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Source : MNHN, Paris

AINTTES

INTERNATIONAL JOURNAL OF BATRACHOLOGY

March 1998 Volume 15, N° 4

Alytes, 1998, 15 (4): 137-144. 137

Dieta larval de Phyllomedusa tetraploidea

Pombal & Haddad, 1992

en la provincia de Misiones (Argentina)

Rafael C. LasMANoOvICH * & Julian FAIVOVICH **

* Becario del CONICET, Instituto Nacional de Limnologia (CONICET),

José Maciä 1933, 3016 Santo Tomé (Santa Fe), Argentina

++ Divisiôn Herpetologia. Museo Argentino de Ciencias Naturales

“Bernardino Rivadavia”, Casilla Postal 220, Suc. 5, 1405 Buenos Aires, Argentina

The diet of larval Phyllomedusa tetraploidea was studied from sam-

ples of stages 35 to 38 collected in Misiones (Argentina). The relative

frequencies of food items (% FO) and numerical frequence percentages

(% EN) as well as content’s diversity were determined. Results show that P.

tetraploidea larvae have a herbivorous trophic spectrum composed mainly

by green algae, diatoms, detritus and vegetal rest. Finally it is suggested

that feeding in these larvae probably occurs mainly on the bottom of the

water body rather than near the surface.

INTRODUCCION

Los contenidos intestinales de las larvas de anfibios son buenos indicadores de la

composicién floristica del ambiente ya que actüan como alimentadores continuos, aprove-

chando al mäximo los recursos disponibles (FARLOWE, 1928; KAMAT, 1962), pudiendo ser la

causa de sübitas reducciones y alteraciones de las mismas (DICKMAN, 1968; SEALE et al. 1975;

SEALE, 1980), influenciando en los niveles trôficos superiores que se soportan en las comuni-

dades perifiticas.

Phyllomedusa tetraploidea fue descripta por POMBAL & HADDAD (1992); el status de esta

especie en Argentina fue tratado por LANGONE & CARRIZO (1995). En la descripciôn original

los autores incluyeron datos sobre la biologia del adulto y describieron la larva, no existiendo

a nuestro saber informaciôn sobre su biologia y ecologia larval.

El objetivo de este trabajo es brindar un aporte al conocimiento de la composiciôn

cuali-cuantitativa de la dieta de P. tetraploidea en ambientes acuâticos semipermanentes de la

provincia de Misiones (Argentina), basändose en el estudio del contenido intestinal, de la

diversidad y del amplitud del nicho.

Bibliothèque Centrale Muséum

3 3001 00023937 il MNHN, Paris

138 ALYTES 15 (4)

MATERIAL Y MÉTODOS

Se estudiaron 18 contenidos intestinales de larvas elegidas al azar de un lote colectado el

16 de enero de 1995, en un cuerpo de agua cuadrangular de 7 m de lado y 1 m de profundidad,

con märgenes cubiertos por ciperâceas y marginalmente por manintâceas, en la provincia de

Misiones, Dto. Guarany, km 1272 ruta nacional 14, campo anexo INTA “Cuartel Rio

Victoria” (26°55'S, 54°25/W). El sitio se halla a 534 m sobre el nivel del mar. Las larvas fueron

colectadas con red durante la noche y fijadas inmediatamente en formol 10 % neutralizado.

No se registré vegetaciôn flotante.

La vegetaciôn de la provincia Misiones corresponde a la provincia fitogeogräfica “Para-

naense”, del Dominio Amazénico (CABRERA, 1971). El clima es cälido y hümedo, con

precipitaciones durante todo el año que oscilan entre 1500 y 2000 mm anuales. La tempera-

tura media anual varia entre 20°C y 21°C.

En la coleccién de la Divisién Herpetologia del Museo Argentino de Ciencias Naturales

“B. Rivadavia”, se conserva bajo el nümero MACN 36201 un lote de ejemplares de referencia

colectado junto con los utilizados en este estudio.

Las larvas se agruparon por estadio de desarrollo segün la tabla de GosnER (1960). Se

seccionaron los tubos digestivos en forma completa, y bajo lupa binocular se extrajeron los

contenidos para su determinaciôn y cuantificaciôn mediante microscopio de 400 X. Para

calcular los porcentajes de frecuencia numérica de las algas cuantificables en los contenidos

intestinales, se aplico un método indirecto, homogeneizando y diluyendo cada muestra en una

proporcién conocida (1:10), contändose 3 alicuotas de 1 cm” que se evaluaron por el método

de la gota (microtransecta) de LACKEY (1938) que permite calcular el nämero de organismos

por ml segün la siguiente ecuaciôn:

N°mI=CxTA/AXS XV,

donde TA es el ärea del cubreobjetos en mm?, A el ärea de 1 hilera (transecta en el

cubreobjetos) en mm?, C el nümero de organismos contados, S el nümero de hileras contadas

y V el volumen de la muestra bajo el cubreobjetos.

Se obtuvieron los porcentajes de frecuencia de ocurrencia (% FO) y los totales (N) de las

distintas categorias alimentarias cuantificables. En las categorias no cuantificables, se aplico

una escala de abundancia arbitraria (que se detalla en la tabla 2), basada en los porcentajes de

ocurrencia de las distintas categorias alimentarias.

Para determinar la diversidad trofica se siguié el criterio de HURTUBIA (1973), que

consiste en calcular la diversidad tréfica (H) para cada individuo utilizando la férmula de

BRILLOUIN (1965):

H = (1/H) x (log, N - log, Nil),

donde N es el nümero total de organismos hallados en el intestino de cada individuo y Nies

el nümero total de organismos de la especie i en cada intestino.

Source : MNHN, Paris

LAJMANOVICH & FAIVOVICH 139

Tab. 1. - Dieta de Phyllomedusa tetraploidea de la provincia de Misiones. Categorias

cuantificables. FO: porcentaje de frecuencia de ocurrencia de cada categoria en el total.

FN: porcentaje de frecuencia numérica en el total de categorfas. ni: no identificado.

Algae

Euglenophyta

Euglenophyceae

Phacus spp.

Trachelomonas spp.

Chrysophyta

Bacillarioficeae

Navicula spp.

Fragilaria spp.

Pinnularia spp.

Diatoma spp.

Eunotia spp.

Surillela spp.

Caloneis spp.

Chlorophyta

Chlorophyceae

Closterium spp.

Closterium cf. turgidum

Euastrum spp.

Scenedesmus spp.

Stauroneis spp.

Ankistrodesmus spp.

Restos animales

Ciliados (ni)

Tecamebianos

Arcella spp.

Se calculé la diversidad media (H) y la diversidad tréfica acumulada (Hk). La amplitud

tréfica del nicho se obtuvo mediante el indice de LEVINS (1968):

Nb= EP)!

donde P;; es la probabilidad del item i en la muestra j.

RESULTADOS

Las larvas estudiadas se encontraban en los estadios de desarrollo 35 a 38 de la tabla de

Goser (1960), presentando una longitud de cuerpo media de 18,56 mm (desviaciôn eständar

= 0,9). La totalidad de los tubos digestivos analizados (n = 18) contuvieron alimento.

Source : MNHN, Paris

140 ALYTES 15 (4)

Otros (1,8 %)

Chrysophyta

(33,6 %)

Chiorophyta

(47,6 %)

Euglenophyta

(16,8 %)

Hongos

Algas cenobiales

Algas filamentosas

Restos Minerales

Restos Animales

Restos Vegetales

Detritus

0 20 40 60 80 100%

Fig. 1. - Composiciôn de la dieta de Phyllomedusa tetraploidea en ambientes semi-permanentes de la

provincia de Misiones. (a) Categorias cuantificables: frecuencia numérica. (b) Categorias no

cuantificables: frecuencia de ocurrencia.

Source : MNHN, Paris

LAJMANOVICH & FAIVOVICH 141

Tab. 2. - Dieta de Phyllomedusa tetraploïdea de la provincia de Misiones. Categorias no

cuantificables. FO: porcentaje de frecuencia de ocurrencia de cada categoria en el total.

Rangos de abundancia: MA, muy abundante; A, abundante; ES, escaso; R, raro

Taxon Abundancia

Cyanophyta

Cyanophyceae

Merismopedia spp.

Chlorophyta

Chlorophyceae

Oedogonium spp.

Detritus

Restos vegetales

Restos animales

Restos minerales

Hongos

Hyphomycetidae

Los especimenes de P tetraploidea examinados presentaron un espectro tréfico inte-

grado por detritus, 5 divisiones de algas, restos vegetales, animales, minerales y hongos (fig. 1,

tab. 1-2). En las algas cuantificables (tab. 1), la mayor representaciôn fue de las cloréfitas

(47,6 %) (en especial por el aporte numérico de Closterium cf. turgidum), seguido por las

criséfitas (33,6 %) y euglenôfitas (16,8 %) y con porcentajes menores al 3 % en el resto de las

categorias.

En lo referente a las categorias no cuantificables (tab. 2), las algas fueron escasas con una

mayor representaciôn de restos vegetales, animales y detritus.

La diversidad media (H) fue de 3,15 (desviacién eständar = 0,31). La diversidad

acumulada (Hk) fue de 3,52 y con la suma de las 18 muestras la curva tiende a la estabilizaciôn

con la acumulaciôn de 3 intestinos (fig. 2). Este valor indicaria, segün HURTUBIA (1973), que

la muestra minima de ejemplares requeridos para este tipo de estudio es de tres intestinos. La

amplitud del nicho (Nb) fue de 7,56.

DIsCUSIÔN

La dieta de las larvas de anuros se basa fundamentalmente en algas, sobre las que se

alimentan indiscriminadamente (FARLOWE, 1928; JENSEN, 1967; SEALE & BECKVAR, 1980),

restos de plantas superiores en maceraciôn, protozoos y rotiferos y cadäveres en descompo-

sicién, siendo en algunos casos predadores de otros renacuajos.

En la especie estudiada, el espectro tréfico es principalmente herbivoro, integrado

mayoritariamente por cloréfitas, en especial por Closterium cf. turgidum, organismo de

Source : MNHN, Paris

142 ALYTES 15 (4)

1 2 3 4 5 6 7 8 9 1011 12 13 14 15 16 17 18

Némero de intestinos

Fig. 2. - Curva de diversidad tréfica acumulada versus nümero de intestinos analizados. La flecha indica

el punto aproximado de estabilizaciôn.

tamaño considerablemente mâs grande (aprox. 1500 um) respecto a otras algas registradas

como alimento en larvas de anuros (VIERTEL, 1992; LAIMANOvVICH, 1994). Por las caracteris-

ticas ecolôgicas de las desmideäceas se infiere un tipo de alimentacién ligada al fondo en

donde se encuentra, por su tamaño, este taxon de algas.

Los valores de diversidad y amplitud tréfica del nicho calculados permiten inferir un

espectro alimentario homogéneo con algunas categorias mayoritarias. Los valores de estos

indices son similares a los hallados por LAIMANOVICH (1997) en otras larvas de anuros; esto

reafirma que las estrategias alimentarias se basan en una gran diversidad de taxones, lo que se

corresponde con la abundancia de recursos de los ambientes temporales en que se desarrollan

las larvas.

En el cuerpo de agua en donde fueron capturadas las larvas se observé que éstas se

encontraban ubicadas préximas a la superficie, suspendidas con la cabeza hacia arriba,

manteniéndose con râpidos movimientos del extremo flageliforme de la cola. Este compor-

tamiento fue constatado en otras especies del género Phyllomedusa por WASSERSUG (1973),

LAVviILLA (1983) y BRANCH (1983).

Es interesante notar la no correspondencia que surge de confrontar la ubicacién de estas

larvas en el cuerpo de agua con los resultados de este estudio, que indican que la alimentaciôn

esta compuesta principalmente por algas ligadas al fondo.

Source : MNHN, Paris

LAJMANOVICH & FAIVOVICH 143

En el cuerpo de agua muestreado se ha registrado que aproximadamente a la media

noche las larvas se sumergen masivamente, disminuyendo de manera notable la densidad de

individuos en la superficie. Esta informaciôn es preliminar y sugiere que las larvas de la especie

estudiada poseen un patrôn de distribucién espacial complejo, el cual podria explicar el

fenémeno descrito. Si esto fuese asi, seria en parte congruente con los datos obtenidos por

BRANCH (1983). Este autor observé que las larvas de Phyllomedusa vaillanti se hallaban

suspendidas a media agua o cerca de la superficie durante la noche y que durante el dia

estaban mâs activas y se encontraban a mayor profundidad; igualmente, también reporté la

presencia de algas de fondo en la dieta de las larvas que estudi6. No obstante, el patrôn de

actividad observado por BRANCH (1983) es temporalmente inverso al que se describe para P.

tetraploidea.

Este trabajo se considera el primer aporte al conocimiento de la dieta de esta especie y es

necesario realizar estudios de distribuciôn espacial en las larvas de P. tetraploidea que puedan

conciliar estas observaciones.

RESUMEN

La dieta de las larvas de Phyllomedusa tetraploidea fue estudiada en material proveniente

de la provincia de Misiones (Argentina), entre los estadios 35 y 38 de Gosner (1960). Se

calcularon las frecuencias de apariciéôn de las distintas categorias alimentarias (% FO) y los

porcentajes de frecuencia numérica (% FN) segün el método de LACKEY (1938); se evalué la

diversidad de los contenidos mediante el indice de LEVINS (1968). De acuerdo a los resultados,

las larvas de P tetraploidea presentan un espectro tréfico herbivoro, integrado mayoritaria-

mente por cloréfitas, diatomeas, detritus y restos vegetales.

Finalmente se sugiere que la alimentacién de estas larvas estä asociada principalmente al

fondo y no a las proximidades de la superficie.

AGRADECIMIENTOS

A Esteban LAviLLA y Néstor Basso por la lectura critica del manuscrito y a A. VINOCUR por la

corroboraciôn de las especies de algas.

LITERATURA CITADA

BrAncH, L. C., 1983. — Social behavior of the tadpoles of Phyllomedusa vaillanti. Copeia, 1983 (2):

420-428.

BRILLOUIN, L., 1965. — Science and information theory. New York, Academic Press: 1-320.

CABRERA, A., 1971. — Fitogeografia de la Repüblica Argentina. Bol. Soc. arg. Bot., 14 (1-2): 1-42.

DickMaAN, M., 1968. - The effect of grazing by tadpoles on the structure of periphyton community.

Ecology, 49 (6): 1188-1190.

FARLOWE, V., 1928. — Algae of ponds as determined by an examination of the intestinal contents of

tadpoles. Biol. Bull., 55: 443-448.

Source : MNHN, Paris

144 ALYTES 15 (4)

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

identification. Herpetologica, 16: 183-190.

HurTuBIA, J., 1973. - Trophic diversity measurement in sympatric predatory species. Ecology, 54 (4):

885-890.

JENSEN, T. A., 1967. — Food habits of the green frog, Rana clamitans, before and during metamorphosis.

Copeia, 1967: 214-218.

KAMAT, N. D. 1962. - On the intestinal contents of tadpoles and algae of small ponds. Current Sci., 321

(7): 300-310.

LackEy, J. B., 1938. - The manipulation and counting of river plankton and changes in some organisms

due to formalin preservation. Pub. Health Rep., 53: 20-80.

LaJMANOVICH, R. C., 1994. - Contribuciôn al conocimiento de la alimentaciôn de larvas de la rana criolla

Leptodactylus ocellatus (Amphibia: Leptodactylidae) en el Paranä, Argentina. Stud. neotrop.

Fauna Environ., 29 (1): 55-61.

— 1997. — Alimentaciôn de larvas de anuros en ambientes temporales del sistema del rio Paranä,

Argentina. Doñana Acta Vertebrata, en prensa.

LANGONE, J. A. & CaRRIZO, G. R., 1995. - Confirmaciôn de la presencia en la Repüblica Argentina de

Phyllomedusa tetraploidea Pombal Jr. & Haddad, 1992 (Amphibia, Anura, Hylidae). Resimenes

IX Reuniones de Comunicaciones Herpetolégicas: 18.

LAviLLA, E. O., 1983. - Contribuciôn al conocimiento de los estados larvales de anuros argentinos. Acta

2001. lilloana, 37 (1): 5-13.

LEVINS, R., 1968. — Evolution in changing environment. New Jersey, Princeton, Univ. Press: 1-120.

PoMBaL, J. P. Jr. & HaDDaD, C. F. B., 1992. — Espécies de Phyllomedusa do grupo burmeisteri do Brasil

oriental, com descriçâäo de uma espécie nova (Amphibia, Hylidae). Rev. brasil. Biol., 52 (2):

217-229.

SrALE, D. B,, 1980. — Influence of amphibian larvae on primary production, nutrient flux, and competi-

tion in pond ecosystem. Ecology, 61: 1531-1550.

SeaLe, D. B. & BECK VAR, N., 1980. - The comparative ability of anuran larvae (genera: Hyla, Bufo, and

Rana) to ingest suspended blue-green algae. Copeia, 1980: 495-503.

SEALE, D. B.; RODGERS, E. & Boraas, M. E., 1975. — Effects of suspension feeding frog larvae on

limnological variables and community strusture. Verh. int. Ver. Limnol., 19: 3179-3184.

VIERTEL, B., 1992. - Functional response of suspension feeding anuran larvae to different particle sizes at

low concentrations (Amphibia). Hydrobiologia, 234: 151-173.

WASsERSUG, R. J., 1973. — Aspects of social behavior in anuran larvae. Jn: VIAL, J. L. (ed.), Evolutionary

biology of the anurans, Columbia, Missouri, Univ. Missouri Press: 273-297.

Corresponding editor: Ulrich SINSCH.

Source : MNHN, Paris

Alytes, 1998, 15 (4): 145-157. 145

Food and feeding habits in a population

of common spadefoot toads

(Pelobates fuscus)

from an island

in the lower Danube floodplain

Dan COGÂLNICEANU, Florin AIOANEI, Constantin CIUBUC

& Anghelutä VADINEANU

Bucharest University, Faculty of Biology, Splaiul Independenfei 91495, 76201 Bucharest, Romania

Analysis of the food of a population of Pelobates fuscus inhabiting an

island in the lower Danube floodplain revealed high prey diversity. Most of

the animals studied were juveniles and subadults because they represent the

largest part of the population. The high percentage of empty stomachs

(38 %) suggests that feeding is variable in time. Most of the prey items

consumed (75 %) were small, with a body length from 2 to 6 mm. A

significant positive correlation was found between toad body length and

prey length. Significant positive correlations were also found between toad

body length and the average, maximum, and minimum prey sizes. Insects

represented more than half (58.8 %) of the prey categories consumed. The

dominant groups of prey taxa were snails (Gastropoda, Pulmonata), ants

(Hymenoptera, family Formicidae), and beetles (Coleoptera). These three

taxa represent half (50.6 %) of the prey categories consumed, with the rest

of the taxa representing less than 10 % each.

Prey abundance in the environment and diversity of prey consumed

were positively correlated. There was a positive correlation between the

relative abundance in the environment and the frequency of occurrence.

Two preference indices (Manly’s Alpha Index and lvlev’s) were tested, but

both were of limited value in discriminating between prey categories

preferred or avoided. The present data suggest that Pelobates fuscus is a

generalist feeder with a broad trophic niche.

INTRODUCTION

The common spadefoot toad, Pelobates fuscus, is a strictly nocturnal, burrowing species.

Despite being distributed over most of Europe (ARNOLD & BURTON, 1992), little is known

about its feeding habits and food preferences (MEDVEDEV, 1974; SCZERBAK & SCZERBAN,

1980; NÔLLERT, 1984). It is not yet known if Pelobates fuscus is a generalist or a specialist

feeder, and if it is a sit-and-wait or an active foraging predator.

During a study on the dynamics of the amphibian communities of an island in the

lower Danube floodplain, data were gathered to allow a detailed analysis of feeding habits

Source : MNHN, Paris

146 ALYTES 15 (4)

and food choice of Pelobates fuscus. Data on the availability and diversity of litter inverte-

brates were also available.

MATERIALS AND METHODS

STUDY AREA

The study site is the island of Chiriloaia, situated south of the town of Bräila (fig. 1). The

island is flat (less then 1 m above sea-level), and is 9 km long by 4 km wide. Several large,

shallow lakes exist in the interior of the island. The vegetation cover is extremely dense and

diverse, ranging from reeds (Phragmites sp.) and cattails (Typha sp.) near the lakes and

channels, to natural willow tree forests and willow and poplar tree plantations. Old willow

trees provide shelter and large amounts of wood debris. Clearcuts and pastures represent less

than 10 % of the island'’s area.

Life on the island is controlled by the periodic floods of the Danube, as suggested by the

flood-pulse concept (BAYLEY, 1995). During the spring floods water levels rise more than 4 m.

At times of high waters, the entire island is submerged. Most of the animals survive these

often prolonged periods of flooding by seeking refuge on the large amounts of floating debris

and on large, old willow trees.

SAMPLING PROGRAM

During 1994 and 1995, 70 one- and two-liter pitfall traps were used in the field. Sampling

started in July 1994, when the water level began to decrease after the spring flood. The traps

were filled with 40 % ethylene glycol and were emptied twice a month. During the high water

levels in the spring of 1995, some of the traps were flooded and the captured animals lost.

Aside from invertebrates and other amphibians, 162 juvenile and subadult Pelobates fuscus

were captured in these traps, primarily during their migrations (COGÂLNICEANU et al., 1997).

Eight adults (seven males and one female) and 20 juveniles were captured at night during

torch surveys. In total, 190 spadefoot toads were captured, measured to the nearest 0.1 mm

with dial calipers, and their stomach contents were preserved in 70 % alcohol for identifica-

tion. Prey items were classified by taxon (to family when possible) and life stage, and their

length was measured to the nearest 0.1 mm using a binocular microscope. The invertebrates

captured in the pitfall traps were also counted and identified.

DATA ANALYSIS

The relative abundance of the various prey categories was estimated from the stomach

content analysis. The frequency of occurrence was determined by dividing the number of

stomachs that contained a particular prey type by the total number of stomachs with prey.

Source : MNHN, Paris

COGÂLNICEANU et al. 147

Fig. 1. - Location of the study site (island of Chiriloaia) in the lower Danube floodplain.

The trophic diversity was estimated with Brillouin’s index (HB) and evenness (E)

(MAGURRAN, 1988):

128 = MINE and

E=HBIHB,

where N is the total number of prey individuals and n, is the number of individuals of prey

type i.

The Berger-Parker (d) index of dominance was also used, as a measure of the degree of

specialization:

d= 11 max

where d varies between 1/N and 1. A value of d close to 1 indicates a high specialization in

feeding, while a low value close to 1/N is characteristic of a generalist feeder.

The different prey resource states were compared with the relative numerical abun-

dance in the environment and the selectivity in feeding was estimated using Manly’s Alpha

prey preference index (MANLY et al., 1972; CHESSON, 1978):

Source : MNHN, Paris

148 ALYTES 15 (4)

No. of individuals

= Captured

in traps

60 Ge

# Stomach

content

50 analysed

July Aug. Sep. Oct. Nov. Dec. Jan. Feb. March Apr. May June July Aug. Sep.

1994 1995

Fig. 2. - Numbers of individuals of Pelobates fuscus captured monthly in traps and numbers of stomach

contents analyzed.

where n, and n, represent the abundance of prey taxa i and j in stomach contents, while r; and

r; represent their abundance in the environment. If a; > 1/T, where T is the total number of

prey categories, then the prey type i is considered preferred. Selectivity in feeding also was

estimated according to IVLEV (1961):

where E can vary between -1 and 1.

RESULTS

The frequency of capture in traps and field observations indicated that the highest

number of active spadefoot toads was observed when newly metamorphosed juveniles began

migrating away from water (fig. 2). Increased activity was also observed during or after rain.

Most of the animals studied were juveniles and subadults (fig. 3a), because they represented

the largest part of the population (COGÂLNICEANU et al., 1997).

Each of 119 individuals of the 190 studied (62 %) contained at least one prey item in its

stomach. The average number of prey items per stomach was 5.5, ranging between 1 and 17

Source : MNHN, Paris

COGÂLNICEANU et al. 149

No. of individuals

85 40 50

Body length (mm)

300 -

260 +

No. of individuals

B à

L 1

0 a

0 10 20 30 40 50

Prey body length (mm)

Fig. 3. - (a) Size distribution of the 190 individuals of Pelobates fuscus studied. (b) Size distribution of the

664 prey items identified in stomach contents.

Source : MNHN, Paris

150 ALYTES 15 (4)

(fig. 4). A total of 664 invertebrates from 36 prey categories were identified from the stomach

contents (tab. 1). The average number of individuals per prey category was 17 + 3.7. The

number of prey categories encountered increased with the number of stomachs analyzed,

approaching the upper limit at 100 stomachs. Overall, the average number of prey categories

per stomach was low (1.4 + 0.08).

Most of the prey consumed (75 %) were small, with a body length between 2 and 6 mm

(ig. 3b). The largest prey taxa belonged to four categories: Annelida, Diplopoda, Chilopoda

and larvae of Lepidoptera. A significant positive correlation was found between toad body

length and prey length (r = 0.29, P < 0.001, n = 634). Significant positive correlations were also

found between toad body length and average, maximum and minimum prey size (fig. 5). The

lower prey size limit for Pelobates fuscus is around 0.8-1 mm, while the upper limit seems

constrained only by the width of the mouth.

Insects represented more than half (58.8 %) of the prey categories consumed, with ants

(Formicidae) and beetles (Coleoptera) being dominant. These two insect taxa together with

snails (Pulmonata) comprised half (50.6 %) of the prey categories consumed. The remaining

taxa represented less than 10 % each (tab. 1). Spadefoot toads had a broad trophic niche, with

little specialization in feeding. The mean number of individuals per stomach was greater than

2.0 for snails (Pulmonata), ants (Formicidae) and fly larvae (Muscidae). These three prey taxa

either are slow moving or are distributed locally in the environment, indicating that Pelobates

Juscus is actively foraging on them. Prey diversity was high, with a Brillouin diversity index of

2.83, and evenness of 0.83. The Berger-Parker index of dominance has a low value (4 = 0.18),

with the dominant prey taxa being ants (Formicidae).

Pitfall traps are commonly used for capturing cursorial invertebrates and in their action

they may be regarded as analogous to a sit-and-wait predator (CoRNIsH et al., 1995). Catch

size is dependent on both abundance and activity (THIELE, 1977) and can correlate well with

prey encounter rate for spadefoot toads. A total of 55,364 invertebrates from 53 taxa were

captured with pitfall traps and identified during the study period. The relative abundance of

prey types in stomach contents and in the environment and the values of the two selectivity

indexes are presented in tab. 2. The analysis of the relative abundance of the different taxa

from the environment showed that five categories represented more than 70 % of the identi-

fied prey taxa. These were: Crustacea, Acari, Collembola, Formicidae and Carabidae. Seven-

teen taxa present only occasionally in the pitfall trap captures were not found amongst the

food of spadefoot toads.

The frequency of occurrence of two of the dominant taxa (Hymenoptera and Coleo-

ptera) correlated well with their relative abundance in the environment. For Gastropoda, the

estimated abundance from trap captures is biased, due to the low mobility of this group. The

most abundant taxon in traps is Collembola, a group of tiny, soil dwelling insects.

The relative abundance of the different prey taxa in the environment and in the stomach

contents of Pelobates fuscus is positively correlated (r = 0.71, P < 104) (fig. 6). There is also a

positive correlation between the relative abundance in the environment and the frequency of

occurrence (r = 0.69, P < 10‘).

Source : MNHN, Paris

COGÂLNICEANU et al. 151

Tab. 1. - Prey taxa removed from stomachs of 119 spadefoot toads. n: number of individuals per taxon. A,:

percent of the total number of invertebrates accounted for by the particular prey type. FO: frequency of

occurrence, i.e. number of stomachs containing at least one prey item, divided by the total number of

stomachs with prey (119), multiplied by 100. ANIS: average number of individuals/stomach.

Platyhelminthes

Annlida, Oligochaeta

Crustacea, Isopoda, Oniscoidea

Gastropoda, Pulmonata

Diplopoda, Juliformia

Chilopoda, Lithobiomorpha

Collembola, Entomobryidae

Collembola, Sminthuridaë

Total Collembola

Dermaptera, Forficula sp.

Homoptera, Aphididae

Homoptera, Cicadidae

Total Homoptera

Hymenoptera, undetermined

Hymenoptera, Formicidae

Total Hymenoptera

Diptera, adults undetermined

Diptera, larvae undetermined

Diptera, Muscidae, adults

Diptera, Muscidae, larvae

Diptera, Culicidae

Total Diptera

Coleoptera, undetermined

Coleoptera, Carabidae, adults

Coleoptera, Carabidae, larvae

Coleoptera, Chrysomelidae

Coleoptera, Curculionidae

Coleoptera, Coccinellidae

Coleoptera, Elateridae

Coleoptera, Staphylinidae

Coleoptera, Syiphidae

Total Coleoptera

BraownannaRs Hsrssaw

Source : MNHN, Paris

25-

20 +

Frequency

2 4 6 8 10 12 14 16

No. of prey/stomach

Fig. 4. — Frequency distribution of the number of prey found per stomach in the 119 individuals of

Pelobates fuscus with at least one prey item in their stomach.

# average length

16) [a minimumlength | à @%

R=0.35 P<0.0001

R=0.39 P<0.001

R=0.35 P<0.005

Log {prey body length]

T T T

140 145 150 185 160 165 170

Log (body length Æ/oéates fscus)

Fig, 5. - Relationships between spadefoot toad body length and average, maximum and minimum prey

body length.

Source : MNHN, Paris

COGÂLNICEANU et al.

153

Tab. 2. - Prey abundance in the stomach contents (sc) of spadefoot toads and in the environment (env),

expressed as number of individuals (7) and percentage of the total number (A). Also shown are the

values for the two measures of prey preference: Manly's Alpha index (&) and Ivlev's index (E).

‘ Preferred prey taxa (a, > 0.035); ” Preferred prey taxa (E, > 0.5); ” Avoided prey taxa (E, < -0.5).

Prey taxa

Annelida, Oligochaeta

Crustacea, Oniscoidaea

Gastropoda, Pulmonata

Diplopoda, Juliformia

Chilopoda, Lithobiomorpha

Arachnida, Acari

Arachnida, Araneae

Arachnida, Opilionidae

Collembola

Dermaptera

Hemiptera

Lepidoptera, larvae

Orthoptera

Homoptera, Aphididae

Homoptera, Cicadidae

Other Homoptera

Hymenoptera

Hymenoptera, Formicidae

Diptera

Coleoptera, larvae

Coleoptera, Carabidae

Coleoptera, Chrysomelidae

Coleoptera, Curculionidae

Coleoptera, Coccinellidae

Coleoptera, Elateridae

Coleoptera, Staphylinidae

Coleoptera, Sylphidae

As (%)

0.77 28

8.22 6589

19.72 1363

5.59 1799

1.24 495

1.86 4837

4.50 3726

0.31 315

7.29 | 10543

0.62 il

0.46 369

341 148

0.46 122

2.63 520

1.86 310

5.12 830

2.48 1014

19.25 | 11354

5.12 1663

1.08 2002

4.03 6349

0.93 76

0.93 52

0.31 7

0.46 22

0.93 396

0.15 341

DISCUSSION

The feeding habits of any species lies somewhere between the two extremes of foraging

patterns: sit-and-wait and active-foraging. The population of Pelobates fuscus studied is

nearer the active foraging mode. This can be inferred from the large numbers of prey types

with relative low mobility (Gastropoda, Crustacea) and isolated distributions in the environ-

ment (Formicidae and larvae of Muscidae) consumed. Individuals in the studied population

of Pelobates fuscus appear to be opportunistic feeders with a wide trophic niche and a lack of

selectivity in feeding.

Few papers present data on the food and feeding habits of spadefoot toads. Most of the

existing studies are based on the analysis of a small number of animals. For example,

Gurowski & KRZYSZTOFIAK (1988) analyzed the stomach contents of only two animals, and

Source : MNHN, Paris

154 ALYTES 15 (4)

Log {4bundance in the environment]

Log (Abundance in stomach contents)

Fig. 6. - Correlation between prey abundance in the environment and abundance in stomach contents.

other authors have ignored small prey taxa, like Collembola and Acari (MEDVEDEV, 1974;

SCZERBAK & SCZERBAN, 1980). Contrary to the present results, KUZMIN (1995) reported that

spadefoot toads preferred crawling invertebrates while ants are consumed less frequently.

Among Coleoptera, this author stated that beetles belonging to the families Carabidae and

Tenebrionidae are preferred, while Juszczyk (1974) stated that spadefoot toads prefer

Carabidae, Elateridae and Staphylinidae.

MEDVEDEV (1974) analyzed the stomach contents of 25 spadefoot toads and identified

100 prey items, of which 9 % were spiders and 81 % were Coleoptera. Ants were completely

lacking from the diet, although they were present in the food of the other amphibian species

from the same location. The Berger-Parker index was 0.23 when the most abundant carabid

species (Harpalus distinguendus) was considered, and 0.64 when all carabids were pooled. It is

possible that because smaller animals were not counted these results were skewed. SCZERBAK

& SCZERBAN (1980) identified 48 prey items in the stomach contents of 25 spadefoot toads.

The most abundant taxon was also Carabidae (Berger-Parker index d = 0.23). They also

reported the presence of several flying insects like Noctuidae (12.6 %) and Ichneumonidae

Source : MNHN, Paris

COGÂLNICEANU et al. 155

(4.2 %). Ants were represented by only two individuals (4.2 %). The small proportion of ants

reported by these authors suggests that, although generally abundant in the environment, ants

are usually avoided. In the present study, Pelobates fuscus inhabited a highly variable,

unpredictable environment, and ants were consumed possibly because of a limited choice of

other prey during floods. A study of the feeding ecology of two genera of North American

spadefoot toads (Scaphiopus and Spea) showed that these are generalized arthropod preda-

tors, with ants representing a significant component of their diet (PUNZO, 1991).

The striking variations in food profiles of spadefoot toads reported by different authors

may be explained by differences in prey availability and may support the idea that Pelobates

fuscus is a generalist feeder. No further comparisons are possible due to the lack of informa-

tion on prey availability and the fact that prey taxa identification was done at different

taxonomic levels.

Many papers are devoted to the study of prey availability in relation to selective

predation (CHESsON, 1978; GRIFFITHS, 1975; HURTUBIA, 1973; SMITH, 1982; WINEMILLER &

PiankA, 1990). Of the various approaches proposed, two of the simplest measures of prey

preference (Manly’s Alpha index and Ivlev’s index) were tested in the present study (tab. 2).

Manly’s Alpha index is not very useful because the preferred taxa, Annelida, Gastro-

poda, and Lepidoptera larvae, have low mobility and a low probability of capture in traps, so

their relative abundance in the environment is probably higher than estimated. Another taxon

that appeared to be preferred (Dermaptera) yielded low numbers of individuals in the

stomach content analysis and in the trap captures, so no valid conclusion can be drawn. The

same caution must be taken when considering the various preferred families of Coleoptera.

The selectivities computed using this index tended to give counterintuitive weight to some

insignificant taxa. Therefore, no definite conclusions about prey preference can be drawn from

using this index.

Ivlev’s index measures not only the preference for a prey type but also the lack of

preference. Positive values indicate preference while negative values suggest avoidance of the

prey. À threshold of + 0.5 was considered, indicating the preference (> 0.5) or avoidance (<

—0.5) for a particular prey taxon. Only three prey taxa were avoided (Acari, Coleoptera larvae

and Sylphidae), with E; values smaller than — 0.5. Acari and Coleoptera larvae have low

mobility and live mainly in the litter, so they are consumed only accidentally. Sylphidae are

present in very low numbers and are active mainly during daylight, contrary to spadefoot

toads that feed only during the night.

Ten taxa had positive values greater than 0.5, suggesting a preference in prey choice. Of

these, eight were also considered preferred by the Manly’s Alpha index. The other two taxa

(Cicadidae and unidentified Homoptera) are mainly plant dwelling insects and are seldom

caught in traps. Although Ivlev’s index discriminates better in analyzing selectivity in feeding,

most of the results are biased by the method of estimating relative abundance in the

environment of the particular taxa.

A high number of juveniles, up to 96 % of the entire population, was also recorded in a

Pelobates fuscus population from an artificial island surrounded by the Danube, near Vienna

(JEHLE et al., 1995). The large number of empty stomachs encountered, together with the large

stomach volume of this species, suggests that feeding varies in time depending on weather

Source : MNHN, Paris

156 ALYTES 15 (4)

conditions, rain and air temperature. During the study period, spadefoot toads were active

and feeding during most of the year, except for the three-month period of frost from

December 1994 to February 1995 (fig. 2). According to KUZMIN (1995), spadefoot toads use

active search foraging tactics, searching up to 200 m° each night.

Theslight positive correlation between the size of spadefoot toads and prey size, together

with the diversity of prey categories encountered, suggested that larger animals eat larger

prey, which is consistent with GRiFFITHS’s (1975) predictions that predators can eat only a

certain size range of prey organisms. The low occurrence of Collembola in food is possibly

due to their small dimensions that make them difficult to locate and not energetically

rewarding to pursue. BoomsA & ARNTZEN (1985) found that the proportion of Collembola in

the stomach contents in a population of Bufo calamita varied sharply during the year, their

relative contribution being negatively correlated with the size of the toad. However, this does

not limit the breadth of the trophic niche, and larger animals consumed a higher diversity of

food. Partitioning of resources between size classes is thus possible, as previously observed by

JoLy & GIACOMA (1992).

LITERATURE CITED

ARNOLD, E. N. & BURTON, J. A., 1992. - Reptiles and amphibians of Britain and Europe. London, Collins.

BAYLEY, P. B., 1995. - Understanding large river-floodplain ecosystems. BioScience, 45 (3): 153-158.

Boomsa, J. J. & ARNTZEN, J. W., 1985. - Abundance, growth and feeding of natterjack toads (Bufo

calamita) in a 4-year old artificial habitat. J appl. Ecol., 22: 395-405.

CHessoN, J., 1978. - Measuring preference in selective predation. Ecology, 59: 211-215.

COGÂLNICEANU, D., CRISTOFOR, S. & VADINEANU, À., 1997. - The dynamics of amphibian communities

on two islands of the lower Danube floodplain. Preliminary results. In: W. BÔHME, W. BISCHOFF &

T. ZIEGLER (ed.), Herpetologia Bonnensis: 71-80.

CorNisH, C. A., OLDHAM, R. S., BULLOCK, D. J. & BULLOCXK, J. A., 1995. - Comparison of the diet of

adult toads (Bufo bufo L.) with pitfall trap catches. Herp. J., 5: 236-238.

GRIFFITHS, D., 1975. — Prey availability and the food of predators. Ecology, 56: 1209-1214.

Gurowski, J. M. & KRzYSZTOrIAK, L., 1988. - Materials for the investigation of food composition in

anuran amphibians in north-eastern Poland. Przeglad Zool., 32: 225-235. [In Polish].

Hurrugia, J., 1973. — Trophic diversity measurement in sympatric predatory species. Ecology, 54:

885-890.

IVLEV, V. S., 1961. — Experimental ecology of the feeding of fishes. New Haven, Connecticut, Yale

University Press.

JERLE, R., Hôpz, W. & THONKE, A., 1995. — Structure and dynamics of central European amphibian

populations: a comparison between Triturus dobrogicus (Amphibia, Urodela) and Pelobates fuscus

(Amphibia, Anura). Aust. J. Ecol., 20: 362-366.

JoLy, P. & GracoMa, C., 1992. - Limitation of similarity and feeding habits in three syntopic species of

newts (Triturus, Amphibia). Ecography, 15: 401-411.

Juszczvk, W. 1974. — Plazy i gady krajowe. Warsaw. [In Polish].

KUzMIN, S. L., 1995. — Die Amphibien Russlands und angrezender Gebiete. Magdeburg, Westarp Wis-

senschaften: 1-274.

MAGURRAN, À. E. 1988. - Ecological diversity and its measurement. London, Croom Helm.

Many, B.F., MILLE, P. & Cook, M., 1972. - Analysis of a selective predation experiment. 4m. Nat.,

106: 719-736.

MEebvEDEv, S. I., 1974. - Data on study of amphibians’ food in the region of the middle flow of the

Seversky Donets river. Vestmik Zool., 1: 50-59. [In Russian].

NÔLLERT, A., 1984. — Die Knoblauchkrôte. Wittenberg Lutherstadt, Zimsen Verlag, Die Neue Brehm-

Bucherei, 561.

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COGÂLNICEANU et al. 157

Punzo, F, 1991. - Feeding ecology of spadefooted toads (Scaphiopus couchi and Spea multiplicata) in

western Texas. Herp. Rev., 22 (3): 79-80.

SCZERBAK, N. N. & SCZERBAN, M. I. 1980. — Amphibians and reptiles of Ukrainian Carpathian mountains.

Kiev, Naukova Dumka Publ. [In Russian].

SMITH, E. P., 1982. — Niche breadth, resource availability, and inference. Ecology, 63: 1675-1681.

THieLE, H. U., 1977. - Carabid beetles in their environments. Springer-Verlag, Zoophysiology and Ecology,

10.

WINEMILLER, K.. O. & PIANKA, E. R., 1990. - Organization in natural assemblages of desert lizards and

tropical fishes. Ecol. Mon., 60 (1): 27-55.

Corresponding editor: Janalee P. CALDWELL.

Source : MNHN, Paris

Alytes, 1998, 15 (4): 158-170.

Aspectos de la reproducciôn

in-vitro y del desarrollo larval

de Melanophryniscus stelzneri

(Wevenbergh, 1875)

(Anura, Bufonidae),

con comentarios acerca

del érgano de Bidder

Dinorah D. ECHEVERRIA

Universidad de Buenos Aires, Facultad de Ciencias Exactas y Naturales,

Departamento de Ciencias Biolôgicas, Laboratorio de Vertebrados, 1428 Buenos Aires, Argentina

Fax: 54.1.782.0620 - E-mail: echeverria@biolo.bg.fcen.uba.ar

Some aspects of the ontogenetic development of Melanophryniscus

stelzneri from in-vitro fecundation to metamorphosis are described,

emphasizing the larval development. Unfertilized egg diameter varies from

1400 to 1600 um, eggs are arranged in clusters, and their colour is

polarized. The animal hemisphere is brown and the vegetative hemisphere

is light yellow or white. The eclosion takes place in stage 21 at 23 + 1 °C.

The embryos and exotrophic larvae are hazel brown pigmented. No evi-

dence of Bidder’s organ is found in any larval, postmetamorphic, juvenile,

preadult or adult stage.

INTRODUCCION

FERNANDEZ (1927) efectué las primeras observaciones del desarrollo embrionario de

Melanophryniscus stelzneri (Weyenbergh, 1875) proveniente de una poblacién de las Sierras

de Cérdoba. Hizo referencia a la pigmentaciôn castaña que presentaban los huevos y

embriones, sin describir el individuo completo ni la larva. BOKERMANN (1967) y STARRETT

(1967) describieron la larva de M. moreirae. PRIGIONI & LANGONE (1990) realizaron la

descripciôn de la larva de M. orejasmirandai, señalando las diferencias y similitudes externas

mäs conspicuas con las larvas de M. stelzneri montevidensis y M.s. stelzneri sobre la base de las

descritas por GARRIDO (1989) y FERNANDEZ (1927), respectivamente. PRIGIONI & LANGONE

(1990) manifestaron que las descripciones se realizaron de manera incompleta debido a que

no disponian de suficiente informaciôn en el caso de M.s. stelzneri. ECHEVERRIA (1992) aporté

evidencias acerca de la microanatomia bucal de la larva de M. stelzneri y coincidié con

FERNANDEZ (1927) en la férmula dentaria de M. stelzneri. LANGONE (1994) expresé que

algunas subespecies de M. stelzneri han sido consideradas especies plenas: es el caso, por

Source : MNHN, Paris

ECHEVERRIA 159

ejemplo, de M. stelzneri de las Sierras de Cordoba y de San Luis en Argentina y de M.

montevidensis, que estä presente en Argentina, Uruguay y Brasil.

El propésito de este trabajo es contribuir al conocimiento del desarrollo larval de M.

stelzneri y aportar nueva informacién acerca del 6rgano de Bidder de dicha especie.

MATERIAL Y MÉTODOS

Se capturaron cuatro hembras adultas y dos machos de Melanophryniscus stelzneri de la

localidad de El Trapiche (provincia de San Luis) para producir una induccién de la ovulaciôn

y fertilizaciôn in-vitro. La induccién de la ovulaciôn se Ilev6 a cabo con hipéfisis no homéloga

de un macho de Bufo arenarum procedente de la provincia de Buenos Aires (localidad de José

C. Paz). Se utilizo una hipéfisis de Bufo arenarum macerada en 5 ml de Holtfreter 0,1 N. A

cada hembra de M. stelzneri se le inyecté en forma subcutänea 1 ml (0,5 ml por lado) en la

regiôn ventral del abdomen. Cada hembra fue alojada en un recipiente cuyo fondo se

mantuvo hümedo.

Para efectuar la fertilizaciôn se utilizaron los testiculos de M. stelzneri. Cada testiculo fue

macerado en 10 ml de Holtfreter 0,1 N y, sin diluir, se puso en contacto con los oocitos. El

tiempo de contacto para la inseminaciôn fue de 10 minutos.

El desarrollo se Ilevé a cabo en bandejas de plästico de 80 mm de profundidad, con

abundante medio liquido. Durante las primeras 48 horas los huevos se desarrollaron en

liquido de Holtfreter 0,1 N a una temperatura de 20°C. Luego, se cambi6 paulatinamente a

agua mineral natural y los embriones continuaron su desarrollo a una temperatura ambiente

de 23 + 1 °C. Los estadios del desarrollo fueron identificados segün la nomenclatura

propuesta por GosER (1960). El cuidado de los embriones y la alimentaciôn de las larvas se

Ilev6 a cabo segün ECHEVERRIA & FIORITO (1981). Los juveniles y adultos de M. stelzneri

fueron alimentados con insectos vivos (Drosophila melanogaster), isépodos terrestres

pequeños (adultos y/o crias de Armadillium sp. y Porcellio sp.), y colémbolos. A los ejemplares

adultos de M. stelzneri y de B. arenarum se les administraron ademäs trozos de higado de vaca

fresco cada dos dias.

A efectos de realizar comparaciones de coloracién y, en los casos que fuera necesario,

verificar la presencia de la génada y/o del érgano de Bidder, se conté con larvas (vivas y fijadas

en formol 10%) de Bufo arenarum y de Melanophryniscus stelzneri en distintos estadios,

capturadas en la naturaleza. Se efectuaron observaciones con microscopio estereoscopico

in-vivo y en material fijado en formol 10 %, que ilustran la ontogénesis y la anatomia interna

en la regiôn abdominal. Se efectuaron observaciones con microscopio éptico de los cortes

histolégicos de las génadas en larvas, juveniles y adultos.

En busca de evidencias de la presencia del 6rgano de Bidder y del desarrollo de las

gônadas, se analizaron 7 ejemplares durante la etapa larval y 12 juveniles y adultos. El

material examinado es el siguiente: 5 ejemplares de M. stelzneri montevidensis de la colecciôn

del Laboratorio de Investigaciones Herpetolôgicas de la Universidad de Buenos Aires

(LIHUBA 2039, 3 machos y 2 hembras, provenientes de la localidad de Sierra de La Ventana),

y 7 ejemplares de M. stelzneri stelzneri de la colecciôn del Laboratorio de Vertebrados de la

Source : MNHN, Paris

160 ALYTES 15 (4)

Fig. 1. - (a) Dos oocitos de Melanophryniscus stelzneri sin fecundar, en la câpsula (C). Escala: 2 mm.

(b) Dos embriones de M. stelzneri en estadio 20. Escala: 2 mm.

(c-d) Estadio 42 de M. stelzneri. Escala: 5 mm. (c) Vista ventral. S: orificio del espiräculo. (d)

Vista dorsal.

Source : MNHN, Paris

ECHEVERRIA 161

Fig. 2. - Vista dorsal y ventral de M. stelzneri (a, c) y de Bufo arenarum (b, d) en estadio 44. Escala: 5 mm.

Source : MNHN, Paris

162 ALYTES 15 (4)

Tab. 1. - Registro de las cuatro hembras de A4. s. stelzneri. LT: largo total (mm), NOO: nümero de oocitos

ovulados; TO: tipo de oocitos. E: oocitos ovulados y hallados en el exterior del cuerpo; O: oocitos

retenidos en el ovisaco.

Facultad de Ciencias Exactas y Naturales de la Universidad de Buenos Aires (LVFCENUBA

4, 6,8, 12, 16, 17 y 78, provenientes de las localidades de El Trapiche y de Tanti). En 3 larvas

(estadio 36, 42 y 45) se realizaron cortes histolôgicos transversales y longitudinales.

Las técnicas Ilevadas a cabo para las observaciones en microscopio 6ptico son las

utilizadas por ECHEVERRiA (1988). Los cortes se realizaron a 7 um. En los juveniles y adultos

se extrajo el opistonefros con la génada y el cuerpo graso correspondientes al lado izquierdo.

La coloracién utilizada es hematoxilina-Masson. Se tomaron fotografias de los huevos,

embriones, larvas y de la regién urogenital de juveniles y adultos disecados, con microscopio

estereoscopico.

RESULTADOS

La ovulacién se produjo dentro de las 21 horas después de la inyecciôn subcutänea de

macerado de hipôfisis de Bufo arenarum. La respuesta de las mismas se expresa en la tab. 1.

Los oocitos recién liberados y sin fecundar se mantienen formando un racimo. Cada uno

estä rodeado por una sustancia gelatinosa. Los huevos con su câpsula gelatinosa individual se

unen por medio de filamentos de adherencia que dificultan el manipuleo de los mismos. El

diâmetro medio de los oocitos recién ovulados y de los huevos oscila entre 1400 y 1600 um.

Antes de la fecundacin los oocitos presentan los hemisferios de diferente coloraciéôn. La

pigmentacién estä polarizada: el hemisferio animal es de color castaño y el hemisferio

vegetativo es de color blanco amarillento. En el polo animal la pigmentacién es de color

blanco amarillento rodeado de castaño (fig. 1a). La coloraciôn castaña se mantiene en el

hemisferio animal de los huevos fecundados y en los embriones de los primeros estadios de la

etapa embrionaria (estadios 1 al 8).

Los embriones presentan un color castaño claro, que es homogéneo desde el estadio de

placa neural hasta el estadio de circulacién branquial inclusive (13 al 20).

La eclosiôn se produce aproximadamente a las 50 horas de la fecundaci6n, cuando los

embriones se hallan en el estadio 21.

En el estadio 21 y 22 el color del cuerpo y de la musculatura del cola se mantiene castaño

claro; las aletas dorsal y ventral de la cola son semitransparentes y carecen de la coloraciôn

castaña del resto del cuerpo (fig.1b).

Source : MNHN, Paris

ECHEVERRIA 163

Tab. 2. - Registro de datos meristicos (mm) en una larva de M. s. stelzneri en estadio 34 y otra en estadio 44.

Estadio

Largo total

Distancia rostro-espiräculo

Distancia espiräculo-posterior

Largo cuerpo

Largo cola

Altura müsculos cola

Altura aletas

Altura mâxima cuerpo

Ancho mâximo cuerpo

Ancho cuerpo a nivel ojos

Distancia naso-ocular

El esbozo de miembros posteriores se evidencia en el estadio 25 con pequeñas protube-

rancias blanquecinas. En los estadios 26 al 30, la region dorsal del esbozo del miembro

posterior toma una coloraciôn castaña homogénea, y la regiôn ventral del mismo es blanque-

cina. El cuerpo, los müsculos caudales y el iris presentan una pigmentacién homogénea de

color castaño. Las aletas caudales son transparentes.

En el estadio 34 el cuerpo de la larva es globoso, y los ojos y las narinas son dorsolate-

rales. El ancho mäximo estä ubicado en el cuerpo. El hocico es redondeado en vista dorsal y

lateral. El disco oral es triangular, de posiciôn subterminal ventral y con una emarginaciôn

comisural. Las papilas orales son simples y conicas; estän localizadas ünicamente en las

regiones supra e infracomisurales. Presenta amplias brechas rostral y mental. Los orificios

nasales son dorsales, de forma levemente oval, con el borde anterior dirigido hacia la linea

media. El espiräculo, de posicién ventrolateral, se halla sobre el lado izquierdo del cuerpo. La

abertura del tubo espiracular estä elevada. El tubo cloacal es mediano. La aleta caudal

representa el 41,79 % de la longitud total. La aleta dorsal nace sobre el cuerpo. La aleta

ventral surge a continuaciôn del tubo cloacal. La regiôn mediana de la aleta ventral presenta

escaso pigmento; los extremos distal y proximal son transparentes. El margen posterior de la

aleta caudal es redondeado (fig. 4).

A medida que en el basipodio se diferencian los dedos de la mano o del pie, la regiôn

ventral del mismo y los extremos de los dedos permanecen incoloros. Los miembros anterio-

res mantienen el mismo patrôn de coloraciôn que los miembros posteriores. La regiôn ventral

de los basipodios anterior y posterior y de los respectivos dedos (palma de la mano y del pie),

se mantiene incolora hasta el estadio 46 inclusive (fig. 1c).

Desde el estadio 28 al 30, la coloraciôn dorsal del cuerpo se torna ms oscura y la ventral

blanquecina. Esta diferencia en la coloraciôn se incrementa durante el climax metamérfico

(fig. 1d y 2). La longitud total de los juveniles que completaron la metamorfosis en el

laboratorio alcanzé 0,8 mm. En las figuras 3, 4 y 5 se ilustran algunos estadios del desarrollo

de M. stelzneri.

Source : MNHN, Paris

164 ALYTES 15 (4)

Fig. 3. - Esquema de un oocito y de dos estadios del desarrollo larval en periodo prometamérfico de M.

stelzneri. (A) Oocito sin fecundar. (B) Embriôn en estadio 20. (C) Embriôn en estadio 21. Escala:

2mm.

Fig. 4. Periodo premetamérfico de M. stelzneri: esquema del estadio 34. Vistas ventral (v), dorsal (d) y

lateral izquierda (t), y detalle de miembro posterior izquierdo (p). Escala: 2 mm.

En la tabla 2 se registran los valores meristicos en una larva en estadio 34 y otra en estadio

44.

OBSERVACIONES DE LA ANATOMÏA INTERNA

Los cortes histolégicos efectuados en las larvas, juveniles y adultos revelaron la ausencia

del érgano de Bidder o de su esbozo. Sélo se hallé la presencia de la pregénada (fig. 6).

Source : MNHN, Paris

ECHEVERRIA 165

Fig. 5. - Esquema de algunos estadios del desarrollo larval en climax metamérfico de M. stelzneri: (A)

estadio 42; (B) estadio 43; (C) estadio 45. Vista dorsal (d), ventral (v) y lateral izquierda (t); detalle

de miembro anterior derecho en vista ventral (m); detalle de miembro posterior izquierdo en vista

lateral interna (p). Escala: 2 mm.

En ejemplares de estadio 42, el higado se observa de color amarillo. No se hallan vestigios

del érgano de Bidder, ni de las génadas. La pigmentaciôn negra del higado, caracteristica de

los adultos, se hallé en los ejemplares en metamorfosis avanzada (estadios 45 y 46).

En los machos adultos, el peritoneo que cubre los testiculos puede observarse totalmente

blanco, negro o combinado; los testiculos pueden presentar forma oval o levemente esférica,

destacändose sobre la regiôn anterior de los riñones (fig. 7).

DISCUSIÔN

La coloracién de los oocitos no fecundados y de los huevos de Melanophryniscus

stelzneri es castaña en el hemisferio animal y amarillenta en el hemisferio vegetativo en las

hembras adultas provenientes de El Trapiche, de Tanti y de la Sierra de la Ventana. Esto no

coincide con lo manifestado por FERNANDEZ (1927) quien mencioné que, en los ejemplares

Source : MNHN, Paris

166 ALYTES 15 (4)

Fig. 6. - Corte transversal a nivel urogenital anterior, en una larva de M. stelzneri en estadio 45. A: aorta

dorsal; G: pregénada; P: vena pstcava; R: opistonefros. Escala: 240 um.

por ella estudiados, procedentes de Tanti, los huevos mostraron un color homogéneo marrôn

claro. Quizäs esta diferencia esté en relaciôn con el estado de maduracién de los oocitos

(Roca & ECHEVERRIA, 1996).

Los huevos examinados de M. stelzneri son 1,4 a 1,6 mm de diâmetro. Esta variaciôn

podria depender de factores intrinsecos de cada hembra. Roca & ECHEVERRiA (1996) halla-

ron en M. stelzneri oocitos ovulables de 1,4 mm con la coloraciôn polarizada, y MCDIARMD

(1971) mencioné “huevos con el polo animal negro y el polo vegetativo claro”, de 1,2 mm de

diâmetro. Estos resultados no coinciden con FERNANDEZ (1927) quien mencioné 1 mm de

diâmetro y coloraciôn homogénea para los oocitos recién puestos de M. stelzneri provenientes

de una poblacién de Tanti, Cérdoba.

A partir de las observaciones efectuadas en el laboratorio, en M. stelzneri he compro-

bado que la eclosién se realiza en el estadio 21 a 23 + 1 °C. El desarrollo hasta la

metamorfosis se realizé en 4 semanas. Es posible que en la naturaleza el lapso posterior a la

eclosiôn se reduzca o se prolongue, segün las condiciones ambientales del momento. Estas

suelen ser muy drästicas y variadas durante el verano, desde octubre a febrero, época en que

se reproduce M. stelzneri en San Luis y Cordoba.

DEL CONTE & SILRIN (1951) y ECHEVERRIA & FIORITO DE LOPEZ (1981) realizaron el

desarrollo in-vitro de Bufo arenarum a 18 y 23 °C respectivamente, alcanzando la liberaciôn de

Source : MNHN, Paris

ECHEVERRIA 167

Fig. 7.- Regiôn urogenital de un ejemplar macho adulto de M. stelzneri. E: cuerpo graso; R: opistonefros;

T: testiculo. Escala: 4 mm.

la gelatina, en el estadio 17, en 76 horas a 18°C y la eclosiôn, en el estadio 18, a las 100 horas

a 23°C. En Melanophryniscus stelzneri la eclosién se produjo a las 50 horas de la fecundaciôn;

por lo tanto se puede considerar que el periodo embrionario de M. stelzneri es corto,

implicando una râpida velocidad inicial de desarrollo. Esta caracteristica le permite el uso de

recursos tales como cuerpos de agua temporarios, de escaso volumen y/o profundidad.

En coincidencia con BUDGETT (1899; fide MCDiaRMID, 1971), en M. stelzneri cada huevo

estä rodeado por una câpsula gelatinosa. Esto mismo ocurreen M. moreirae, en que los huevos

se hallan rodeados por câpsulas gelatinosas individuales (BOKERMANN, 1967; STARRETT, 1967).

Los huevos de M. stelzneri se unen unos a otros por medio de filamentos de adherencia.

Las larvas exotrôficas de M. stelzneri presentan un tipo ecomorfolégico benténico (en el

sentido utilizado por ALTIG & JOHNSTON, 1989) que frecuentan ambientes lôticos tempora-

Source : MNHN, Paris

168 ALYTES 15 (4)

rios de escasa o nula corriente. LANGONE (1994) menciona las caracteristicas distintivas de la

larva de M. montevidensis, coincidiendo con M. stelzneri, fundamentalmente, en la férmula

dentaria y en la posicién del tubo cloacal.

WirsCHi (1933) y Davis (1936) expresan que el érgano de Bidder est mejor desarrollado

en los juveniles que en los adultos de algunas especies de Bufo. ECHEVERRIA (1990) observa la

presencia del érgano de Bidder en las larvas, en los juveniles de ambos sexos y en los machos

adultos de Bufo arenarum.

En la diagnosis del género Melanophryniscus, GALLARDO (1961a-b) no menciona la

presencia del érgano de Bidder; sin embargo, éste es incluido por MCDIARMID (1971: 41; 1972:

20) y hace referencia a la “presencia del érgano de Bidder y del peritoneo testicular negro o no

pigmentado”. DUELLMAN & TRUEB (1985) expresan que todas las larvas de los bufénidos

presentan un érgano de Bidder, y que la presencia de éste es ünica entre ellos, aunque en

algunos adultos de Dendrophryniscus puede faltar. WAKE (1980) señala que el érgano de

Bidder estä presente en los embriones de las especies de Nectophrynoïdes pero que no persiste

en los adultos, y sugiere una tendencia evolutiva hacia la reorganizaciôn y pérdida secundaria

del érgano de Bidder en esta linea de bufénidos.

RoEssLer et al. (1990) consideran al érgano de Bidder como una extensién del ovario,

afirmaciôn con la cual no coincido, sobre la base de las observaciones efectuadas en Bufo

arenarum (ECHEVERRIA, 1990) y en otras especies del género Bufo (KING, 1908; TANIMURA &

IwasaWA, 1986), puesto que el 6rgano de Bidder se desarrolla en la larva antes de que la

génada se organice.

En el caso de M. stelzneri he comprobado la ausencia del érgano de Bidder tanto en las

larvas como en los juveniles y adultos de los ejemplares del Trapiche en las Sierras de San Luis

y en los adultos de la Sierra de La Ventana. Por lo tanto, las afirmaciones de GRiFriTHs (1959;

fide McDiarMip, 1971) con respecto a la presencia del érgano de Bidder en Melanophryniscus

stelzneri quedarian descartadas.

Con referencia a los 6rganos internos, cabe destacar que la pigmentaciôn oscura del

higado, caracteristica de los adultos, se hallé en las larvas de estadios en avanzada metamor-

fosis (estadios 45 y 46). La génada permanece indiferenciada por lo menos hasta que la

metamorfosis se completa.

La coloracién del peritoneo que cubre el testiculo varia desde el color blanco al negro o

presenta ambos colores. Esta variaciôn puede depender de la edad y del estado de distensiôn

del peritoneo debido al ciclo del testiculo.

M. stelzneri muestra caracteristicas en la puesta y en el desarrollo que merecen ser

destacadas: los huevos se organizan en racimos, en vez de disponerse en ristras como en Bufo,

la cantidad de oocitos que constituyen una puesta es relativamente menor que en otros

bufénidos, los oocitos ovulables y los huevos presentan pigmentaciôn polarizada, la puesta es

“intermitente” y cada pareja requiere varios dias para completarla. Estas caracteristicas,

sumadas a la falta de érgano de Bidder, y a hallazgos como la presencia de alcaloides en la piel

de M. stelzneri (GARRAFFO et al., 1993) y la tendencia en los adultos a alimentarse bâsica-

mente de formicidos (FILIPELLO & CRESPO, 1994), me inducen a formular un interrogante:

podrian ser éstas, suficientes evidencias para modificar la afinidad conocida entre los buféni-

dos (MCDiARMiD, 1971; CANNATELLA, 1986) o con ellos ?

Source : MNHN, Paris

ECHEVERRIA 169

CONCLUSIONES

Las hembras de M. stelzneri pueden ser inducidas a la ovulaciôn utilizando macerado de

hipôfisis de macho adulto de Bufo arenarum.

Los huevos presentan una pigmentacién polarizada, castaña en el hemisferio animal y

blanco amarillenta en el hemisferio vegetativo. El diâmetro medio de los huevos ovulables

varia de 1.400 a 1.600 um. La eclosiôn se realiza en el estadio 21 a 23 + 1°C, tardando

aproximadamente 50 horas desde la fecundacién.

Los embriones y las larvas son de color castaño.

El desarrollo hasta la metamorfosis se Ileva a cabo, en las condiciones de laboratorio, en

4 semanas.

No se registré la presencia del érgano de Bidder en ningün estadio larval ni en los

ejemplares postmetamérficos, juveniles, preadultos o adultos. De acuerdo con este estudio es

posible concluir que la afinidad de M. stelzneri con los bufénidos podria quedar reducida.

RESUMEN

En este estudio se describe el desarrollo ontogenético de Melanophryniscus stelzneri,

desde la fecundacién in-vitro hasta la metamorfosis, con énfasis en el desarrollo larval. El

diâmetro de los huevos sin fecundar varia desde 1400 a 1600 um, se organizan en racimos y

presentan una coloracién polarizada. El hemisferio animal es castaño y el vegetativo es ocre

o© blanco amarillento. La eclosiôn se realiza en el estadio 21 (GosneR, 1960) a 23°C. Los

embriones y las larvas exotrôficas presentan un color castaño. No se registré la presencia del

érgano de Bidder en ningün estadio larval ni en los ejemplares postmetamérficos, juveniles,

preadultos o adultos.

AGRADECIMIENTOS

A la Lic. Maria F. Roca y al Sr. Gabriel Rosa por la asistencia técnica.

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©ISSCA 1998

Source : MNHN, Paris

Alytes, 1998, 15 (4): 171-175. 171

Tolerance of high electrolutic

and non-electrolytic osmolarities

in Bufo arenarum

premetamorphic tadpoles

under organism density stress

Lucrecia FERRARI

Applied Ecophysiology Program, Basic Sciences Department, National University of Lujan,

CC 221, (6700) Lujän, Buenos Aires, Argentina (prodea @ mail.unlu.edu.ar),

and Scientific Investigations Commission of Buenos Aires Province, CIC, Argentina

The mean (+ standard error) lethal osmolarity was determined in

electrolyte (NaC!) and non-electrolyte (mannitol) media for premetamorphic

Bufo arenarum tadpoles. The effect of organism density stress over the

survival capacity within high osmolarity media was evaluated. Acute toxicity

tests were performed in accordance with the Standard Methods guidelines

at two densities: high (1 tadpole per 4 ml of solution) and standard

{1 tadpole per 20 ml of solution). Experimental solutions were obtained by

adding NaCI and D-mannitol to distilled water. The osmolarity range ran

from 141 to 271 mOsm. Control medium was artificial fresh water at

5 mOsm. There are no differences in survival between an electrolytic and a

non-electrolytic medium and survival is not affected by high organism

density. À mean lethal osmolarity was found at 279.0 + 9.6 and 220.5 +

2.0 mOsm for 48 and 144 h respectively.

The conditions under which the first life stages take place are of vital importance for the

ecological success of a species. In general the aquatic environment under which most

amphibians spend their larval stages is changing both in salinity, osmotic pressure, oxygen

content and organism density, among other stress factors. The way in which some of these,

and particularly organism density, may affect the growth and metamorphosis of Bufo

arenarum tadpoles, has been widely studied (MURRAY, 1990; MIRANDA & PISANO, 1993;

KHER, 1994). However, none of these works has taken into account the effect of osmotic stress

combined with an increase in organism density.

Anuran tadpoles have registered tolerance at higher osmolarities than those of their

natural medium. Except for a few species, tadpoles died when the internal medium became

hyposmotic to the surroundings. Chemical composition of incubation media is not indifferent

to tadpoles; because of their incapability to increase their plasmatic osmolarity by organic

osmolytes synthesis, the compensatory response in hyperosmotic media appears to be almost

entirely due to plasmatic NaCI (BALINSKY, 1981). If the incubation medium is a non-

electrolytic solution, hydric regulation is not possible and they react as limited osmocon-

Source : MNHN, Paris

172 ALYTES 15 (4)

APW (1/4 —1/20)

204 mOsm (1/4)

204 mOsm (1/20)

247 mOsm (1/4)

247 mOsm (1/20)

271 mOsm (1/4)

271 mOsm (1/20)

+05»-DmD4

Survivorship (%)

mannitol

Time (Hs)

Fig. 1. - Survivorship as a function of time for Bufo arenarum premetamorphic tadpoles (stage 26)

exposed to different osmolarities of mannitol at 20° C and two conditions of density. 1/20:

1 tadpole/20 ml solution; 1/4: 1 tadpole/4ml solution.

formers (KATZ, 1987). In the specific case of Bufo arenarum tadpoles, exposed to osmotic

stress, the existence of a dual behavior has been demonstrated, in which both highly saline

environments are tolerated and survival in aionic media does not lead to important changes

(FERRARI, 1995).

The aim of this work was to assess the survival response of Bufo arenarum tadpoles

submitted to electrolytic and non-electrolytic osmolarity stress when they were exposed to

organism density stress.

Semistatic assays were conducted following procedures of the American Public Health

Association (ANONYMOUS, 1992). Tadpoles obtained by “in vitro” fertilization were used at

the first larval stage (GOsNER, 1960). AIl tests were conducted with animals acclimatized 48 h

before the beginning of the experiment at constant temperature (20°C) and photoperiod (12

L:12 D), and which remained under the same conditions throughout the experiments. All tests

were conducted in duplicate at the rate of 1 larva/20 ml (1 g organism/l; ANONYMOUS, 1992),

and, to test the effect of density stress, 1 larva/4 ml (5 g organism/l).

The following osmolarities (in mOsm) were tested for electrolytic solutions (NaCI) as

well as for non-electrolytic solutions (mannitol): 141, 204, 247 y 271. The control of artificial

pond water (APW) of 5 mOsm was run (ALVARADO & JOHNSON, 1966). Solutions were

renewed daily. Tadpoles were examined at 24 h intervals during 144 h. Those exhibiting

Source : MNHN, Paris

FERRARI 173

APW (1/4)

APW (1/20)

204 mOsm (1/4)

204 mOsm (1/20)

247 mOsm (1/4)

247 mOsm (1/20)

271 mOsm (1/4)

271 mOsm (1/20)

+0bDHOD44

Survivorship (%)

(NacI)

Time (Hs)

Fig. 2. - Survivorship as a function of time for Bufo arenarum premetamorphic tadpoles (stage 26)

exposed to different osmolarities of NaCl at 20°C and two conditions of density. 1/20: 1 tadpole/20

ml solution: 1/20: 1 tadpole/4ml solution.

no heartbeat or which did not respond to gentle prodding were considered dead and were

removed from assay containers.

Mean lethal osmolarity (LO 50) was calculated by Probit analysis after adjusting for

mortality among tadpoles in the control treatment with the Abbott’s correction. Ninety-fice

percent confidence limits for each estimate were calculated by Fieller’s theorem (FINNEY,

1971).

Both in the NaCI as in the mannitol tests, in the two tested densities (1 tadpole/20 ml and

1 tadpole/4 ml) the mortality rate in controls (APW 5 mOsm) was always less than 5 %. Figure

1 shows the survival curves as a function of time for each of the non-electrolytic osmolarities

(mannitol) assayed at both densities. The survival curves are similar at both densities. At 24

hours of exposure, mortality was less than 10 % in all the solutions, as of that moment

mortality increased with osmolarity and time. At 271 mOsm, the fall in survival was very

pronounced between 24 and 48 hours and then declined remarkably. At 247 mOsm, survival

diminished continuously, with a slope close to 1, while at 204 mOsm mortality did not reach

30 % after five days of the test.

Figure 2 shows the survival curves as a function of time for each of the NaCI solutions

assayed at both densities. In this case also, the effect upon survival began to appear 48 hours

after exposure, and after this time mortality increased continuously with time, but

Source : MNHN, Paris

174 ALYTES 15 (4)

Tab. 1. - Mean lethal osmolarity (LO 50) of mannitol and NaCI in premetamorphic Bufo arenarum

tadpoles under two conditions of density (1 tadpole/4 ml and 1 tadpole/20 ml). Degrees of

freedom: 3. r = 40 tadpoles/solution.

À 252.2-278.8

1 tadpole 2 236.7-256.8

/4 ml . 220.4-237.3

208.8-233.4

À 262.1-274.4

1 tadpole É 234.5-247.5

/20 ml . 223.4-236.4

215.0-234.9

k 272.6-297.8

1 tadpole 243.9-255.2

/4 ml # 230.3-240.2

209.5-220.2

À 285.2-364.1

1 tadpole ï 265.6-289.8

/20 ml k 256.0-273.8

211.1-228.5

differently to that observed with mannitol. The survival curve gradient between the two

densities was different for 247 and 271 mOsm. At the end of the bioassay, however, values

found were very similar for both densities. At 204 mOsm, the final mortality rate was around

30 %.

Table 1 shows the results of Probit analysis for NaCI and mannitol at the two assayed

densities. Since the mean lethal osmolarity values (LO 50) for mannitol and NaCI show

overlapping confidence limits for all the times of the assay and for both densities, we can

conclude that, under these experimental conditions, the chemical composition and the high

organism density have not effect on the survival response of Bufo arenarum tadpoles. This

allows to calculate an LO 50 (mean value + standard error) for 48 and 144 hours of 279.0 +

9.6 and 220.5 + 2.0 mOsm respectively. These values are higher than those recorded for

anuran tadpole plasma (DEGANI & NEVO, 1986). PADHYE & GHATE (1992) determined a

mean lethal concentration (LC 50) of NaCI and KCI for different stages of embryos and

tadpoles of Microhyla ornata. They reported a LC 96 of NaCI of 0.69 %; in hind limb

tadpoles stage, a value very close to the one reported here. The results obtained show that Bufo

arenarum premetamorphic tadpoles present a similar response as concerns survival in an

environment highly osmotic which is independent of the electrolytes. The low levels of (Na +

K)-ATPase detected in anuran tadpoles (KAWADA et al., 1969) suggest that Na exchange with

the medium is only passive (WARBURG & ROSENBERG, 1990); under the assay conditions, Bufo

arenarum tadpoles are probably able to compensate the osmotic gradient by keeping their

Source : MNHN, Paris

FERRARI 175

plasma slightly hyperosmotic with reference to the incubation medium through a relative

increase of the NaCI (SHOEMAKER, 1992; BALINSKY, 1981). The values of LO 50 found in this

study could indicate the possible limits of such a compensation.

ACKNOWLEDGMENTS

The results reported here were part of the author’s Doctoral Dissertation carried out under the

guidance of Dr. A. SALIBIAN at the University of Buenos Aires. The technical cooperation of Roberto

YosHIHARA and Javier KATz is acknowledged. This work received financial support from the CONICET

and from the National University of Lujan.

LITERATURE CITED

ANONYMOUS, 1992. — Standard methods for examination of water and wastewater. 18th ed. Washington,

D.C., APHA-AWWA, WPCF: i-xxxiv + 8.1-8.83.

ALVARADO, R. J. & JOHNSON, S. R., 1966. - The effects of neurohypophysial hormones on water and

sodium balance in larval and adult bullfrog (Rana catesbeiana). Comp. Biochem. Physiol., 18:

549-561.

BALINSKY, J. B., 1981. - Adaptation of nitrogen metabolism to hyperosmotic environment in amphibian.

J exp. Zool., 215: 335-350.

DeGani,G. & NEVO, E., 1986. - Osmotic stress and osmoregulation of tadpoles and juveniles of Pelobates

syriacus. Comp. Biochem. Physiol., 83 (A): 365-370.

FERRARI, L., 1995. — Equilibrio hidromineral de larvas de Bufo arenarum: respuestas compensatorias al

“stress” osmôtico y al cadmio. Doctoral Dissertation, Facultad de Ciencias Exactas y Naturales,

Universidad de Buenos Aires: 1-162.

FINNEY, D. J., 1971. — Probit analysis. Cambridge University Press: 1-333.

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

identification. Herpetologica, 16: 183-190.

KATZ, U., 1987. - The effect of salt acclimation on the uptake and osmotic permeability of the skin of the

toad (Bufo viridis, L.). J. Physiol., 82: 183-187.

KAwWADA, U., TAYLOR, R. E., Jr. & BARKER, S. B., 1969. - Measurement of Na-K-ATPase in the separated

epidermis of Rana catesbeiana frogs and tadpoles. Comp. Biochem. Physiol., 20: 965-975.

KHEr, A. I., 1994. — Density-dependent response in tadpoles of Bufo arenarum (Anura, Bufonidae).

Physis, (B), 49: 31-36.

MIRANDA, L. A. & PIsANO, A., 1993. — Efecto de la densidad poblacional en larvas de Bufo arenarum

producido a través de señales visuales. A/ytes, 11: 64-76.

MurrAY, D. L., 1990. - The effects of food and density on growth and metamorphosis in larval wood

frogs (Rana sylvatica) from central Labrador. Can. J. Zool., 68: 1221-1226.

PADHYE, À. D. & GHATE, H. V., 1992. - Sodium chloride and potassium chloride tolerance of different

stage of the frog, Microhyla ornata. Herp. J,, 2: 18-23.

SHoEMAKER, V. H., 1992. - Exchange of water ions and respiratory gases in terrestrial amphibians. In: M.

E. FepEr & W. W. BURGGREN (eds.), Environmental physiology of the amphibians, Chicago, Univ.

Chicago Press: 125-150.

WarBURG, M. R. & M. ROSENBERG, 1990. — Ion and water balance and their endocrine control in the

aquatic amphibians Zn: W. HANKE (ed.), Biology and physiology of amphibians, Stuttgart & New

York, Gustav Fisher Verlag: 385-403.

Corresponding editors: Ulrich SinscH & Alain Dugois.

Source : MNHN, Paris

Alytes, 1998, 15 (4): 176-204. Book review

Mapping European amphibians and reptiles:

collective inquiry and scientific methodology

Alain DuBois

Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

Jean-Pierre GAsC, Antonia CABELA, Jelka CRNOBRNJA-ISAILOVIC, Dag DOLMEN, Kurt GROSSENBACHER,

Patrick HAFFNER, Jean LESCURE, Harald MARTENS, Juan Pablo MARTINEZ-RICA, Hervé MAURIN,

Maria Elisa OLIVEIRA, Theodora S. SOFIANIDOU, Michael VerrH & Annie ZUIDERWUK (editors). —

Atlas of amphibians and reptiles in Europe. Paris, Societas Europaea Herpetologica & Muséum

National d'Histoire Naturelle (IEGB/SPN), 1997: 1-496.

ABBREVIATIONS

AFIE: Association Française des Ingénieurs Ecologues.

ASIH: American Society of Ichthyologists and Herpetologists.

CITES: Convention on International Trade in Endangered Species of Wild Fauna and Flora.

IEGB: Institut d’Ecologie et de Gestion de la Biodiversité.

IUCN: International Union for the Conservation of Nature.

L-E: Rana lessonae-esculenta population system.

MNAN: Muséum National d'Histoire Naturelle, Paris, France.

NMW: Naturhistorisches Museum, Wien, Austria.

P-G: Rana perezi-grafi population system.

R-E: Rana ridibunda-esculenta population system.

SEH: Societas Europaea Herpetologica.

SFF: Secrétariat de la Faune et de la Flore.

SHF: Société Herpétologique de France.

SPN: Service du Patrimoine Naturel.

SSAR: Society for the Study of Amphibians and Reptiles.

UTM: Universal Transverse of Mercator mapping system.

INTRODUCTION

In 1983, shortly after its founding, the Societas Europaea Herpetologica (SEH) established a

Mapping Committee, which was entrusted with the charge of preparing a distribution atlas of amphibi-

ans and reptiles in Europe. Under the leadership of Jean-Pierre GasC (Paris), and with the technical

support of a service of the Paris Museum (MNHN) first known as “Secrétariat de la Faune et de la Flore”

(SFF) and renamed in 1995 “Service du Patrimoine Naturel” (SPN), this Committee worked for 15 years

Source : MNHN, Paris

Dugois 177

and produced the Atlas of amphibians and reptiles in Europe, which was published in July 1997. This

500-page volume is the result of a truly international work which involved several hundred persons all

over Europe for the collection of original field data, and about a hundred authors for the writing of the

texts devoted to the species. Altogether, this Atlas is based on 85,067 “species/square/period” data

concerning 3940/4362 (i.e. 90 %) of all the 50 X 50 km UTM squares covering Europe (within the limits

defined by MERTENS & WERMUTH, 1960). Of these data, 41,704 (i.e. 49 %) concern the 62 amphibian

species and 43,363 (i.e. 51 %) the 123 reptile species recognized as valid in this work.

The first part of the volume contains a general presentation of the methodology used to prepare this

Atlas, and overviews of the European climate and vegetation, of the paleogeography of the European

herpetofauna, and of problems posed by the conservation of this fauna. This is followed by analytical

data concerning the species: each species account consists of a map of reported occurrences (except for

five marine turtle species), and of basic information, comments and bibliographic references. Although

each account is signed by one or several author(s), it is ciear that only the written part of the account is

to be credited to the latter, while the maps are the result of the collective inquiry (actually, each map bears

a mention of copyright by “MNHN/SPN & SEH”), and should therefore be quoted as being by Gasc et

al. (1997). These accounts are followed by three appendices, i.e. updatings of the lists of European

amphibians and reptiles, and a table presenting the official conservation statuses and levels of threats of

European herpetological species. Finally, a bibliography of about 2500 references and an index to taxa are

provided.

The book, of format 21 X 29.7 cm, is soft-covered, which may be appropriate for a volume to be read

once or twice, but less so for a book of frequent use. It is well printed, with nice maps in white, grey and

blue. Printing mistakes are rather numerous, and suggest that reading of the final proofs has been too

quick (e.g., just in the introductory chapters and the appendices: p. 4, “french”; p. 6, “Europran”: p. 11,

13: “Oural”; p. 11, “In particular, was necessary”; p. 15, “data was”; p. 16, “many various”; p. 21, italics:

p. 29-30, several misprints; p. 31, one paragraph repeated, “Portugese”; p. 31-32: “systematic”, “speciesi”;

P. 405, “collectind”, “beguen”; p. 406, “hybridisation”; p. 407, “to determinate”; p. 412, “occuring”).

Unfortunately, the book was edited and printed in France, and the editors did not properly care for the

way words should be hyphenated at the right margin of lines, so that they were so according to the French

rules, not the English ones (see e.g., WooLr, 1974; SUMMERS, 1995; PROCTER, 1995): thus, in English, the

proper division is “men-tioned”, not “mentio-ned” (p. 8), “standard-ised”, not “standar-dised” (p. 9),

“chal-lenge”, not “challenge” (p. 9), “famil-iar”, not “fami-liar” (p. 9), “pro-duce”, not “produ-ce”

(p. 10), etc. Black and white drawings of some species are provided, but on p. 4 of the book the authors

of part of them only are acknowledged: e.g., the nice drawing of Discoglossus pictus of p. 494 is not

credited to its proper author, namely Jean-Jacques MoRÈRE (this authorship had already been ignored in

the original publication of this drawing, on the cover of Bull. Soc. herp. France, 5, January 1978).

Interestingly, according to this Atlas, there is a significant difference between the distribution of

species richness of the two studied zoological groups all over Europe: while the combined map for all

amphibian observations (p. 34) clearly points to a larger species diversity in central Europe, the combined

map for all reptiles (p. 160) not less clearly shows a higher diversity in southern Europe, mostly in the

Mediterranean region. Unfortunately, this finding is not discussed at all, in particular in the light of the

following question: does this difference reflect a genuine biological fact, which would then call for a

scientific explanation (climatic-ecological, historical, or both), or does it simply reflect a different

distribution of observers of both groups all over Europe?

The Mapping Committee of SEH is to be commended on having been able to carry this collective

undertaking to its term. Altogether, the amount of work which has been necessary for the production of

this book is impressive, as are the wide geographical and political scope of the inquiry, covering about 40

different countries from the Atlantic to the Ural and to the Caucasus, and the high number of

collaborators involved in it. No doubt this book will become a major reference for a number of European

governmental and official bodies, who need basic documentation about the distribution and conservation

status of native animal species in order to be able to take administrative and legal decisions concerning

their management, collecting, transport, commerce and protection. The genuine interest of such official

bodies for this kind of works is emphasized by the fact that this Arlas was largely supported financially by

the French Ministry of the Environment, who had already funded the production and publication of the

two editions of the French distribution atlas of amphibians and reptiles (CASTANET, 1978; CASTANET &

GUYÉTANT, 1990).

Source : MNHN, Paris

178 ALYTES 15 (4)

But the potential interest and impact of such a work is much wider. In the Preface of the European

Atlas (p. 9), Wolfgang BÔHME rightfully writes: “The careful documentation of distribution data is the

most important prerequisite for evaluating the situation of animal species in a given geographical frame.

This frame (...) provides invaluable zo0-geographical information, from both historical and ecological

points of view, on the taxon concerned. This helps us understand the history, and estimate the future, of

animal populations.” In other words, reliable chorological data are answers to “what” questions (see

Mayr, 1997) that provide information irreplaceable for answering the “how” and “why” questions that

phylogenetic or ecological research ask, and for being able to properly deal with the threats that many

European herpetological species are currently facing. According to the geographical area covered and to

the scale chosen, distribution maps can provide different kinds of information. On the scale of a region,

and especially if the latter represents a significant geographical unit for the organisms studied, such maps

can help to better understand the ecological requirements of species, phenomena of competition,

altitudinal limits and some conservation problems. On a national scale, maps can contribute to deter-

mining the responsibilities of states regarding their natural heritage. Finally, on a continental scale, maps

can provide an interesting light on the biogeographies of species, or even of genera or families. Needless

to say, in all these cases, to be able to play correctly their role, distribution maps must be produced with all

the care and rigour usually required for scientific works. Are these conditions met with in this European

Atlas? 1 will consider this question under several points of view. As this review is written for readers of a

batrachological journal, I will concentrate here mostly on examples taken in the amphibians, but most of

the statements below are also valid for the reptile sections of the volume.

TAXONOMY AND NOMENCLATURE

In the short anonymous text entitled “How to use the Atlas” (p. 31-32), one can read: “The

nomenclature used in the atlas is the one prevailing at the time the texts were written. Where there is no

consensus, the author of the text is responsible for choosing the nomenclature used. The scientific name

may designate a species complex, according to current knowledge (1997). (p. 31). However, in the

methodological introduction (p. 11-16), H. MAURIN, P. HAFFNER, H. DA Cosra & J.-F. BRULARD provide

a slightly different information, since they state that the 185 species recognized as valid in the Atlas were

so “according to the nomenclature as it stood in 1995” (p. 11), and that subsequent changes could not be

taken into account: “because time was very short, it was not possible to process the newly described or

newly found species within the standard procedure. There was no time to collect the data necessary for the

distribution maps or to find authors willing to write the accompanying species’ reviews.” (p. 13). Some

comments regarding these changes were therefore added in some of the species accounts and in the special

appendices “updating the lists of species”, by A. OHLER and I. INEICH, that appear at the end of the book

(p. 404-407).

Even if the imprecise “1995” Iandmark is to be understood as “1st January 1995”, it was not always

respected in the book, and still more so if it is “31 December 1995”. Some species described or recognized

as valid well before the beginning of 1995 were not duly considered in the body of the Adlas, and are only

listed in the appendices (p. 404-407). Among amphibians, the most striking case is Rana pyrenaica, whose

original description (SERRA-CoBo, 1993) was published in March 1993, and included a detailed distribu-

tion map of the known localities of the species, which could well have been integrated in the Atlas after

transcription into UTM squares. Other amphibian examples include Triturus carnifex, Triturus dobrogi-

cus and Triturus karelinii (BUCCI-INNOCENTI et al., 1983; MACGREGOR et al., 1990), Rana cerigensis and

Rana cretensis (B#ERLI et al., 1994), and a few other taxa (Salamandra corsica, Bufo verrucosissimus, Hyla

sarda, Rana bergeri), which have recently been considered valid species by some authors, although

published evidence for such taxonomic decisions is lacking (see DuBois & OHLER, 1995a and OHLER’s

appendix to the Arlas). Similar problems exist in reptiles, not all of which were mentioned in the appendix:

to give just one example, the oriental populations long referred to Hemidactylus turcicus are now referred

to other taxa (see e.g. DELAUGERRE & CHEYLAN, 1992: 57), which is ignored in the contribution by U.

GRUBER (p. 211).

The /nternational Code of Zoological Nomenclature (ANONYmous, 1985; quoted below as “the

Code”) was not always understood or respected by contributors to the book, so that the latter contains a

Source : MNHN, Paris

Dugois 179

number of nomenclatural mistakes. In amphibians, several of them concern the green frogs of the

subgenus Rana (Pelophylax), whose nomenclature was reviewed by Duois & OHLER (1995a-b) and

CrocHer et al. (1995), a fact which is only briefly mentioned in the appendix of the Atlas (p. 404-405) but

should rather have been taken into account in the body of the book itself: thus, the species reported in the

book as Rana balcanica, if valid (see BEERLI, 1994; BEERLI et al., 1996), should be known as Rana

kurtmuelleri. The book does not include any discussion or reference concerning the nomenclatural

problems which have recently been raised regarding the genus of Plethodontidae successively known as

Geotriton, Hydromantes and Speleomantes. In several works, B. LANZA argued that the European and

American salamanders of this group should be placed in two distinct genera (LANZA & VANNI, 1981;

Lanza et al., 1996). In these works as well as in the Ar/as, this author decided to use the name

Hydromantes for the European species of this genus, apparently because he considered that the Interna-

tional Commission on Zoological Nomenclature would “almost certainly” take the decision which had

his preference (see LaANZA et al., 1996: 17, 21). But this prediction proved wrong, as the Commission

decided that the European species, if considered generically distinct, should bear the name Speleomantes

(Anonymous, 1997). Possibly the parts of the Arlas concerning this genus were written before publication

of the Commission’s Opinion in March 1997, but then the text should have been corrected before

publication, in order to follow this decision which has force of law for all zoologists worldwide,

irrespective of their personal tastes. Even before the Commission had voted on this case, the authors of

the Atlas should have used the name Spe/eomantes for the European species, since, as had been shown by

SaLvinio (1995) and Dusois (1995b), no general “current usage” could be claimed to exist in this case, as

two parallel usages were in force after the publication of Dugois’s (1984a) paper: while most North

American authors continued to use the name Hydromantes for salamanders of this group, a clear

tendency developed in Europe, including in several “official lists”, to replace it by the name Speleomantes.

This mistake, and even worse, the fact that the Ar/as does not discuss this case at all, is unfortunate, as this

volume will become an important international reference and will contribute to the spreading of an

incorrect nomenclature and to the continuation of a regrettable situation of nomenclatural confusion. An

example of this confusion is to be found in the Atlas itself: in J. P. MARTINEZ RiCA’s contribution on

climate and vegetation, both names Hydromantes and Speleomantes are used as valid names in different

paragraphs of p. 21!

Some mistakes are also to be found in the Aflas regarding the valid spelling of scientific names, the

nomenclatural availability of scientific names, the author’s names and dates of nominal taxa, or the

inclusion of these names and dates in parentheses. The name “Chamaeleontidae” is properly written in

pages 6 and 201, but misspelt “Chamaeleonidae” in pages 26-27 (J.-C. RAGE), the name “’Trionychidae”

is misspelt “Tryonichidae” in page 27 (J.-C. RAGE), and the subspecific name “Triturus alpestris bukkien-

sis” is misspelt “hükkiensis” in pages 72 (A. ZUIDERWUK) and 492 (index). The name “ Molge syriacus

Valenciennes, 1877”, listed by L. J. BorkiN in p. 86 among the synonyms of Triturus vittatus, has no status

in nomenclature: it was first published by LATASTE (1877: 365) as a synonym of Triton vittatus, and was

not adopted as a valid name before 1961, so that by virtue of Art. 11.e of the Code it is not an available

name, and has no therefore no type-specimen, contrary to the statement of THIREAU (1986: 74-76).

Several cases of incorrect authorship and date can be pointed out in amphibians: the nominal species

Alytes muletensis was created by SANCHIZ & ADROVER (1979), not by “SANCHIZ & ALCOVER (1977) (no

publication corresponds to this reference), as written by J. P. MARTINEZ RICA in p. 92; the nominal species

Rana balcanica by SCHNEIDER & SINsCH (1992), not by SCHNEIDER, SINSCH & SOFINANIDOU (1993), as

stated by T. S. SOFIANIDOU in p. 130 (see DuBois & OHLER, 19954: 179-180); and the nominal species Rana

dalmatina by FITZINGER in BONAPARTE (1838), not by BONAPARTE (1840), as written, after many others,

by K. GROSSENBACHER in p. 134 (see Dumois, 1984b: 117-118). In reptiles, according to Art. 50 of the

Code, the author of the nominal subspecies Podarcis hispanica cebennensis is FRETEY (1986: 81), not

“GUILLAUME & GENIEZ in FRETEY (1986)”, as stated by C. P. GUILLAUME in p. 278; the mention by

GUILLAUME & GENIEZ (1986) of a specimen figured in FRETEY (1986) as the “holotype” of this subspecies

results in the designation of a lectotype for the latter (Art. 74.b) and in a restriction of the type-locality to

Valros (Hérault). The Code’s principle of coordination requires that the nominative subspecies of

Agkistrodon halys bear the same author and date as the species, which is ignored in the Arlas by L. S.

DaREvskY (p. 378). The authors’s names and dates of Bufo bufo verrucosissimus, Eryx jaculus turcicus,

Eryx miliaris miliaris, Natrix tessellata heinrothi and Macrovipera lebetina obtusa (if the latter genus is

recognized as valid) should be enclosed in parentheses, as these nominal species-group taxa were created

Source : MNHN, Paris

180 ALYTES 15 (4)

in other genera, which is ignored in the accounts dealing with these taxa. Finally the Ar/as contains several

cases of confusion between two terms of the Code having distinct meanings and uses, i.e. “nominotypi-

cal” and “nominal” (see ANONYMOUS, 1985): in amphibians, the latter is used for the former in the

contributions on Triturus helveticus (A. ZUIDERWUK, p. 78), Triturus marmoratus (A. ZUIDERWUK, p. 82)

and Bufo viridis (P. ROTH, p. 122), while the correct term “nominotypical” is used under Triturus

superspecies cristatus (J. W. ARNTZEN & L. BORkIN, p. 76) and Bufo bufo (L. J. Borkin & M. VEITH,

p. 118).

Although for reptiles most recent generic taxonomic changes were duly considered, for amphibians

the generic taxonomy used in the At/as is the “traditional” one, found e.g. in the checklist of MERTENS &

‘WERMUTH (1960). Subgenera are not recognized, not even discussed. However, several taxonomists have

proposed or adopted a subgeneric classification for some genera of European amphibians and reptiles,

including the amphibian genera Hydromantes (see discussion above), Triturus, Alytes, Bombina and

Rana. In two cases at least (genera Triturus and Rana), presentation of the species under their respective

subgenera would have been useful: mixing all European Ranidae in alphabetical order under the generic

name Rana is much less enlightening for the reader than would have been their separate listing under the

subgeneric names Aquarana, Pelophylax and Rana (see DuBois, 1998a).

Several other examples clearly stress the little concern of the authors and editors of the Atlas for

taxonomic and nomenclatural matters. The first one is that of the so-called partial “synonymies”’ which

are provided at the head of each species account, under the heading “Main synonyms”, These are stated

to include “the synonyms most frequently used in literature” (p.31). According to the Code (ANONYMOUS,

1985: 266), a synonym is “each of two or more scientific names of the same rank used to denote the same

taxon”. In this definition, a “scientific name” is to be understood as a Latin name validly published under

the Code to designate a new taxon. Posteriorly to its original publication, such a name is liable to be

modified, in its spelling, combination or onymorph (see SMrr & PEREZ-HIGAREDA, 1986), but this does

not result in the creation of a new name: such modified names are not synonyms of the original name, but

merely different “name-forms” of the latter, which have no independent status in nomenclature. They

should therefore not appear in a synonymy sensu stricto. They may appear in a synonymy and chresonymy

(see SMirH & SMITH, 1973) or chreso-synonymy, either complete or partial, but the difference between the

two should be clearly understood and mentioned (see e.g. Duois, 1997b: 184-185). Complete confusion

exists in the Arlas regarding the status of the partial “synonymies” provided: over 62 amphibian species

recognized as valid in the book, only 27 % of the “synonymies” provided deserve the qualification of

genuine synonymies, while the other 73 % are partial chreso-synonymies (31 %), mere partial synonymies

(34 %) or wrong synonymies (8 %). Here are the details for each of these categories:

(1) Genuine synonymies (total 17). (a) Complete synonymies (including cases where no synonymy is

provided and no synonym is known to exist) (total 11): Chioglossa lusitanica, Salamandra atra, Salaman-

dra lanzai, Altes cisternasü, Pelobates syriacus, Hyla meridionalis, Rana epeirotica, Rana iberica, Rana

latastei, Rana macrocnemis and Rana perezi. (b) Partial synonymies (total 6): Bufo bufo, Bufo viridis, Rana

catesbeiana, Rana dalmatina, Rana ridibunda and Rana temporaria.

(2) Partial chreso-synonymies (total 19): all species of the genera Salamandrella (1), Pleurodeles (1)

and Salamandrina (1); Éuproctus platycephalus, Salamandra salamandra, Triturus boscai, Triturus helve-

ticus, Triturus vittatus, Triturus vulgaris, Alytes obstetricans, Bombina bombina, Discoglossus sardus,

Pelobates cultripes, Pelobates fuscus, Bufo calamita, Hyla arborea, Rana arvalis, Rana k]. esculenta and

Rana lessonae.

(3) Mere partial chresonymies (total 21): all species of the genera Speleomantes (as “Hydromantes”)

(6), Mertensiella (1) and Pelodytes (2); Euproctus asper, Euproctus montanus, Triturus alpestris, Triturus

cristatus, Triturus italicus, Triturus marmoratus, Triturus montandoni, Alytes muletensis, Discoglossus

montalentii, “ Rana balcanica”, Rana italica and Rana shgiperica.

(4) Wrong synonymies (total 5). (a) Names of valid subspecies listed as synonyms (total 3): Bombina

variegata, Discoglossus galganoi and Discoglossus pictus. (b) No synonymy provided, although synonyms

are known to exist (see e.g. MERTENS & WERMUTH, 1960) (total 2): Proteus anguinus and Rana graeca.

The account for Proteus anguinus by J. DURAND starts with the following statement (p. 50): “Main

synonyms: None”. Then, a few lines below, one can read: “Fitzinger (1850) described 7 species of Proteus

(...). These species are today invalidated; nevertheless Mertens & Wermuth (1960) still mention 12

different names. We may consider there is one species and possibly 2 to 3 subspecies (...).” Such statements

Source : MNHN, Paris

Dugois 181

can be understood under the pen of authors of species accounts who are not taxonomists, but it would

clearly have been the responsibility of the editors of the Arlas to care for the quality and homogeneity of

the information. As this has clearly not been done, users of this book should be warned not to rely on

these “synonymies”, but to rather use other works providing serious synonymies (e.g., for European

amphibians: MERTENS & WERMUTH, 1960; Dugois, 1995a; Dugois & OHLER, 1995a-b, 1997b).

A similar warning of caution can be made for another section that appears in all species accounts,

under the heading “Terra typica”. Beside the fact that this designation is not that recognized by the Code,

which uses the formula “type locality”, two major problems appear regarding this section. First, the

type-locality is provided only for the nominal taxon whose name is currently the valid one of the

taxonomic taxon. However, this information has only nomenclatural, not biological, value and interest,

and, if given for the valid name, should also be provided for its synonyms (for more details, see DUBOIS,

1987b: 104-107). Second, the authors of the Aflas have taken for granted so-called “restrictions of

type-localities” which were not accompanied by lectotype or neotype designations, although such

restrictions are clearly invalid under the Code (see Dugois & OHLER, 19954: 146, 1997a: 312-313; MYERS

& BÔHME, 1996: 17-18). As long as no such type designations have been made, such invalid restrictions

may be “provisionally retained”, and, if this proves possible, for the sake of stability it may be justified to

“validate them a posterori” through lectotype or neotype designation (Dugois & OuLer, 19974: 313). But

this is not always possible or desirable and, at any rate, as soon as a valid objective restriction of

type-locality through lectotype or neotype designation has occurred, neither the “original type-locality”

nor subsequent invalid restrictions are in force any more. Thus, the designation by DuBois & OHLER

(1997b: 334) of a figured specimen as lectotype of Rana arborea Linnaeus, 1758 restricted the type-

locality of this nominal species to the region of Zürich (Switzerland), and the original type-locality of

LiNNAEUS (1758: 213) (“sub foliis arborum Europae, Americae”) is not valid any more, contrary to the

statement of A. STUMPEL on p. 124 of the Aflas; similarly, the type-locality of Rana kl. esculenta

Linnaeus, 1758 is Nürnberg (Germany) through the designation by Dugois & OHLER (1995a: 149) of a

figured specimen as lectotype, not through the two successive so-called “restrictions” by MERTENS &

MÜLLER (1928: 19; 1940: 18), still recognized as valid by R. GÜNTHER on p. 138 of the Arlas (despite his

citing Dugois & OHLer, 1995a). According to the Code (Art. 72.h), the type-locality of a nominal

species-group taxon is the “place of capture or collection” of its name-bearing type, not any other locality

where it may possibly have come from, except in the case of unnatural transportation by man: therefore,

the type-locality of Proteus anguinus is the Cerknisko jezero (lake Cerknica) in Slovenia south of

Ljubljana (see Hagic, 1993), the place where had been collected the single specimen (holotype) on which

LAURENTT's diagnosis (1768: 37) and figure (1768: pl. 4 fig. 3) were based, and the subsequent so-called

“emendation” of this locality by FEJERVARY (1926) is invalid, as is the multiple type-locality given for this

species by J. DURAND in p. 50 of the Arlas.

“Common names” in several languages are provided for all species considered valid in the Arlas. In

fact, such names are not “common”, “current” or “vernacular” names at all, as most of them were coined

specially for a recent book (STUMPEL-RIENKS, 1992) and have not yet been significantly used in the

respective countries where these languages are spoken. For the time being, and until they are widely used

in popular literature, they should rather be regarded as proposals, not as “official” names. Then, some

other proposals, some of which (Duois, 1982b-c; MaTz & WEBER, 1983) are anterior to STUMPEL-

RiENKS’s (1992) and some others (DuBois & OHLER, 1995a) cover species not considered in the latter work

and with a different rationale for selection of names, should also have been mentioned in the Arlas.

The last paragraph of taxonomic relevance provided for each species concerns their “European

subspecies”. This paragraph also is quite unsatisfactory for any reader interested in taxonomic and

evolutionary problems. Why were only “European” subspecies mentioned for all species? Europe is a

political, not a natural zoogeographical unit, and mention of data concerning extra-European range and

subspecies of the “European” herpetological species would be very useful in such a book. No homoge-

neity exists in this volume concerning the information provided for the “European” subspecies. The

authors and dates of the subspecific names are given in most cases, but not always: in amphibians, this

information is wanting for Triturus alpestris (A. ZUIDERWUX, p. 72), Triturus vulgaris (S. L. KUZMIN & A.

ZunerwuxK, p. 88), Discoglossus pictus (M. Vera & H. MARTENS, p. 104) and Bufo viridis (P. ROTH,

p. 122). Type-localities and areas of distribution of the different subspecies are given for a few species only.

This is all the more strange as the concept of subspecies, at least as used by modern taxonomists (e.g.,

Mayr & AsHLoCKk, 1991), is equivalent to the older concept of “geographical race”, and is eminently

Source : MNHN, Paris

182 ALYTES 15 (4)

“mappable”: if well defined, the subspecies of a given species have different, allopatric or parapatric,

distribution areas, that can easily be shown on a map. On the maps of the Atlas, it would have been useful

and enlightening to use different symbols to show the occurrence of different subspecies and of possible

intermediate populations or hybrid zones, or to add, e.g. as broken lines, the known or supposed limits of

the subspecies areas, and contact or hybridization zones between them. This simple mapping would have

helped pointing to the existence of taxonomic problems concerning the validity of some of the currently

accepted subspecies: if mapping of subspecies limits appears difficult, it may well be an indication that the

subspecies are poorly defined and need revision. For example, it would be most enlightening to map the

so-called subspecies of Bombina variegata, with their type-localities and supposed ranges (see ARNTZEN

1978), in order to see what comes out concerning the so-called “subspecies” scabra and kolombatovi

which are supposed distinct but whose type-localities are very close. Finally, beside bringing information

about geographic variation, it would have been particularly important to pay more attention to subspecies

in this volume for two major reasons: (1) for conservation problems (see below the Triturus alpestris case);

(2) because many taxa currently regarded as subspecies are likely to be considered species in the future (see

Dusois, 1998a): the existence of maps for the subspecies would then have been readily available to future

authors as a first evaluation of the range of these species.

The importance of taxonomic and nomenclatural problems pointed out above in the “well-known°?

animal group of European amphibians may appear strange to some readers. The fact is that the taxonomy

of this group, like those of reptiles and of many other animal and plant groups on our planet, is still far

from being “finished and stabilized”, and that a lot of work, and of novelties, can still be expected in this

field (for more details, see DuBois, 1998a). At any rate, to be valid and useful, any chorological work must

be based on a reliable and up-to-date taxonomy and nomenclature, and on a good knowledge of the taxa

to be mapped (see Dugois, 1998b). In the absence of a serious, professional, taxonomic basement, any

z00geographical work is bound to encounter other kinds of problems, which will directly affect the

validity of the zoogeographical data themselves, as we will now see.

CHOROLOGICAL DATA

Even a cursory survey of the Atlas immediately shows that the distribution maps presented are of

various quality, accuracy and completeness. Some, especially those of species with a limited distribution,

were apparently prepared on the basis of excellent field, literature and/or collection data and seem quite

reliable. But this is not the case of all maps. I will concentrate here on a few examples taken in Ranidae,

but unfortunately these are not the only ones, and the methodological problems raised by these examples

are important enough to throw a shadow of doubt over the entire book, as a reliable source for

chorological data on the European herpetofauna.

The first example is that of European green frogs of the subgenus Rana ( Pelophylax). For sure, the

evolutionary status and taxonomy of these frogs is a complex one, which has only recently started to be

disentangled (GüNTHER, 1979; GRAF & PoLLS PELAZ, 1989: OGiELska et al., 1995). However, if a

distribution atlas is to be of some genuine scientific help and significance, it is precisely in such difficult

cases! The least that can be expected from such a book in such complex situations is to point out problems

and difficulties, rather than “erasing” them under seamingly accurate maps based on wrong data and

contributing in fact to spread confusion and misunderstandings. It is clear that, for the time being,

identification of live specimens of European green frogs is difficult, if not impossible in the field without

having recourse to bioacoustics or to laboratory techniques such as protein electrophoresis or morpho-

metrics, and use of all these methods requires quite specialized knowledge and experience. For this

reason, distribution data on these species based on written answers to questionnaires should be accepted

only with considerable caution. In most cases, and even when the information came from well-known and

serious observers, the only possible serious use of such data is to regard them as mere evidence of presence

in the surveyed region of “green frogs” of the subgenus Rana ( Pelophylax), without further precision. In

many cases, electrophoretic or morphometric study of specimens has revealed the presence in some

regions of forms or species of green frogs unsuspected in these areas. This problem has become

particularly serious because, especially since the development of deep-freezing food techniques, green

Source : MNHN, Paris

Dugois 183

frogs are more and more used as a source of human food in Europe, which has resulted in wide-scale

commerce and transportation of these frogs, and also of the American species Rana (Aquarana)

catesbeiana: the problems of genetic pollution (Dusois & MorÈRE, 1979, 1980; Dugois, 19834, 1990), and

of faunistic pollution (Dusois, 1983c), that this new commercial development has caused, were analysed

in detail by Dugois (1976, 1977, 1983c, 1985a). Without reference to the previous works, these problems

were recently “rediscovered” by REINERT (1991) and ARANO et al. (1995). This large-scale displacement of

frogs contributes to obscure the patterns of distribution of green frogs in Europe and explains that, for

zoologists interested in the intriguing problems posed by the evolution of this exceptional complex of

species and kleptons, it would seem imperative to oppose all projects of commercial exploitation of the

populations of these frogs, which do not constitute a traditional or first-priority food for most Europeans.

Unfortunately this attitude has not been shared by all individuals and associations concerned with

herpetological matters (see Dugois, 1983c, 1985a), so that these problems will undoubtedly take a

growing importance in the forthcoming years, and that the “original” distribution of green frogs in

Europe, before their transportation by humans, will probably never be possible to trace, at least in all

details.

Authors of a scientific distribution atlas must be aware of these problems (1) of identification of

specimens and (2) of genetic and faunistic pollution, and should at least mention them in the discussions

of such a work. Unfortunately, except in one case (see below), such discussions are badly wanting in the

European Atlas. Beautiful distribution maps of green frogs in Europe are provided, but without the

necessary information or warning concerning these problems. Let us consider some of these maps.

First, let us compare the maps provided for Rana kl. esculenta (p. 138), Rana lessonae (p. 148) and

Rana ridibunda (p. 154). In his comments of the former map, R. GÜNTHER (p. 139) rightfully writes: “R.

kl. esculenta’s range is almost identical with that of R. lessonae.” However, comparison of the maps

provided for these two species shows that they display considerable differences: the first species is reported

from 1172 squares and the second from 767; furthermore, the reverse situation also exists (R. lessonae but

not R. kl. esculenta being reported for some squares), so that on the whole the overlap between the two

maps is less than 767 squares. Strictly taken, these data would suggest that in more than 35 % of the

squares where it is present (405/1172), the klepton R. kl. esculenta occurs there without R. lessonae. In

part of these, R. ridibunda is also reported, but in most of them R kl. esculenta alone is shown, so that in

these squares, according to the Arlas, green frogs appear to be represented only by pure populations of R.

kl. esculenta. Such populations are indeed known to exist, especially in central and northern Europe, but

it is unlikely that they occur in all the squares where R. kl. esculenta alone is reported in the Arlas.

Particularly striking in this respect are all the spots on the map of R. kl. esculenta in southern and

south-western France, as is aptly underlined by R. GÜNTHER in his accompanying text (p. 139). Looking

at the map of p. 138 gives the misleading impression that the distribution of R. kl. esculenta virtually

covers all the territory of France, and may be limited in the South-West by the chain of the Pyrenees.

Actually, all published evidence available for the time being suggests that both R lessonae and R. kl.

esculenta, at least as natural populations, are absent in south-western France, and are replaced there by

the “P-G system”, ie. mixed populations of Rana perezi and Rana KI. grafi (see CROCHET et al., 1995). In

this respect, the Atlas seems much more reliable on the Spanish than on the French side of the Pyrenees.

This may be due to different methodologies followed by the national coordinators of the inventory in

these two neighbouring countries. The data for France closely resemble those published in the French

Atlas (CASTANET & GUYÉTANT, 1990: 86), which were clearly unsatisfying as the different kinds of green

frogs had not been distinguished by most observers. In this respect, R. GÜNTHER is fully correct when he

writes in the Arlas (p. 139): “records of occurrences by inexperienced people in questionnaire actions are

doubtful, because water frogs are difficult to identify”. However, this comment has a much wider reach

than it seems from this modest sentence, as green frogs are not the only species difficult to identify for

“inexperienced people”, and data in the Aslas concerning some countries (such as France) came mostly

from “questionnaire actions”.

The maps provided in the Arlas for R. kl. esculenta, R. lessonae and R. ridibunda are therefore most

unreliable and cannot be used for scientific analysis. In such a case, rather than mapping separately these

three species, and even without going into the details of the nine population types that can be recognized

in these frogs (see e.g. RyBACKI, 1995: 346), it would have been useful to present at least five maps: two of

“L-E system” and “R-E system” populations (UZZELL & BERGER, 1975), and three of pure R. lessonae,

R ridibunda and R. ki. esculenta populations. Of course, such maps could not be prepared by “inexpe-

Source : MNHN, Paris

184 ALYTES 15 (4)

rienced people”, and could be so only on the basis of laboratory work or of bioacoustic survey by

experienced researchers: the total number of spots that could currently be obtained this way would be

much lower than that presented in the Arlas, but this would be “better than nothing”. We here touch a

basic methodological question regarding this kind of atlases, which will be considered in more details

below: what is “better than nothing”? Is it a nice but unreliable map covered with hundreds of spots, or

a reliable map with only a few dozen spots based on scientifically reliable data?

Several other green frog maps are open to the same questions. The distinction between R. ridibunda

and R. perezi also requires good experience or laboratory techniques, so that the parts of the maps in the

areas where both species are stated to occur (southern half of France) are also highly doubtful. Strangely,

while the Arlas took a lot of information from questionnaires as granted, it did not include many

data already published by professionals and based on reliable laboratory techniques. Thus, although

they were only recently given Latin scientific names (Dusois & OHLER, 19954; CROCHET et al., 1995),

beside R. kl. esculenta, two other kleptons have been known for a long time to oceur in Europe, and

published data are available about their distribution (GÜNTHER, 1979; GRAF & POLLS PELAZ, 1989;

OGLska et al., 1995): however, in the Atlas, distribution data for one of them (Rana kl. hispanica) are

lumped with those of R. kl. esculenta, and those available for the second one (Rana kl. grafi) are

completely ignored.

The Atlas is supposed to provide information on introductions, and such data are important indeed

to point out the potential problems of genetic and faunistic pollution alluded to above. However,

establishing that a population is of alien origin deserves careful work and information, and the Atlas is

also disappointing in this respect. The introduced populations of R. lessonae in southern England

mentioned on p. 149 in R. GÜNTHER'S text are not shown on the map of p. 148, and the introduced

populations of R. catesbeiana, R. lessonae, R. ridibunda and R. kl. esculenta reported by ARANO et al.

(1995) are ignored in the respective maps of these species, despite the fact that these data are referred to

on p. 153 in the text on R. perezi by M. GaRCiA Paris. On the other hand, no reference or comments are

provided for the statement that some populations from Italy and Denmark are composed of or derived

from introduced specimens of “Rana balcanica”, ï.e. of Rana kurtmuelleri or, if the latter species is not

valid, of Rana ridibunda. Inversely, no reason is given for considering only two of the three spots credited

to the snake species Natrix maura in Corsica as introduced, although all observations of this species in

this island were reported in the same publication (FONS et al., 1991) and are most probably the result of

introductions from Sardinia (DELAUGERRE & CHEYLAN, 1992: 84).

Let us now leave the green for the brown frogs, and look at the map of Rana dalmatina (p. 134). This

map shows a continuous presence of this species all along the Pyrenean chain, except in the French

eastern part of the chain. This information is highly surprising, and would call for an explanation, but K.

GROSSENBACHER, in his accompanying text (p. 135), does not discuss it in detail, just writing that “old

records from Cataluña could not be confirmed in recent years”. However, Dugois (1982c: 62-64) provided

a detailed analysis showing that, although this species is present in the Landes and in the Garonne valley,

and can probably reach the foot of the chain, no serious data are available to ascertain its presence in the

Pyrenean chain itself, at least on its French side. He pointed to several misidentifications by previous

authors and suggested that most, if not all, of the older reports of this species in the chain were based on

specimens of Rana temporaria, particularly of long-legged specimens which he proposed to call provi-

sionally “Gasser’s frog”. On the basis of an extensive survey of 3220 publications (i.e. roughly one third

more than all those cited in the European Alas) dealing with the chorology of the French herpetofauna

(PARENT, 1982), PARENT (1981) proposed a distribution area of R. dalmatina excluding three French

Pyrenean departments (Basses-Pyrénées, Hautes-Pyrénées, Pyrénées Orientales) and including two other

ones (Ariège, Haute-Garonne), but possibly on the basis of extra-Pyrenean populations. À few pieces of

evidence support the idea that the range of the species extends to the first foothills of the chain, at least in

some areas: ZUIDERWUK & VEENSTRA (1984) reported the species from several localities of the Basque

provinces of Alava and Navarra south of the Pyrenean chain proper, and Pierre-André CROCHET

{personal communication) has seen typical eggs, larvae and adults of R. dalmatina in Ariège (Plantaurel

chain). I personally had the opportunity to see only one specimen of “agile-like” frog in the Basque

country, with Annemarie OHLER and Miguel VENCES in 1997 in a locality where a local naturalist had

reported having seen R. dalmatina: this specimen (shown here in fig. 1) is a long-legged Pyrenean R.

temporaria, i.e. a Gasser’s frog (DuUBois, OHLER & VENCES, unpublished data). I know of no other

convincing published data or specimens supporting the presence of R. dalmatina in the French Pyrenees.

Source : MNHN, Paris

Fig. 1. - Specimen (MNHN 1997.4446) of long-legged Rana temporaria (“Gasser’s frog”) from along

Hasquette river north of Hasparren, Pyrénées-Atlantiques, France, 28 October 1997 (photo Pierre-

André CROCHET).

However, and without any discussion, CASTANET & GUYÉTANT (1990) mapped the species as present in

several parts of this chain, even at high altitude (which is quite impossible), and their data seem to have

been uncritically incorporated in the European Atlas. It is likely that most, if not all, of the Pyrenean

spots credited to R. dalmatina in the latter book are based on observations of R. temporaria. Nevertheless,

the state of the art concerning our knowledge of the amphibians of this region (see Dupois, 1982c, 1983a;

SerRA-CoBo, 1993) is not such that the presence of R. dalmatina at low elevations in the Pyrenean chain

can be completely ruled out. This is a typical example of a situation where, if correctly carried out, an

international inquiry could bring interesting new data. But, to be useful in this context, the data received

from questionnaires should be carefully evaluated: we here touch basic methodological questions that will

be discussed in more detail below.

Absence of distinction by some observers between R. temporaria and R. dalmatina raises other

problems. The general distribution areas of both species widely overlap in western and central Europe, as

it clearly appears on the maps of p. 134 and 158 of the Arlas. However, all those who have field experience

know that, at least in western Europe (e.g. in most of France), both species are only rarely found together

in the same localities, even in plain habitats which look superficially quite similar: for example, in the Paris

Source : MNHN, Paris

186 ALYTES 15 (4)

region, R. dalmatina alone is found in the Fontainebleau forest and R. temporaria alone in the Carnelle

forest, and, in forests where both occur (e.g., the Rambouillet or the Compiègne forests), they breed

together in some ponds only, while others only harbour one of the two species (DuBois, MORÈRE, OHLER,

PAYEN & VACHARD, unpublished data). Careful analysis of such facts would allow to know better the

histories and ecological requirements of both species, and to be more efficient in our conservation

strategies. In western France, each species seems to be absent from rather large areas where the other one

is present (Dumois, unpublished data), and a careful mapping of the occurrence of both species would be

most interesting and useful. However, it is clear that, if and when someone wants to undertake such a

study, the latter should be started from the beginning, as the data of the two French and of the European

atlases are not reliable: even if the number of spots based on misenditifications between the two species is

low, there is no way for the reader to know which spots are wrong, as no voucher specimens can be

re-examined. In such areas, the European Atlas will be of little help to solve scientific questions and to

help taking decisions regarding conservation matters.

CONSERVATION PROBLEMS

One of the stated purposes of this Arlas is to serve as a source for information on conservation

problems facing European amphibians and reptiles. A brief introduction to this question by K. CORBETT

is provided at the beginning of the book (p. 29-30), and the third appendix of the volume, by M. E.

OLIVEIRA, P. DASZKIEWICZ & B. GAUVRIT (p. 408-412), presents data on the conservation status and the

level of threat of each species in Europe, under the form of a table giving their coded categories in the

species lists of the Habitat and Species Directive, of the Bern Convention, of the CITES Convention and

of the IUCN Red Lists. Most unfortunately, only the codes of the categories are provided in this table,

without their definitions or descriptions, and even without bibliographic references to such information,

so that this table will be of little help to many users of the Atlas.

By themselves, chorological data on current and past distribution of species can be a precious help

regarding the recent evolution of their population status and a guide for future conservation actions, but

of course, to be so usable, these chorological data must first be reliable. Furthermore, additional problems

must be considered, among which two are of particular importance and will now be discussed: (1) the

need for a good knowledge of the existing literature, particularly concerning old data on distribution and

population status of the studied species; (2) the taxonomic and genetic heterogeneity of species, which

results in the particular significance and importance of threats on some populations.

The first problem will be discussed in the light of the example of the species Pelobates fuscus. The

map presented in the Alas (p. 110) shows a rather “logical” distribution, reminding in many respects

those of other species, such as Bufo viridis: according to this map, P fuscus appears to be a species widely

distributed and apparently common in eastern and central Europe, but whose distribution ends quite

abruptly west of the Alps, of Lorraine and of eastern Benelux. The author of the accompanying text for

this map, A. NôLLERT (p. 111) seems to consider this map to show, not only the current situation of

populations of this species in western Europe, but also its “potential” area of distribution in this region,

since he writes that Alsace and eastern Benelux are the “western distribution limit” of the species. With

such ideas in mind, he is rightfully surprised by the presence in the Arlas’s map of an isolated spot in

central-western France, and he writes about it: “Another isolated (doubtful?) locality is situated in Central

France (Buzançais).” This isolated spot, which was also the only one shown west of Lorraine in the

French Atlas (CASTANET & GUYÉTANT, 1990), was based on three tadpoles reported by Dumois (1984c),

who discussed the status (introduced or not) of this population and specified that these tadpoles were kept

in the collections of the Paris Museum under the numbers MNHN 1984.448-450. If the authors and

editors had “doubts” about this observation, which, quite unlike most other data of the Arlas, was

accompanied by voucher specimens, why didn't they examine them? Posterior to the 1984 observation,

mating calls of the species were heard in the same locality on 15 April 1985 (Dupois, 1985b, unpublished

data), and an adult female and a young one photographed (fig. 2), and then released, on 2 May 1986

(Dusois & EVRARD, unpublished data), so that the population is known to have existed in this locality at

least until 1986. But this is not the most important point in this case.

Source : MNHN, Paris

Dugois 187

Fig. 2. - Specimen (released) of Pelobates fuscus from a small pond near Sainte-Gemme, Indre, France,

2 May 1986 (photo Philippe EVRARD).

Although Dugois’s (1984c) observation is the only one from northern France mapped in CASTANET

& GUYÉTANT (1990) and in the European Arlas, these data are not the only recent ones from this region:

Morère (unpublished lecture cited in Dumois, 1984c) reported survival of the species in several other

French localities, but unfortunately he never published these important data. However these recent data

are nothing compared with the numerous older data, especially from the 19th century, documenting the

presence and distribution of P fuscus in northern France. On the basis of a critical survey and evaluation

of the existing literature (PARENT, 1982), PARENT (1981) synthesized the then available and reliable data:

he listed the species as having been reliably reported in 16 departments of northern France and doubtfully

in 17 additional ones. On the basis of these data, he mapped the southern limits of the 19th century

distribution of this species in northern France. He also stressed the fact that this species was currently

suffering regression in Belgium. Regression of populations of this species in France is an important fact

during our century. One hundred years ago, P. fuscus was a rather common species in northern France,

even close to Paris, where it was repeatedly reported by such famous batrachologists as DUMÉRIL &

BiBRON (1841: 480), LATASTE (1876: 12), HÉRON-ROYER (1886: 75-76) or BOULENGER (1897: 203-204), and

from where specimens are kept in the collections of the London Museum (BOULENGER, 1882: 438, 1898:

346), of the Paris Museum (MNHN 4551, Bondy, 19 April 1875, coll. DEGUÉ, MNHN 8063, 8066, CD.56,

neighbourhood of Paris, no date) and of the Wien Museum (NMW 6567, Paris, 1879, coll. LATASTE), but

now this species seems to be totally extinct in all the Paris region (see Dusois & OHLER, 1988). Regression,

and in fact almost total extinction, of Pelobates fuscus fuscus in northern France during our century, while

the same subspecies seems to have remained quite healthy in central and eastern Europe, is a major fact

that (1) calis for a scientific explanation and (2) should have been stressed in a distribution Atlas of

European amphibians. This appears indispensable in order to allow this fact to be properly taken into

account in international conventions and other official lists, all documents which until now have ignored

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188 ALYTES 15 (4)

the high threat level of this subspecies in this part of its range, while much more attention was paid to the

subspecies Pelobates fuscus insubricus in northern Italy: thus, in CORBETT"s introductory chapter to the

Atlas, in the appendix by OLIVERA et al. and in CorBerT’s (1989) book, only the latter subspecies is

mentioned regarding conservation matters, and the subspecies Pelobates fuscus fuscus is not even cited,

although it is clearly the most endangered taxon of the whole French amphibian fauna. Probably the

second species in this respect is Bombina variegata, many populations of which have become extinct in

several parts of France during the last decades (BREUIL & PAILLETTE, 1983; DuBois, unpublished data),

but this fact also is completely ignored in the European Atlas. Both species P. fuscus and B. variegata are

in France in the westernmost part of their range, which may in part account for their fragility in these

regions.

This example shows that, to be useful for dealing correctly with conservation problems, a distribu-

tion survey cannot rely only on recent data, but should also incorporate a good (ï.e., not only complete,

but also critical) knowledge of all the older literature and collection data. The qualification of “critical”

is important. In systematics and faunistics, like in many other scientific fields, some mistakes can have a

very long life, through their repeated copy from the original publication to a second, then a third one, etc.

It may be difficult to break such chains of repetitions, as is well exemplified again in the Arlas: in his text

on Algyroides fitzingeri (p. 219), once again B. SCHNEIDER gives credit to the “legend” of the islets

Bocognanco, Cauro and Orezza, although the latter have never existed, as was already stressed by LANZA

(1983: 733) and DELAUGERRE & CHEYLAN (1992: 66). Critical analyses of the data in the older literature

are therefore of great importance. Probably, for the time being, the most thorough survey of the

chorological herpetological literature in Europe is that of PARENT (1981, 1982) for France and Benelux,

which unfortunately has not been duly taken in consideration by authors and editors of the European

Atlas. Hopefully this important work will be consulted by future workers on the European amphibians

and reptiles, and hopefully also similar works will be prepared and published concerning other parts of

Europe. Such serious and critical surveys of available older data, and also of museum and other

herpetological collections, will be the only way to have objective information on the past distribution of

species in Europe, and, by comparison with recent data, to obtain reliable estimates of the recent changes

in the status of the populations and species, and of threats hanging over them. In the absence of such

objective information, part of the decisions regarding conservation of the European species (inscription

on official lists, allocation to threat levels, legal restrictions to their transport or commerce, etc.) will be

based exclusively on subjective “feelings” by a few people, so that only the species which happen to be

personally well-known of these persons will be properly dealt with, while others, like the northern French

populations of Pelobates fuscus just discussed, or some populations of Triturus alpestris discussed below,

Will be ignored or their status and threats will be incorrectly assessed.

The second example is meant at stressing the fact that species are not “black boxes” of identical

individuals or populations, but display internal variability, and particularly geographical variation that

may in some cases be worth of being highlighted through taxonomic recognition of subspecies. In such

cases, in the frame of an international conservation policy, special attention should be paid to some

subspecies having a very limited distribution and/or being particularly threatened with extinction. The

cies Triturus alpestris is a good example of this situation. Although the distribution of this

s illustrated on the map of p. 72 of the European Atlas, covers a large part of western, central

and southern Europe, this distribution shows discontinuities, and several subspecies are currently

recognized within this species. The conservation status of these different subspecies is not the same, and

this fact must be taken into account when considering legal and administrative decisions. The nominative

subspecies Triturus alpestris alpestris has a very wide distribution area with numerous populations: as

such, this subspecies is not particularly threatened with extinction, although, like all other European

species, destruction or modification of aquatic habitats clearly results in the regression or extinction of

many local populations. But the situation is much more critical for other subspecies currently recognized

in this species. Thus, A. ZUIDERWUX is correct when she writes, in her accompanying text of T. alpestris

(p. 71): “The subspecies T. a. inexpectatus is rare and endangered and any collecting of specimens means

a serious threat to this subspecies.” Actually, this subspecies, which some consider to deserve species

status as Triturus inexpectatus (BREUIL, 1983, 1986; ANDREONE, 1990), is currently known from only four

populations (Dugois & BREUIL, 1983; Dumois, 1983b, 1993; Gracoma et al., 1988), some of which are

threatened with extinction (Dugois, 1983b), and the absence of any mention of this subspecies in all

current official lists of the European fauna (see OLIVEIRA et al.’s appendix to the A/as) is a serious lack,

Source : MNHN, Paris

Dugois 189

for which the SEH Conservation Committee clearly has some responsibility. This subspecies is not at all

mentioned in CoRBETT"s introductory chapter to the Arlas or in CORBETT”s (1989) book. Other subspecies

of T. alpestris also deserve more attention than they are given in the Atlas.

Of particular importance for amphibians, especially for species or populations that spend a large

part of the year in water, are the problems posed by the introduction of fish in closed water bodies (see e.g.

Duois, 1990, 1991, 1994). For several European amphibian species, this factor of population’s regres-

sion or extinction is certainly as severe as, if not much more so than, “persecution” or “predation by

domestic cats”, but, contrary to the latter, it is not even once mentioned in CORBETT's text in the Arlas or

in several species accounts where they should have been so, like Triturus alpestris. The importance of this

threat on some amphibian species seems therefore to be underestimated by several European herpetolo-

gists, and may then deserve a special discussion. Species of the genus Triturus are particularly vulnerable

to this factor, especially in their populations where newts spend most or all of the year in the water, like

many mountain populations. This can be highlighted by several examples.

On 22 August 1978, I had the opportunity to visit the Prokosko jezero (Bosnia-Herzegovina),

type-locality of the nominal subspecies Triturus alpestris reiseri, and I saw thousands of these large-

headed newts standing on the bottom in the clear water, at the rate of several ones per square meters all

around the lake (Dugois, unpublished data), but I was not allowed by the guards of the lake to collect even

a single specimen: although disappointed, 1 was satisfied with the impression that this unique population

was carefully protected. I informed Michel BREUIL, who applied for and obtained an official collecting

permit for some newts, and visited again the locality in September 1981 and August 1982, but had then the

bad surprise (BREUIL, 1985) to realize that, seemingly as early as in 1972, trouts had been introduced in the

lake, and that the type-population of this nominal subspecies was almost extinct, just a few specimens

having escaped trout predation in a few small zones of difficult access or in neighbouring small ponds.

BREUIL (1985) described similar situations for many other mountain populations of T. alpestris, including

the type-populations of several other nominal subspecies, and BREUIL & PARENT (1988a-b), in their

interesting study (not cited in the Arlas) of the taxonomic, distribution and conservation status of the

subspecies Triturus alpestris veluchiensis, insisted on the dangers that could result for this subspecies from

salmonid introductions. Similar threats have concerned T. alpestris in the French Alps, e.g. in the Parc

National des Ecrins, where introduction of trouts was followed by total extinction of some populations

(Breuil, 1985). This problem is there of a particular significance, since in this area the subspecies Triturus

alpestris alpestris and Triturus alpestris apuanus meet (BREUIL, 1986): extinction of natural populations

following trout introductions will preclude any further study, e.g. by protein electrophoretic methods or

by study of DNA microsatellites, of fine genetic structure of these interesting populations, to reconstruct

migration and introgression phenomena involving the two taxa.

For a species like Triturus alpestris, which often inhabits mountain lakes where most or all of the year

cycle may take place in the water (especially in populations with a high percentage of paedomorphic

specimens), introduction of fish may be a very rapid and irreversible factor of extinction of populations.

The problem also exists for plain species or populations of newts, especially when associated with another

threat factor, duly mentioned by CoRBETT in the Atlas (p. 29), namely habitat fragmentation. In some

parts of the Paris region for example, ponds and other breeding habitats suitable for amphibians have

become so rare that many populations of these animals may be regarded as inhabiting continental islands

completely isolated one from another, and particularly vulnerable. In a growing number of small isolated

ponds, local people have introduced cyprinids, not for fishing purposes but apparently solely for the

purpose of seeing red fish in the water: in a number of these ponds, these fish, probably through predation

on the eggs, have led local populations of amphibians, and particularly of newts, to extinction, and as

these populations are now separated from other neighbouring conspecific populations by impassable

zones of monocultures, built areas or roads, they cannot be colonized again (DuBois, unpublished data).

In such regions, newt populations may become extinct one after another, each local extinction con-

tributing to weaken even more the remaining neighbouring populations and leading ultimately to

complete extinction of some species over a growing area.

This important factor of fish introductions should therefore be given proper attention: diffusing

information on this problem and trying to introduce in international and European legislative texts severe

regulations against uncontrolled introductions of fish are among the first actions European batracholo-

gists are entitled to expect from a European herpetological society, but unfortunately this question is not

tackled even once in CoRBETT's introduction to the European Alas. Nevertheless, a number of European

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190 ALYTES 15 (4)

batrachologists are aware of this problem, as is made clear by the fact that it is mentioned in passing by

several of them in their accompanying texts in the Arlas, in the following species accounts: Euproctus

asper, Euproctus platycephalus, Triturus italicus, Triturus montandoni, Triturus vulgaris, Bombina bom-

bina, Pelobates cultripes, Pelobates fuscus, Rana iberica and Rana temporaria. This question is also briefiy

but repeatedly mentioned in several chapters of CoRBETT's (1989: 15, 45, 130-131, 136, 139, 155, 160-161,

171, 175-176, 202, 208, 256) book about conservation of European amphibians also published under the

umbrella of SEH. The absence of any general statement and international strategy of SEH regarding this

problem is all the more difficult to understand. Possibly the old tradition of always considering

amphibians and reptiles together, under the general discipline of “herpetology”, may contribute to

obseure the biological particularities of amphibians that require distinct conservation strategies for these

animals. As a matter of fact, the particular problem posed to amphibians by fish introductions was

among the examples mentioned to support the need for recognizing batrachology as a distinct scientific

discipline (Dugois, 1991).

METHODOLOGY OF THE INQUIRY

The fact that in the Ar/as some of the spots credited to Rana dalmatina were almost certainly based

on observations of Rana temporaria shows that the critical evaluation of data before their incorporation

in the maps was insufficient. These two species show superficial resemblances but nevertheless any

experienced naturalist can distinguish one from another by simple examination of the external phenotype

(Dusois, 1984b). If an observer providing basic data to the inquiry cannot tell R. femporaria from R.

dalmatina, there is a strong possibility that the same observer will also have identification difficulties in

many other cases, such as all other frogs of the genus Rana, Triturus helveticus versus Triturus vulgaris,

Bufo calamita versus Bufo viridis, Hyla arborea versus Hyla meridionalis, or even Alytes versus Pelodytes

or Pelobates, or Discoglossus versus Rana ( Pelophylax) — not to mention the lizards.

Such a statement can easily be confirmed by any zoologist who has examined numerous museum

collections: no major collection worldwide is free from specimens badly identified, even if the work was

done by professional scientists. To mention here only examples from the Paris Museum collection, which

has had a continuous curating by professional zoologists since 1793, here is a non-limitative list of

identification mistakes concerning western Palearctic species which I or other colleagues found while

cursorily looking into the collections since 1977: Rana temporaria under the names of Rana dalmatina

(MNHN 1971.343; see Dupois, 1982c: 63), of Rana ridibunda perezi (MNHN 1973.64-67; see DUBOIS,

1982c: 63) or of Rana gr. esculenta (MNHN 1987.832-914); Discoglossus pictus scovazzi under the name

of Rana ridibunda perezi (MNHN 1961.52, 1961.56, 1961.58, 1961.70-71); Alytes obstetricans maurus

under the name of Discoglossus pictus (MNHN 1908.111, 1994.1894-1897; see Dumois, 1998a). Similar

gross mistakes can be found in the Paris Museum collections from other countries, and actually in most

large herpetological collections all over the world. At least, museum collections have an important

advantage over other kinds of data: specimens remain available for study, re-examination and correction

of identification. This is not the case of distribution data based on field observations for which no voucher

specimen was kept: in such cases one is bound to rely on the validity of the identifications made in the field

by observers. We all know examples of gross identification mistakes made by people not closely

acquainted with the zoological group considered, or even by people who should, according to their

responsibilities, avoid such errors. Such problems are not new: identifications of specimens by a number

of authors of the past, who had a particularly bad knowledge or “feeling” about amphibians and/or

reptiles, cannot be taken for granted, and, before using their data, their specimens must be examined

again. Several names could easily be mentioned in this respect, and are well-known of all experienced

taxonomists.

These examples are not given in order to throw “shame” on any particular persons, but to really

stress, for the many people who do not seem to be aware of this problem, how the identification of

European amphibians (and reptiles) may in many cases be difficult without proper feeling, training,

experience and sometimes sophisticated techniques. Some think that this problem can be solved by the

publication of books and identification keys aiming at helping “amateurs” (or some professionals”) to

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Dusois 191

recognize the species, or by special training courses like those organized by some herpetological societies

in Europe. This is certainly in part true. However, several of the existing books contain mistakes of

various magnitudes, and training courses are useful only if organized by truly competent naturalists. But

this may not be the most severe problem: a 35-year experience has convinced me that identification of

many of these animals requires, beside theoretical knowledge, a certain amount of “feeling” that some

people will always lack. This statement will be well understood by all good field naturalists, who know

that no book or training course will ever replace the intuitive knowledge of some people in the field, who

willimmediately know where to go to look for certain mushrooms in a forest or certain marine animals at

low-tide, even if they are unable to “explain” how they found them, while others will spend the full day

with them but find nothing.

In a sense, books, field guides and keys may play a rather negative role. Providing seemingly simple

keys using just a few characters may give inexperienced people the misleading impression that identifi-

cation of European amphibians and reptiles is a simple and rather mechanical process, rather than a

scientific action: “Actually, putting a Latin name on a specimen is a scientific, not technical, activity.

Giving a name amounts in fact to making a scientific hypothesis, that of conspecificity of this specimen

with the one that originally beared this name, i.e., in nomenclatural terms, its name-bearing type”

(Dumois, 1998b). Rather than a single-step process based on a few characters, identification of a specimen

must rely on a synthetic appraisal of all characters (phenotype, behaviour, mating call, etc.). The existence

of “keys” may contribute to perpetuate a typological conception of species, according to which intra-

specific variability is ignored or grossly underestimated. There are few “diagnostic” characters that are

not liable to vary within a species, and most of the characters used in identification keys can in some cases

be misleading. Here are a few examples, all based on my personal observations, of such “diagnostic”

characters of European amphibian species that may vary in some individuals or in some population and

might lead to uncorrect identifications by inexperienced observers: in Rana temporaria, although usually

the leg is shorter than in Rana dalmatina, in some populations (Gasser’s frog and Rana temporaria

honnorati) it may be almost as long as in the latter species, the heel extending beyond snout tip when the

leg is folded along the body (Dumois, 1982c); in Hyla arborea, although usually a dark stripe is present on

flank, this stripe may in some individuals be very weak or absent, like in Hyla meridionalis: in the genus

Bufo, although usually a yellow mid-dorsal stripe is present in the species calamita and absent in the

species viridis, exceptions to these “rules” can be observed in some specimens or populations of both

species; in A/ytes obstetricans obstetricans, although usually three tubercles are present on palm of hand,

rare individuals may have only two tubercles, like in A/ytes cisternasii, in Triturus alpestris, although

usually the throat is unspotted in the subspecies alpestris and spotted in the subspecies apuanus, it can be

spotted in some individuals and particularly in some populations of the nominative subspecies; similarly,

Triturus helveticus, unlike Triturus vulgaris, normally has an unspotted throat, but some individuals may

have black gular spots, usually surrounded with white. In all these cases correct identification of

specimens generally raises no real problem if the phenotype is considered as a whole and not as a

collection of artificially isolated “diagnostic” characters.

AIl the seemingly pessimistic statements above are not meant at stating that all information from

questionnaires should be banned from a distribution inquiry, but that such data should be used with

considerable caution and after critical analysis. In other word, in order to carry out an international

distribution inquiry, a serious reflection on methodological matters is in order. Let us now examine more

closely these methodological questions !.

1. In all what precedes, I have assumed that, if identification mistakes were made by some observers, they were

so involuntarily, but this may not be the case, as stressed by Frank GLAW (personal communication, 16 January

1998) in his comments on the manuscript of this paper: “Beside the incompetence of observers there are some

other aspects 10 be considered: people can consciously provide wrong data, for example for political reasons.

They may state that endangered ‘red list species’ occur in a given habitat just to have better arguments to protect

“their habitats’ as nature reserves, However, it is nearly impossible to find hard evidence for such kind of fraud.

Itis even possible that people introduce specimens from another locality or that they provide voucher specimens

with wrong locality data. Biological inventories provided by commercial bureaus sometimes seem to produce

their species lists just by looking at the habitat. They then write down species that ‘must’ occur there (like Rana

temporaria), although they were actually never found. Another problem is that of ‘psychopaths’, who try to

make themselves interesting by providing rather spectacular data. And of course even voucher specimens can be

very misleading when locality data are confused. Such voucher specimens with wrong locality data can produce

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192 ALYTES 15 (4)

Any distribution inquiry makes sense only if spots are based (1) on correct identification of observed

specimens and (2) on exact and precise locality (and, although less importantly, date) of observation.

Any reflection on the methodology of such an inquiry should therefore start with a careful evaluation of

the problems posed by the scientific validation or control of these basic data. This is indeed what is

found in serious methodological reflections on distribution inquiries (e.g. PARENT, 1974: 81-88,

1976, 1979: 10-15, 1981: 86-87, 1982: 373-390; ALCHER et al., 1979; BREUIL et al., 1982; Dumois &

More, 1983; DELAUGERRE & CHEYLAN, 1992: 16-17). Surprisingly, this question does not seem to have

been at the center of the reflections of the SEH Mapping Committee. The presentation of methodological

aspects of the Atlas by H. MAURIN et al. (p. 11-16) is very enlightening: it only deals with technical matters

of coding information in questionnaires, of optical reading of the latter and of computer processing

of the data leading to the building of maps, but almost nothing is said about scientific control of the

data. The few words mentioning this aspect are very vague: “According to the objectives set by the

Mapping Committee and also because of the way the work was organised, based upon a network of

responsible persons, but also of ‘loose’ collaborators, it was necessary to choose a very simple methodol-

ogy. This methodology had to be as free as possible from any language problems and well adapted to the

type of available data.” (p. 13). “Co-ordinators regularly sent the filled in questionnaires for processing.

These were checked and then digitised. When questionnaires were sent directly to the SFF/SPN by

collaborators, these were first sent back to co-ordinators for approval before being entered in the

computer.” (p. 15). Therefore, it appears that scientific control of the validity of the data computerised

and used for drawing the maps was not cared for by the Mapping Committee but by the national

co-ordinators chosen by this Committee. The Mapping Committee does not seem to have prepared

guidelines for this scientific control, so that each national unit of the inquiry was apparently free of

developing its own scientific methodology. Viewed under this light, this international inquiry therefore

appears more like the technical juxtaposition under a single mapping system of several distinct inquiries

having slightly or strongly different scientific methodologies for the collect and scientific control of the

basic field data. Heterogeneity in the scientific reliability of the results is not surprising under such a

methodology.

A similar lack of concern and information on methodological problems of scientific control of the

validity of the observations is also striking in various other texts presenting the SFF, and later SPN,

working methodology (DE BEAUFORT & MAURIN, 1985; MAURIN, 1989, 1994; MAURIN et al., 1993) or the

SHF inquiry on the distribution of amphibians and reptiles in France which has largely served as a model

for the European inquiry (CASTANET, 1978; CASTANET & GUYÉTANT, 1990; Gasc et al., 1994). DuBois

(1982a) and Dusois & MorÈRE (1983) have shown that the methodology of the latter inquiry (by a

posteriori “global validation” of computer-produced maps, with simple suppression of some “unlikely”

spots without going back to the original questionnaires, rather than by a priori “spot by spot” critical

control of basic data) was scientifically unsatisfying. A different, stricter methodology was advocated by

several batrachologists, first within (ALCHER et al., 1979), then outside (BREUIL et al., 1982) the SHF

inquiry, but this new inquiry did not result in a final publication, for lack of financial support and staff,

and despite the fact that about 3500 amphibian distribution data from France had been gathered (PAYEN

& DAUM, 1988): rather than lose all these data, it would now seem logical to incorporate them in the

computerised data base of SPN and SEH, but before doing so, it may be useful to discuss in more detail

these methodological questions.

Technical problems of collect, computerisation and mapping of the data are of course important for

any enterprise of the magnitude of the European Ar/as. But these technical questions should not obscure

the scientific ones. As everybody knows, computers will only give you back what you have fed them: if the

basic data (spots of observation of species) are wrong, the final maps will be incorrect, misleading and

unuseful. How can reliable scientific distribution data on organisms be obtained? Zoologists have worked

for two centuries for ascertaining the geographic distribution of species and mapping them, well before

the introduction of computers, data bases and automatic mapping. Two major methods were used in this

respect: field observations, and capture, fixation and conservation of specimens in permanent collections.

Clearly the second method is the only one that meets the requirements of scientific research: in all fields

much confusion (there are numerous examples from Madagascar), since they are generally considered to be more

reliable than the observations of any observer.” These very justified comments provide an additional reason for

paying a close attention to methodological questions in any collective inquiry.

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Dugois 193

of science, repeatability and independent evaluation of data by different researchers is a prerequisite for

acceptability by the scientific community of these data as genuine scientific results. Since distribution data

are not only geographic data, but also historical ones, repeatability of observation is not possible later if

no voucher specimens, or at least photographs, paintings, drawings or detailed descriptions, are available.

What can we do with old data when no specimen or precise information about them was kept? If we do

not want to just discard these data, which may play a crucial role in some cases as testifying to the past

presence of a species in a region where it is now lacking, we are bound to evaluate the reliability of the old

observation through (1) evaluation of the risk of taxonomic mistake at the time of the observation and (2)

evaluation of the taxonomic competence of the observer.

The risk of taxonomic mistake is of course much larger when several similar species are likely to

occur in the area of the observation: if this is the case, and if particular characters are of importance for

the correct identification of species, it will be useful to see if the observer mentioned having checked these

characters in the reported specimens. But, of course, the problem will be almost insoluble if the taxonomy

has changed since the time of the observation: if several species are now recognized in what was then

believed to be a single species, and if several of these species may be expected to have lived in the

observation’s locality, it will usually be impossible to allocate a posteriori this observation to a species,

and the spot must be abandoned altogether, at least at species level (it may remain as an evidence of the

occurrence of an unidentified species of a given genus or species-group).

As for the taxonomic competence of the observer, this point is rarely stated in full words in scientific

publications, perhaps because it sounds “politically incorrect”. However, it is a reality, and science, if it is

to remain a reliable reference for the knowledge of reality, cannot accept all data in order not to upset

anybody. As tackled above, all taxonomists know that not all their colleagues are similarly reliable in their

identifications. In most cases, I personally will have no hesitation (except when there has been a recent

change in the taxonomy of the group, as just mentioned) to accept field identifications, even when not

documented by voucher specimens, from confirmed field naturalists like L.-F. HÉRON-ROYER, F. LATASTE

or G. A. BOULENGER, but I will be much more careful with data from P. CHABANAUD (who could e.g.

identify a Pelobatidae as a Bufonidae: see DuBois, 1980: 174; see also PARENT, 1976, 1981: 86), P.

CANTUEL (see e.g. PARENT, 1981: 86, 1982: 82) or E. AxL (who could e.g. describe the same species as new

under 10 different names: see GorAM, 1974: 157). Even when very good naturalists are at stake, prudence

may be justified, for example when identifications were based on tadpoles or on mating calls: a famous

case is that of the albino tadpole, first identified as Pelodytes punctatus by LATASTE (1878), which was an

Alytes obstetricans (HÉRON-ROYER, 1878, 1887; LATASTE, 1880). Even the great BOULENGER was not free

from mistakes, since, unlike HÉRON-RoYER, he refused to accept the validity of taxa which are now

recognized as valid under the names Pelobates fuscus insubricus, Discoglossus pictus auritus, Hyla

meridionalis or Rana temporaria honnorati. The conclusion of all this discussion is that the greatest care

should be taken before using field data undocumented by voucher specimens. Of course, in areas or

countries poorly explored and for which data are scarce, mapping of amphibians may be in part based on

sighting of specimens in the field even without capture, or on recording or hearing of mating calls (see e.g.

Dusois, 1974; AMIET, 1983), but this can be done only by experienced naturalists, and usually, even for the

latter, it is much more reliable to catch and examine the specimens in the hands and to keep them for

further laboratory study.

Does this mean that data obtained from questionnaires are totally unreliable and that the thousands

of data gathered this way for the European Atlas should be completely discarded? I am not suggesting

this, but rather that control of these data should be much more careful, which is possible, as shown by

some excellent distribution surveys published in the recent years. However, most of these works were of

a much lower magnitude than the European Ar/as. Careful control of the data is more realistic (which

does not mean easy and quick!) in the case of surveys covering a much smaller geographical area (see e.g.

the excellent distribution atlas of Corsican herpetofauna by DELAUGERRE & CHEYLAN, 1992) or only a

given taxon (see e.g. the contributions of the Catalogue of American amphibians and reptiles published

first by ASIH and now by SSAR, whose quality is due to a very careful, species by species, publication

program). Of course, for a work of the magnitude (in terms of numbers of species and of observers, and

of political heterogeneity of the geographical coverage) of the European Atlas, imposing stringent

methodological requirements on the collect and treatment of data and on their analysis would have a cost,

in terms of financial funding, of staff, of working time and of delays before publication. Whether or not

this cost would be justified is another question that will be examined below.

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194 ALYTES 15 (4)

How should the validity of observations be ideally controlled? A minority of observers do mention

in some questionnaires the existence of additional information on a given observation report, such as

photographs, drawings, descriptions or even voucher specimens (e.g., specimens found dead in the field):

in such cases, any doubt on the identification is liable to be removed by study of these documents or

specimens. But in the vast majority of cases no such additional information is available. In such cases, the

only way to assess the scientific reliability of data is indirect. It can then rely on two major kinds of

evidence: (1) an evaluation of the likeliness of the observation of a given species in a given area, or of the

risk of misidentification in this given case; (2) an evaluation of the competence of the observer.

Evaluation of the risk of misidentification requires knowledge of several important facts: whether

the species reported in the questionnaire is sufficiently similar to another or several other ones to allow

confusion by inexperienced observers; whether this confusability exists for all specimens or only for one

sex or at some stages (e.g., egg, tadpole, imago, adult) or for some characters (e.g., mating call); whether

in the area of the observation two or more such confusable species are likely to occur. This evaluation of

risk should therefore be entrusted to specialists who should ideally have a good knowledge of both the

region of the observation and of all confusable species likely to be present there. Finding such specialists

may sometimes prove difficult. In some regions or countries, there may exist for the time being no good

specialist of some herpetological groups: in such cases, it may be necessary to entrust the responsibility of

the local inquiry to someone from another region or country. Reluctance to do so may be “politically”

understandable but may result in poor scientific results.

Even more difficult, of course, is the evaluation of the competence of observers. For this, the best is

clearly the existence of good personal contacts between the responsibles of the inquiry and the observers,

ideally involving personal meetings and common field work. Contact can also be developed by mail or

phone. Finally, if direct contacts are lacking, some evidence can be obtained from careful analysis of the

questionnaires. Examining altogether all the questionnaires sent by an observer, before their possible

distribution to species specialists or their computerisation, can be an efficient way to point to possible

identification mistakes or difficulties. For example, if an observer sent numerous questionnaires from

different localities in the Paris region mentioning the presence of Triturus vulgaris but none of Triturus

helveticus, or the contrary, it will be likely that this observer did not distinguish both species; a similar

warning of caution may come from seeing only Rana temporaria, but no Rana dalmatina, or only RanaKkl.

esculenta, but no Rana lessonae, in questionnaires from this region. Various other kinds of information

can be obtained through a detailed survey of all questionnaires sent by an observer, which can tell us a lot

about the reliability of the data submitted. A similar kind of evidence can be obtained, without seeing

specimens, through detailed analysis of publications: thus, a careful reading of the paper by Spitz (1971)

suggests that this author’s report of Lacerta viridis and Lacerta agilis being often caught together in the

same traps, in a locality where only the former species (rather now Lacerta bilineata; see RYKENA, 1991)

is known to occur (J.-P. BARON, personal communication), was based on misenditifications where only

male Lacerta viridis were recognized as such, while females were mistaken for Lacerta agilis.

What should be done when careful analysis of the data, under the lines suggested above, throws

doubts on the validity of some identifications? The best is certainly not to simply “suppress” the data

altogether, as the possibility always exists that a species, ‘“‘unlikely” to occur in an area, was introduced in

this region: ignoring such data would result in losing an interesting information. If possible, direct contact

should be taken with the observer, which will sometimes allow, through a discussion, to find the source of

the problem. In some cases, it will even be possible to correct a posteriori an identification, so that the data

will not be lost for the inquiry. Only in cases when doubts remain after this effort, should the data of the

questionnaire be considered unreliable, and discarded before computerisation. But in such cases, the fact

that a given observer misidentified some specimens should be kept in memory, and the possibility that

other misleading data were sent by the same observer should be considered seriously, even if the other

data by this observer “look reliable”: we should always remember that, as much as an “unlikely”

observation can be correct, a “likely” observation can be wrong.

This leads us to a final striking methodological problem. Data on the geographic distribution of

animals on our planet are based on two major kinds of information: field collected specimens, with

information on their collection date and place; and data based on scientific judgement, i.e. taxonomic

allocation of these specimens. Only the second kind of information is liable to change with time: as

taxonomy of a group evolves, or as misidentified specimens are re-examined, the names given to

specimens may change. But the specimens remain the same, and their place and date of origin also. When

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Dusois 195

a taxonomist re-examines a collection and changes some names, the corresponding specimens do not

“disappear” from the chorological data, they only shift from one taxon to another: the spot on a map

corresponding to a given specimen remains, only the scientific name associated to it changes. Further-

more, in any “professional” taxonomic, faunistic or chorological work, such re-allocation of names to

specimens cannot be done “silently”, it must be accompanied by a scientific justification given in full

words when the change is introduced: new taxon, new synonym, correction of misidentification, etc. Just

changing names of taxa on distribution maps, or mere “suppression” of some spots on the maps without

written explanation, is not a serious scientific process. However, this is precisely what can be observed in

the series of atlases published by SFF/SPN, of which the European Arlas is the last production.

Detailed comparisons of the successive maps provided for many species in the two successive

versions of the French atlas (CASTANET, 1978; CASTANET & GUYÉTANT, 1990) and in the French part of

the maps of the European Aflas show important differences. Not surprisingly, many of these differences

are increases in the distribution assigned to a species: one expects such an increase as more and more data

are collected. But other changes are the reverse way: for some species, the distribution in France

recognized in these successive books shows a significant decrease. The only possible explanation of such

facts would appear to be re-evaluation of the basic data and new taxonomic allocation of the spots to

other species. But no written explanation of these changes were given with these successive versions of the

maps. Let us consider the species Rana arvalis. To be sure, most of the spots credited to this species in the

first French Atlas (CASTANET, 1978: 63) were completely outside the known range of the species (ARNOLD

& BURTON, 1978; PARENT, 1981), and were most likely based on misidentifications. Suppression of these

spots in the second version (CASTANET & GUYÉTANT, 1990: 82) is not surprising, but not a single word is

provided to explain this: were these spots just erased, or transferred to other species after correction of

identification? Concerning now Bombina variegata, an isolated spot north of Nantes has disappeared

without any explanation between the maps in CASTANET (1978: 41) and in CASTANET & GUYÉTANT (1990:

58); in the European Atlas (p. 98), another unexplained suppression concerns an isolated spot in

Normandy, although Bombina variegata was recently and reliably documented from this region by LEMÉE

(in CoLLEaAu, 1986: 3). As for Salamandra atra, one of the two spots shown in CASTANET (1978: 23) has

disappeared without explanation in CAsrANET & GUYÉTANT (1990: 38), and all three spots shown in the

latter map are absent in the European Atlas (p. 64). Other striking “silent” spot suppressions between the

books of CASTANET (1978) and CASTANET & GUYÉTANT can be found in the maps of the species Triturus

vulgaris, Alytes obstetricans, Pelobates fuscus, Pelobates cultripes, Bufo viridis and Hyla meridionalis;

while in Pelobates fuscus and Bufo viridis the suppressed spots were indicated as “doubtful” in the first

atlas, this was not the case for the other four species. The absence of any explanation for suppressions of

spots from one atlas to the next one and of information on the fate of the “suppressed” spots (allocation

to other taxa or complete discarding of the data) is not compatible with the claim that such atlases are

scientific works: these changes are incomprehensible for the reader and, above all, as such undocumented

changes have occurred already over three successive atlases, there is no reason to think that the next

version of the European Atlas will not include new mysterious changes!

“BETTER THAN NOTHING”"?

The European Atlas, a major collective international endeavour and realisation, is disappointing

in its results, as the scientific validity of the basic data on which the maps were based is open to question.

Clearly, some spots shown on the maps were based on erroneous identifications of specimens, some basic

bibliographic references and museum specimens were ignored, and some texts contain important

mistakes or omissions regarding either the distribution data or their interpretation, particularly in terms

of conservation; additionally, this book contains a number of errors concerning taxonomy and

nomenclature of European amphibians and reptiles. Although these weaknesses were clearly documen-

ted above, what is much more difficult to evaluate is their quantitative importance. When basic data are

voucher specimens, which is the case for most taxonomic and distribution surveys of amphibians and

reptiles over most of the planet (and particularly in tropical countries), mistakes can eventually be

corrected whenever these specimens are examined again. But here the basic data are questionnaires, not

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196 ALYTES 15 (4)

specimens. Whether these questionnaires will now be available to the international scientific community

for critical study, as are usually museum specimens, is not stated in the Atlas. But, even if it is the case,

re-assessment of the reliability of data in these questionnaires would be difficult, for several reasons

analysed above: any questionnaire by itself may often be insufficient for this work, and additional

information may be needed from direct contact with the observers, or at least through comparative study

of all questionnaires sent by a given person. Doing again this work for all 85,000 basic data used in the

Atlas would be at least as time- and energy-consuming as has been the original work which led to the

production of this book. Clearly the methodological reflection should have been deepened further before

Starting the work.

Of course, it is clear that no scientific work is free from errors, and one cannot expect a large-scale

taxonomic or chorological survey, involving hundreds of collaborators and thousands of data, to be so.

However, in order for such a work to deserve the qualification of “scientific”, one should expect the rate

of errors and omissions to be below a certain level: I have suggested elsewhere (Dupois, 1987a-c) that, in

this domain like in other scientific fields, an acceptable standard rate of errors and omissions (“EO rate”)

should be below 5 %. Is this rate respected in the European Atlas? For the time being, too little

information is available to allow to appreciate quantitatively the amount of errors and omissions in this

work (except in the case of synonymies examined above, 27 % only of which are “genuine synonymies”).

The Atlas provides no information on whether, for a given species, the author of the text has seen the

original questionnaires or was only provided the final map, whether all the older relevant literature was

examined, critically evaluated and computerised, whether data on specimens kept in all major museums

were incorporated in the data base, etc. What seems clear is that the way the basic data were obtained and

critically studied before computerisation was heterogeneous. Different methodologies were apparently

used according to the country, and perhaps also to the taxon studied. While it is nice to see that this work

was truly collective and involved several hundred persons, perhaps in a way there were 100 many people

involved to obtain a homogeneous high scientific level result. On another hand, despite this high number

of collaborators, one is struck by the total absence in the lists of observers and authors of several

prominent herpetologists, some of whom have produced significant contributions to European herpetol-

ogy, such as distribution atlases, field guides, books or scientific papers, and are largely cited in the list of

reference at the end of the volume, or of this review: clearly this book was the result of the work of a part

only of the community of European herpetologists.

The problems raised above are probably due to two major kinds of causes: deficiencies in the

methodological reflection before starting the inquiry, and time shortage. This latter problem can be

guessed from some statements in the book itself (p. 11: “Because time was pressing”; p. 13: “because time

was very short”). It is not unique to this work, rather it is a common problem in current research and

scientific publication (see e.g. DuBois, 19876). In particular, this “time shortage” question is often raised

for collective books, as publishers do not like to wait undefinitely for completion of the final manuscript

and tend to impose precise (and usually close) deadlines to editors and authors. Although this is not

justified scientifically, such a hurried attitude is understandable when the publisher is a private company

with commercial constraints. Should it be the same when the publishers are a non-profit scientific

association (SEH) and state organisms (French Ministry of Environment and SPN)? In such cases, one

would expect the major criterion to be scientific quality, not speed of publication (see also DuBois, 1987b:

ii).

Possibly, for the production of such a volume, the motivations of state organisms are different from

those of scientists. In the recent years, state organisms like the French Ministry of Environment have

tended to support financially the publication of distribution atlases, checklists or other documents having

some connection with conservation problems. In some cases, when one considers the scientific quality of

works so produced (often under very short time constraints), one cannot help from wondering whether

the primary goal of such publications was scientific accuracy or simply “to have a document”, whatever

it may be. In several European countries, laws now require that, before undertaking some major works

like building a road, a railway, a dam, etc.), a public inquiry be made on the impact that this work is likely

to have on the environment and on living species. However, these legislative texts usually only require “to

have a study”, not that it be scientifically irreproachable or that its conclusions have a binding effect on the

conception of the works to be done. In such a context, “having an atlas” might appear as a sufficient goal

for such organisms, irrespective of its scientific rigour and quality. Should scientists and naturalists adopt

the same goal?

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Dusois 197

In the recent years, I have talked with many colleagues in different countries and I know that all

do not share my attitude on these problems. Some think that it is better to have an imperfect atlas

than no atlas at all, or an imperfect checklist than no checklist at all (see DuBois, 1987a-b). This is

in part due to a laudable general positive attitude towards such works, with the idea that the result is

“better than nothing”, and also to the fact that, as each person of course knows personally well only a

part of the data covered by such huge endeavours (be these taxonomic or chorological), it is impossible

for any of us to detect all mistakes occurring in such collective works. Quite significantly however,

when one talks with people who tend to support such works, in many cases they will tell you that

the book is good and reliable, except precisely in the given field (be it taxonomic or chorological)

of their particular competence, and in this limited field they will point to mistakes or omissions;

often, probably through a nice “act of faith”, they will assume that such errors are not as common

in the other parts of the book. However, experience shows that exactly the contrary is true: pointing to

specific mistakes in the necessarily limited field of one’s particular competence (as I have done above)

usually allows to disclose the existence of more general methodological problems that will affect all the

work.

The question that must seriously be asked regarding important collective works such as checklists or

atlases is “what is ‘better than nothing’?” Is it a seemingly complete work including numerous mistakes,

or in incomplete work with a low rate of mistakes? I contend that only the second situation qualifies for

the characterization of “better than nothing”, while the first one, in some cases, may be “worse than

nothing”.

The personal responsibility of any researcher when carrying out a scientific work is to make all

possible efforts to produce a scientifically irreproachable result, given the material means that have been

put at his/her disposal to carry out the work. These efforts should bear on all aspects of the work, i.e.

carefully defining the research methodology, rigorously applying this methodology to obtain and analyse

the results, honestly and competently discussing these results and drawing conclusions, and clearly

presenting all these data in a final publication. Although it is clear that a researcher should try his/her best

to obtain proper funding and staff support for the research project, he/she cannot be taken responsible for

deficiencies in this respect, while he/she can be blamed for bad methodology or insufficiently rigorous

work. Science is supported by society as a whole, although of course, in the detail, this financial and

human support is provided through various channels, from international to state and to private ones. The

support currently given in our societies to scientific research is quite different according to the scientific

field at stake, clearly reflecting disparities in the importance that is afforded by our societies to these

different research fields. Can one imagine that, a space probe sent to Mars missing the target by a few

thousand kilometers, or a dam keeping its water for some years after building and then breaking out, or

a HIV-test detecting the presence of the virus in human blood in some cases only, the comments would be:

“it was better than nothing”? I am choosing three caricatural examples on purpose. What is common to

them is that the aim of the work is considered important for mankind, or at least for some people. On the

other hand, why are many zoologists apparently ready to accept that publishing incorrect taxonomic or

chorological data is “better than nothing” and should not be criticised? Possibly because, even among

zoologists themselves, a poor rating is given to these activities, and to their potential consequences in the

real worid. What can be the consequences of publishing an incorrect distribution map of Pelobates

fuscus? These will include an incorrect basic understanding on the history and ecology of the species, i.e.

a consequence which “merely” concerns our scientific knowledge of a “negligible” part of nature on our

planet, and possibly, as a result, inadapted conservation measures concerning this “obscure” species.

Frogs and salamanders are not elephants or whales and, except for a few spectacular ones such as

Mantella, Dendrobates or Bufo periglenes, they elicit little interest among non-specialists. Who cared for

the virtual extinction of Triturus alpestris reiseri? Who will care if Triturus (alpestris) inexpectatus

becomes extinct? Needless to say, to many people and social groups in our society, such problems are of

very weak importance or of no importance at all, so that, for them, an imperfect atlas, rather than “better

than nothing”, might be regarded as “good enough” for its purpose. Should zoologists share this

attitude? If they decide to do so, they should not expect other social groups in our societies to support

what should be their own concern. À number of current zoologists whose major activities are in the

“traditional” fields of taxonomy, faunistics or inventories seem to be almost “ashamed” of their own

work, perhaps because they are impressed by other more recent developments of biology, such as

molecular research, phylogenetic analysis or evolutionary ecology (all works which, of course, are of

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198 ALYTES 15 (4)

great theoretical and practical interest, but which deal with other questions). If zoologists do not struggle

for these “out-fashioned” activities, who will care for the inventory of biodiversity on our planet before

large parts of it are extinct (see Dupois, 1997a)?

In many respects, the importance of the realisation of such collective works as the European Atlas

commands admiration, when one considers the efforts produced by many individuals to produce the basic

data. Most of these observers were amateurs, who had to support personally all the costs implied for them

by this inquiry. Is this situation “normal” and desirable? Are space probes sent to Mars, or molecular

researches carried out, by enthusiastic amateurs, at their own cost? If the 85,000 basic data of the Atlas

had to have been collected by competent professional scientists with normal salaries and paid field work

expenses, the cost of the inquiry would have been much higher. Of course, according to the current

priorities of our societies, such an idea may seem completely irrealistic, if not crazy. Why doesn't it appear

irrealistic or crazy to spend incommensurably higher funds to send space probes to Mars? Is it because

exploration of space is of much more immediate need and importance for mankind than inventorying,

evaluating and conserving biodiversity on our planet? Or is it because the latter aim is regarded of very

low priority by most people in charge of taking major decisions in our societies? Questions like this

should be seriously considered by those who think that mediocre works should be accepted as “better

than nothing”, without discussion, in our field of research, or “good enough” for the later, rather than

struggling for much more funds (for research, collections, publications), much more academic and

non-academic laboratories, jobs of researchers and technicians, high level courses and diplomae, for the

inventory and study of biodiversity. Otherwise, present and future complaints about the impoverishment

of this biodiversity, and about the consequences of this fact on the environment, and ultimately on

mankind, will be completely hypocritical and inefficient.

ACKNOWLEDGEMENTS

For information provided by them and for their comments on a previous version of this paper, I am

grateful to Franco ANDREONE (Torino), Roger Bour (Paris), Pierre-André CRocHer (Montpellier),

Patrick DAviD (Paris), Michel DELAUGERRE (Bastia), Frank GLAW (München), Britta GRILLITSCH

(Wien), Heinz GriLLrTsCH (Wien), W. Ronald HEYER (Washington), Annemarie OHLER (Paris) and

Olivier PAUWELS (Bruxelles).

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202 ALYTES 15 (4)

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INDEX TO SCIENTIFIC NAMES

Agkistrodon halys: 119

Algyroides fitzingeri: 188

Alytes: 180, 190

Alytes cisternasii: 180, 191

Alytes muletensis: 179-180

Alytes obstetricans: 180, 193, 195

Alytes obstetricans maurus: 190

Alytes obstetricans obstetricans: 191

Aquarana: 180

Bombina: 180

Bombina bombina: 180, 190

Bombina variegata: 180, 182, 188, 195

Bombina variegata kolombatovici: 182

Bombina variegata scabra: 182

Bufo: 191

Bufo bufo: 180

Bufo bufo verrucosissimus: 179

Bufo calamita: 180, 190-191

Bufo periglenes: 197

Bufo verrucosissimus: 178

Bufo viridis: 180-181, 186, 190-191, 195

Bufonidae: 193

Chamaeleontidae: 179

Chioglossa lusitanica: 180

Dendrobates: 197

Discoglossus: 190

Discoglossus galganoi: 180

Discoglossus montalentii: 180

Discoglossus pictus: 177, 180-181, 190

Discoglossus pictus auritus: 193

Discoglossus pictus scovazzi: 190

Discoglossus sardus: 180

Eryx jaculus turcicus: 179

Eryx miliaris miliaris: 179

Euproctus asper: 180, 190

Euproctus montanus: 180

Euproctus platycephalus: 180, 190

Geotriton: 179

Hemidactylus turcicus: 178

Hydromantes: 179-180

Hyla arborea: 180, 190-191

Hyla meridionalis: 180, 190-191, 193, 195

Hyla sarda: 178

Lacerta agilis: 194

Lacerta bilineata: 194

Lacerta viridis: 194

Source : MNHN, Paris

204 ALYTES 15 (4)

Macrovipera lebetina obtusa: 179

Mantella; 197

Mertensiella: 180

Molge syriacus: 179

Natrix maura: 184

Natrix tessellata heinrothi: 179

Pelobates: 190

Pelobates cultripes: 180, 190, 195

Pelobates fuscus: 180, 186-188, 190, 195, 197

Pelobates fuscus fuscus: 187-188

Pelobates fuscus insubricus: 188, 193

Pelobates syriacus: 180

Pelobatidae: 193

Pelodytes: 180, 190

Pelodytes punctatus: 193

Pelophylax: 180

Plethodontidae: 179

Pleurodeles: 180

Podarcis hispanica cebennensis: 179

Proteus: 180

Proteus anguinus: 180-181

Rana: 180, 190

Rana arborea: 181

Rana arvalis: 180, 195

Rana balcanica: 119-180, 184

Rana bergeri: 178

Rana catesbeiana: 180, 184

Rana cerigensis: 118

Rana cretensis: 178

Rana dalmatina: 179-180, 184-186, 190-191, 194

Rana epeirotica: 180

Rana gr. esculenta: 190

Rana ki. esculenta: 176, 180-181, 183-184, 194

Rana graeca: 180

Rana ki. grafi: 176, 183-184

Rana kl. hispanica: 184

Rana iberica: 180, 190

Rana italica: 180

Rana kurtmuelleri: 179, 184

Rana latastei. 180

Rana lessonae: 176, 180, 183-184, 194

Rana macrocnemis: 180

Rana perezi: 176, 180, 183-184

Rana pyrenaica: 178

Rana ridibunda: 176, 180, 183-184

Rana ridibunda perezi: 190

Rana shgiperica: 180

Rana temporaria: 180, 184-186, 190-191, 194

Rana temporaria honnorati: 191, 193

Rana (Aquarana): 180

Rana ( Aquarana) catesbeiana: 183

Rana ( Pelophylax): 179-180, 182, 190

Rana ( Rana): 180

Ranidae: 180, 182

Salamandra atra: 180, 195

Salamandra corsica: 178

Salamandra lanzaï: 180

Salamandra salamandra: 180

Salamandrella: 180

Salamandrina: 180

Speleomantes: 179-180

Trionychidae: 179

Triton vittatus: 179

Triturus: 180, 189

Triturus alpestris: 180-182, 188-189, 191

Triturus alpestris alpestris: 188-189, 191

Triturus alpestris apuanus: 189, 191

Triturus alpestris bukkiensis: 179

Triturus alpestris inexpectatus: 188

Triturus (alpestris) inexpectatus: 197

Triturus alpestris reiseri: 189, 197

Triturus alpestris veluchiensis: 189

Triturus boscai: 180

Triturus carnifex: 178

Triturus cristatus: 180

Triturus superspecies cristatus: 180

Triturus dobrogicus: 178

Triturus helveticus: 180, 190-191, 194

Triturus inexpectatus: 188

Triturus italicus: 180, 190

Triturus karelinii: 178

Triturus marmoratus: 180

Triturus montandoni: 180, 190

Triturus vittatus: 179-180

Triturus vulgaris: 180-181, 190-191, 194-195

Source : MNHN, Paris

AINTES

International Journal of Batrachology

published by ISSCA

EDITORIAL BOARD

Chief Editor: Alain Dupois (Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France). At

Deputy Editor: Janalee P. CALDWELL (Oklahoma Museum of Natural History, University of Oklahoma,

Norman, Oklahoma 73019, USA). |

Editorial Board! Jean-Louis ALBARET (Paris, France); Ronald G. ALTIG (Mississippi State University, USA);

Franco ANDREONE (Torino, Italy); Günter GOLLMANN (Wien, Austria); Tim HALLIDAY (Milton Keynes,

United Kingdom); W. Ronald Hever (Washington, USA); Karen R. Lirs (Canton, USA); Masafumi

Marsui (Kyoto, Japan); John C. PoyNroN (London, England); Ulrich Sinscx (Koblenz, Germany); Erik

R. Win (Dubuque, USA). .

Technical Editorial Team (Paris, France): Alain Duois (texts); Roger BouR (tables); Annemarie OnLe (figures).

Index Editors: Annemarie OHLer (Paris, France); Stephen J. RICHARDS (Townsville, Australia).

SHORT GUIDE FOR AUTHORS

(for more detailed Instructions to Authors, see Alytes, 1997, 14: 175-200)

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 tab. 4. 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 (BOURRET, 1942; GRAF & POLLS PELAZ, 1989;

INGER et al., 1974). References in the literature cited section should be presented as follows:

Bourne, R., 1942. — Les Batraciens de l'Idochine. Hanoï, Institut Océanographique de l'indochine: rx +

Grar, J-D. & PoLLs PeLAz, M., 1989. - Evolutionary genetics of the Rana esculenta complex. In: R. M. DAWLEY

& J. P. BoGART (ed.), Evolution and ecology of unisexual vertebrates, Albany, The New York State

Museum: 289-302.

InGeR, R. F., Voris, H. K. & Voris, H. H., 1974. - Genetic variation and population ccology of some Southeast

Asian frogs of the genera Bufo and Rana. Biochem. Genet, 12: 121-145.

Manuscrits should be submitted in triplicate either to Alain Duois (address above) if dealing with

amphibian morphology systematics, biogcography evolution, enetis or developmenta biology, o 1 Janalee

P. CALDWELL (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 referces.

If possible, afer acceptance a copy of the final manuscripton a oppy dik (3 or 5) should be sent

to the Chief Editor. We welcome the following formats of text processing: (1) preferably, MS Word (1.1 to 6.0,

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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 Duois.

Numéro de Commission Paritaire: 64851.

FSC TIPS Source : MNHN, Paris

Alytes, 1998, 15 (4): 137-204.

Contents

Rafael C. LarmaANoOvICH & Julian FAIVOVICH

Dieta larval de Phyllomedusa tetraploidea Pombal & Haddad, 1992

en la provincia de Misiones (Argentina) ............................. 137-144

Dan COGÂLNICEANU, Florin AIOANEI, Constantin CruBuc & Anghelutä VÂDINEANU

Food and feeding habits in a population of common spadefoot toads

(Pelobates fuscus) from an island in the lower Danube floodplain ..……. 145-157

Dinorah D. ECHEVERRIA

Aspectos de la reproduccién in-vitro y del desarrollo larval

de Melanophryniscus stelzneri (Weyenbergh, 1875) (Anura, Bufonidae)

con comentarios acerca del érgano de Bidder ........................ 158-170

Lucrecia FERRARI

Tolerance of high electrolytic and non-electroplytic osmolarities

in Bufo arenarum premetamorphic tadpoles

UDC OPSA ISIN STE rte ee me 171-175

Book review

Alain Duois

Mapping European amphibians and reptiles:

collective inquiry and scientific methodology ........................ 176-204

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: 1% trimestre 1998.

Source : MNHN, Paris: