine newt (Triturus al-

pestris, Amphibia Urodela) during the aquatic phase. Biol. Behav., 14 : 116-131.

MarTor, B.S., 1953. - Territoriality in the green frog. Rana clamitans. Ecology, 34 : 165-174.

NACE, G.W. & MANDERS, E.K., 1982. - Marking individual amphibians. 7. Herpetol., 16 : 309-311.

NisHikAWA, K.C. & SERVICE, P.M., 1988. — À fluorescent marking technique for individual recogni-

tion of terrestrial salamanders. Ÿ. Herpetol., 22 : 351-353.

RAFINSKI, J.N., 1974. — Studies on the genetic structure of the Alpine newt, Triturus alpestris (Laur.)

populations. Acta Biol. Cracov., Ser. Zool., 17 : 51-58.

ee 1977. - Autotransplantation as a method for permanent marking of Urodele Amphibians (Am-

phibia, Urodela). 7. Herpetol., 11 : 241-242.

RANEY, E.C., 1940. - Summer movements of the bull-frog, Rana catesbiana Shaw, as determined by

the jaw-tag method. Amer. Midl. Nat., 23 : 733-745.

us 1941. — Attempts at tagging small salamanders in life history studies. Science, 93 : 578.

RickER, W.E., 1956. - Use of marking animals in ecological studies : the marking of fish. Ecology, 37 :

666-670.

Twirry, V.C., 1966. — Of Scientists and Salamanders. San Francisco, Freeman & Co. : 1-178.

WisNiEWski, P.J., PAULL, L.M., MERRY, D.G. & SLATER, F.M., 1980. — Studies on the breeding mi-

gration and intramigratory movements of the common toad (Bufo bufo) using Panjet dye-

marking techniques. Brit. J. Herpetol., 6 : 71-74.

Corresponding editor : Alain DuBois.

Source : MNHN, Paris

Alytes, 1989-1990, 8 (1) :17-21. 17

Studies on the biology of the tree-frog Hyla arborea

during the breeding season in North Western Italy

(Amphibia, Anura, Hylidae)

Ivana PAVIGNANO

Via San Carlo, 8, 10010 Piverone (TO), Italy

Some parameters on the breeding phenology and larval period of Hyla arborea

during 1988 and 1989 spawning seasons were investigated in six ditches in North West-

ern Italy.

Spawning period lasted from 31 to 36 days (from 15 April to 22 May) with two

peaks of oviposition (one in April and one in May), and an interruption ranging from 5

to 10 days. Number of egg masses laid was positively correlated with water tempera-

ture.

The mean number of eggs estimated in 100 egg masses was 59 + 23 SD. 93%

of observed eggs were fertile. The percentage of unfertile eggs was significantly greater

in larger masses.

Larval period ranged from the first days of May to the middle of July, with an

average duration of 66-67 days.

INTRODUCTION

Although the reproductive biology of European Anuran Amphibians has been and

continues to be object of numerous studies, only a few publications refer to the tree frog

Hyla arborea (Linnaeus, 1758). This species shows a wide distribution in Europe (ARNOLD

& BURTON, 1978), and in the recent past it was rather common. Now FH. arborea is a vul-

nerable species in Northern Europe (STUMPEL & HANEKAMP, 1986) as well as in North

Western Italy, where it shows a localized distribution (PAVIGNANO & GIacoMA, 1986).

H. arborea is a rather terrestrial species; it only lives in aquatic habitats during the re-

productive period. The aquatic habitat consists of ponds in grasslands and marshes. Extent

of aquatic vegetation cover, surrounding terrestrial habitat, level of human interference are

discriminant parameters for this species in choosing breeding sites (PAVIGNANO et al., 1989

a). Alteration of these ecological parameters is probably the cause of current sporadic dis-

tribution of species. Where the habitat is suitable, H. arborea occurs usually with a numer-

ous breeding population (PAVIGNANO et al., 1989 b); this is probably due to the several re-

productions during the breeding season. The reproductive period of Hyla is in fact a long

one (DIAZ-PANIAGUA, 1986; GARTON & BRANDON, 1975; PERRIL & DANIEL, 1983); in

Northern Italy two spawn periods of H. arborea have been observed (PAVIGNANO, 1990).

There is no specific work on reproductive biology of H. arborea; in this study I in-

vestigated some parameters of the breeding phenology and larval period of H. arborea in six

Source : MNHN, Paris

18 ALYTES 8 (1)

ponds in North Western Italy during the years 1988 and 1989. Particular emphasis was given

to gathering precise information concerning the duration of spawning period, the number of

eggs per clump, the number of fertile eggs, and duration of metamorphosis of tadpoles.

MATERIALS AND METHODS

The study area has been described in detail in a former paper (PAGIGNANO, 1989) and

consists of a series of temporary draining ditches in fields, of which six sites were populated

by H. arborea.

Observations were made every day during the breeding period.

For the estimation of the total spawn, all the egg masses at each pond were counted

during every visit and the newly laid egg masses were marked on maps. The total number

of eggs per clump was calculated by direct count from randomly selected egg masses and in

each clump the number of fertile eggs was determined by stereo-microscope.

Tadpole populations in the ponds were sampled once every two days. The larvae, cap-

tured for each sample with a dipnet in different location in the ponds, were counted and

larval stages (according to GOSNER, 1960) of each species was determined. Tadpoles were

then released. À small number of eggs and tadpoles was fixed in 60 % ethanol for micro-

scopical observation of exact larval stage, and then photographed.

Maximum and minimum water and air temperature was recorded daily by a maxi-

mum/minimum thermometer placed in each pond, together with general conditions (i.e. if

it had rained or not, etc.). The local climate is classified as continental with rainfall maxima

in the middle of April (mean = 113mm) and November (mean = 79 mm); minima occurs

in January (mean = 32 mm) and July (mean = 79 mm) (DURIO et al., 1983).

RESULTS

In Northern Italy, H. arborea begins to spawn between the middle and end of April.

During the two years of my observations the spawning season lasted 36 days in 1988 (from

15 April to 20 May) and 31 days in 1989 (from 22 April to 22 May). In the 1988 breeding

season a total of 404 laid egg masses (mean = 67.33 + 22.06 SD for each pond) was re-

corded, and 354 (mean = 59.00 + 27.63 SD) in 1989. No oviposition occurred when min-

imum water temperature was below 8°C and air temperature below 6°C. During the spawn-

ing period average water temperature varied between 10-22°C. Number of egg masses laid

was positively correlated with average water temperature (r = 0.469, p < 0.005). In both

years there were two peaks of oviposition (fig. 1) : 15 to 21 April and 15 to 18 May for 1988

(in those periods 82% of egg masses were laid, 60% of which in April) ; 25 to 29 April and

13 to 17 May for 1989 (84% of egg masses, 59% of which in April). The maximum spawning

activity took place in April. An interruption of oviposition for about ten days in 1988 (5 to

14 May) and five days in 1989 (9 to 13 May) was observed. The laid egg masses were ob-

served either before or after rainy days; no significant correlation was recorded between the

number of laid egg masses and rain.

Source : MNHN, Paris

PAVIGNANO 19

lis.

]

(1)

—|

8 8 5

NUMBER OFEGG MASSES

| is dll.

dll uns Nb

Fig. 1. - Number of egg masses per day laid in 1988 (a) and 1989 (b) spawning seasons.

The egg masses were laid in small groups, near the pond edges, at a depth of 5 to 10

cm, fixed on the vegetation. The shape of the egg masses was oval, the colour of the eggs

was pale with grey animal hemisphere.

In the first days of October a further breeding activity was observed, revealing itself

only by male calls. Males called from aquatic vegetation, but in a shorter and less frequent

way than in April.

The number of eggs per mass determined by counting in 100 randomly selected egg

masses is shown in Table I. The mean number of eggs per mass in the two years was 59 (+

23 SD) eggs. The difference in the mean number of eggs in the two years was not statisti-

cally significant (Student’s t = 1.75, p > 0.05). 60% of egg masses included 60-100 eggs.

Very small egg masses (20-30 eggs) were laid at the end of the spawning period and com-

prised 30% of total; there were also a few very large egg masses of more than 100 eggs com-

prising 5% of total. 93% of observed eggs were fertile; the percentage of unfertile eggs was

significantly (Student’s t = 0.78, p < 0.05) greater in large masses. In masses with 20-30

eggs, they were all fertile.

Table I. - Number of eggs per mass observed in 1988 and 1989 spawning seasons (*number

of fertile eggs).

Year N Max Min M SD

1988 50 125 20 57 28

*120 20 54 27

1989 50 118 25 62 18

*110 25 56 12

Total 100 125 20 59 23

*120 20 55 20

Source : MNHN, Paris

20 ALYTES 8 (1)

Average larval period duration was of 67 days + 1.02 SD in 1988 and 66 days + 1.32

SD in 1989 (from the first days of May to the middle of July), from the first tadpoles leaving

stage 25 (last embryonic stage according to GOsNER, 1960) until the last tadpoles completed

their metamorphosis. Larval populations were composed of individuals at different devel-

opment stages, because the various egg masses were laid in different times. There was a tem-

poral overlap, lasting about 36 days, between tadpoles of different age and size classes. When

the development of tadpoles from first ovipositions was between stage 34 and 40, there were

tadpoles between stage 28 and 32; when the tadpoles from first ovipositions completed their

metamorphosis, there were still tadpoles at stages 36 and 40.

DIscUSsION

The reproduction of Amphibians is usually related to weather factors, such as tem-

perature and rainfall. H. arborea belongs to the type of Anurans which have a long spawning

season; a first consequence of this is a relation with weather conditions. Although there were

two oviposition peaks (about one month one after the other), this species seemed not to be

particularly dependent on the rainfall, while no oviposition below 8°C water and 6°C air tem-

perature was observed, and the observed oviposition was well correlated with temperature.

Breeding activity not dependent on rain has also been observed in other species of Hyla (DIaz-

PANIAGUA, 1986).

A second consequence of the long spawning period was the temporal overlap of larval

populations composed of individuals at different ages and sizes. The length of the larval pe-

riod of Hyla is related to water temperature (DIAZ-PANIAGUA, 1986; STUMPEL & HANEKAMP,

1986); an increase of 5 — 8°C water temperature causes a shortening of about 10 days in H.

arborea’s larval period (PAVIGNANO, 1990).

The variation in eggs number per mass shown by my results is in good agreement with

LanZA (1983). The percentage of fertile eggs was related to the number of eggs per mass.

Probably, as it happens in other Anuran species (HAAPENEN, 1982; SOFIANIDOU & KYRIA-

KOPOULOU-SKLAVOUNOU, 1983), the number of eggs laid by a female is dependent on its size.

Quantitative information concerning the number and size of eggs from individual females,

and relations with body size require further investigations.

LITERATURE CITED

ARNOLD, E.N. & BURTON, J.A., 1978. — À field guide to the Reptiles and Amphibians of Britain and Eu-

rope. London, Collins : 1-267.

Draz-PANIAGUA, C., 1986. — La reproduccién de Hyla meridionalis en el suroest de Espana. Doñana

Acta Vertebrata, 13: 5-20.

Durio, P., Mori, D. & PEROSINO, G.C., 1983. — Aspetti limnologici del lago di Candia. Riv. Piem.

St. Nat., 4: 137-169.

GARTON, J.S. & BRADON, R.A., 1975. - Reproductive ecology of the green tree frog Hyla cinerea in

Southern Illinois. Herpetologica, 31: 150-161.

Gosner, K.L. 1960. — A simplified table for staging anuran embryos and larvae with notes on identi-

fication. Herpetologica, 16: 183-190.

Source : MNHN, Paris

PAVIGNANO 21

HAAPANEN, A., 1982. — Breeding of the common frog (Rana temporaria L.). Ann. Zool. Fennici, 19:

75-79.

Lanza, B., 1983. - Anfbi, Rettili (Amphibia, Reptilia). Guide per il riconoscimento delle specie animali

delle acque interne italiane. Consiglio Nazionale delle Ricerche, 27: 1-196.

PAVIGNANO, I., 1989. — Rilevamento dei siti e analisi del ciclo riproduttivo degli Anfibi presenti in un-

’area dell’Anfiteatro Morenico d’Ivrea. Boll. Mus. reg. Sci. nat. Torino, 7 (2), in press.

PAVIGNANO, I., 1990. — Niche overlap in tadpole populations of Pelobates fuscus insubricus and Hyla

arborea at a pond in North Western Italy. Boll. Zool., 57 (1), in press.

PAVIGNANO, I. & GIacoma, C., 1986. — Osservazioni sulla distribuzione e sul comportamento ripro-

duttivo degli Anfibi present in un’area della pianura piemontese. Riev. Piem. St. Nat., 7: 153-171.

PAVIGNANO, I., MATTIOLI, M., PIGNONE, A. & GracoMa, C., 1989 a. - Censimento di anfibi in un’area

della cintura torinese. Riv. Piem. St. Nat., 10: 183-194.

PAVIGNANO, L., GIACOMA, C. & CASTELLANO, S., 1989 b. — Multivariate analysis of amphibian com-

munities in North Western Italy. Amphibia-Reptilia, submitted.

PeRRIL, S.A. & DANIEL, R.E., 1983. - Multiple egg clutches in Hyla regilla, H. cinerea and H. gratiosa.

Copeia, 1983: 513-516.

SoFIANIDOU, T.S. & KYRIAKOPOULOU-SKLAVOUNOU, P., 1983. — Studies on the biology of the frog Rana

dalmatina Bonaparte during the breeding season in Greece (Amphibia : Anura : Ranidae). Am-

bhibia-Reptilia, 4: 125-136.

STuMPEL, A.H.P. & HANEKAMP, G., 1986. — Habitat and ecology of Hyla arborea in the Netherlands.

In: Z. RotEK (ed.), Studies in Herpetology, Prague, Charles University Press: 409-412.

Corresponding editor: Pierre JOLY.

Source : MNHN, Paris

Alytes, 1989-1990, 8 (1): 22.

Dates de publication du journal Alytes

(1989)

Alain Dugois

Laboratoire des Reptiles et Amphibiens,

Muséum National d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

Cette liste fait suite à celles que nous avons déjà publiées (Dugois, 1988, 1989) pour

les années 1982-1987, et a été préparée de la même manière. A ce sujet, il nous faut signaler

ici une erreur qui s’est glissée dans la première de ces listes (DuBois, 1988) : le fascicule 4

du volume 1 d’Alytes a été publié en fait le 30 décembre 1982, et non pas le 31 décembre

1982.

Volume Fascicule Pages Date figurant Date réelle

sur le fascicule de publication

7: 2 45-76 Juin 1988 27 février 1989

7 3 77-124 Septembre 1988 11 juin 1989

7 4 125-168 Décembre 1988 3 novembre 1989

3-4 Index i-xii 1984-1985 3 novembre 1989

RÉFÉRENCES BIBLIOGRAPHIQUES

Dusoïs, A., 1988. — Dates de publication du journal Alytes (1982-1987). Alytes, 6: 116.

—— 1989. - Dates de publication du journal Apres (1988). Alpes, 7: 75.

Source : MNHN, Paris

Alytes, 1989-1990, 8 (1): 23. 23

International Society for the Study

and Conservation of Amphibians (ISSCA)

The ISSCA, International Society for the Study and Conservation of Amphibians

(International Society of Batrachology), a non-profit association, was founded in Paris in 1988.

It is now fastly growing in all parts of the world, under the Presidence of Raymond F.

LAURENT (Tucumän, Argentina).

Its aims are to contribute worldwide to the study and knowledge of Amphibians, to

their conservation and that of their environment, to the establishment, reinforcement and

facilitation of communication between batrachologists, and to the promotion of batrachology

as a distinct scientific discipline. The ISSCA will organize international symposia and be active

at international level in the fields of research, conservation and information relating to

Amphibians.

The ISSCA is now the publisher of Alytes (International Journal of Batrachology) and

Circalytes (Information Bulletin). The journal Alytes now accepts papers in English, French

and Spanish and now publishes color photos, despite a subscription price which has remained

very low. The more subscribers it has, the more papers and pages it can publish for the same

subscription price.

The ISSCA is opened both to individual members from all countries in the world, and

to local sections (from various countries or regions) and thematical or working groups

(specialized on given subjects). These sections and groups retain their full independence for

all their activities, but they are entitled to use the journals Alytes and Circalytes for their

internal and external communications and for the publication of their works.

Subscription rates for 1989 and 1990 are as follows (preferably by direct postal transfer

to our postal account, “ISSCA”, N° 1-398-91-L, Paris, or in FF or in U.S. $, by cheques

payable to “ISSCA”, sent to the address above; if you use “Eurocheques”, please add 20.00

FF or 4.00 $ to the sum chosen below):

Individual direct subscription to ISSCA + Alytes + Circalytes :

230.00 FF or 46.00 $.

Individual subscription to ISSCA + Alytes + Circalytes for members of groups

or sections of the ISSCA :

195.00 FF or 39.00 $.

Individual subscription to Alytes alone :

150.00 FF or 30.00 $.

Institutional subscription to Alytes :

300.00 FF or 60.00 $.

Source : MNHN, Paris

Alytes, 1989-1990, 8 (1): 24.

Application for membership of the ISSCA and/or

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subscription to Alvtes

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Send this application to : ISSCA, Laboratoire des Reptiles et Amphibiens, Muséum national

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Source : MNHN, Paris

AINTES

International Journal of Batrachology

published by the ISSCA

EDITORIAL BOARD FOR 1989

Chief editor : Alain Dugois (Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire

naturelle, 25 rue Cuvier, 75005 Paris, France).

Deputy editor : Pierre JoLy (Laboratoire de Biologie animale et Ecologie, Université Claude Bernard

Lyon I, 69622 Villeurbanne Cedex, France).

Other members of the Editorial Board : Jean-Louis AmIEr (Yaoundé, Cameroun) ; Stephen D. Busack

(Ashland, U.S.A.) ; Günter GOLLMANN (Wien, Austria) ; Tim HaLipay (Milton Keynes,

United Kingdom) ; William R. HEYER (Washington, U.S.A.) ; Walter HôDL (Wien, Austria) ;

Milos KaLezrC (Beograd, Yugoslavia) ; Raymond F. LAURENT (Tucumén, Argentina) ; Borja

Sanc1z (Madrid, Spain) ; Dianne B. SEALE (Milwaukee, U.S.A.).

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 ar-

ticles, comments and replies.

The title should be followed by the name(s) and address(es) of the author(s). The text should be

organised 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 :

KaLezié, M.L., Duxié, G., CRNOBRNJA, J. & TVRTKOVIÉ, N., 1987. — On the Triturus vulgaris schrei-

beri problem : electrophoretic data. Alytes, 6 : 18-22.

— when in a multi-authors book :

GarCIA-PARIS, M. & MARTIN, C., 1986. — Amphibians of the Sierra del Guadarrama (1800-2430 m al-

titude). In : Z. RoëEK (ed.), Studies in herpetology, Prague, Charles University Press : 135-138.

— when a book :

BOULENGER, G.A., 1882. - Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British

Museum. London, Taylor & Francis : i-xvi + 1-503, pl. I-XXX.

Manuscripts should be submitted in triplicate to Alain Dugois (address above) if dealing with

amphibian systematics, biogeography, evolution, genetics or developmental biology, or to Pierre JoLY

(address above) if dealing with amphibian ecology, ethology, life history or physiology.

Acceptance for publication will be decided by the editors following review by at least two referees.

No page charges are requested from the author(s), but the publication of color photographs is

charged. For each published paper, 25 free reprints are offered by Alytes to the author(s). Additional

reprints may be purchased by multiples of 25.

Published with the support of

the Muséum national d'Histoire naturelle (Paris, France)

and of the Société Batrachologique de France.

Directeur de la Publication : Alain DuBois.

Numéro de Commission Paritaire : 64851.

9 JUIL. 1990

Alytes, 1989-1990, 8 (1): 1-24.

Contents

Alain Dugoïs & Pierre JoLY

DE FE TE Ro Rs PE EE PRET POUR 1

Jean-Louis AMIET

Images d’Amphibiens camerounais.

I. Sacs vocaux et postures de chant 4... 5

Pierre Jouy & Claude MrAuD

Tattooing as an individual marking technique in urodeles 11

Ivana PAVIGNANO +

Studies on the biology of.the tree-frog Hyla arborea during the breeding

season in North Western Italy (Amphibia, Anura, Hylidae) 17

Alain Dugols

Dates de publication du journal Alytes (1989) :.............................. 22

International Society for the Study and Conservation of Amphibians

(ÉRROAE RRA e T PA RS bre ÉD TES ES GE D DDR ERP 23

Application for membership of the ISSCA and/or subscription to

PET CRETE DR RE CN HR Po Re AIS EN PACE TOO TE 24

Imprimerie Fotek, St.-Niklaas, Belgique.

Dépôt légal: 2ème trimestre 1990.

Source : MNHN, Paris