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 1992

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

General Secretary: Alain DUBoIs (Paris, France).

Treasurer: Dominique PAYEN (Paris, France).

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

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

Deputy Secretary. outside Europe: David B. WAKE (Berkeley, U.S.A.).

Deputy Treasurer, outside Europe: Janalee P. CALDWELL (Norman, U.S.A.).

Other members of the Board: Jean-Louis FISCHER (Paris, France); Roy W. McDiarmip (Washington,

U.S.A.); James I. MENZIES (Boroko, Papua New Guinea).

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Source : MNHN, Paris

AIMTES

INTERNATIONAL JOURNAL OF BATRACHOLOGY

December 1992 Volume 10, N° 4

Alytes, 1992, 10 (4): 113-116. 113

Sperm competition in European water frogs

Leszek BERGER & Mariusz RYBACKI

Research Centre for Agricultural and Forest Environment,

Polish Academy of Sciences, Bukowska 19, 60-809 Poznañ, Poland

Sperm from Rana lessonae was mixed with sperm from Rana ridibunda or

Rana kl. esculenta. Eggs from Rana lessonae or Rana ridibunda were

introduced into sperm suspensions producing progeny with Rana lessonae,

Rana ki. esculenta, or Rana ridibunda phenotypes. Spermatozoa with Rana

lessonae genomes always fertilized many more eggs than those with Rana

ridibunda genomes.

Bibliothèque Centrale Muséum

3001 00111599 6

Central Europe is inhabited by three forms of water frogs: Rana ridibunda Pallas, 17/1

and Rana lessonae Camerano, 1882, two Mendelian species, and Rana kl. esculenta Linné,

1758, a hybridogenetic hybrid between the two (BERGER, 1988 a; GRAF & POLLS PELAZ,

1989). Gametes of hybrids usually contain genomes of one of their parental species and

genomes of the other species are excluded from the germ line premeiotically (UZZELL,

Horz & BERGER, 1980; TUNNER & HEPPICH, 1981). In the Poznañ region of Poland the

sperm of Rana kl. esculenta contains the Rana ridibunda genome (UzZELL & BERGER, 1975).

INTRODUCTION il

In Central Europe, water frogs usually breed in May with the males appearing first

at the breeding site. Their distribution at the breeding arena is not random (BLANKEN-

HORN, 1977): in mixed Rana lessonae — Rana esculenta populations (L-E system, UZZELL

& BERGER, 1975) most Rana kl. esculenta males gather in the center of the arena, while Rana

lessonae males gather along the margins. AII males compete for females, but Rana lessonae

males have a higher probability for successful pair formation. AIl pairs in amplexus wander

to the center of the arena (BLANKENHORN, 1977; BERGER & RYBACKI, pers. obs.) where they

release gametes and further competition between spermatozoa may take place.

Our knowledge of sperm competition in Amphibia is confined to species with internal

fertilization (HALLIDAY & VERRELL, 1984; HOUCK & SCHWENK, 1984). There are no pub-

lished data on sperm competition between Amphibia with external fertilization, although it

is likely to occur in foam-nesting frogs of the family Rhacophoridae (FUKUYAMA, 1991).

Source : MNHN, Paris

114 ALYTES 10 (4)

MATERIAL AND METHODS

Five females and ten males provided gametes for crossing experiments (Table I)

carried out according to standard methods (BERGER, 1988 b) during the breeding time of

water frogs. Males were killed and both testes were carefully crushed in equal volumes of

aged tap water in separate petri dishes;, spermatozoa were very mobile and always in

masses. The testes of Rana ridibunda were approximately three times larger (about 120

mm) than those of Rana lessonae (about 40 mm°), and those of Rana kl. esculenta were

intermediate. Sperm suspensions of two selected males were poured into one dish and

mixed, and ovulated eggs of Rana lessonae or Rana ridibunda were then squeezed into this

mixture. In cross 6x/91 we used 131 eggs from Rana lessonae and 141 eggs from Rana

ridibunda. When larvae reached stage 25 (SHUMWAY, 1940), numbers of survivors were

assessed and many of them were reared further in aquaria and basins outdoors. After

metamorphosis, all progeny were killed and preserved in 3 % formalin. Their genotype was

inferred from morphological phenotype (BERGER, 1964; BERGER, ROGUSKI & UZZELL,

1978).

Table I. — Viability and genomic composition of water frog progeny.

Parents. — N/y: successive number of female (N) and year of laying the eggs (y);

letters designate the male (or males) used in crosses with the same female; les: Rana

lessonae; rid: Rana ridibunda; esc: Rana kl. esculenta.

Tadpoles. — N: number of individuals; survived: survived to the end of

experiment; phenotype of the progeny: LL, lessonae, RL, esculenta, RR, ridibunda.

| Parents Progeny ___ Rearing of tadpoks

ist Stage Sur Phenotype Ratio of

N/y Mothers Fathers Eggs 25 ON vived LL RL RR P

age N genomes

% % Number Re

2/TTa les! lesl+ridl 945 980 886 185 149 129 20 1

2911 less ls3+rid2 241 992 884 113 87 59 28 2:1

39e ridl les3ærid2 349 99.1 974 140 123 106 17 6:1

3/90k _les2 les2+escl 145 924 221 32 30 28 2 14:1

2/91m_les3 les3+esc2 287 99.3 767 220 153 149 4 37:1

3/14 ridl les3+ese2 331 99.1 970 120 114 114 114:0

lg ridl lesd+esc3 191 99.5 859 164 127 123 4 31:1

Gx91 lest+ridl lesdtrid3 272 1000 94.5 158 147 56 91 19

3P91b ridl rid3 103 1000 971 —

AUSEUM

PARIS

Source : MNHN, Paris

BERGER & RYBACKI 115

RESULTS AND DISCUSSION

Egg cleavage and zygote viability was very high in all crosses except one (cross 3/90k).

The progeny in all crosses contained many more individuals fathered by Rana lessonae

than those fathered by Rana ridibunda or Rana kl. esculenta. Spermatozoa with ridibunda

genomes (R) fertilized eggs with /essonae genomes (L) more frequently than eggs with

ridibunda genomes (R). Rana ridibunda individuals (RR) were not numerous or even absent

in the progeny.

Cross 6x/91 exemplifies this phenomenon. Given nearly equal numbers of ova

(131:141) and assuming equal chances of fertilization by sperm of Rana lessonae and Rana

ridibunda, the expected ratio of LL:RL:RR offspring in this cross should be approximately

1:2:1; there were, however, only progeny with /essonae (LL) and esculenta (RL) phenotypes

in the 147 individuals, in a ratio of about 1:2 (56:91). Cleavage of all eggs and very high

viability of offspring in cross 6x/91 suggests that individuals which might have received a

R genome each from both parents were not eliminated during development. The very high

viability of zygotes derived from the same Rana ridibunda pair, which developed in a

separate petri dish (cross 3/91b), shows that both parents were normal and capable to

create Rana ridibunda (RR) individuals. Scarcity of RR progeny is even more surprising

because Rana ridibunda males and their testes were about three times larger than Rana

lessonae ones and contained about three times more gametes (BERGER & RyBACKI,

unpubl.). At present we cannot explain the causes underlying this phenomenon, but we

assume that it is connected with the presence of Rana lessonae gametes in the sperm

suspension.

Our data from all other crosses corroborate this conclusion. In three crosses (3/91c,

3/91d, 3/91g), about 50 % of ridibunda (RR) individuals are expected for equal numbers

of spermatozoa with lessonae (L) and ridibunda (R) genomes and random fertilization, but

the RR genotype appeared in only two crosses and in low frequency. Activity of

spermatozoa with R genomes apparently is connected with their origin: spermatozoa of

Rana ridibunda males fertilized more eggs (13.3-32.2 %) than spermatozoa of Rana kl.

esculenta males (0-6.7 %) which also contain ridibunda (R) genomes (BERGER, 1988 a).

CONCLUSION

The European water frogs are promising objects for studies on gamete competition.

AI crosses between Mendelian forms in this group are easily achieved and give viable

progeny with characters intermediate between parental types (BERGER, HOTZ & UZZELL,

unpubl.). It is easy to determine their genetic origin from morphology (BERGER et al., 1978,

1982, 1986) or from electrophoresis (HoTz & UZZELL, 1982). We anticipate that further

studies will promote understanding of the mechanisms responsible for gamete competition.

Source : MNHN, Paris

116 ALYTES 10 (4)

ACKNOWLEDGEMENTS

We thank Dr. H. G. TUNNER, Dr. T. R. HALLIDAY, and especially Dr. H. Horz, for reading the

manuscript and making important suggestions.

LITERATURE CITED

BERGER, L., 1964. — Is Rana esculenta lessonae Camerano a distinct species? Ann. Zool., 22: 45-61.

1988 a. — On the origin of genetic systems in European water frog hybrids. Zool. Polon., 35: 5-32.

1988 b. — Principles of studies of European water frogs. Acta Zool. Cracov., 31: 563-580.

BerGER, L., Horz, H. & RoGuski, H., 1986. — Diploid eggs of Rana esculenta with two Rana

ridibunda genomes (Amphibia, Salientia, Ranidae). Proc. Acad. nat. Sci. Phila., 138: 1-13.

BerGER, L., ROGUSkI, H. & Uzzeuc, T., 1978. — Triploid F, progeny of water frogs (Rana esculenta

complex). Folia Biol. (Krakôw), 26: 135-152.

BERGER, L., UZZeLL, T. & Horz, H., 1982. — Crossing experiments between some western Palearctic

species of water frogs (Salientia, Ranidae). Vertebr. hung., 21: 33-45.

BLANKENHORN, H. J., 1977. — Reproduction and mating behavior in Rana lessonae — Rana esculenta

mixed population. /n: D. H. TAYLOR & S. I. GUTTMAN (eds.), The reproductive biology of

amphibians, Plenum Press, New York and London: 389-410.

FukuYAMA, K., 1991. — Spawning behaviour and male mating tactics of a foam-nesting treefrog,

Rhacophorus schlegelii. Anim. Behav., 42: 193-199.

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, AÏbany,

The New York State Museum: 289-302.

HALLIDAY, T. R. & VERRELL, P. A., 1984. — Sperm competition in amphibians. /n: R. L. SMITH (ed.),

Sperm competition and the evolution of animal mating systems, Academic Press: 487-508.

Horz, H. & UzzeLi, T., 1982. — Biochemically detected sympatry of two water frog species: two

different cases in the Adriatic Balkans (Amphibia, Ranidae). Proc. Acad. nat. Sci. Phila., 134:

50-79.

Houck, L. D. & ScHwenk, K., 1984. — The potential for long-term sperm competition in a

plethodontid salamander. Herpetologica, 40: 410-415.

SHumway, W., 1940. — Stages in the normal development of Rana pipiens. I. External form. Anat.

Record, Philadelphia, 78: 139-144.

TUNNER, H. G. & HEPPICH, S., 1981. — Premeiotic genome exclusion during oogenesis in the common

edible frog, Rana esculenta. Naturwiss., 68: 207-208.

UzzeLi, T. & BERGER, L., 1975. — Electrophoretic phenotypes of Rana ridibunda, Rana lessonae, and

their hybridogenetic associate, Rana esculenta. Proc. Acad. nat. Sci. Phila., 127: 13-24.

Uzzeui, T., Horz, H. & BERGER, L., 1980. — Genome exclusion in gametogenesis by an interspecific

Rana hybrid: evidence from electrophoresis of individual oocytes. J. exp. Zool., 214: 251-259.

Corresponding editor: Günter GOLLMANN.

Source : MNHN, Paris

Alytes, 1992, 10 (4): 117-129. 117

The amphibian fauna of the French

Upper-Rhône floodplain.

The Lavours marsh and the Jons sector

Pierre JOLY

URA CNRS 1451 Ecologie des Eaux douces et des grands Fleuves,

Université Claude Bernard Lyon 1, 43 bd du 11 novembre 1918,

69622 Villeurbanne Cédex, France

The amphibian fauna of two sectors of the French Upper-Rhône was

investigated at 38 sites. These sectors and sites were chosen to represent the

braided or anastomosed floodplain. Among the 14 taxons inhabiting this

district, 11 bred within the floodplain. The most constant taxons were Triturus

helveticus, Rana dalmatina and Rana synkl. esculenta. Bufo calamita, Hyla

arborea and Pelodytes punctatus were observed mainly in temporary sites in

the vicinity of the active channel. Bombina variegata occurred in shallow

overforested sites at the border of the floodplain. The distribution of taxons is

discussed in connection with the patch dynamics concept.

INTRODUCTION

Although European amphibian communities in agrosystems are relatively well known

(STRUBOSCH, 1979; PAVIGNANO, GIACOMA & CASTELLANO, 1990), information on those that

inhabit floodplains is still scarce. Floodplains are of great interest in understanding

determinants of amphibian occurrence. In floodplain sectors where the geomorphology of

the valley makes meandering possible, the diversity of aquatic sites may be very high

(Amoros et al., 1987, 1988). Diversity concerns both site configuration and the modes of

water retention. Amphibian breeding sites are mainly semi-aquatic or permanent, with

standing water or low current velocity. The semi-aquatic sites may originate according to

three different processes (AMOROS, RICHARDOT-COULET & PAUTOU, 1982). Firstly, depres-

sions may be formed in pebble beds or in alluvial side bars by the mechanical erosion of

floods (or by human excavation). The bottom of such sites is not silted and water is

retained for rather short periods (some weeks). The second group includes old oxbow lakes

(“parapotamon” of AMOROS, RICHARDOT-COULET & PAUTOU, 1982), which have been

partially filled with alluvium or autogenic organic matter. Their silted bottom retains water

for several months. The third group includes marshes, which are the last stage of transition

towards terrestrial habitat. Depending on water table level, water may be found more or

less permanently in local depressions. Oxbow lakes, man-made excavations for peat

extraction or drains are permanent aquatic sites, which may host breeding amphibians.

Source : MNHN, Paris

118 ALYTES 10 (4)

The terrestrial surroundings of these different kinds of sites also vary greatly (PAUTOU et

al., 1985). In the French Upper-Rhône alluvial valley (south-eastern France), sites situated

in pebble beds are mainly surrounded by moorland with Phalaris arundinacea, Salix alba

and Ainus incana. Several species of Salix, Alnus incana, Populus nigra and Quercus robur

characterize the alluvial forest on the side bars. The marshes are most often peat-moors

colonized by A/nus glutinosa. The main system investigated in the present study was the

Lavours marsh. We compared the amphibian fauna of this marsh with that of more

juvenile sites situated in the braided sector of Jons. In this sector, the sites sampled were

a connected oxbow lake and more or less temporary ponds located in alluvial side bars

near the stream.

MATERIAL AND METHODS

STUDY DITES

The Lavours marsh

The Lavours marsh (fig. 1) is located 70 km downstream from Geneva. It occupies a

1200 ha hollow between the rivers Rhône and Séran just upstream from their confluence.

The present description briefly summarizes the detailed ecological and historical studies by

AÏN & PAUTOU (1969), BRAVARD (1987) and PAUTOU et al. (1991). The marsh originated

in a post-glacial lake. This lake was progressively filled with sediment. When the Rhône

breached through the moraine at Massignieu, the water level decreased, leading to a great

productivity of the vegetation along the lake shores. Both sediment and peat then filled the

northern part of the lake (the peat layer is 8 metres thick in the centre of the marsh). The

lake was reduced until it reached its present shape, that of the present Lake Bourget, south

of the marsh. Water flow from the river to the marsh was checked by man-made

embankments, mainly erected during the XIXth century. The marsh is now overflowed

only in decennial floods, when the discharge of the Rhône reaches 1500 m°/s. The flow

regime is of nival type, with high water in spring, whatever the amount of rain. Water from

the Séran is then pushed back by the high level of the Rhône and floods the western border

of the marsh. Such repeated floods have formed two alluvial side bars. The Rhône alluvial

side bar is mainly composed of argillous alluvium and may be up to 1 km wide. The Séran

side bar is more silty and its width ranges from 100 to 200 m. The water table is level with

the peat surface in the middle of the marsh. The water mainly originates from the

neighbouring mountains, as revealed by numerous exurgences and resurgences. The

vegetation is strongly influenced by the depth of the water table and the substrate

composition. Crops are grown in the north, in the vicinity of the Rhône. A remnant of the

original fluvial forest occupies the southern part of the Rhône side bar (Lavours Forest).

A typical alluvial forest occupies the banks of the Séran. This forest is characterized by

Alnus glutinosa, Acer campestre, Fraxinus excelsior. Some Quercus robur may be found on

the outcrops. To the east, the forest is gradually replaced by a moor with Salix cinerea,

Betula verrucosa and Alnus glutinosa. The moor then becomes more open with Frangula

alnus, Carex elata, Molinia coerulea and Filipendula ulmaria. The central area of the marsh

Source : MNHN, Paris

JoLy 119

F{ LavouRs marsH

Fig. 1. — Geographic situation of studied sectors and studied sites. Sites in the Jons sector : 1, 2, 3,

4: Lône des Pêcheurs oxbow lake; 5, 10, 11: permanent ponds; 6, 7, 8, 9, 12: temporary ponds.

Sites in the Lavours marsh: (1) sites in closed habitat: 11, 19: puddles; 15: drain; 16: peatbog;

20: Séran oxbow; (2) sites in open habitat: 2, 3, 4, 9, 10, 12, 13, 24: puddles; 1, 5, 7, 14, 21, 25:

drains; 6, 8, 17, 18, 22, 23: peat excavations.

is occupied by the peat-bog characterized by Schoenus nigricans and Cladium mariscus. The

vegetation was for a long time influenced by hay exportation, which hampered the

colonization of the marsh by trees. Over the last thirty years, hay exportation has gradually

disappeared and A4/nus glutinosa is now observed everywhere. Crop covers a third of the total

marsh area. Another third is managed as a Natural Reserve. It concerns the north-western

part of the marsh (northern reserve) and a plot of land in the south-western part (southern

reserve). Several experiments, including pasturing by horses and Highland Cattle in an

enclosure, have been carried out to restrict colonization by the forest. At present, the rest

of the marsh is managed according to traditional agricultural practices.

Twenty-six aquatic sites were visited during the spring of 1987 and 1991 (fig. 1). These

sites were classified according to their shape and to their degree of cover as follows :

(1) Temporary ponds or puddles. These shallow sites with very gentle slopes were

sampled in both open and covered environments. The density of aquatic vegetation in

these ponds depended on the cover. In a forest, they were, for instance, vegetationless ruts

Source : MNHN, Paris

120 ALYTES 10 (4)

in pathways. In the open marsh, they might also be ruts, or natural depressions in the peat

where underground water appeared. In most of them, water was not permanent, the

lowering of the water table causing most of the puddles to dry up at the end of July. The

watertight bottom of the covered sites and their protection from sunlight ensured longer

water retention, as confirmed by the presence of dense colonies of Tubifex.

(2) Drains. A dense network of man-made drains criss-crosses the whole marsh. In

most of them, the water was almost stagnant and the banks abrupt. The connection of

these drains with the Rhône or the Séran made fish colonization possible, especially by

sticklebacks (Gasterosteus aculeatus).

(3) Large ponds. Their origins are diverse:

(a) In the open marsh, they are remains of peat excavations. The largest one, named

Etang Delastre, measures 2 ha, but the area of some of them does not exceed a few square

metres. They may be several metres deep and their banks are always abrupt. They are

colonized by Nymphea alba.

(b) In the Séran alluvial side bar, an old oxbow lake is semi-permanent and 1.5 m deep

when filled, with gentle slopes. Its vegetation is diverse (Glyceria stagnatilis, Nymphea alba,

Nuphar luteum).

The Jons sector

The Jons sector is situated approximately 80 km downstream from the Lavours

marsh. Numerous oxbow lakes and side arms in this sector are the remains of former

meanders (JUGET et al., 1976). The present study took place along a partly-closed side arm

named “Lône des Pêcheurs”, which opens into the river downstream. Several hydrobio-

logical studies have been carried out on this arm (JUGET et al., 1976, 1979). Twelve study

sites were selected. Six of them (J6 to J11), located in the stretch of land between the arm

and the Rhône, are overflowed almost annually. Sites J6, J7 and J8 were relatively silted,

and retained water throughout spring. Their banks were colonized by Phalaris arundina-

cea. J9 was a recent unsilted and vegetationless depression in the pebble bed, with great

variations in water level, in relation with the discharge of the river. J10 and J11 were deep

and large permanent ponds ringed with Typha latifolia. Five other sites (J1 to J5) were in

the side arm bed. J5 was a pond located underbush and isolated 100 m upward from the

side arm, whereas the others (J1 to J4) were located properly in the side arm. One site (J12)

lies on the land side of an embankment, which prevents it from being overflowed. This site

only contains groundwater. The terrestrial environment of all these sites ranges from a

moorland with willows of the Salix cinerea group to an alluvial forest with A/nus glutinosa,

Salix alba and Populus nigra on the more elevated bars.

METHODS

The sites were visited from two to four times during spring. Adult anurans were

detected both visually and by listening. Amphibian larvae and adult newts were sampled

with a landing-net. Abundance was arbitrarily estimated according to a semi-quantitative

Source : MNHN, Paris

JoLy 121

scale with four levels : 0 = absence; 1 = only one observation of a given taxon; 2 = taxon

observed several times in the site but irregularly; 3 = regular and numerous observations

in the site. When small and close to one another, sites of the same type were grouped as

one site in the results. In order to draw an overview of the amphibian fauna in the alluvial

valley, an index of presence was computed as follows: for a given habitat type à, the

presence index P(i) of a taxon was: P(i) = £(a X n,) | N,, where a is the abundance

of the taxon estimated with the scale described above, n, is the number of sites where the

taxon was present and N, the total number of sites in category i.

In France, discrimination among green frog taxons is not possible on the basis of

morphological characters because of the undisrupted distribution of parameter values

from Rana lessonae to Rana ridibunda pure types (ZUIDERWUK, 1980; H. G. TUNNER, pers.

comm.). The determination of Rana ridibunda was only accepted on the basis of calling

according to the guide of GUYÉTANT & ROCHÉ (1987) or on basis of colour and shape of

metatarsal tubercule according to ARNOLD, BURTON & OVENDEN (1978). Rana lessonae and

Rana ki. esculenta could not be distinguished by means of biometry. Consequently, they

are grouped under the name Rana ‘“esculenta”’. When precise determination of green frog

was not accomplished, the term Rana synkl. esculenta was used (see DuBois & GÜNTHER,

1982).

In addition to the authors own observations, this study includes those of Yves

MAICHRZAK, Alain MORAND and Hubert TOURNIER.

RESULTS

LAVOURS

At Lavours, 10 taxons were observed (11 if Rana ‘‘esculenta’’ and Rana ridibunda are

discriminated) (Table 1). The highest richness (7 taxons) was reached in the puddles in an

open environment. Among these puddles, the richest site (6 taxons) was pool L3. The

particularity of this site is its area, which is approximately 2 ha. The puddles in the

southern reserve contained 4 taxons. The most frequent taxons in open environments were,

in decreasing order, Rana dalmatina, Rana synkl. esculenta and Triturus helveticus. Bufo

bufo, Bufo calamita, Rana temporaria and Hyla arborea were occasional in this type of site.

Triturus alpestris occurred in the Séran oxbow lake together with the three commonest

taxons described above. This site was the only one in which this species was observed.

Most of the excavations were very poor, except for site L23, which was inhabited by

3 taxons: Rana dalmatina, Rana synkl. esculenta and Bufo bufo. Note that this site was the

only man-made excavation in alluvium. The other excavations were in peat bogs and

contained only one taxon, exceptionally two and sometimes none. Rana synkl. esculenta

was the most regular taxon there. Rana dalmatina and Hyla arborea were only occasionally

observed. Amphibians were rather scarce in drains where Rana synkl. esculenta was the

most constant taxon. Rana dalmatina and Triturus helveticus occurred in drains when

depth was less than 0.5 m.

Source : MNHN, Paris

Table I. — Abundance of amphibian taxons in the sampled sites (semi-quantitative scale).

Jons sector

Lavours: closed habitat

Habitat type

(Site number

T. helvericus

TT. alpestris

IB. variegata

P. punctatus

B. bufo

B. calamira

IH. arborea

IR. remporaria

IR. dalmatina

IR. synkl. esculenra

Oxbow

J3 J4

Perm. ponds

J5 JI0 Ji1

Temporary ponds

JT J8 J9

D ©

Temp. ponds|Drains [Perm. ponds

J12| 11 19 15 16 20

2 3

2. 1

Lavours: open habitat

Habitat type

Site number

IT. helveticus

T. alpestris

B. variegata

IP. punctatus

1B. bufo

IB. calamira

IH. arborea

LR. temporaria

R. dalmatina

[R. synkl. esculenta

DORE

Temporary ponds

9 10

12 13 24

Drains

5 7 14 21

1e 1

Excavations

25 2616 8 17 18 22 23

TTI

(+) O1 SALATV

Source : MNHN, Paris

JoLy 123

The underbush puddles hosted some clutches of Rana dalmatina. Some puddles were

inhabited by small groups of Bombina variegata, particularly in the Séran alluvial forest.

These puddles were also colonized by juvenile R. synkl. esculenta in summer.

Figure 2 shows the dispersion of the different taxons over the marsh. Rana dalmatina,

Rana synkl. esculenta and Triturus helveticus were the only ones to be distributed

throughout the marsh. Triturus alpestris, Bombina variegata, Pelodytes punctatus, Bufo

bufo and Rana temporaria were found only in the vicinity of forested areas. Data

concerning Bufo calamita and Hyla arborea were scarce. Both species were found in the

open marsh. When partial determination was possible within the Rana kl. esculenta

synklepton, Rana “’esculenta’’ was identified in shallow puddles near the pasture enclosure

(L12, L13) whereas Rana ridibunda was found in deep drains near the forest (L17) and in

large peat excavations (Etang Delastre: L18).

Jons

In the Jons sector, the amphibian fauna was similar to that observed at Lavours

except for the absence of Bombina variegata and Bufo bufo (Table I). Highest richness was

reached in temporary sites. Diversity and abundance were very low in permanent eutrophic

ponds and in the side arm. In these sites, Triturus helveticus and Rana synkl. esculenta were

the only nearly constant taxons.

J9 stood out among the temporary ponds as it hosted only Bufo calamita and

Pelodytes punctatus. This site was characterized by its oligotrophy and short water

retention. Both Bufo calamita and Pelodytes punctatus appeared in this site immediately

after water rise.

As at Lavours, Triturus helveticus, Rana dalmatina and Rana synkl. esculenta were the

most constant taxons in all the sites. However, they were more abundant in temporary

ponds, where Triturus alpestris also occurred frequently. Bufo calamita and Hyla arborea

were relatively more abundant at Jons than at Lavours.

HABITAT TYPES

When analysing presence indices, the site categories were reduced to the nine habitat

types in Table I. Both taxons and habitat types were rearranged in order to bring together

similar habitats and similar taxons (fig. 3).

The habitats were first classified according to cover. The less covered was the

environment, the richer was the breeding amphibian fauna. Temporary ponds from the

two sectors were the richest habitats, whereas the underbush drains and side arm contained

the poorest fauna, with only two taxons. Rana synkl. esculenta, Rana dalmatina and

Triturus helveticus were the most constant taxons. In contrast, Bufo calamita, Pelodytes

punctatus and Hyla arborea were mostly found in temporary ponds. Bombina variegata was

characteristic of underbush temporary ponds.

Source : MNHN, Paris

124 ALYTES 10 (4)

Triturus helveticus

Pelodytes punctatus

Hyla arborea Bufo calamita

Rana dalmatina Rana "esculenta" Rana ridibunda Rana synkl. esculenta

Fig. 2. — Dispersion of amphibian taxons over the Lavours marsh.

Source : MNHN, Paris

Joy 125

En: CLOSED HABITAT + OPEN HABITAT FE

ES Es 8 E.

£ On x 52, Ses 8 Ses $e

3 £ ES Eve ÉvTO £ 3 1

SE 8% 885 sea Ë Sa Às

& à 6 #83 83 ë 83 PR

Bombine variegata

Rana synkl.esculenta

Rana delmatina

Triturus helveticus

Rana temporaria

Bufo buto |

Triturus alpestris | |

Hyla arborea ]

Pelodytes punctatus |

Bufo calamita

1 1

PRESENCE INDEX

Fig. 3. — Classification of both habitat types and taxons through ordering

of presence index (see text).

Rana ‘esculenta”’ was identified only in very shallow pools of the peat bog at Lavours.

Rana ridibunda was the only green frog found at Jons, both in temporary ponds and in

deeper sites, such as the side arm and the regularly overflooded ponds J10 and J11. At

Lavours, Rana ridibunda was less abundant and was identified in the large peat bog (L18),

in large drains and in the Séran oxbow lake.

DISCUSSION

Both at Lavours and at Jons, highest richness was reached in temporary shallow

ponds in open environments. Diversity and abundance of amphibians decreased as ponds

became more covered or showed more abrupt slopes (as in drains or peat excavations).

Amphibians also became scarce as fish became abundant, for instance in large deep ponds

or in the side arm connected with the river. With regard to the floodplain typology of

Source : MNHN, Paris

126 ALYTES 10 (4)

BRAVARD, AMOROS & PAUTOU (1986), great amphibian diversity is found in temporary

functional units belonging to both plesiopotamic and paleopotamic functional sets.

Juvenile temporary ponds in gravel beds highly influenced by river discharge are

characteristic of the units of the plesiopotamic set. In the present study, they were

represented by the temporary ponds at Jons and some sites (L3, L9, L24) near the Séran

at Lavours. In contrast, temporary units of the paleopotamic set are mature sites mildly

influenced by river discharge, such as sites L2, L4, L10, L12, L13 at Lavours.

Triturus helveticus, Rana dalmatina and Rana synkl. esculenta form a group of taxons

that is frequently observed in the south-east of France where it may be regarded as the

basis of the amphibian fauna (ZuIDERWIWK, 1980). Therefore, these taxons are not

characteristic of the floodplain. Because of its small body size, Triturus helveticus occupies

a rather small spatial niche restricted to gently-sloping banks rich in vegetation. It may be

considered as a dispersive pioneer species because its range of aquatic sites is broader than

that of other taxons (unpubl. data). In the study area, the coexistence of Rana dalmatina

and Rana synkl. esculenta may be the consequence of niche partitioning. Rana dalmatina

is a precocious breeder that spawns at the beginning of March, while Rana synkl. esculenta

spawns from the end of April to the beginning of June. When reproduction occurs in the

same site, Rana dalmatina tadpoles develop before those of Rana synkl. esculenta, provided

that water temperature is sufficiently high to allow their metamorphosis before June. Such

a succession can be observed when the environment receives sufficient sunlight. Except

during the breeding period, adults of Rana dalmatina were predominantly observed in the

alluvial forest, where their life is terrestrial. Inversely, Rana synkl. esculenta inhabits pond

banks and is typically amphibious. At Lavours, it is noticeable that Rana dalmatina breeds

on the first days of mild weather in sites which can be up to 1 km from the forest,

suggesting that frogs hibernate either in or near such sites.

In contrast to the first group of taxons, Bufo calamita, Pelodytes punctatus and Hyla

arborea are characteristic of shallow temporary ponds. This group of species was also

found frequently in isolated puddles in large riverbeds along the Rhône in the vicinity of

Montélimar (south of France) by ZUIDERWUK (1980). In Bufo calamita and Pelodytes

punctatus, the date of reproduction is not strongly determined by temperature or

photoperiod but may occur over a long period of time (DiAZ-PANIAGUA, 1986; SINSCH,

1988). These species may adjust their breeding date to temporal variations of pond filling,

that depend on relatively unpredictable river discharge. On the other hand, in these species,

the diet of the tadpoles can be more diversified than in other species because of both

behavioural and anatomical adaptations (DiaZ-PANIAGUA, 1985, 1989; ViERTEL, 1987,

1990). Consequently, we may assume these species to be more efficient in exploiting

oligotrophic sites and regard them as typical of the floodplain.

Rana temporaria, Bufo bufo, Triturus alpestris and Bombina variegata were always

found in the close vicinity of forest areas. These species become abundant and dominant

in the forested hills surrounding the river. In the floodplain, they seem to colonize the more

elevated parts of the alluvial forest.

Among the 14 amphibian taxons of the French Upper-Rhône region (CASTANET &

GUYÉTANT, 1989), 11 were found in the floodplain. The missing species were Salamandra

salamandra, Triturus cristatus and Alytes obstetricans. Reproduction of both Salamandra

Source : MNHN, Paris

Jouy 127

salamandra and Alytes obstetricans has evolved towards a greater independence from the

aquatic environment (ovoviviparity for Salamandra, egg carrying for Alytes), their

fecundity is low. We may assume that the selective pressures which led to such an

adaptation to terrestrial life also involved the loss of capacity for aquatic life. The absence

of these species from the regularly overflowed area may be a consequence of intolerance

to prolonged submersion. The causes explaining the absence of Triturus cristatus may be

different. This species chooses deep (depth > 1.5 m) fish-free ponds for breeding (DOLMEN,

1988; PAVIGNANO, 1989). In the alluvial valley, most of the deepest ponds do not dry up

because of the high level of the water table and many fish remain in such sites after floods

(Copr, 1989). The occurrence of convenient ponds for Triturus cristatus is more likely

outside the flooded area.

In a braided or anastomosed sector of the floodplain, the semi-aquatic sites may be

classified according to a gradient of river influence from the active channel(s) to the

floodplain borders. The mechanical impacts of floods (by erosion and sedimentation) are

more intense in the vicinity of active channels, where pond turnover is expected to be

relatively rapid (less than 10 years). The bottom of such ponds is not silted, which leads

to relatively unpredictable water filling depending on river discharge. The mechanical

impact of the floods decreases as the distance from the active channel increases. On the

border of the floodplain, natural ponds are formed by oxbow fragmentation following

filling up with both sediment and organic matter. The same mechanism also leads to the

disappearance of ponds. This process may be slackened by underground water exurgence.

Such ponds may persist for several decades. Their bottom is silted by inorganic and

organic matter, leading to a retention of both flood- and rain-water during several months.

The sites close to the river are characterized by high temporal and spatial variability,

whereas those at the border of the floodplain (last stages of the paleopotamon) show a

greater stability, both within space and time. According to the patch dynamics concept

(TownNsEND, 1989), the first type should host communities whose principal mode of

regulation lies in colonization processes, whereas the second type should be inhabited by

competition-regulated communities. The present data partially corroborate this model and

lead us to the following assumptions: Bufo calamita, Pelodytes punctatus, Rana ridibunda,

and, to a smaller extent, Hyla arborea, may be considered as pioneer species. Rana

dalmatina, Rana synkl. esculenta, Bufo bufo and Rana temporaria may coexist in more

stable ponds, where niche partitioning occurs among tadpoles. At the end of the aggrading

process, when depth becomes very shallow, water presence becomes temporary. This last

stage may be colonized by Bombina variegata, a site-faithful species with spawning-date

flexibility (BARANDUN, 1990). These assumptions and the validity of the patch dynamics

concept will be the basis of a detailed study of amphibian communities in the French

Upper-Rhône floodplain.

ACKNOWLEDGEMENTS

The English version of the manuscript was revised by E. PATTEE and G. THOYRON. I thank

Günter GOLLMANN and two anonymous referees for their comments on the first version of this paper.

Tam grateful to Alain MORAND, Raymond GRUFFAZ and Yves MAJCHRZAK, from the Natural Reserve

of Lavours, for their assistance.

Source : MNHN, Paris

128 ALYTES 10 (4)

LITERATURE CITED

Aïn, G. & PAuTOU, G., 1969. — Etude écologique du marais de Lavours (Ain). Documents pour la

carte de végétation des Alpes, Université de Grenoble, 7: 25-64.

AMoRoS, C., RICHARDOT-COULET, M. & PAUTOU, G., 1982. — Les “ensembles fonctionnels”: des

entités écologiques qui traduisent l’évolution de l’hydrosystème en intégrant la géomorphologie

et l’anthropisation (exemple du Haut-Rhône français). Rev. Géogr. Lyon, 57: 49-62.

Amoros, C., Roux, A. L., REYGROBELLET, J. L., BRAVARD J. P. & PAUTOU, G., 1987. — A method

for applied ecological studies of fluvial hydrosystems. Regul. Rivers, 1: 17-36.

AMoRoOS, C., BRAVARD, J. P., REYGROBELLET, J. L., PAUTOU, G. & Roux, A. L., 1988. — Les concepts

d’hydrosystème et de secteur fonctionnel dans l'analyse des systèmes fluviaux à l'échelle des

écocomplexes. Bull. Ecol., 19: 531-546.

ARNOLD, E. N., BURTON, J. A. & OVENDEN, D. W., 1978. — 4 field guide to the reptiles and amphibians

of Britain and Europe. London, Collins: 1-272.

BARANDUN, J., 1990. — Reproduction of yellow-bellied toads Bombina variegata in a man-made

habitat. Amphibia-Reptilia, 1: 277-284.

BRAVARD, J. P., 1987. — Le Rhône, du Léman à Lyon. Lyon, Ed. La Manufacture: 1-45].

BRAVARD, J. P., Amoros, C. & PAUTOU, G., 1986. — Impact of civil engineering works on the

successions of communities in a fluvial system; a methodological and predictive approach

applied to a section of the Upper Rhône River, France. Oikos, 47: 92-111.

CASTANET, J. & GUYÉTANT, R., 1989 (eds). — Atlas de répartition des amphibiens et reptiles de France.

Paris, Société Herpétologique de France: 1-191.

Copp, G. H., 1989. — The habitat diversity and fish reproductive function of floodplain ecosystems.

Environ. Biol. Fishes, 26: 1-27.

Diaz-PANIAGUA, C., 1985. — Larval diets related to morphological characters of five anuran species

in the biological reserve of Doñana (Huelva, Spain). Amphibia-Reptilia, 6: 307-322.

—— 1986. — Reproductive period of amphibia in the Biological Reserve of Doñana (SW Spain). /n:

Z. Rocek (ed.), Studies in herpetology, Praha: 429-432.

ee 1989. — Larval diets of two anuran species, Pelodytes punctatus and Bufo bufo, in SW Spain.

Amphibia-Reptilia, 10: 71-75.

DoLMEN, D., 1988. — Coexistence and niche segregation in the newts Triturus vulgaris (L.) and T.

cristatus (Laurenti). Amphibia-Reptilia, 9: 365-374.

Dunois, A. & GÜNTHER, R., 1982. — Klepton and synklepton: two new evolutionary systematics

categories in zoology. Zool. Jb. Syst., 109: 290-305.

GUYÉTANT, R. & ROCH, J.-C., 1987. — Batraciens de France. Guide sonore des crapauds et grenouilles.

La Mure, Sittelle. Tape recorder cassette.

JUGET, J., AMOROS, C., GAMULIN, D., REYGROBELLET, J. L., RICHARDOT, M., RICHOUX, P. & Roux,

C., 1976. — Structure et fonctionnement des écosystèmes du Haut-Rhône français. II. Etude

hydrologique et écologique de quelques bras morts. Premiers résultats. Bull. Ecol., 7: 479-492.

JUGET, J., Yi, B. J., Roux, C., RICHOUX, P., RICHARDOT-COULET, M., REYGROBELLET, J. L. & AMOROS,

C., 1979. — Structure et fonctionnement des écosystèmes du Haut-Rhône français. VIL. Le

complexe hydrographique de la Lône des Pêcheurs (un ancien méandre du Rhône). Schweiz. Z.

Hydrol., 41: 395-417.

PAUTOU, G., DECAMPS, H., AMOROS, C. & BRAVARD, J. P., 1985. — Successions végétales dans les

couloirs fluviaux: l'exemple de la plaine alluviale du Haut-Rhône français. Bull. Ecol., 16:

203-212.

PAUTOU, G., MAICHRZAK, Y., MANNEVILLE, O., GRUFFAZ, R. & MOREAU, D., 1991. — Dynamique

de la végétation et gestion de la réserve naturelle du marais de Lavours (Ain). Rev. Géogr. Lyon,

66: 61-70.

PAVIGNANO, L., 1989. — A multivariate analysis of habitat determinants for Triturus vulgaris and T.

carnifex in north-western Italy. Alytes, 7: 105-112.

PAVIGNANO, I., GIACOMA, C. & CASTELLANO, S., 1990. — A multivariate analysis of amphibian habitat

determinants in north western Italy. Amphibia-Reptilia, 1: 311-324.

Source : MNHN, Paris

Jouy 129

SINsCH, U., 1988. — Temporal spacing of breeding activity in the natterjack toad, Bufo calamita.

Oecologia, Berlin, 76: 399-407.

STRUBOSCH, H., 1979. — Habitat selection of amphibians during their aquatic phase. Oikos, 33:

363-372.

TOWNsEND, R. S., 1989. — The patch dynamics concept of stream community ecology. J. n. amer.

Benthol. Soc., 8: 36-50.

VaerTEL, B., 1987. — The filter appararus of Xenopus laevis, Bombina variegata and Bufo calamita

(Amphibia, Anura). À comparison of different larval types. Zool. Jb. Anat., 115: 425-452.

— 1990. — Suspension feeding of anuran larvae at low concentrations of Chlorella algae (Amphibia,

Anura). Oecologia, 85: 167-177.

ZUIDERWUK, A., 1980. — Amphibian distribution patterns in western Europe. Bijd. Dierkunde, 50:

52-72.

Corresponding editor: Günter GOLLMANN.

Source : MNHN, Paris

Alvtes, 1992, 10 (4): 130. Announcement

An Important New Book

HERPETOLOGY

CONTEMPORARY RESEARCH ON THE

BIOLOGY OF AMPHIBIANS AND REPTILES

HERPETOLOGY is at once an old and yet also a newily revitalized field of research. I was

founded in the 18h and 19h centuries on taxonomic, anatomical, and distributional studies,

£ | but during he Last century, ends in biology and science fragmented the study of amphibians

Sand repiles into narrower and more disconnected disciplines. The recent renaissance of

bolistic approaches has belped to transcend these disciplinary boundaries, enabling berpetologists to become active participants

and even leaders in this rebirth of te integrated study of animals, as the authors in this new volume demonstrate,

‘The “New Herpetology" was fonmalized in 1989 at the First World Congress of Herpetology, beld in tbe United Kingdom,

which brought together more than 1000 specialisis on amphibians and reptiles from some 60 countries. Ecologists, ana1omists,

conservationists, geneticists, physiologists, palcontologists, and others wbo span the spectrum of biology met 10 share ideas and

information. There was a general acknowledgment of the value of broadlÿ integrated approaches 19 Lbe study of à group of

organismes, and the recognition that modem berpetology represents a vigorous and cobesive discipline of enduring significance,

The main feature of the Congress was a series of plenary lectures chosen 10 emphasize the range of current herpetological

studies and to highlightthose topics in which research on amphibians and repüiles have made major contributions 10 biology. This

book contains the revised and updated versions of those lectures by authors who are among the world's leaders in their respective

fields. Together, these essays demonstrate the breadth of modem berpetology and its continuing vitality as a discipline.

Tbe volume also contains a detailed, illustrated summary of tbe meeting, with ist ofall participants in symposia, workshops,

roundtables, poster sessions, and other events. Included is a complete list of delegates and their addresses. This volume represents

the summary of record for the First World Congress of Herpetology and a synopsis of current research in our discipline. is an

excellent introduction 10 modem herpetology for students and others interested in the biology of amphibians and repiles.

Specifications: 225 pages, format 8 " by 11 inches (22 by 28 cm), 20 photographs, numerous tables and graphs, clothbound

in library-grade buckram. To be published December 1992.

TABLE OF CONTENTS

CARL GANS (USA): “The Status of Herpetology”

ILYA S. DAREVSKY (Russia): “Evolution and Ecology of Parthenogenesis in Reptiles”

LINDA MAXSON (USA): “Tempo and Patem in Anuran Speciation and Phylogeny: An Albumin Perspective”

RUSSELL A. MITTERMEIER (USA) AND OTHERS: “Conservation of Amphibians and Reptiles”

TM HALLIDAY (UK): “Sexual Selection in Amphibians and Reptiles: Theoretical Issues and New Directions”

ARMAND DE RICQLÈS (France): “Paleoherpetology"

8. DONALD BRADSHAW (Australia): “Ecophysiology of Desert Repiles””

ERIC R. PIANKA (USA): “The State of the Art in Community Ecology"

DAVID B. WAKE (USA): “An Integrated Approach to Evolutionary Studies of Salamanders”

SUMMARY OF THE FIRST WORLD CONGRESS OF HERPETOLOGY

LIST OF CONGRESS DELEGATES WITH ADDRESSES

Prices and Ordering

“The pre-publication price (before 15 November 1992) 1 SSAR members and to Delegases of he First Congress is 520. The price 10 all

other persons and to institutions is 528, (Packing and shipping per volume: USA add 52, other countries add 54).

Numbers of copies _ at $___ per copy, plus $__ total packing and shipping: total amount enclosed: $__.

Send orders to: Dr. Robert D. Aldridge, SSAR Publications Secrelary, Deparment of Biology, St. Louis University, SL Louis,

Missouri 63103, USA. (Telephone 314-658-3916; fax 314-658-3117). Overseas customers must pay in USA funds using a draft

drawn on American banks (include an additional amount to cover bank conversion charges) or by International Money Order.

MasterCard or VISA are accepted (provide account number and expiration date); a 5% bank charge will be added 1o your account.

A complete list of SSAR publications and applications for Society membership are available on request from Dr. Aldridge.

Source : MNHN, Paris

Alytes, 1992, 10 (4): 131-136. 131

Prey choice behaviour in light

and darkness in a facultative cave dweller,

the Pyrenean salamander Euproctus asper

Franz UIBLEN !

Zoologisches Institut und Zoologisches Museum der Universität Hamburg,

Martin Luther King Platz 3, 2000 Hamburg 13, Germany

In prey choïce experiments, seven larvae of the Pyrenean salamander,

Euproctus asper, were confronted in light and in darkness with two glass

microaquaria each filled with either 10 dead or 10 living chironomids. One

longside of each microaquarium was closed with a net to allow the free

exchange of water with the surroundings. In light, all seven E. asper chose the

microaquarium containing the living chironomids. During prey choice, one

individual directed the snout at the net. AI others directed their snouts either

at one of the glass walls or at the mirror image produced by the chironomids

underneath the microaquarium. In darkness, all E. asper located the prey with

three individuals choosing the living and four choosing the dead prey. AI

individuals directed their snouts at the net. E. asper prey choice behaviour in

light is clearly dominated by vision. In darkness, this facultative cave dweller

forages successfully but is less choosy than permanent cave dwellers.

INTRODUCTION

When offered a choice between prey types with differential activity, predators

frequently prefer the more active ones (e.g., FREED, 1980). Active prey provide additional

visual, mechanical, and chemical cues reducing search times and enhancing the foraging

efficiency in visually as well as in non-visually hunting predators (MARTIN, WITHERSPOON

& KEENLEYSIDE, 1974; HETHERINGTON, 1989; UiBLEIN et al., 1992). For instance, the blind

cave salamander, Proteus anguinus, shows a significant preference for active prey if offered

a choice between water conditioned with dead and living chironomid larvae (DURAND,

PARZEFALL & RICHARD, 1982). Under the same conditions however, the epigean

salamander Necturus maculosus, the closest relative of Proteus, showed no significant

preference (DURAND, PARZEFALL & RICHARD, 1982).

The Pyrenean salamander Euproctus asper facultatively lives in caves (CLERGUE-

GAZEAU, 1971) and adjusts behaviourally both to epigean and hypogean foraging

conditions. Experimental exposure to living or dead chironomids in light or darkness

showed that E. asper detects and captures living prey in light faster than dead prey in light

1. Present address: Forschungsinstitut Senckenberg, Senckenberganlage 25, 6000 Frankfurt 1, Germany.

Source : MNHN, Paris

132 ALYTES 10 (4)

or living prey in darkness (UIBLEIN et al., in press). In light therefore, E. asper should

prefer living to dead chironomids. In darkness however, foraging efficiency is reduced and

living prey may be less preferred.

The preference for living prey in light and in darkness was tested experimentally

confronting E. asper with a choice between living and dead chironomids.

MATERIAL AND METHODS

Seven larvae of Euproctus asper, two years old and with total lengths between 31 and

37 mm, were tested individually. AIl experimental animals had former feeding experience

with living and dead chironomids and had been reared in the cave of the Laboratoire

Souterrain du C.N.R.S. at Moulis, France, under controlled conditions. The experiments

were also carried out in that cave between 10.00 and 20.00 h and at a constant temperature

of 11°C.

180 minutes before the experiments started, one hungry E. asper (four days without

food) was introduced into the experimental glass tank (30 x 15 x 20 cm height). At the

same time, two empty glass microaquaria (48 X 20 *X 28 mm height) used in the

subsequent choice experiments were added to accustom the animal to the test conditions.

One longside of each microaquarium was closed with a fine net (fig. 1) to allow the free

exchange of water with the sourroundings. One broadside had a small slot for introduction

and extraction of chironomids. This slot could be tightly closed by a rubber plug (fig. 1).

Before the experiments started, the microaquaria were removed and filled each with

either ten living or ten dead chironomid larvae (freshly killed by deepfreezing and then

defrosted). Then the microaquaria were put back into the experimental tank at opposite

positions and at a distance of 11 cm between each other with the side covered by the net

directed towards the center of the tank. A distance of at least 7.5 cm was left between the

microaquaria and the margins of the tanks. The lateral positions of the microaquaria

(either left or right) were determined by a random pattern.

Each experimental animal was tested in two successive experiments, one in weak light

(3 lux), and one in complete darkness. In the latter experiment, the observations were done

with an infrared equipment (UIBLEIN et al., in press). The experiments started immediately

after introduction of the microaquaria filled with chironomids into the experimental tank.

With regard to the low sample size (n = 7), the following strict criterium (examined by

preliminary testing) was used to determine a clear choice: the salamander remains with the

snout in close contact with one of the walls of one microaquarium for at least one minute.

For each choice, three alternative orientations of the snout were recorded (fig. 1): (1) the

snout remains at the net; (2) the snout remains at one of the glass walls; (3) the snout

remains at one of the side walls with the head held down and the snout directed at the

mirror image produced by the chironomids underneath the microaquarium. In addition,

the time period between the start of the experiment and the first occurrence of a clear

choice (“latency”), and the number of snaps directed at the respective microaquarium were

recorded.

Source : MNHN, Paris

UIBLEIN 133

rubber plug

glass wall

A XX?

XRONE

RL,

ANS

RAR

RASE

NN x

NO p

0e ns

glas wall

Fig.l. — Scheme of the type of glass microaquarium used in the choice experiments. The three

possible orientations of the snout during prey choice are indicated.

Source : MNHN, Paris

134 ALYTES 10 (4)

RESULTS

In light, £. asper showed a significant preference for living prey. AI seven individuals

chose the microaquarium with the living chironomids (P = 0.008, one-tailed binomial test

[ZAR, 1984]; Table I). The mean latency was 9.24 mn. The salamanders showed

interindividual differences in snout orientation during prey choice (Table I): one individual

directed the snout at the net, three individuals directed the snout at one of the glass walls,

and three individuals directed the snout at the mirror image of the chironomids underneath

the side with the net. AII seven individuals held their positions for at least one minute with

five of them snapping at a rate of between one and ten snaps per minute (Table I).

Table 1. — Choice of prey type, latency, orientation of the snout, and number of snaps per minute

in light (L) and in darkness (D). Seven Euproctus asper (a-g) were tested individually in two

successive experiments. See text for further explanations.

Prey type chosen Latency (mn) Snout orientation Snaps/mn

D L D L D L D

a dead 17.67 11.42 mirror net 0 2

b dead 9.77 16.35 mirror net 1 0

d dead 1.33 7:53 mirror net 0 0

e dead 9.40 52.67 net net 2 0

f living 5.68 6.78 glass net 2 4

g living 1.67 2.97 glass net 10 6

In darkness, no significant preferences for either prey type were detected with three

individuals choosing living chironomids and four selecting the dead ones (P = 0.5,

one-tailed binomial test; Table 1). The median latency was 17.94 mn. Six of the seven

E. asper showed longer latencies in darkness than in light (Table I). For the total set

however, no significant differences between latencies in light and darkness could be

detected (P = 0.09, Wilcoxon test [CAMPBELL, 1989]). AII seven individuals directed

their snouts at the net during prey choice (Table I). This is significantly different from

the number of individuals at the net in light (P = 0.002, Fisher’s exact test [SOKAL &

RouLF, 1981]). Three individuals showed a snapping response (Table 1), which is not

significantly different from to the number of individuals snapping in light (P = 0.3,

Fisher’s exact test).

Source : MNHN, Paris

UIBLEIN 135

DISCUSSION AND CONCLUSION

In light, E. asper clearly preferred the living chironomids showing a choice behaviour

guided mainly by vision. Only one of the seven individuals tested directed the snout at the

net. Three individuals approached the microaquarium at the side with the net but oriented

their snouts at the mirror image instead of maintaining a multisensory contact with the

chironomids at the net. These data are in accordance with earlier observations of E. asper

foraging behaviour demonstrating the dominance of vision over non-visual perception in

light (UIBLENN et al., in press).

In darkness, all E. asper located the chironomids and showed a clear choice within

several minutes. However, no consistent preference for either prey type was detected. E.

asper lives in caves only facultatively and may be less sensitive to non-visual prey cues than

permanent cave dwellers. Proteus anguinus, for instance, approaches living chironomids in

darkness significantly faster than E. asper (UIBLENN et al., in press) and clearly preferred

living to dead prey in choice experiments (DURAND, PARZEFALL & RICHARD, 1982; UIBLEIN

& PARZEFALL, in press). À similar choice behaviour has been found in a cave dwelling

population of the fish Astyanax fasciatus (KLIMPEL & PARZEFALL, 1990).

A first crucial step towards efficient predation in caves is the shift from a visually

directed to a more “widely”, chemically and mechanically guided foraging mode (UIBLEIN

et al., in press). A second step may be to choose those prey types which can be approached

faster and are more profitable in terms of nutritional quality. E. asper is capable of

performing the first step at present (UIBLEIN et al., in press). However, no indications have

been found that this species may also realize the second step (see also LENGVENUS &

PARZEFALL, 1992). To further evaluate the current status of E. asper adaptation to cave

foraging conditions, a thorough comparison of the behavioural ecology of epigean and

cave populations is needed.

ACKNOWLEDGEMENTS

Many thanks to J. PARZEFALL and C. JUBERTHIE for supporting my work in numerous aspects.

J. Dupuy and M. R. TECHENÉ provided invaluable technical assistance. 1. ScuLurr, U. SiNscH, H.

WINKLER, and an anonymous reviewer commented on former drafts of the manuscript. This study was

made possible by an E. SCHRÜDINGER fellowship of the Austrian F.W.F., project No. J0610-BIO.

Travel expenses were covered by a PROCOPE program No. 311-pro-ca 10 J. PARZEFALL.

LITERATURE CITED

CAMPBELL, R. C., 1989. — Statistics for biologists. Cambridge, Cambridge University Press: 1-446.

CLERGUE-GAZEAU, M., 1971. — L'euprocte pyrénéen. Conséquences de la vie cavernicole sur son

développement et sa reproduction. Ann. Spéléol., 26: 825-960.

Source : MNHN, Paris

136 ALYTES 10 (4)

DURAND, J. P., PARZEFALL, J. & RICHARD, B., 1982. — Etude comparée de la détection chimique des

proies par Proteus anguinus, cavernicole, et son parent de surface Necturus maculosus

(Proteidae, Urodela). Behav. Process., 7: 123-134.

FREED, A. N., 1980. — Prey selection and feeding behaviour of the green treefrog (Hyla cinerea).

Ecology, 61: 461-465.

HETHERINGTON, T. E., 1989. — Use of vibratory cues for detection of insect prey by the

sandswimming lizard Scincus scincus. Anim. Behav., 37: 290-297.

Kuimpez, B. & PARZEFALL, J., 1990. — Comparative study of predatory behaviour in cave and river

populations of Astyanax fasciatus (Characidae, Pisces). Mém. Biospéléol., 17: 27-30.

LENGVENUS, W. & PARZEFALL, J., 1992. — The role of the visual reaction in the behaviour of an

epigean and a cave living population of Euproctus asper Dugès (Salamandridae, Urodela).

Mém. Biospéléol., 19: 111-115.

MARTIN, J. B., WITHERSPOON, N. B. & KEENLEYSIDE, M. H. A., 1974. - Analysis of feeding behaviour

in the newt Notophthalmus viridescens. Can. J. Zool., 52: 277-281.

SokaL, R. R. & ROHLF, F. J., 1981. — Biometry. Second edition. San Francisco, W. H. Freeman &

Company: 1-859.

UtBLEIN, F., DURAND, J. P., JUBERTHIE, C. & PARZEFALL, J., in press. - Predation in caves: the effects

of prey immobility and darkness on the foraging behaviour of two salamanders, Euproctus

asper and Proteus anguinus. Behav. Process. 28, in press.

UIBLEIN, F., EBERSTALLER, J., PôCKL, M. & WINKLER, H., 1992. — Effects of differential prey mobility

on the foraging behaviour of a cyprinid fish, Vimba elongata. Ethology, Ecology & Evolution,

4: 293-297.

UnBLeiN, F. & PARZEFALL, J., in press. — Does the cave salamander Proteus anguinus detect mobile

prey by mechanical cues? Mém. Biospéléol., 20, in press.

ZaR, J. H., 1984. — Biostatistical analysis. Second edition. Englewood Cliffs, N. J., Prentice-Hall:

1-718.

Corresponding editor: Ulrich SINSCH.

Source : MNHN, Paris

Alytes, 1992, 10 (4): 137-143. 137

Microscopia electrônica de barrido

del aparato bucal

de las larvas de Melanophryniscus stelzneri

(Weyemberg, 1875) (Anura, Bufonidae)

Dinorah Diana ECHEVERRIA

Facultad de Ciencias Exactas y Naturales (UBA),

Depto. de Ciencias Biolôgicas, Laboratorio de Vertebrados,

1428 Buenos Aires, Argentina

Scanning electron microscopy shows that the buccal apparatus of

Melanophryniscus stelzneri stelzneri (Anura, Bufonidae) larvae has a well

developed homy beak with teeth with large main cusp, multicusped labial

teeth with 10 to 12 denticles in each paddle, paddles with short necks, and an

oral disc with only few (12 to 16 on each side) supramarginal and inframar-

ginal conical papillae.

INTRODUCCION

FERNANDEZ (1927) efectu6 por primera vez una descripcién general de la morfologia

del aparato bucal externo de la larva de Melanophryniscus stelzneri (Weyemberg, 1875)

(Anura, Bufonidae), observando que la disposiciôn de las hileras de dientes labiales o

queratodontos determinaban la formula dentaria 2/3. GALLARDO (1987) coincide con

FERNANDEZ (1927) en la distribucion de las hileras dentarias. Basändose en datos

recopilados por Cet (1980), ALTIG & JOHNSTON (1986) organizaron la descripciôn sintética

de la anatomia externa revelada por el microscopio estereoscopico.

PRIGIONI & LANGONE (1990) destacan la existencia de diferentes formas de distribu-

ciôn de las hileras de dientes labiales en las larvas de tres especies del género

Melanophryniscus y solo efectüan una breve caracterizacion de la morfologia de los

rostrodontos y queratodontos en Melanophryniscus orejasmirandai.

Las referencias acerca de la anatomia bucal de las larvas de M. stelzneri stelzneri son

escasas (FERNANDEZ, 1927; CE1, 1980), siendo nula toda referencia acerca de la

conformaciôn ultraestructural del aparato bucal.

El propésito de este trabajo es poner en evidencia la conformaciôn anatémica de los

componentes del disco oral de las larvas de M. stelzneri stelzneri que revela el microscopio

electrônico de barrido, permitiendo mayor precision en la descripcién de algunos

Source : MNHN, Paris

138 ALYTES 10 (4)

caracteres larvales y contribuir asi al conocimiento de la anatomia larval comparada con

una nueva gama de caracteres morfolégicos de posible aplicaciôn taxonémica.

MATERIALES Y MÉTODOS

Veinticinco larvas de Melanophryniscus stelzneri stelzneri fueron obtenidas en la

provincia de Buenos Aires (Sierra de la Ventana), en la provincia de San Luis (Sierras de

San Luis, El Trapiche) y en la provincia de Cérdoba (Sierras de Cordoba, Dique Los

Molinos; Tanti y El Tigre, Calamuchita).

Los ejemplares fueron fijados en Ancel y Vitemberguer (RoUGH, 1962) o formol al

4% y al 10%.

Se consideraron los ejemplares en los estadios con el aparato bucal completamente

desarrollado y funcional (estadios 27 al 37, segün Gosner, 1960). Se seleccionaron quince

ejemplares para efectuar las observaciones con microscopio electrénico de barrido (MEB),

y el resto se destiné para las observaciones con microscopio estereoscopico.

El tratamiento aplicado al material destinado a MEB fué basado en el método de

FIORITO DE LOPEZ & ECHEVERRIA (1984), pero en la mayoria de los casos no se Ilev a cabo

la desecaciôn final al vacio, sino que se modifico para poder realizar una deshidratacion

lenta: (1) en alcohol etilico (70 %, 24 h; 80 %, 12 h; 96 %, 6 h; 100 %, 3 h); (2) en una

mezcla de alcohol 100 % — acetona, en distinta proporcion: (a) 3:1, 3 h; (b) 1:1, 2 h; (c)

1:3, 2 h; (3) en acetona pura: por lo menos tres horas; (4) secado al aire y montaje.

En el caso que el material no quedara totalmente seco a los pocos segundos después

de separarlo de la acetona, debi6 colocarse nuevamente en acetona pura durante una hora,

como minimo.

La terminologia utilizada para la descripcién del aparato bucal y de los dientes estä

basada en la propuesta por VAN Duxk (1966), ALTIG (1970) y DEUNFF & BEAUMONT (1959).

RESULTADOS

El disco oral presenta forma triangular cuando se halla en posiciôn de descanso, con

una brecha papilar dorsal y otra central muy amplias. En la region comisural (o angular),

el disco es emarginado. Presenta papilas marginales ünicamente en la region comisural,

dispuestas en dos grupos netos: suprangulares e infrangulares. Cada grupo està formado

por seis a ocho papilas, y los grupos estän separados por una emarginaciôn profunda. Las

papilas marginales son simples, cônicas, mäs largas que anchas y con borde libre liso

(fig. 1).

No se hallaron papilas intramarginales.

El pico cérneo estä bien desarrollado y queratinizado con una coloraciôn castaño

oscuro o negro. La porcin superior forma un arco amplio y convexo. La porciôn inferior

presenta una configuracion en forma de V.

Source : MNHN, Paris

ECHEVERRIA 139

El pico corneo se muestra profundamente aserrado. Las piezas dentarias (rostrodon-

tos) se disponen en empalizada, con la cüspide alargada (fig. 2). La altura de cada diente

es, por lo menos, el doble del ancho de la base del mismo. En ejemplares correspondientes

al estadio 28, se ha observado que cada diente del pico côrneo puede alcanzar un largo

mäximo de tres veces el largo de su base. El äpice de cada rostrodonto puede presentarse

agudo o romo (figs. 2 y 4).

Los pliegues labiales se diponen en cinco hileras: dos anteriores y tres posteriores,

sosteniendo a los queratodontos y permitiendo el movimiento de las hileras de dientes.

Las hileras A-1, A-2, P-1 y P-2 se extienden desde un extremo al otro de las märgenes

laterales del disco. La P-3 es la hilera de menor extensiôn (fig. 1).

FiG. 1. — Vista general del aparato bucal de la larva de Melanophrynisc

hilera anterior; A-2, segunda hilera anterior; P-1, primera hilera posterior; P-2, segunda hilera

posterior; P-3, tercera hilera posterior; PI, parte inferior del pico cérneo: PS, parte superior del

pico corneo; m, papila marginal. Escala: 100 um.

FiG. 2. — Detalle del pico cérneo de la larva de Melanophryniseus stelzneri. Escala: 100 pm.

stelzneri. A-1, primera

FIG. 3. — Queratodontos de la larva de Melanophryniseus stelzneri. Escala: 10 pm.

FIG. 4. — Queratodontos de A-1 de la larva de Melanophryniscus stelzneri por desprenderse y

suprarostrodontos con äpices desgastados. Escala: 100 um.

Source : MNHN, Paris

140 ALYTES 10 (4)

La distribuciôn de hileras de queratodontos determina la férmula dentaria 2/3. En

sélo tres casos (en estadios 31, 32 y 33), se observé una pequeña escotadura mediana en

la P-1 (fig. 1).

Cada vez que emerge una camada de queratodontos, ellos presentan una espätula

corta pero ms larga que ancha, con denticulos o cüspides en nümero variable de 10 a 12,

generalmente con 6 a 8 cüspides centrales, bien desarrolladas y separadas. El cuello es

corto (fig. 3). En los casos en que se observan aflorando dos camadas de dientes, una por

debajo de la otra, los mäs distales pueden presentar el cuello alargado, pero nunca mäs

largo que la espâtula (fig. 4).

Los pliegues labiales soportan los queratodontos espatulados, presentan varias

cüspides funcionales al mismo tiempo (aproximadamente entre 8 y 10) y con una

proporciôn ancho mäximo espätula / largo de espätula de 1/2.

DISCUSIÔN

(1) Los caracteres morfolôgicos aqui descritos para la larva de Melanophryniscus

stelzneri stelzneri determinan la formula dentaria 2/3 en coincidencia con FERNANDEZ

(1927) y con GARRIDO (1989) para M. stelzneri montevidensis.

FERNANDEZ (1927) y GALLARDO (1974) señalan que el pico corneo de las larvas de M.

stelzneri es débil, quizäs como sinénimo de pequeño, como lo describen KEHR & WILLIAMS

(1990). En la muestra estudiada se observé que el pico côrneo estä bien desarrollado y

queratinizado, pero es angosto en sentido anteroposterior.

Por otra parte, si tomamos a manera de ejemplo la extensiôn relativa que ocupa el

pico corneo superior en el disco oral desde un extremo al otro, veremos que, en el caso

de la muestra estudiada, representa el 50 % de la extensiôn total del disco tomada entre

ambas comisuras, a nivel de las emarginaciones. En el caso observado por FERNANDEZ

(1927: 277, fig. 1), la distancia entre los extremos del pico corneo superior representa sélo

el 33 %. Esta diferencia podria estar asociada con causas intrinsecas del ünico ejemplar

estudiado por FERNANDEZ y en cuyo diagrama se basan la mayoria de los autores que

coinciden con ella.

(2) Las variaciones morfolôgicas detectadas en los rostrodontos y en los queratodon-

tos no implican la existencia de heterodoncia dimensional permanente, sino que se deben

al desgaste de los mismos.

A medida que los dientes afloran, envejecen: la espâtula y el cuello se alargan,

pudiendo cambiar la forma general hasta que se desprenden y caen. Esto fué comprobado

anteriormente en Bufo arenarum (ECHEVERRIA et al, 1987; FIORITO DE LOPEZ &

ECHEVERRIA, 1989).

El grado de envejecimiento del diente aumenta con el tiempo de utilizaciôn del mismo;

se pueden producir leves modificaciones en las piezas corneas, siendo mäs notables en los

queratodontos que en los rostrodontos. En ellos las cüspides tienden a hacerse romas hasta

que, a veces, sus extremos se acortan o se pierden. En los queratodontos el cuello puede

alargarse y afinarse cuando estän proximos a desprenderse.

Source : MNHN, Paris

ECHEVERRIA 141

Por ello, para poder obtener una fiel descripcion de la morfologia dentaria, ser

conveniente tomar como base de referencia a las piezas dentarias observadas en los

estadios 25 o 26 y en aquellos inmediatamente después de cada recambio. Es decir, cuando

aflora la primera camada de queratodontos, o eliminando mecänicamente los dientes

modificados, para hallar los que se encuentran inmediatamente debajo de ellos, que

conservan intacta la morfologia original.

(3) En cuanto a la escotadura detectada en la P-1, considero que es probable que ésta

sea una disposiciôn transitoria y que esté relacionada con el crecimiento del aparato bucal,

en especial de los pliegues labiales que contienen las columnas de dientes en desarrollo,

como es el caso de Bufo arenarum (FIORITO DE LOPEZ & ECHEVERRIA, 1989).

(4) La forma general de los dientes labiales se acerca, por ejemplo, a la observada por

FIORITO DE LOPEZ & ECHEVERRA (1984) en Bufo arenarum, de amplia distribuciôn en la

Argentina, que, en la época de reproduccién, frecuenta aguas quietas permanentes o

semipermanentes.

En las larvas de M. stelzneri stelzneri examinadas, la posiciôn de los queratodontos

y de las cüspides de los mismos se presentan marcadamente curvados hacia la regiôn bucal

(mäs pronunciado se observa en las hileras posteriores). Esto podria estar asociado al tipo

de captura del alimento que Ilevan a cabo, por raspado o rastrillado, reteniendo el

alimento y acumuländolo entre las hileras (dientes tipo Rana, segün ALTIG & JOHNSTON,

1989).

La forma de V del pico inferior podria contribuir al corte y/o retenciôn de restos

vegetales, früstulos de diatomeas, etc., como se estima que ocurre en otras especies.

Llama la atenciôn que, en la porciôn anterior del intestino, se hallaron largos

früstulos (de mäs de 200 um) y partes rotas (de aproximadamente 100 um) de diatomeas

plancténicas como Synedra ulna y su forma de resistencia, quizâs recogidos del fondo del

cuerpo de agua semipermanente con corriente intermitente. Es posible que por la

morfologia del pico cérneo superior, que permite una abertura bucal amplia, quede

facilitado el ingreso a la cavidad bucal de estructuras rigidas como los largos früstulos o

las diatomeas vivas de Synedra ulna, en gran cantidad y enteras.

Considerando que en un cuerpo de agua las fuentes de alimentos pueden ser varias

y que cada una de ellas estä sujeta a ciclos biolôgicos, si los renacuajos de M. stelzneri se

comportaran como oportunistas alimentarios, y de acuerdo con la morfologia bucal ya

comentada, tendrian fâcil acceso a alimentos de gran tamaño que quizâs provean en

conjunto, mayor contenido calérico ütil para el desarrollo que, segün FERNANDEZ (1927),

es rapido.

CONCLUSIONES

(1) Los queratodontos tienen espätulas cortas, con 10 a 12 denticulos o cüspides,

generalmente con 6 a 8 cüspides centrales bien desarrolladas y separadas. El cuello es

corto.

Source : MNHN, Paris

142 ALYTES 10 (4)

(2) El pico cérneo es bien desarrollado y queratinizado, profundamente aserrado.

(3) Hay escasas papilas, sélo marginales, comisurales (12 a 16 por lado).

(4) Las variaciones morfolôgicas en los dientes se deben al desgaste.

(5) La férmula dentaria observada es 2/3.

RESUMEN

El aparato bucal de las larvas de Melanophryniscus stelzneri stelzneri presenta un pico

côrneo bien desarrollado y queratinizado. Los rostrodontos son unicuspidados, con

cüspides pronunciadas. El infrarostrodonto presenta una configuracién en V en estado de

reposo y es poco visible. El suprarostrodonto es convexo, muy extenso.

Los dientes labiales son espatulados, multicuspidados (10 a 12 cüspides), con cuello

corto, nunca mâs largo que la espätula. Hay papilas comisurales ünicamente, inframar-

ginales y supramarginales, desde 12 hasta 16 por cada lado.

AGRADECIMIENTOS

AI Dr. Raymond F. LAURENT por la lectura critica del manuscrito. A la Lic. Ana M. FILIPELLO

y al Prof. Gustavo CaRRIZO, por contribuir con parte del material de estudio.

A los señores Héctor CHiocHio y Angel FUSARO del Servicio de Microscopia Electrônica de

Barrido, del Instituto de Neurobiologia dependiente del Consejo Nacional de Investigaciones

Cientificas y Técnicas, por la asistencia técnica.

LITERATURA CITADA

ALTIG, R., 1970. — A key to the tadpoles of continental United States and Canada. Herpetologica,

26: 180-207.

ALriG, R. & Jonnsron, G. F., 1986. — Major characteristics of free-living anuran tadpoles.

Smithsonian Herpetological Information Service, 67: 1-15.

1989. — Guilds of anuran larvae: relationships among developmental modes, morphologies and

habits. Herpetol. Monogr., 3: 81-109.

Cri, J. M., 1980. — Amphibians of Argentina. Monit. zool. ital., Monogr. 2: 1-609.

Deunrr, F. & BEAUMONT, A., 1959. — Histogénèse des dents et du bec cornés chez les larves de

, FIORITO DE Lopi L. E., VACCARo, O. B. & FiLiPELLO, A. M., 1987. —

Consideraciones acerca de las férmulas dentarias de las larvas de Bufo arenarum Hensel (Anura,

Bufonidae). Cuad. Herp., 3 (3): 33-39.

FERNANDEZ, K., 1927. — Sobre la biologia y reproducciôn de Batracios argentinos. Bol. Acad. nac.

Cienc. Côrdoba, 29: 271-328.

FioRITO DE Lopez, L. E. & ECHEVERRIA, D. D., 1984. — Morfogénesis de los dientes larvales y pico

côrneo de Bufo arenarum (Anura, Bufonidae). Rev. Mus. arg. Cs. nat. Bernardino Rivadavia,

Zool., 13 (60): 573-578.

Source : MNHN, Paris

ECHEVERRIA 143

se 1989. — Microanatomia e histogénesis del aparato bucal en las larvas de Bufo arenarum (Anura,

Bufonidae). Cuad. Herp., 4 (2): 4-10.

GaLLARDo, J. M., 1974. — Anfibios de los alrededores de Buenos Aires. Editorial Eudeba: 1-231.

— 1987. — Anfibios Argentinos. Guia para su identificaciôn. Buenos Aires, Ediciones Libreria

Agropecuaria S.A.: 1-98.

GarIDO, R. R., 1989. — Descripciôn de la larva de Melanophryniscus stelzneri montevidensis

(Philippi, 1902) (Anura, Bufonidae). Bol. Soc. Zool. Uruguay, 5: 7-8.

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

identification. Herpetologica, 16: 183-190.

Kemr, A. I. & WILLIAMS, J. D., 1990. — Larvas de anuros de la Repüblica Argentina. Cuad. Herp.,

(ser. monogr.), 2: 1-44.

PRIGIONI, C. M. & LANGONE, J. A. 1990. — Descripciôn de la larva de Melanophryniscus

orejasmirandai Prigioni y Langone, 1986 (Amphibia, Anura, Bufonidae). Com. Zool. Mus.

Montevideo, 12 (173): 1-9.

RouGH, R., 1962. — Experimental embryology. Techniques and procedures. Burguess Publishing Co.

VAN Du, D. E., 1966. — Systematic and field keys to the families, genera and described species of

the Southern African anuran tadpoles. Ann. Natal Mus., 18 (2): 231-286.

Corresponding editor: Alain Dusois.

BIBL.DU

\PARE

Source : MNHN, Paris

Alytes, 1992, 10 (4): 144. Announcements

Meetings

TERCEIRO CONGRESSO LATINO-AMERICANO DE HERPETOLOGIA

Universidade Estadual de Campinas, Säo Paulo, Brasil.

12-18 December 1993.

For information please contact:

Comissäo Organizadora Central do 3° CLAH, Departamento de Zoologia, Univer-

sidade Estadual de Campinas, I. B. 13081-970, Campinas, Säo Paulo, Brasil.

Phones: 00(55)-192-39-7022; 00(55)-192-39-8492.

Fax: 00(55)-192-39-3124.

SECOND WORLD CONGRESS OF HERPETOLOGY

University of Adelaide, South Australia, Australia.

29 December 1993 - 6 January 1994.

For information please contact:

Michael J. TYLER, Department of Zoology, University of Adelaide, Box 498, G.P.O.

Adelaide S.A. 5001, Australia.

Phone: 61-8-228-5333.

Fax: 61-8-223-5817.

Source : MNHN, Paris

AIVTES

International Journal of Batrachology

published by ISSCA.

EDITORIAL BOARD FOR 1992

Chief Editor: Alain Dusois (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.); Alain COLLENOT (Paris, France); Tim HaLLDay (Milton Keynes, United

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

JoLy (Lyon, France); Milos KALEZIé (Beograd, Yugoslavia); Raymond F. LAURENT (Tucumän,

Argentina); Petr ROTH (Libechov, Czechoslovakia); Dianne B. SEALE (Milwaukee, U.S.A.);

Ulrich SNscH (Bonn, Germany).

Index Editor: Annemarie OHLER (Paris, France).

GUIDE FOR AUTHORS

Alytes publishes original papers in English, French or Spanish, in any discipline dealing with

amphibians. Beside articles and notes reporting results of original research, consideration is given for

publication to synthetic review articles, book reviews, comments and replies, and to papers based

upon original high quality illustrations (such as color or black and white photographs), showing

beautiful or rare species, interesting behaviors, etc.

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

be typewritten or printed double-spaced on one side of the paper. The manuscript should be organized

as follows: English abstract, introduction, material and methods, results, discussion, conclusion,

French or Spanish abstract, acknowledgements, literature cited, appendix.

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, 1948; So & So, 1987;

EveryBopy et al., 1882). 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. & Pos 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:

x + 1-547, pl. I-IV.

Manuscripts should be submitted in triplicate either 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.

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

referees. If possible, after acceptance, a copy of the final manuscript on a floppy disk (3 1/2 or 5 1/4)

should be sent to the Chief Editor. We welcome the following formats of text processing:

(1) preferably, MS Word DOS and Windows, WordPerfect (4.1 to 5.1) or WordStar (3.3 to 5.5);

(2) less preferably, formated DOS (ASCII) or DOS-formated MS Word for the Macintosh. ;

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.

Published with the support of AALRAM

(Association des Amis du Laboratoire des Reptiles et Amphibiens

du Muséum National d'Histoire naturelle, Paris, France).

Directeur de la Publication: Alain Dumois.

Numéro de Commission Paritaire: 64851.

Alytes, 1992, 10 (4): 113-144.

Contents

Leszek BERGER & Mariusz RYBACKI

Sperm competition in European water frogs............................ 113

Pierre JOLY

The amphibian fauna of the French Upper-Rhône floodplain.

The Lavours marsh and'theJons sector...........1......40.0.eeanee 117

Franz UIBLEN

Prey choice behaviour in light and darkness

in a facultative cave dweller,

the Pyrenean salamander Euproctus asper .............................. 131

Dinorah Diana ECHEVERRA

Microscopia electrônica de barrido del aparato bucal

de las larvas de Melanophryniscus stelzneri (Weyemberg, 1875)

(ARUTA ES BUTONITAE) Er nee iunrmamenhuee dt e nan 137

Announcements

Herpetology. Contemporary research on the biology

Offamphibians andirepliles 2-0 uen rec be ceci 130

Meetings .

Alytes is printed on acid-free paper.

Alytes is indexed in Biosis, Cambridge Scientific Abstracts, Current Awareness in Biological

Sciences, Pascal, Referativny Zhurnal and The Zoological Record.

Imprimerie F. Paillart, Abbeville, France.

Dépôt légal: 4°" trimestre 1992.

©ISSCA 1992

Source : MNHN, Paris