International Society for the Study

and Conservation of Amphibians

(International Society of Batrachology)

SEAT

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

25 rue Cuvier, 75005 Paris, France

BOARD FOR 1993

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

General Secretary: Alain DUBOIs (Paris, France).

Treasurer: Dominique PAYEN (Paris, France). À

Deputy Secretaries: Günter GOLLMANN (Wien, Austria); Stephen J. RICHARDS (Townsville, Australia);

David B. Wake (Berkeley, U.S.A.).

Deputy Treasurers: Janalee P. CALDWELL (Norman, U.S.A.); Annemarie OHLER (Paris, France).

Other members of the Board: Sergius L. KuzMN (Moscow, Russia); Roy W. MCDIARMD

(Washington, U.S.A.).

TARIFFS FOR 1993

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Museum of Natural History, University of Oklahoma, Norman, Oklahoma 73019, U.S.A.

Source : MNHN, Paris

AIDTTES

INTERNATIONAL JOURNAL OF BATRACHOLOGY

March 1993 Volume 11, N° 1

Alytes, 1993, 11 (1): 1-15. 1

Descripciôn de una nueva especie europea

de rana parda (Amphibia, Anura, Ranidae)

liothèque Centrale Muséum

CL LL

Jordi SERRA-COBO 3. 3001 0011158

Instituto Pirenaico de Ecologia (C.S.I.C.), Apartado 64, 22700 Jaca, España

This paper describes a new species of brown frog, discovered on the

fringes of the Ordesa and Monte Perdido Park (Huesca, Spain). It is a slender,

agile, long-legged and medium sized species with usually a light coloration, a

small tympanum and well-separated nostrils, and it favours torrent habitats. It

lives in clear, cold, oxygenated and reasonably fast-running waters where .it can

find refuge under stones or in tiny fissures. In most cases it is sympatric with

Euproctus asper and, in some cases, with Rana temporaria. Its morphology and

ecology distinguish it from Rana temporaria, Rana iberica and Rana dalmatina.

Its area of distribution is limited, so far, to the Spanish side of the central and

westem Pyrenees.

A finales de verano de 1990, con motivo de un estudio faunistico sobre los

vertebrados del Parque Nacional de Ordesa y Monte Perdido (provincia de Huesca,

España) y su zona periférica, el autor capturé, por primera vez, la especie aqui descrita.

Las ranas se hallaron en un torrente del valle de Ordiso. A partir de la referida fecha, se

realizaron multiples estudios y prospecciones periédicas para determinar y conocer el

alcance del hallazgo. En invierno de 1991 se logré encontrar diversas puestas de dicha

especie, siendo fotografiadas y trasladadas al laboratorio para su ulterior estudio

ontogénico. Posteriormente, la realizacion de campañas prospectivas permitié hallar

diecisiete localidades distintas para la nueva rana, algunas de ellas muy alejadas entre si.

Los resultados obtenidos después de dos años de estudios, indican que las ranas halladas

pertenecen a una nueva especie.

El presente articulo describe la morfologia y algunas de las caracteristicas ecolégicas

de la nueva rana, destacando sus principales rasgos distintivos. Su morfometria, ecologia,

biologia, diferencias aloenzimäticas, cariotipo y desarrollo larvario se trataraän mäs

exhaustivamente en oportunas ulteriores publicaciones, algunas de ellas ya en curof see DU

MUSEUM

PARIS

Source : M

2 ALYTES 11 (1)

(articulo en preparacién sobre morfometria elaborado por DuBois, OHLER & SERRA-

CoBo).

Abreviaciones utilizadas. — MNCN: Museo Nacional de Ciencias Naturales de Madrid

(CSIC); MNHN: Muséum National d'Histoire Naturelle, Paris; MZB: Museu de Zoologia

de Barcelona; DZV: Departament de Biologia Animal, Fac. Biologia, Universitat

Barcelona; IPE: Instituto Pirenaico de Ecologia (CSIC), Jaca-Zaragoza. La notaciôn

utilizada para cada variable, viene indicada en el pie de la Tabla I.

Material de comparaciôn. — Dieciocho adultos de Rana iberica (MNHN 1992.4852-4853,

4857, 4859-4860, 4863, 4865, 4870-4873, 4875, 4878, 4881-4883, 4885-4886) de la Laguna

Grande de Gredos (Âvila, España). Nueve machos adultos de Rana temporaria (MNHN

1979.7510-7514, 7516-7517, 7520, 7522) del Col du Pourtalet (Pyrénées Atlantiques,

Francia). Diez machos de Rana dalmatina (MNHN 1980.1009-1016, 1019-1020) de la

Forêt de Fontainebleau (Seine-et-Marne, Francia).

Metodologia estadistica utilizada. — Todas las variables, excepto SVL, han sido

transformadas en tanto por mil de la longitud hocico-ano antes de ser procesadas

estadisticamente. La Tabla I se ha elaborado a partir de las medidas del holotipo y de los

paratipos (8 hembras y 15 machos). Para comparar las distintas especies se ha considerado

como unidad de tratamiento los individuos adultos de una misma poblacién y mismo sexo

(machos capturados en la Espata: MNCN 16661; MNHN 1993.2502, 2504, 2509-2511,

2514-2516), utilizando el test estadistico de la U de Mann-Whitney (Dupois, 1984a; SIEGEL,

1985; OHLER & KAZADI, 1990).

Rana pyrenaica sp. nov.

(fig. 1-4)

Holotipo. — MNCN 16661, macho adulto capturado el 12 de octubre de 1992, por Jordi

SERRA-CoBo, en la Espata (Villanua, Huesca, España).

Medidas del holotipo. — SVL = 45,3 mm; HW = 339; HL = 337; IFE = 174; FLL =

245; TL = 546; FOL = 574; IN = 86; EN = 71; EL = 91; TYD = 35; TFL = 124;

IMT = 42, ITL = 124; 2TL = 209; 4TL = 515; TT = 253; ABL = 195; LCD = 497;

E = 62.

Paratipos. — Siete hembras adultas (MNHN 1992.5234-5235, 1993.2501, 2507, 2512-2513;

DZV 2766) y siete machos adultos (MNHN 1993.2502, 2504, 2509-2511, 2514-2515)

capturados el 14 de julio de 1992, y un macho adulto (MNHN 1993.2516) capturado el

12 de octubre de 1992, por Jordi SERRA-CoBo, en la Espata (Villanua, Huesca, España).

Dos machos adultos (MNHN 1993.2506, 1992.5236) capturados el 10 y el 12 de octubre

de 1992, respectivamente, por Jordi SERRA-CoBo, en el valle de Bujaruelo (Torla, Huesca,

España). Un macho adulto (MNHN 1993.2505) capturado el 11 de setiembre de 1990, dos

machos adultos (MNHN 1993.2503, 2508) capturados el 15 de julio de 1992, y una hembra

adulta (MZB 92.0167) y un macho adulto (IPE 1992.4040) capturados el 12 de octubre de

1992, por Jordi SERRA-CoBo, en el valle de Ordiso (Torla, Huesca, España).

Etimologia del nombre especifico. — Hace referencia a la cordillera donde se descubri6 la

especie. À

LA Source : MNHN, Paris

-GEe?

>n2

Tabla I. — Resumen de los datos biométricos obtenidos para 8 hembras y 15 machos de Rana pyrenaica.

X, media; 5, desviaciôn estandard; V;,, coeficiente de variaciôn de Haldane (HALDANE, 1955; DELAUGERRE & Dumois, 1985); U, resultado

del test de Mann-Whitney (comparaciôn hembras-machos); P, probabilidad asociada al valor de U para un test bilateral. Variables

estudiadas: SVL, longitud morro-cloaca; HW, anchura de la cabeza; HL, longitud de la cabeza; IFE, separaciôn entre la parte anterior

FLL, longitud del antebrazo (desde el codo al borde proximal del tubérculo palmar externo); TL, longitud de la pierna

(desde la articulacién tibio-tarsal a la tibio-femoral); FOL, longitud del pie (desde el borde proximal del tubérculo metatarsiano interno

hasta el extremo del cuarto dedo); IN, separaciôn entre orificios nasales; EN, distancia entre borde anterior del ojo y orificio nasal;

EL, longitud del ojo; TYD, diâmetro del timpano; TFL, longitud tercer dedo de la mano (de la base primer tubérculo al extremo del

dedo); IMT, longitud del tubérculo metatarsiano interno; ITL, longitud primer dedo del pie; IMT/ITL, tanto por mil del tubéreulo

metatarsiano interno respecto a la longitud del primer dedo del pie; 2TL y 4TL, longitudes del 2° y 4° dedos del pie, respectivamente

(medidos desde el borde distal del tubérculo metatarsiano interno, hasta el extremo del dedo); TT, longitud del talôn (desde articulaciôn

tibio-tarsal al borde proximal del tubéreulo metatarsiano interno);, ABL, longitud del brazo (medido por la cara ventral); LCD,

de los ojos;

longitud del muslo (de la cloaca a la rodilla); EE, distancia interorbitaria.

Hembras Machos

Variable Rango x C2 Mediana Va Rango x C2 Mediana Va U P

svL 355.510 433 10,69 33,445,7 420 331 si soi | si >ou0

Hw 323410 350 an 305.365 350 16.97 353 93 | 485 >o10

HL 329-368 346 429 309-357 34 145 337 400 | 36 >o10

IFE 162-198 1 12 166-191 ms 820 174 476 | 45 >010

FLL 212274 243 9.4 222281 258 16,96 262 668 | 37 >010

TL 518-588 557 sn 543619 565 22,87 557 a12 | 525 >ou0

FOL 497-582 550 5.88 539.642 579 2328 519 a | 25 <oos

IN 88-101 93 5.53 84100 9 so 90 sa | 4 >o10

EN 7179 1 447 6284 n 5418 mn 72 | 4 >o10

EL 96-123 109 918 88-124 16 12,4 105 1,85 | 505 >0.10

TD 3148 si a NE 2647 36 578 36 1632 | 33-010

TEL 126-144 136 135,5 an 110-139 Ds 83 128 662 | 27 <oos

IMT 33-50 a 405 14,02 33:51 a sn #2 155 | 56 >010

mL 90-124 ut 15 1023 93-128 u7 102 m1 88 | 375 >0.10

IMTATL 279-480 370 385.5 1818 260-494 36 6324 351 157 | 6 >o10

TL 173-228 212 214 EE 182-239 2 1604 27 735 | 355 >o10

sTL 465-563 su si7 627 480.569 523 24 528 548 | 41 >o10

TT 220-273 251 252.5 543 239-298 257 1460 251 528 | 52 >o10

ABL 162-225 201 206,5 10,38 206 22,82 209 11,26 50 >0,10

LoD 469.565 sir 5155 642 510 2571 509 513 | s5 >o10

EE 57.93 n n 16,05 13 898 mn msi | 55 >o1

OH0D-VHUES

Source : MNHN, Paris

4 ALYTES 11 (1)

Tabla 11. — Comparaciôn morfométrica entre 9 machos adultos de Rana pyrenaica y 18 machos

adultos de Rana iberica. Se considera P la probabilidad asociada al valor de U para un test

bilateral.

Especie Rango X 5 Vu Mediana U P

Longitud del antebrazo (en % de SVL)

Rpyrenaica 245-277 260 13,08 5,17 262

R iberica 202-304 236 21,10 9,36 234 13 <0,002

Diämetro mäximo del timpano (en % de SVL)

R. pyrenaica 26-47 38 5,89 15,93 39

R iberica 42-70 52 7,73 15,08 51,5 8 <0,002

Longitud del tercer dedo de la mano (en %o de SVL)

Rpyrenaica 110-139 125 8,24 6,78 127

R. iberica 143-178 164 8,74 5,48 164,5 0 <0,002

Longitud del primer dedo del pie (en %o de SVL)

R. pyrenaica 93-128 116 11,18 9,91 121

R. iberica 112-151 133 8,58 6,54 133,5 13 <0,002

Longitud del segundo dedo del pie (en % de SVL)

R pyrenaica 182-239 216 18,06 8,59 217

R iberica 222-259 244 987 4,10 247,5 Il <0,002

Longitud del talôn (en % de SVL)

R pyrenaica 243-298 259 1705 6,77 251

R iberica 228-281 245 13,21 5,47 242,5 38 <0,02

Distancia interorbitaria (en % de SVL)

R. pyrenaica 62-86 71 8,42 12,19 72

R: iberica 85-112 95 6,41 6,84 95 Ll <0,002

Diagno — Especie de cuerpo grâcil y talla mediana, inferior a Rana temporaria y R.

dalmatina (Tablas III y IV). Orificios nasales mâs separados entre si y proporcionalmente

mäs distantes del borde anterior del ojo que en R. temporaria y R. dalmatina (Tablas III

y IV). Mancha temporal poco conspicua, menos que en R. iberica, R. temporaria y R.

dalmatina. Timpano muy pequeño y dificil de distinguir (Tabla I). Hocico menos

acuminado que en R. iberica. Antebrazo y pierna relativamente largos respecto a la talla

del cuerpo (Tabla I). Tercer dedo de la mano relativamente menor que en R. iberica y R.

Source : MNHN, Paris

SERRA-COBO

Tabla II. — Comparaciôn morfométrica entre 9 machos adultos de Rana pyrenaica y 9 machos

adultos de Rana temporaria. Se considera P la probabilidad asociada al valor de U para un test

bilateral.

Especie

Rango X 5 Vu Mediana U P

R pyrenaica

R. lemporaria

R. pyrenaica

R. temporaria

R. pyrenaica

R. temporaria

R. pyrenaica

R. temporaria

R. pyrenaica

R lemporaria

R: pyrenaica

R: temporaria

R pyrenaica

R. temporaria

Longitud del hocico-ano (en % de SVL)

390457 43,2 2,50 5,95 43,8

589-703 65,8 3,20 5,00 65,9 0 <0,002

Separaciôn entre la parte anterior de los ojos (en % de SVL)

167-176 172 3,76 2,25 174

133-150 145 5,03 3,57 146 0 <0,002

Longitud de la pierna (en %o de SVL)

543-564 552 6,97 1,30 552

513-555 533 15,84 3,05 530 11 <0,02

Separaciôn entre orificios nasales (en %o de SVL)

86-100 90 4,58 5,23 89

68-80 72 4,22 6,02 72 0 <0,002

Distancia entre orificio nasal y borde anterior del ojo (en %o de SVL)

62-79 71 4,93 7,14 70

54-68 63 4,19 6,84 64 6 <0,002

Diämetro mäximo del timpano (en %o de SVL)

26-47 38 5,89 15,93 39

56-77 70 6,12 8,99 70 0 <0,092

Longitud del segundo dedo del pie (en %o de SVL)

182-239 216 18,06 8,59 217

230-280 257 1892 7,57 257 4 <0,002

dalmatina (Tablas II y IV). Pierna relativamente mäs larga que en R. temporaria (Tabla

JT) y menos que en R. dalmatina (Tabla IV). Articulaciôn tibio-tarsal rebasando el

extremo del hocico cuando se dobla la extremidad posterior hacia adelante. Longitud del

primero y segundo dedos del pie menor que en R. iberica (Tabla II). Dedos de las manos

y de los pies redondeados en su extremo distal. Coloraciôn de la garganta normalmente

jaspeada gris ténue y sin linea media clara como en R. iberica. Coloracion dorsal entre

canela crema y gris olivâceo. Canto grave y débil, mäs flojo que en R. temporaria.

Source : MNHN, Paris

ALYTES 11 (1)

Tabla IV. — Comparaciôn morfométrica entre 9 machos adultos de Rana pyrenaica y 10 machos

adultos de Rana dalmatina. Se considera P la probabilidad asociada al valor de U para un test

bilateral.

Especie

Rango X LA Vu Mediana U P

R. pyrenaica

R. dalmatina

R. pyrenaica

R. dalmatina

R pyrenaica

R dalmatina

R. pyrenaica

R. dalmatina

R pyrenaica

R. dalmatina

R pyrenaica

R. dalmatina

R pyrenaica

R. dalmatina

R pyrenaica

R. dalmatina

R pyrenaica

R. dalmatina

R pyrenaica

R. dalmatina

R pyrenaica

R dalmatina

Longitud del hocico-ano (en % de SVL)

390457 432 2,50 595 43,8

489614 56,1 3,84 7,02 56,7 0 <o,002

Separaciôn entre la parte anterior de los ojos (en %o de SVL)

167-176 172 3,76 2,25 174

147-163 154 5,64 3,75 154 0 <0,002

Longitud de la pierna (en % de SVL)

543-564 552 697 1,30 552

607-631 618 824 1,37 618,5 O0 <0,002

Longitud del pie (en %o de SVL)

539-591 567 15,97 2,13 574

571-603 588 12,21 2,13 588,5 13 <0,02

Separaciôn entre orificios nasales (en %o de SVL)

86-100 90 4,58 5,23 89

71-78 74 23 3,09 74 0 <0,002

Distancia entre orificio nasal y borde anterior del ojo (en %o de SVL)

62-79 71 4,93 7,14 70

60-72 65 3,99 6,29 65 13,5 <0,02

Diämetro mäximo del timpano (en %o de SVL)

26-47 38 5,89 15,93 39

71-88 78 6,43 8,45 75,5 0 <0,002

Longitud del tercer dedo de la mano (en %o de SVL)

110-139 125 8,24 6,78 127

130-143 136 5,15 3,88 136,5 6,5 <0,002

Longitud del cuarto dedo del pie (en %o de SVL)

480-542 510 19,06 3,84 505

513-560 537 15,94 3,04 541 LE) <0,02

Longitud del muslo (en %o de SVL)

476-524 503 14,66 3,00 502

539-585 570 16,44 2,96 576,5 O0 <0,002

Longitud del talôn (en %o de SVL)

243-298 259 1705 6,77 251

268-292 279 7,01 2,58 278 12 <0,02

Source : MNHN, Paris

SERRA-COBO 7

Descripciôn morfolôgica del adulto. — Es un anuro de talla mediana (Tabla I) y de cuerpo

esbelto. La cabeza es ligeramente mäs ancha que larga (Tabla I). La distancia entre los

orificios nasales es superior a la separaciôn interorbitaria (Tabla I). Las oberturas nasales

estän mâs cercanas al extremo del hocico que al borde anterior del ojo (fig. 1). El hocico

es corto, no acuminado, proyectändose ligeramente por delante de la obertura bucal. La

mancha temporal es poco conspicua, prolongändose, en franja estrecha, por delante del

ojo hasta alcanzar el extremo del morro. La parte posterior de la referida mancha estä

delimitada por dos pliegues prominentes: uno supratemporal y otro subtemporal. Los

ojos, no muy saltones, se disponen lateralmente. La pupila es oval. El canthus rostralis es

romo. Sobre el labio superior un repliegue de coloraciôn clara se prolonga hacia la region

subtemporal, rebasando la comisura bucal y, en algunos casos, alcanzando la base de

las extremidades anteriores. En la mayoria de individuos dicha franja clara se inicia

cerca del extremo del hocico. El timpano es pequeño y de dificil observacion (Tabla I),

rasgo caracteristico de la especie (fig. 1-2). Presenta dos series de dientes vomerianos

situadas detrâäs de las coanas y en posiciôn oblicua respecto al eje del cuerpo, mäs

separadas entre si en su extremo distal que en su extremo proximal. La separacion entre

el extremo distal de los dientes vomerianos y las coanas es menor que la longitud de las

series dentarias.

Los miembros anteriores son relativamente largos (sobre todo el antebrazo) y

permiten el levantamiento del torax y la cabeza. El antebrazo es mâs largo que el brazo

(Tabla I). El tercer dedo de la mano es corto (Tabla I), aproximadamente la mitad del

antebrazo. El extremo distal de los dedos de la mano es redondeado. Los tubérculos

subarticulares son romos o ligeramente acuminados, circulares y estän bien desarrollados.

Las manos presentan dos tubérculos palmares alargados (fig. 3).

Las extremidades posteriores son relativamente largas, sobre todo la pierna, respecto

a la talla del cuerpo. AI doblarlas hacia adelante, la articulaciôn tibio-tarsal apenas rebasa

el extremo del hocico. El pie (desde el borde proximal del tubérculo metatarsiano interno

hasta el extremo del cuarto dedo) es aproximadamente igual a la longitud de la pierna

(Tabla I). Los dedos del pie presentan su extremo distal redondeado. La membrana

interdigital es fina y està muy desarrollada, incluyendo en ella la mayor parte de todos los

dedos, excepto la porciôn distal del cuarto dedo. El tubérculo metatarsiano interno es

pequeño, romo, oval y de consistencia blanda, siendo su longitud alrededor de 1/4 a 1/2

de la talla del primer dedo. Los tubérculos subarticulares estän, en general, bien

desarrollados y son circulares, romos o ligeramente acuminados (fig. 4).

La piel es lisa, fina y muy viscosa en animales vivos. Los repliegues dorso-laterales son

estrechos, prolongändose desde la parte posterior del ojo hasta la region anal.

Coloraciôn. — La coloracién dorsal varia entre color canela crema y gris olivâceo,

presentando discretas manchas verdosas. Sin embargo, las referidas moteaduras son poco

conspicuas en animales vivos, ora en la region dorsal, ora en las extremidades anteriores

y posteriores. Algunos ejemplares pueden presentar una mancha en forma de V invertida

en la espalda. Las hembras suelen tener ciertos tonos rojizos en los pârpados, alrededor

de los repliegues dorso-laterales y en la regiôn dorsal de las extremidades posteriores, y su

cuerpo suele ser mäs voluminoso. No obstante, en algunas ocasiones se han observado

machos con tonalidades rojizas, si bien éstas eran ms tenues (holotipo MNCN 16661). El

Source : MNHN, Paris

ALYTES 11 (1)

Fig. 1.- Rana pyrenaica sp. nov., visiôn lateral de la cabeza (MNHN 1992.5236).

5smm

Fig. 2. — Rana pyrenaica sp. nov., vision dorsal de la cabeza (MNHN 1992.5236).

Source : MNHN, Paris

SERRA-COBO 9

Fig. 3. — Rana pyrenaica sp. nov., visiôn ventral de la mano derecha (MNHN 1992.5236).

Fig. 4. — Rana pyrenaica sp. nov., visiôn ventral del pie derecho (MNHN 1992.5236).

Source : MNHN, Paris

10 ALYTES 11 (1)

iris es dorado con pequeñas puntuaciones de color oscuro. La regiôn ventral es muy clara,

existiendo, casi siempre, un ligero jaspeado gris-rosäceo en la garganta, pero en ningün

caso se ha observado una banda clara en la linea media, ni tampoco presenta una banda

transversal. La coloracion ventral a nivel femoral es amarillenta, pudiendo ser azulada en

verano. Durante la época de celo la region femoral de los machos es sonrosada. La

pigmentaciôn tiende a ser bastante homogénea en todos los ejemplares estudiados.

Caracteres sexuales secundarios. — La longitud del tercer dedo de la mano suele ser mayor

en las hembras que en los machos (Tabla I). La longitud del pie tiende a ser mayor en los

machos que en las hembras (Tabla I). Los machos carecen de sacos bucales y su canto es

grave y tenue. Las callosidades palmares de los machos son, generalmente, amarillentas

(fig. 3). Después del letargo, los machos presentan desarrollados pliegues cutäneos en la

region lateral del tronco, en la region dorso-posterior de la cabeza y a nivel femoral, los

cuales devienen conspicuos durante el amplexus.

Puesta y desarollo. — La puesta se efectüa entre finales de invierno y principios de

primavera (febrero-marzo), dependiendo de las caracteristicas climatolôgicas del biotopo,

la altitud de la localidad de puesta y las oscilaciones meteorolôgicas estacionales. Las

puestas son pequeñas, pues suelen estar constituidas por un nümero de huevos

comprendido entre 70 y 110 (se han observado unas 15 puestas), y no forman masas

compactas sino mäs bien racimos. La gelatina envolvente es muy densa, impidiendo la

flotacién. Por el contrario es escaso su espesor protector. La especie tiende a adherir la

puesta bajo las piedras (un 87 % de las puestas observadas), en la vegetacion de fondo o

en las fisuras rocosas de arroyos y, ocasionalmente, sobre fondo lodoso siempre que las

aguas estén quietas. El embriôn es de color pardo-oscuro y relativamente grande respecto

a su envoltura gelatinosa.

La velocidad de desarrollo embrionario estä en funciôn de las condiciones climato-

lôgicas anuales, siendo frecuente el retraso ontogénico como consecuencia de las nevadas

y frios primaverales tardios, tipicos de la regiôn pirenaica occidental. Transcurridas unas

semanas, el embriôn pierde su intensa coloraciôn oscura, adquiriendo un tono grisäceo

claro al Ilegar al estadio en que se forman los esbozos de la cabeza y la cola. Llama la

atenciôn el relativo gran desarrollo de su abdomen y su delgada y delicada piel. La cola

adquiere rapidamente una notable longitud relativa.

Las larvas ya desarrolladas son muy caracteristicas y fâciles de distinguir por su

coloracién oscura, su musculosa aleta caudal y sus manchas de color dorado con

irizaciones (pueden ser mäâs o menos moteadas segün las poblaciones). Tienen aspecto

robusto y a su vez estilizado y elegante. Estän bien adaptadas a los cursos de agua,

disponiéndose en su fondo al amparo de la corriente o nadando a modo de “trucha”.

Distribuciôn y ecologia. — Los adultos y subadultos se alejan poco del agua, situändose

en riberas de arroyos o bien escondiéndose bajo pequeños saltos de agua, piedras situadas

en torrentes o en estrechas fisuras. La especie tiende a ser simpâtrica con Euproctus asper,

siendo muy similares sus residencias ecolôgicas (un 82 % de las 17 localidades donde se ha

hallado a R. pyrenaica también habitaba E. asper). Suele vivir en aguas claras, frias y

oxigenadas, ora pequeños cursos hidricos con poca renovaciôn (siendo simpätrica con R.

temporaria y desovando ambas especies en el mismo lugar), ora en torrentes caudalosos.

Source : MNHN, Paris

SERRA-COBO 11

R. pyrenaica no se ha hallado nunca en aguas estancas. Su gracilidad, agilidad, estrategia

reproductora y anatomia de sus larvas, le permiten poblar biotopos donde a R. temporaria

y al resto de anuros montanos pirenaicos les es dificil residir. Escapa, asi, a una posible

competencia trofica con la rana bermeja, Bufo bufo y Alytes obstetricans, anuros

abundantes en la regin centro-occidental del Pirineo. De las diecisiete localidades

estudiadas, sélo en tres de ellas la especie es simpâtrica con R. temporaria. Los efectivos

de R. pyrenaica suelen dispersarse a lo largo de los cursos de agua, ora referente a los

individuos adultos, ora a las larvas cuyas concentraciones no suelen ser tan elevadas como

en R. temporaria. Dicha menor densidad de renacuajos, cabe atribuirla, en parte, a la

estrategia reproductora seguida por R. pyrenaica, la cual tiende a ser de la K (como en

Alytes obstetricans) y no de la r como en R. temporaria y Bufo bufo, y a la referida

dispersiôn de los individuos adultos reproductores (BEGON, HARPER & TOWNSEND, 1988).

Se ha hallado, a R. pyrenaica, en la franja altitudinal comprendida entre los 1200 y 1700

m s.m. Su distribuciôn se circunscribe, por ahora, a la regiôn pirenaica centro-occidental

española (fig. 5).

Discusiôn. — La ecologia, morfologia y etologia de R. pyrenaica le distinguen bien de R.

temporaria, especie muy abundante en el Pirineo (BOULENGER, 1879, 1898, 1910; LANTZ,

1927; BECK, 1943; BALCELLS, 1956; MERTENS & WERMUTH, 1960; FRETEY, 1975; ARNOLD

& BURTON, 1978; ANDRADA, 1980; PARENT, 1981; DuBois, 1982a-b; MARTINEZ-RICA, 1983;

MARTINEZ-RICA & REINÉ-VINALES, 1988; ESTEBAN, 1990). Menor talla del cuerpo, timpano

mäs pequeño, mancha temporal mucho mäs tenue, parte distal del hocico proporcional-

mente mâs ancha, mayor separaciôn, en relaciôn a la longitud del cuerpo, entre orificios

nasales y entre estos ültimos y el borde distal del ojo, talla relativa de la pierna suele ser

mäs larga (excepto en relaciôn a la rana de Gasser, segün estudio morfométrico realizado

por Duois, OHLER & SERRA-COBO, actualmente en preparaciôn), menor longitud del

segundo dedo del pie respecto a la talla del cuerpo y distinta coloraciôn, son las principales

caracteristicas externas que permiten distinguir R. pyrenaica de R. temporaria (Tabla HI).

A ellas cabe añadir las diferencias existentes entre las puestas y las larvas de ambas

especies. Las puestas de la nueva rana presentan menor nümero de huevos, no flotan (en

R. temporaria si) y el diâmetro de los embriones es relativamente mayor respecto a la densa

envoltura gelatinosa. Por otra parte, los renacuajos de R. pyrenaica siempre son mucho

mäs obscuros, musculosos y con aleta caudal distinta (la parte distal suele ser mâs

lobulada). Igualmente, es diferente la etologia reproductora: R. temporaria suele desovar

en aguas quietas, normalmente estancas, mientras R. pyrenaica es mucho mäs selectiva y

suele poner bajo piedras o en pequeñas grietas de arroyos y torrentes. La mayoria de los

rasgos distintivos, mencionados unas lineas mäs arriba, entre el nuevo taxon y R.

temporaria, sirven también para diferenciar R. pyrenaica (especie mucho mäs pequeña) de

R. temporaria parvipalmata Seoane, 1885, y de la rana de Gasser (DuBois, 1982b, 1983).

No obstante, las referidas diferencias serän tratadas ampliamente en otro articulo (Dugois,

OHLER & SERRA-COBO, en preparaciôn).

Las poblaciones simpätricas de R. pyrenaica y R. temporaria tienen gran interés

ecolôgico y biolôgico, pues confirman la diferenciacion interespecifica. Las localidades

donde cohabitan ambas especies, presentan siempre aguas quietas con cierto flujo

renovador que impide el estancamiento hidrico. Si R. pyrenaica y R. temporaria fuesen

morfos distintos de una misma especie adaptados a diferentes condiciones ecolôgicas, la

Source : MNHN, Paris

12 ALYTES 11 (1)

FRANCE

p»

ZARAGOZA

BARCELONA

100Km

Fig. 5. — Distribuciôn de Rana pyrenaica conocida hasta la fecha. La estrella corresponde a la

localidad tipo, mientras los asteriscos indican las localidades donde se ha hallado la especie. El

asterisco mayor agrupa diversas localidades prôximas entre si.

Source : MNHN, Paris

SERRA-COBO 13

variabilidad de los individuos pertenecientes a poblaciones simpätricas seria escasa, como

resultado de fenémenos de hibridacién (habria flujo génico entre poblaciones). No

obstante, en biotopos donde existe simpatria, los individuos adultos, las puestas y las

larvas de ambos taxones continuan diferenciändose claramente, manteniendo las carac-

teristicas propias de las poblaciones alopätricas. Cabe descartar, asi, que se produzca

hibridaciôn entre las dos especies y que la nueva rana sea fruto del acusado polimorfismo

de R. temporaria. Por otra parte, se ha constatado que en las localidades simpätricas, las

larvas de R. temporaria tienden a eclosionar mäs tarde. Asi mismo, las observaciones

realizadas, tanto “in situ” como en laboratorio, parecen indicar que el desarrollo

ontogénico es mâs lento en R. pyrenaica.

La morfologia de R. pyrenaica también es distinta respecto a R. iberica, especie, esta

ültima, que si bien ha sido citada en repetidas ocasiones en los Pirineos (BELLOC, 1893;

PLANTADA Y FONOLLEDA, 1903; SAGARRA, 1916; MALUQUER, 1916; BoscÀ & CASANOVES,

1918; LANTZ, 1927; BECK, 1942, 1943; ANGEL, 1946, 1947; DOTTRENS, 1963; SALVADOR,

1974; FRETEY, 1975; CASTANET, 1978; CASTANET & GUYÉTANT, 1991; BERTRAND &

CROCHET, 1992), los especialistas actuales ponen en duda su existencia en dicha cordillera

(Dusois, 1982b; MaARTiNEZ-RICA, 1983). Teniendo en cuenta la referida opinion,

actualmente no habrian äreas pirenaicas simpâtricas para ambas especies. En comparacion

con R. iberica, R. pyrenaica presenta el hocico menos acuminado, la mancha temporal no tan

conspicua, el timpano mäs pequeño (Tabla II), la coloraciôn de la garganta menos oscura y

sin linea media clara. En relaciôn a la longitud del cuerpo el tercer dedo de la mano es mâs

corto, el antebrazo es mäs largo, el primer y segundo dedos del pie son proporcionalmente de

menor talla, el talôn tiende a ser mayor y la distancia interorbitaria es menor (Tabla Il).

Lamentablemente, los estudios sobre reproducciôn y desarrollo embrionario de R. iberica

son muy escasos (CRESPO & C1, 1973) e impiden un anälisis comparativo.

Respecto a R. dalmatina, el anuro aqui descrito presenta talla mâs pequeña, mayor se-

paraciôn entre el borde anterior de ambos ojos, orificios nasales mâs separados entre si,

mayor distancia entre orificio nasal y borde distal del ojo, diâmetro timpänico mucho me-

nor, mancha temporal menos conspicua, parte inferior del iris mâs clara (ver DuBois,

1984b), hocico menos acuminado, tercer dedo de la mano mäs corto, menor longitud del

muslo, de la tibia, del pie y del talén, el cuarto dedo del pie mâs corto y, en general, aspecto

del cuerpo menos robusto (Tabla IV). Las puestas contienen menor nümero de huevos. Por

otra parte, R. pyrenaica tiene häbitos mâs acuâticos. Hasta la fecha no se ha comprobado la

presencia de R. dalmatina en los Pirineos, si bien se ha citado repetidamente en la literatura

(DesPax, 1941; BECK, 1942, 1943; ANGEL, 1946, 1947; Hvass, 1972).

El hallazgo, en Europa, de una especie vertebrada completamente nueva para la

ciencia con caracteres morfolégicos que permiten distinguirla en el campo (sin necesidad

de recurrir a técnicas mâs complejas de laboratorio), no deja de ser un hecho sorprendente

en nuestros dias.

RESUMEN

El articulo describe un nuevo anuro, Rana pyrenaica, hallado en la zona periférica del

Parque Nacional de Ordesa y Monte Perdido (Huesca, España). Es una rana parda grâcil,

Source : MNHN, Paris

14 ALYTES 11 (1)

ägil, de mediana talla, coloracin normalmente clara, timpano pequeño, orificios nasales

separados, patas relativamente largas, segundo dedo del pie corto, häbitos torrenticolas y

que suele frecuentar aguas claras, frias, oxigenadas y mâs o menos râpidas, refugiändose

bajo piedras o en pequeñas fisuras. En la mayoria de los casos es simpätrica con Euproctus

asper y en ciertas ocasiones con Rana temporaria. Su morfologia y ecologia le distinguen

de R. temporaria, R. iberica y R. dalmatina. Su reparticiôn se circunscribe, hasta la fecha,

a la vertiente española del Pirineo centro-occidental.

AGRADECIMIENTOS

El autor agradece la colaboraciôn prestada por el Dr. E. BALCELLS, tanto a nivel de prospecciones

de campo, como en seguimientos de desarrollo larvario en el laboratorio. Cabe agradecer también al

Prof. A. Dumois sus inestimables consejos e informaciones facilitadas, asi como reconocer el tiempo

dedicado por A. OHLER, J. MAGRANER y R. BOUR a proporcionar material de comparaciôn,

informaciôn bibliogräfica e informätica. Expresar igualmente gratitud, por el soporte logistico de toda

indole, al Laboratoire d'Ecologie de l'Ecole Normale Supérieure (CNRS URA-258) y en particular

al Prof. R. BARBAULT. À B. AMENGUAL, M. Lopez, C. PEDROCCHI, L. MAS y J. E. GOMEZ agradecer

su colaboraciôn en la labor de campo. Finalmente, citar al P. N. de Ordesa y a la Diputaciôn General

de Aragôn los cuales facilitaron el apoyo logistico que permitiô el hallazgo de R. pyrenaica y al

Ministerio de Educacin y Ciencia (becas posdoctorales de formaciôn en el extranjero) y a la

Comunidad Europea (Human Capital and Mobility Programme, Community training project)

quienes financiaron parte de su ulterior estudio. La sede operativa de la investigaciôn tuvo lugar en

el Centre d'Ecologie Montagnarde de Gabas (Université de Bordeaux I).

LITERATURA CITADA

ANDRADA, J., 1980. — Guia de campo de los anfibios y reptiles de la Peninsula Ibérica. Barcelona,

Omega: 1-159.

ANGEL, F., 1946. — Faune de France. 45. Reptiles et amphibiens. Paris, Librairie de la Faculté des

Sciences: 1-204.

— 1947. — Vie et mœurs des amphibiens. Paris, Payot: 1-317.

ARNOLD, E. N. & BURTON, J. A., 1978. — Tous les reptiles et amphibiens d'Europe en couleurs.

Paris-Bruxelles, Elsevier Séquoia: 1-271.

BALCELLS, E., 1956. — Estudio morfolégico, biolôgico y ecolôgico de Rana temporaria, L. Publ. Ins.

Biol. apl., 1-121.

Beck, P., 1942. — Quelques remarques sur la faune batrachologique du département des

Hautes-Pyrénées. Bull. Soc. zool. Fr., 67: 85-87.

En 1943. — Note préliminaire sur la faune herpétologique des Hautes-Pyrénées. Bull. Sect. sci. Soc.

Acad. Hautes Pyrénées, 1: 48-57.

BEGON, H., HARPER, J. L. & TOWNSEND, C. R., 1988. — Ecology: individuals, populations and

il Barcelona, Omega: 1-886.

BELLOC, E., 1893. — Utilisation des cuvettes lacustres pyrénéennes pour la pisciculture. C. r. Assoc.

Jr. Av. Sci, 21 (2): 516-522.

BERTRAND, À. & CROCHET, P.-A., 1992. — Amphibiens et reptiles d'Ariège. Clermont, Association des

Naturalistes de l'Ariège: 1-137.

Bosc, E. & CASANOVES, E., 1918. — Una nueva forma de anfibio urodelo (Molge Bolivari). Bol. r.

Soc. esp. Hist. nat., 18: 58-61.

BOULENGER, G. A., 1879. — Étude sur les Grenouilles rousses Ranae temporariae et description

d'espèces nouvelles ou méconnues. Bull. Soc. zool. Fr., 4: 158-193.

- 1898. — The tailles batrachians of Europe. Part II. London, Ray Society: 211-376.

- 1910. — Les batraciens, et principalement ceux d'Europe. Paris, Doin: 1-305.

Source : MNHN, Paris

SERRA-COBO 15

CASTANET, J., (ed.), 1978. — Atlas préliminaire des reptiles et amphibiens de France. Montpellier,

Société Herpétologique de France: 1-137.

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

Paris, Société Herpétologique de France: 1-191.

Crespo, E. G. & Cri, J. M., 1973. — El ciclo espermatogenético potencialmente continuo de Rana

iberica en Portugal y el interés de su estudio en äreas pirenaicas de simpatria con Rana

temporaria. Pirineos, 110: 47-49.

DELAUGERRE, M. & DuBois, A., 1985. — La variation géographique et la variabilité intrapopula-

tionnelle chez Phyllodactylus europaeus (Reptilia, Sauria, Gekkonidae). Bull. Mus. nan. Hist.

nat. (4), 7 (A): 709-736.

Despax, R., 1941. — Notes batrachologiques. IV. Présence, dans les Pyrénées, de Rana agilis et de

la forme typica de Salamandra maculosa. Bull. Soc. Hist. nat. Toulouse, 76: 91-92.

DoTTRENS, E., 1963. — Batraciens et reptiles d'Europe. Neuchâtel, Delachaux & Niestlé: 1-261.

Duois, A., 1982a. — A propos de l’article de G. H. Parent sur la répartition des amphibiens et

reptiles en France. A/ytes, 1: 12-15.

À, 1982b. — Notes sur les grenouilles brunes (groupe de Rana temporaria Linné, 1758). I.

Introduction. Alytes, 1: 56-70.

es 1983. — Notes sur les grenouilles brunes (groupe de Rana temporaria Linné, 1758). II. Les

grenouilles du Mont Canigou (Pyrénées Orientales). Alytes, 2: 19-26.

—. 1984a. — Sample-size constraints in the use of the nonparametric Mann-Whitney U test for the

comparison of two independent samples: consequences in anuran amphibians systematics.

Alytes, 3: 20-24.

eu 1984b. — Notes sur les grenouilles brunes (groupe de Rana temporaria Linné, 1758). III. Un

critère méconnu pour distinguer Rana dalmatina de Rana temporaria. Alytes, 3: 117-124.

ESTEBAN, M., 1990. — Evoluciôn del género Rana en la Peninsula Ibérica: estudio de la variabilidad

morfolôgica y genética del complejo Rana temporaria L. Tesis Doctoral, Dep. Biologia Animal

1, Fac. Ciencias Biolôgicas, Universidad Complutense Madrid: 1-211, anexo.

FRETEY, J., 1975. — Guide des reptiles et batraciens de France. Paris, Hatier: 1-238.

HALDANE, J. B. S., 1955. — The measurement of variation. Evolution, 9: 484.

Hvass, H., 1972. — Reptiles and amphibians in colour. London, Blandford Press: 1-153.

Lanrz, L. A., 1927. — Quelques observations nouvelles sur l'herpétologie des Pyrénées centrales.

(Suite). Rev. Hist. nat. appl., 8: 54-61.

MALUQUER, J., 1916. — Noves herpetolôgiques. Bull. Inst. cat. Hist. nat., 13: 111-119.

MARTÎNEZ-RICA, J. P., 1983. — Atlas herpetolôgico del Pirineo. Munibe, 35: 51-80.

MARTINEZ-RICA, J. P. & REINÉ-VINALES, A., 1988. — Altitudinal distribution of amphibians and

reptiles in the Spanish Pyrenees. Pirineos, 131: 57-82.

MERTENS, R. & WERMUTH, H., 1960. — Die Amphibien und Reptilien Europas. (Dritte Liste, nach dem

Stand vom 1. Januar 1960). Frankfurt, Kramer: 1-264.

OuLer, À. & KAzADI, M., 1990. — Description d’une nouvelle espèce du genre Aubria Boulenger,

1917 (Amphibiens, Anoures) et redescription du type d’Aubria subsigillata (A. Duméril, 1856).

Alytes, 8: 25-40.

PaReNT, G. H., 1981. — Matériaux pour une herpétofaune de l'Europe occidentale. Contribution à

la révision chorologique de l’herpétofaune de la France et du Bénélux. Bull. Soc. linn. Lyon, 50:

86-111.

PLANTADA Y FONOLLEDA, V., 1903. — Vertebrats del Vallès. Catäleg dels observats en aquesta

comarca. Bull. Inst. cat. Hist. nat., 3: 111-118.

SAGARRA, I. DE, 1916. — Donatius per al Museu. Butll. Inst. catal. Hist. nat., 13: 108.

SALVADOR, A., 1974. — Guia de los anfibios y reptiles esparñoles. Madrid, Instituto Nacional para la

Conservaciôn de la Naturaleza: 1-282.

SIEGEL, S., 1985. — Estadistica no paramétrica aplicada a las ciencias de la conducta. Barcelona, Trillas:

1-344.

Corresponding editor: Alain DuBois.

Source : MNHN, Paris

Alytes, 1993, 11 (1): 16.

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Source : MNHN, Paris

Alytes, 1993, 11 (1): 17-24. 17

Growth and maturity of brown frogs,

Rana arvalis and Rana temporaria,

in central Poland

Leszek BERGER & Mariusz RYBACKI

Research Center for Agricultural and Forest Environment,

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

Growth of two brown frog species, Rana arvalis and Rana temporaria, was

studied in nature and in captivity. In both species, growth was fastest in the

first two years, maturity was reached in the second year, and gametes were

released after the second hibernation. Individuals of Rana arvalis matured

earlier than those of Rana temporaria, and in some males of Rana arvalis the

sexual features were visible already before the first hibemation.

INTRODUCTION

The two species of brown frogs Rana temporaria and Rana arvalis belong to the

common amphibians in central Europe. Data on their growth and sexual maturity are

numerous, but opinions on this subject are controversial. KRIVOSHEYEV, OPENKO &

SHABANOVA (1960) suggested that they grow most rapidly in the third (Rana arvalis) or in

the fourth year (Rana temporaria), that is, in the year of reaching maturity. Most other

authors state that they grow very quickly in the first and in the second year, whereas

growth decreases after maturation (GIsLÉN & KAURI, 1959; KLEINENBERG & SMIRINA, 1969;

TOMASIK, 1969; VAN GELDER & OOMEN, 1970; LOMAN, 1978; GIBBONS & McCARTHY, 1984;

CHMELEVSKAJA, 1985; RysEr, 1988; among others). They report that the frogs can reach

maturity between the second and the fourth year of their lives and release gametes for the

first time after two to four hibernations.

In 1965 we found a surprisingly small adult Rana arvalis male (37.5 mm, snout-vent

length) in a breeding aggregation, and in 1977 we caught a very small female of Rana

arvalis (39.5 mm) in amplexus, which subsequently laid 404 eggs in an aquarium. These

data prompted us to observe brown frogs more precisely.

In the present paper we review available data and compare them with our results,

which suggest that growth and sexual maturity in these frog species depend not only on

geographical distribution, but also on environmental conditions.

Source : MNHN, Paris

18 ALYTES 11 (1)

MATERIAL AND METHODS

This paper on brown frogs arose as a by-product during the study of European water

frogs (BERGER & BERGER, 1992). The observations are not homogeneous, because they

were carried out in different years (1963-1989), on different populations and in various

environments. The study areas are situated within the drainage of the Warta river in

central Wielkopolska. This region forms a rather homogeneous geographical and climatic

unit (BARTKOWSKI, 1970).

Newly metamorphosed froglets of both species, which are the most common

amphibians in the region (BERGER, 1987), were gathered near water bodies in which they

lived as tadpoles. Individuals were killed and preserved in 3% formaldehyde. They

originated from the vicinity of Poznañ and the Biological Field Station of our Institute in

Turew (about 40 km south of Poznañ), and from Jask6iki near Ostréw Wielkopolski

(about 100 km south-east of Poznan). In two localities the froglets were gathered every two

or three weeks, in the others they were caught during researches of water frogs. We were

not able to gather any data on older brown frogs in nature, because we did not mark them

and it was not possible to estimate their age by their body length.

During some years froglets were reared in captivity. The frogs were measured with an

accuracy of 0.1 mm (snout-vent length) after capture and later on every spring until their

death; some were measured twice in a year (fig. 2). We marked them collectively or

individually by toe-clipping. In 1986, about 20,000 tadpoles of Rana arvalis were put into

10 basins which contained 200-350 1 of water with plants and mud from a pond. The basins

were located in a fenced enclosure of 10 X 10 m in a garden. These tadpoles and frogs

never received any special food during their life-time. It was much easier to keep Rana

arvalis than Rana temporaria. Froglets of the former species were more vital and usually

survived some years in captivity, whereas those of the latter usually perished before or

during the first hibernation.

RESULTS

GROWTH OF FROGS IN NATURE

After metamorphosis, the body length of Rana arvalis and Rana temporaria froglets

was rather similar and averaged about 14-16 mm in the studied localities (fig. 1). In

autumn, individuals of Rana temporaria were much larger than those of Rana arvalis: the

first reached about 35 mm (range 24-42 mm), and the second about 24 mm (range 16-32

mm) in body length. In other localities their body length in autumn was as follows: in

Stawa Wielkopolska near Poznañ (Sept. 14, 1962), Rana arvalis measured 17.2-30.2 mm

(mean 22.45, N = 53), and Rana temporaria 17.8-32.0 mm (mean 24.10, N = 19); in

Zbechy near Turew (Sept. 7, 1978), Rana arvalis measured 22.7-34.0 mm (mean 28.10, N

= 17), and Rana temporaria 24.3-42.5 mm (mean 34.20, N = 28).

Source : MNHN, Paris

BERGER & RYBACKI 19

4 À L B

30} L

20] L

10 L

ñ A à Es à

J À s nn A SSES

month

Fig. 1. — Growth of brown frogs in nature. (A) Rogaczewo near Turew, 1978: Rt, Rana temporaria

(N = 59); Ra, Rana arvalis (N = 130); (B) Poznañ-Naramowice, 1963: Rana arvalis (N = 512).

Mean values of body length are joined by lines, extreme sizes of froglets are indicated by vertical

bars.

Froglets after first hibernation were collected in two localities. In Rogaczewo (May

5, 1978), we gathered 66 froglets of Rana arvalis with body length of 18.7-31.0 mm (mean

25.12; see fig 1), and in Turew park (April 20, 1980), we caught froglets of both species:

Rana arvalis measured 29.0-40.0 mm in body length (mean 35.70, N = 3), and Rana

temporaria 36.0-52.0 mm (mean 44.40, N = 65).

GROWTH OF FROGS IN CAPTIVITY

Froglets from nature (fig. 2)

The froglets from Jaskôïki which were caught in autumn (Sept. 5, 1988) were marked

individually, and the others which were caught during metamorphosis were marked

collectively. The following year, initially smaller individuals grew more rapidly than larger

ones (Table I).

In summer after the first hibernation the sexual features appeared in all surviving

frogs of both species, and after the second hibernation all frogs that were left released

gametes. The three females of Rana arvalis from Poznañ-Junikowo measured 41.5, 42.6

and 46.5 mm in body length and laid 141, 168 and 728 eggs respectively; those from

Zbechy measured 44.0, 49.5, 52.0 and 58.0 mm in body length and laid 692, 1077, 1434

and 2055 eggs respectively.

Source : MNHN, Paris

20 ALYTES 11 (1)

80}

R.arv. R.tem.

70f

years

Fig. 2. — Growth of brown frogs in captivity.

R. arv., Rana arvalis from: (a) Zbechy near Turew, 1978 (N = 26); only two individuals survived

the fifth hibernation; (b) Poznañ-Junikowo, 1970 (N = 32); only two frogs survived the

third hibernation; (c) Jaskôiki near Ostrow Wielkopolski, 1988 (one female only).

R. tem., Rana temporaria from: (a) Jaskôki near Ostrôw Wielkopolski, 1988 (N = 20); 11 frogs

survived the first hibernation; (b) Poznañ, 1974 (N = 30); only five frogs survived the

second hibernation.

Individuals of Rana arvalis from Poznañ-Junikowo and Rana temporaria from Poznañ were

measured twice in the second season. Individuals from Jaskôïki were removed from the

rearing area on July 30.

Among Rana temporaria from Poznañ only one female of 63.0 mm body length laid

972 eggs spontaneously with the only male which survived the winter. The other three

females with body length of 56-58 mm were caught and after dissection we found mature

ova in their ovisacs which were ready to be shed (627, 730 and 821 eggs).

The mean body length of Rana arvalis from Poznañ-Junikowo reached 29.9 mm after

the first hibernation, 42.3 mm in July, and 44.0 mm in the spring of the next year; this

Source : MNHN, Paris

BERGER & RYBACKI 21

Table I. — Growth of Rana temporaria individuals in captivity. The frogs are arranged

according to body length at the day of capture in Jask6iki near Ostréw Wielkopolski.

Females Males

No. Body length Growth No. Body length Growth

offrog Sept. 51988 July 30 1989 mm % offrog Sept. 51988 July 30 1989 mm %

16 272 650 37.8 139 14 25.0 550 300 120

20 218 760 482 174 21 252 630 370 147

2 28.0 710 430 154 27 37.8 770 392 104

15 32.0 710 390 122 13 39.5 610 27.5 70

24 340 680 340 100 23 40.0 660 260 65

26 363 715 352 97 le. ni r

_ Means 309 704 39.5 131 35.5 656 319 101

means that within the first period they grew about 12.4 mm and in the second only 1.7 mm.

The Rana temporaria from Poznañ grew in a similar way. They averaged 37.0 mm in body

length after the first hibernation, 51.0 mm on June 12, and 58.5 mm in the spring of the

next year. Within the first period they grew about 22.6 mm and in the second only

14.0 mm.

Froglets from tadpoles reared in captivity

On April 4, 1986, we brought 23 clumps of spawn of Rana arvalis from

Poznañ-Naramowice pond, which contained 1171-2563 eggs (mean 1737). Sixteen clumps

were kept in containers outdoors, the others in the laboratory. During their development

the temperature decreased to 5°C below zero at night. Percentage of embryos which

reached tadpole stage was as follows: in the laboratory 78.6-97.2 % (mean 88.2 %), and

outdoors 8.5-88.1 % (mean 38.1 %). AII tadpoles were mixed and distributed into basins

in the enclosure in Jaskolki in which there were no brown frogs before. The first

metamorphosed individuals appeared on June 2, but in July tadpoles could still be seen in

basins. At the end of September we caught three froglets (40-45 mm in body length) with

typical male features (blue colour and nuptial pads).

The following year, the frogs formed a breeding aggregation in the largest basin, but

by the end of April there were no eggs. In the basin we found 14 adult Rana arvalis males

with 40.0-48.6 mm in body length (mean 44.8), but there were no females. Juvenile

individuals were also found in the rearing area: 23 females with body length of 19.0-42.8

mm and 14 males with body length of 15.5-39.5 mm. The ovaries of these females were in

bud stage, and in the testes of some juvenile males (33.0-39.5 mm in body length) moving

spermatozoa were present.

Source : MNHN, Paris

2 ALYTES 11 (1)

DISCUSSION

The period of metamorphosis of tadpoles of Rana arvalis and Rana temporaria, which

are explosive early spring breeders, is usually very short (VAN GELDER & OOMEN, 1970;

Ryszkowski & TRUSZKOWSKI, 1975). This suggests that individuals of these species should

form compact age groups (KRIVOSHEYEV, OPENKO & SHABANOVA, 1960; GAJZAUSKIENE,

1966; HEUSSER, 1970). However, their growth rates in nature (VAN GELDER & OOMEN,

1970; LomAN, 1978; CHMELEVSKAIA, 1985; fig. 1) and in captivity (SMIRINA, 1980, 1986) are

quite variable and opinions on their age and sexual maturity are often contradictory.

The data of VAN GELDER & OOMEN (1970) and LoMaAN (1978), and our results suggest

that in nature only the yearlings can be identified by body length, whereas the division of

older frogs into age classes is impossible by body length alone. KLEINENBERG & SMIRINA’S

(1969) skeletochronological studies corroborate this conclusion. They found that individ-

uals of Rana temporaria caught near Moscow in June formed three size classes, but only

one class which contained the smallest individuals (after the first hibernation) was

homogeneous, whereas the other two classes contained individuals which were 2-9 years

old. Other authors (MINA, 1974; ISHCHENKO & LEDENTZOV, 1985, 1986) obtained similar

results.

Our observations show that individuals of both species grew most rapidly at the

beginning of the second season (May-June; fig. 2, Table I), or before reaching maturity.

Within yearlings of Rana temporaria in the Wielkopolska region, however, we observed

considerable differentiation in autumn. Their mean body length ranged from 24.10 mm in

Slawa Wielkopolska to 44.40 mm in Turew (see SMIRINA, KLEVEZAL & BERGER, 1986). The

conditions in our rearing area, in which the yearlings reached intermediate body lengths

(fig. 1-2), are therefore likely to have been close to those in nature. To this extent, our

observations on frogs in captivity can be compared with those obtained by others in wild.

Our data with regard to sexual maturity are also not always in agreement with other

authors. Opinions on this problem are differentiated and data which are reported by many

authors refer mostly to Rana temporaria.

According to GiBBoNs & McCaARTHY (1984), in west Ireland most of the individuals

in the breeding aggregations of Rana temporaria were two years old frogs (84 % of the

males and 52 % of the females). In Britain also numerous such frogs were found in the

breeding time (BEEBEE, 1980; CookE, 1981). However, near Moscow in Russia (CHME-

LEVSKAJA, 1985), near Berne in Switzerland (RYsER, 1986, 1988) and in southern Sweden

(Lomaw, 1976, 1978) so young individuals were extremely rarely present in breeding

aggregations. Some others suggest that individuals of this species reach maturity in the

third (HeUSsER, 1970; Mina, 1974) or in the fourth year (KLEINENBERG & SMIRINA, 1969).

In our rearing (fig. 2, Table I), however, Rana temporaria reached maturity during the

second season and after the second hibernation released gametes as two years old frogs.

These results corroborate ToMasik’s (1969) opinion on Polish Rana temporaria.

The data on Rana arvalis are very scanty. ISHCHENKO & LEDENTZOV (1985, 1986)

found that in the breeding populations near Svierdlovsk in Russia only few individuals

Source : MNHN, Paris

BERGER & RYBACKI 23

(about 1 %) were two years old. Tomasik (1969) also supposes that in Poland Rana arvalis

can reach maturity in the second year. According to our results all individuals of Rana

arvalis in rearing reached maturity in the second season. Other authors suggested,

however, that individuals of this species mature later: in southern Sweden in the third or

in the fourth year (LOMAN, 1976, 1978), and in the Netherlands in the third year (VAN

GELDER & OOMEN, 1970). The latter authors report, however, that at the end of the second

season most frogs had more than 40 mm in body length and at that time it was possible

to distinguish males from females. This statement suggests that such frogs were mature and

took part in the breeding aggregations as two years old animals.

The growth of Rana arvalis and Rana temporaria froglets in the first year is highly

differentiated not only in Sweden (LOMAN, 1976, 1978), but also in the Netherlands (VAN

GELDER & OOMEN, 1970), in Russia (CHMELEVSKAYA, 1985) and in Poland. Near Poznañ

in every locality and nearly in every sample, including froglets after the first hibernation,

there were very large and very small individuals. The situation was, however, reversed in

the second year of their life: the growth of small froglets was much faster than that of large

ones (Table I; see also VAN GELDER & OOMEN, 1970). Similar observations which are

consistent with the “compensation growth phenomenon” in animals (MINA & KLEVEZAL,

1976) have been observed in some other anuran species (JAMESON, 1956; BERGER, 1970;

PLyrycz & BiGaï, 1985; BERGER & RyBACKI, unpublished data), which suggests that this

phenomenon may be rather common in Amphibia.

ACKNOWLEDGMENTS

The authors are grateful to Dr. H. HoTz for reading the manuscript and profitable discussion on

some problems, to Dr. G. GOLLMANN for his valuable suggestions in preparing the manuscript to

press, and to Zygmunt PNIEWSKkI for his assistance in the laboratory work.

LITERATURE CITED

BARTKOWSKI, T., 1970. — Wielkopolska i Srodkowe Nadodrze. [In Polish]. Warszawa, PWN: 1-384.

BEEBEE, T. J. C., 1980. — Amphibian growth rates. Brit. J. Herpet., 6: 107.

BERGER, L., 1970. — Some characteristics of the crosses within Rana esculenta complex in postlarval

development. Ann. Zool., 27: 373-416.

S 1987. — Impact of agriculture intensification on Amphibia. /n: J. J. VAN GELDER, H. STRUBOSCH

and P. J. M. BERGERS (eds.), Proc. Fourth Ord. Gen. Meet. SEH, Nijmegen 1987: 79-82.

BERGER, L. & BERGER, W. A., 1992. — Progeny of water frog populations in central Poland.

Amphibia-Reptilia, 13: 135-146.

CHmeLEvskAIA, L. V., 1985. — Individualnoje mecenie travjanych lagusek (Rana temporaria), rost i

ispolzovanie teritorii. [In Russian]. Voprosy Gerpetologi. VI Vsesoj. Gerpet. Konjer., Taskent

18-20 IX 1985: 219-220.

CookE, A. S., 1981. — Amphibian growth rates. Brit. J. Herpet., 6: 179-180.

GAIZAUSKIENE, L. L., 1966. — Materialy po vozrostnomu sostavu populacii travianoj ljaguski (Rana

temporaria) Litovskoj SSR i jejo morfologii. [In Russian]. Trudy Akad. Nauk Litovskoj SSR,

(5), 1 (39): 91-96.

Source : MNHN, Paris

24 ALYTES 11 (1)

Gros, M. M. & McCarTHy, T. K., 1984. — Growth, maturation and survival of frogs Rana

temporaria L. Holarctic Ecology, 7: 419-427.

Giscén, T. & KAURI, H., 1959. — Zoogeography of the Swedish amphibians and reptiles with notes

on their growth and ecology. Acta Vertebratica, 1: 191-397.

Heusser, H., 1970. — Ansiedlung, Ortstreue und Populationsdynamik des Grasfrosches (Rana

temporaria) an einem Gartenweïher. Salamandra, 6: 80-87.

ISHCHENKO, V. G. & LEDENTZOV, A. V., 1985. — Vnutripopulacjonnaja izmencivost vozrastnoj

truktüry u ostromordoj laguski (Rana arvalis). [In Russian]. Voprosy Gerpetologi. VI Vsesoj.

Gerpet. Konfer., Taskent 18-20 IX 1985: 89-90.

1986. — Dynamics of age structure in a population of the moor frog, Rana arvalis Nilss. In: Z.

Roëex (ed.), Studies in herpetology, Prague: 503-506.

JAMESON, D. J., 1956. — Growth, dispersal and survival of the Pacific tree frog. Copeia, 1956: 25-29.

KLEINENBERG, S. & SMIRINA, E., 1969. — A contribution to the method of age determination in

amphibians. [In Russian]. Zoo!. Zhur., 48: 1090-1094.

KRIVOSHEYEV, V. G., OPENKO, Z. M. & SHABANOVA, E. V., 1960. — Contribution to the biology of

Rana temporaria L. and R. terrestris Andrz. [In Russian]. Zool. Zhur., 39: 1201-1207.

LoMaAN, J., 1976. — Growth in Rana temporaria and R. arvalis. Norw. J. Zool., 24: 232-233.

--- 1978. — Growth of brown frogs Rana arvalis Nilsson and R. temporaria L. in South Sweden.

Ekol. Pol., 26: 287-296.

Mina, M. V., 1974. — Age organization of breeding group of Rana temporaria in a small pond in

the Moscow district. [In Russian]. Zoof. Zhur., 53: 1826-1832.

Mina, M. V. & KLEVEZAL, G. A., 1976. — Rost zivotnych. [In Russian]. Moskow, Ser. Problemy

biologii rozvitja, Nauka: 1-291.

Pryrycz, B. & BiGas, J., 1984. — Preliminary studies on the growth and movements of the

yellow-bellied toad, Bombina variegata (Anura: Discoglossidac). Amphibia-Reptilia, 5: 81-86.

RYSsER, J., 1986. — Altersstruktur, Geschlechterverhältnis und Dynamik einer Grasfrosch-Population

(Rana temporaria L.) aus der Schweiz. Zool. Anz., 217: 234-251.

_—— 1988. — Determination of growth and maturation in the common frog, Rana temporaria, by

skeletochronology. J. Zool., Lond., 216: 673-685.

Ryszkowski, L. & TRUSZKOWSkI, J., 1975. — Estimation of the abundance and biomass of

transformed amphibians in a field pond. Bull. Acad. pol. Sci., ser. Biol., 23: 109-113.

SMIRINA, E. M., 1980. - On growth rate and survival of common frogs (Rana temporaria) during the

first years of life. [In Russian]. Zoo!. Zhur., 59: 1831-1840.

_—— 1986. — Some results of the studies of growth in Anura. Jn: Z. ROëEK (ed.), Studies in

herpetology, Prague: 263-266.

SmiriNA, E. M., KLEVEZAL, G. A. & BERGER, L., 1986. — Experimental investigation of the annual

layer formation in bones of amphibians. [In Russian]. Zool. Zhur., 65: 1526-1534.

Tomasik, L., 1969. — The body sizes of adult gras frog (Rana temporaria temporaria L.) and field

frog (R. arvalis arvalis Nilss.). [In Polish]. Przeglad Zool., 13: 94-98.

VAN GELDER, J. J. & OoMEN, H. C. J., 1970. — Ecological observations on Amphibia in the

Netherlands. I. Rana arvalis Nilsson: reproduction, growth, migration and population

fluctuations. Netherl. J. Zool., 20: 238-252.

Corresponding editor: Günter GOLLMANN.

Source : MNHN, Paris

Alytes, 1993, 11 (1): 25-35. 25

Skin morphology in larval,

paedomorphic and metamorphosed

Alpine newts, Triturus alpestris apuanus

Franco ANDREONE*, Bruno DORE**,

Pasquale Usai** & Adriana PARANINFO**

* Museo regionale di Scienze naturali, Sezione di Zoologia,

via Giovanni Giolitti 36, 10123 Torino, Italy

** Dipartimento di Biologia animale, Università degli Studi di Torino,

via Accademia Albertina 17, 10123 Torino, Italy

Histological data on the skin of larval, paedomorphic and metamorphosed

specimens of the Alpine newt, Triturus alpestris apuanus, have been

and compared with natural history infonmation. The skin of early larvae is

composed of a few cell layers and contains Leydig cells, but lacks exo-

epithelial dermal glands, which appear later during the premetamorphic stage.

In metamorphosed news the skin is typically multilayered and cornified, with

abundant mucous and serous glands. The situation in paedomorphics, which

are branchiate and live in water, is variable: in some cases (corresponding to

immature “giant larvae”’), the skin presents larval aspects (Leydig cells are

present), while in others (usually sexually active and paedogenetic individuals)

it is similar to that of metamorphosed newts. Thus metamorphosis in

paedomorphic newts is not abandoned, but only delayed. The result is a

mosaic of larval and metamorphosed characteristics, which can be related to

the highly plastic ecology of these newts.

INTRODUCTION

The typical life cycle of urodeles is amphibious, with aquatic larvae and metamor-

phosed individuals living in a terrestrial habitat. Nevertheless, paedomorphism (sensu

Dusois, 1987, or neoteny, sensu BREUIL, in press, understood as the retardation in the

development of somatic and/or gonadal features or as the achievement of the sexual

maturity while retaining larval-juvenile characters) often occurs. Paedomorphic newts have

external gills, reach a greater size than larvae, and are totally aquatic. The causes of

paedomorphism are not well known, although ecological constraints are often invoked

(WizBur & CoLLins, 1973). Among the European urodeles, the Alpine newt, Triturus

alpestris (Laurenti, 1768), throughout its range and involving most subspecies, shows a

high incidence of paedomorphism. Recently, as a part of a wider study of paedomorphism

in Italian Alpine newt populations (ANDREONE & Dore, 1991), some preliminary

hypotheses on the ecological basis of the phenomenon were formulated; moreover,

Source : MNHN, Paris

26 ALYTES 11 (1)

information has been gathered on variations in the histology of thyroid glands and gonads

of paedomorphic and metamorphosed individuals (ANDREONE, DORE & Usa, 1991). In the

present paper, as a complement, data on skin morphology during the life cycle are

presented, taking into account that the amphibian integument witnesses the changes from

an aquatic to a terrestrial habitat, acting as a medium for ionic exchange (DUELLMAN &

TRUEB, 1986; Lopi et al., in press).

MATERIAL AND METHODS

The population studied (which belongs to the subspecies Triturus alpestris apuanus),

inhabits an artificial temporary pond located between the towns of Murazzano and

Bossolasco (southern Piedmont, north-western Italy), at an altitude of about 700 m, with

a surface area of about 100 m? and a maximum depth of 1.5 m. No aquatic vegetation

(except Chara algae) is present, while other amphibians living there are the common frog

(Rana temporaria) and the common toad (Bufo bufo). The climate of the area is

Mediterranean (MENNELLA, 1967), with maximum rainfall in May and November. In this

site a remarkable number of aquatic newts are found in the water throughout the year,

even during non-reproductive periods.

Each month, from January 1988 to December 1989, several aquatic newts were caught

by hand-netting (ANDREONE & DORE, 1992). The following categories were examined:

(1) Larvae, ie. branchiate individuals with a total length less than 40 mm, having a

brownish-greenish back, scattered darker spots, and a whitish belly (fig. 1). Sometimes two

age cohorts of larvae may co-exist in the same pond, resulting from two egg depositions

at different times during a year (ANDREONE & DORE, 1992).

(2) Metamorphosed, i.e. sexually mature newts. Such individuals court during the

breeding season, when they display secondary sexual characters (SSC), such as lateral

white stripes, extended dorso-caudal crests and swollen cloacae in males, and a swollen

abdomen and turgid cloaca in females (fig. 2). Although the metamorphosed aquatic

population is mainly represented by adults, juvenile specimens also are found; such

individuals are smaller than sexually mature individuals, do not show SSC and, in some

cases, have a light dorsal line.

(3) Paedomorphics (sensu lato), i.e. individuals with external gills (more or less

developed), larger body size than larvae and with a yellow-orange belly. We refer to

“neoteny” or “paedomorphism” according to KOLLMANN (1884a-c) and Dugois (1987):

“paedomorphic” to indicate branchiate newts without SSC and lacking the courtship

behaviour, corresponding to the “partial neotenic” (sensu BREUIL, in press) (fig. 3), and

“‘paedogenetic” (= “total neotenic”) to indicate the branchiate newts displaying courtship

behaviours and SSC (fig. 4).

After capture, newts were anaesthetized by immersion in a 0.5 % MS 222 Sandoz

solution. Some individuals were dissected to verify their gonadal status, and a portion of

their dorsal and ventral skin was cut off, fixed in cold buffered formalin, dehydrated, and

infiltrated overnight with a glycol metacrylate monomer (“Technovit 7100” Kulzer) in a

Source : MNHN, Paris

Fig. 1. — Larva of Alpine newt (Triturus alpestris apuanus),

characterized by external gills, continuous caudal crests, and

a light brownish scattered back.

Fig. 3. — Paedomorphic (partial neotenic and sexually immature

giant larva) Alpine newt, characterized by a general larval

aspect, undeveloped gonads and by the absence of evident

SsC.

Fig. 2. — Metamorphosed male of Alpine newt during the

breeding season, with well developed secondary sexual char-

acters (SSC), e.g. dorsal and caudal crests, white lateral

stripe, blue back coloration and swollen cloaca.

Fig. 4. — Paedogenetic (totally neotenic) adult male of Alpine

newt, with external gills and well developed secondary sexual

characters.

OANINVUVA # IVSN ‘HO ‘INOHHANY

Source : MNHN, Paris

28 ALYTES 11 (1)

ice bath. Polymerization was carried out at about 6-10°C to preserve enzymatic activities

(Dore & Usai, 1986; ANDREONE & DORE, 1992). Skin sections of 2 um were stained with

acid fuchsin-toluidine blue (DOUGHERTY, 1981), and with Burstone direct coupling method

of substituted naphtols contrasted with methyl green for Alkaline Phosphatase (APH)

activity (MaAZzi, 1977; LOYDA, GossRAU & SCHLEBER, 1979). APH activity may be

correlated with transcutaneous ionic transport (see Lopi et al., in press). Controls were

carried out in the absence of substratum.

RESULTS

LARVAE

The larval skin is composed of a few cell layers (three or four) lying on a dense layer

of dermis (“basement lamella”, Fox, 1977). In all specimens examined the epithelial basal

cells have an elongated nucleus, are irregular in shape, and penetrate with their apical part

among the overhanging Leydig cells. The latter cells, well known in larvae and neotenic

urodeles (see HAY, 1961; RAFFAELLI, 1989) have a poorly known function, but they

probably are glandular (Fox, 1988) and secrete mucus into subsurface extracellular

compartments of the epidermis (DUELLMAN & TRUEB, 1986). Leydig cells are large and

rounded, with a clear cytoplasm, PAS-positive granules and prominent nucleus, and they

occupy the entire epidermal thickness and press against the common epithelial cells with

their convex surface. We did not observe a general uniformity in the morphology and

disposition of cells in the skin of different larvae. However, in early larvae the Leydig cells

are disposed in two layers (basal and sub-apical) and contain abundant PAS-positive

granules, while in older larvae they form only one layer and have a scarce granulation. The

skin surface is not cornified, and is composed of one or two layers of flattened epithelial

cells, while isolated supporting cells are interposed among the underlying Leydig cells.

Neuromasts of the lateral line are constantly present. In early larvae the dermis is thin and

usually not invaded by glandular (serous and mucous) elements. In some large

premetamorphic larvae or in larvae becoming paedomorphic (see later), precocious gland

buds begin to differentiate (fig. 6). Low APH activity is seen, and — when visible — it is

localized in the most external epithelial cells, whereas it is totally lacking in the dermis,

except in the blood vessels of the subcutaneous layer.

METAMORPHOSED SPECIMENS

The epidermis of metamorphosed Alpine newts (either sexually mature or newly

metamorphosed) is similar to that described for Triturus carnifex (LoDi, 1968; Loni &

Bant, 1971), being about four to six cells thick. Migration of deep cells occurs towards the

surface. Cells of the basal or germinative layer are large and irregularly shaped, generally

columnar or cuboidal, and arrayed in a low palisade. Mitoses are often visible in the

germinative layer, and melanophores are commoner in the dorsal dermis than elsewhere,

but some scattered melanophores may be found in the dorsal epidermis (DUELLMAN &

TRUEB, 1986). Flask cells (involved in the sodium transport; see Fox, 1986a-b; ZACCONE

Source : MNHN, Paris

ANDREONE, DORE, USAI & PARANINFO 29

à LEON

Fig. 5. — Histological structure of the skin in an

Alpine newt larva. Two layers of Leydig

cells with PAS-positive granules are evi-

dent, together with plate superficial cells.

Toluidine blue coloration. The bar corre-

sponds to 10 pm.

Fig. 6. — Late larval skin, with Leydig cells, with

a dermal gland bud. Toluidine blue color-

ation. The bar corresponds to 10 um.

Fig. 7. — Histological structure of the skin in a

metamorphosed Alpine newt. Great exo-

epithelial glands are visible, together with

several cell layers and a cornified surface.

Toluidine blue coloration. The bar corre-

sponds to 10 pm.

Fig. 8. — Metamorphosed skin, with flask cells

shown by means of APH and Burstone’s

method of substituted naphtols contrasted

with methyl green. The bar corresponds to

10 pm.

Fig. 9. — Structure of the skin in a paedomor-

phic immature Alpine newt. Empty Leydig

cells (disposed in a single layer) with scarce

PAS-positive granules are visible. Mitoses

are evident in the cells among Leydig cells.

Toluidine blue coloration. The bar corre-

sponds to 10 um.

Fig. 10. — Skin of a paedogenetic Alpine newt:

the structure and morphology — even with

a reduced superficial cornification — is

similar to the metamorphosed one. Tolui-

dine blue coloration. The bar corresponds

to 10 pm.

Source : MNHN, Paris

30 ALYTES 11 (1)

et al., 1986) are distributed more or less regularly in the subcorneous layer (fig. 7). The skin

surface is cornified, with protruded warts, two or three flattened cells thick. Highly

developed mucous and serous glands are always evident in the spongy thick dermis just

below the epithelium either in the ventral or in the dorsal skin. In metamorphosed newts,

APH activity is similar in the dorsal and in the ventral skin, varying from low to moderate

in the cells of corneous and subcorneous layers, but lacking in the deepest layers. Flask

cells usually show an intense APH activity (fig. 8) in their neck region. In the dermis, a

high APH activity is evident, particularly in the blood vessels and around and inside the

glands.

PAEDOMORPHIC NEWTS

In paedomorphic newts, skin morphology is rather variable and not easily general-

ized. In animals showing non-functional gonads and without evident SSC, histological

organization is roughly intermediate between that of larval and metamorphosed skin. The

superficial cells are relatively loose (somewhere separated by empty spaces), neuromasts

can be seen (even if rarer than in larvae), large exo-epithelial glands are present, and

Leydig cells (FOx, 1988) are not so orderly as in larvae, have a lighter cytoplasm, scarce

granules. In these animals mitoses can be observed in cells interposed among Leydig cells

(fig. 9).

In some paedomorphic specimens (GABRION & SENTEIN, 1986), a mosaic situation can

be noticed: the Leydig cells are still present and abundant in some areas, while in others

they are absent. In other branchiate newts (sexually mature and paedogenetic), the skin is

highly similar to that of the metamorphosed ones, although with a lower surface

cornification (fig. 10). It is sometimes possible to observe empty spaces inside the

epithelium, probably derived from the degeneration of Leydig cells. APH activity in the

more superficial layers is usually scarce or even absent, whereas in the dermis it is localized

in the portion just below the basal layer of the epithelium and more evident around the

glands than elsewhere. Flask cells are present in a few paedogenetic specimens and show

a high APH activity in the portion superficial to the nucleus.

DISCUSSION

Analysis of the organization of skin of Alpine newts throughout the life cycle discloses

high variability from larvae to metamorphosed specimens. The skin of paedomorphic

specimens has a structure which varies from larval-like to metamorphosed-like organiza-

tion; sometimes the skin shows both larval characteristics (e.g. Leydig cells, neuromasts)

and metamorphosed characteristics (e.g. dermic glands, horny surface). This intermediate

situation is also evident in the activity of APH, as has been stressed by Lopi et al. (in

press). These data mainly agree with GABRION & SENTEIN (1976), who observed that

Leydig cells vary in number and shape from totally larval to late paedomorphic palmate

newts (Triturus helveticus), in which the skin closely resembles that of meta-

Source : MNHN, Paris

ANDREONE, DORE, USAI & PARANINFO 31

morphosed individuals. In late paedogenetic Alpine newts, the skin rearrangement extends

also to the empty spaces left by the disappearance of the Leydig cells. The mitoses

observed in the intermediate epithelium can be interpreted as part of a regenerative

process.

The variability reported suggests the persistence (and not the arrest) of metamorphic

phenomena even in paedomorphic Alpine newts, and stresses the difficulty in categorizing

branchiate newts. In fact, by external examination it is possible to observe sexually inactive

paedomorphic individuals which resemble larvae, and others (paedogenetic) which exhibit

a typical mating coloration (more visible in males), and are sexually active, behaving

during the courtship like the typical metamorphosed adults (ANDREONE, 1990; BOVERO,

1991). Natural history observations confirm that gill development and length vary among

paedomorphic individuals of the very same population and that these may — at a certain

moment of their life — metamorphose. Thus, it is not possible to consider the

paedomorphics as a whole and uniform category, because they are part of a continuum,

from larval to metamorphosed newts. In this sense it is worth reaffirming, in contrast to

MoLaA & BERTOLANI (1981), that the presence of Leydig cells is not a constant neotenic

character. Perhaps these authors analyzed the skin of larval-like paedomorphic newts

(giant overwintered larvae”) and not that of the typical paedogenetic newts. The

correlation between morphology and function of the skin has been pointed out by Lopi

et al. (in press): a typical larval organization (with Leydig cells) is characterized by the

absence of active sodium transport, that is evident when the epidermis acquires, at least

in part, the metamorphosed characteristics.

All these considerations indicate how the newt life cycle (and more particularly that

of Triturus alpestris) cannot be rigidly schematized, but that plasticity in life history

characteristics is itself adaptive. In fact, the highly aquatic Alpine newts synchronize their

permanence in the water both to altitude differences (and thus to the duration of their

breeding period) and to desiccation-rain alternation in unpredictable habitats (ANDREONE

et al., in press). An outline of the life history of Alpine newts is represented in fig. 11 —

to interpret the observations on skin morphology and on external features (modified from

VERRELL, 1985, DZuxié et al., 1991, and ANDREONE et al., in press). Typically, newts spend

the first part of their existence as aquatic larvae; thereafter metamorphosis induces some

drastic changes (e.g. in the skin structure) as the newts become terrestrial. Growth until

sexual maturity occurs on land (particularly when the breeding site is temporary), but

juveniles may return to water for non-reproductive reasons, as has been reported for other

species by VERRELL (1985) and ANDREONE & GIACOMA (1989). Nevertheless an alternative

strategy is observed in more stable habitats: adulthood is reached through a direct aquatic

development, and larvae may overwinter in the water (especially those hatched in autumn,

see ANDREONE & DORE, 1992), becoming paedomorphic juveniles. Later they may become

paedogenetic, thus displaying SSC and courtship patterns. Throughout this aquatic

development, skin morphology modifies, although the changes are not so drastic as in the

transformation from water to land in the typical life cycle. For this reason a mosaic

situation is seen, and neotenic individuals may present both larval and metamorphosed

characters. Nevertheless metamorphosis may occur any time in the aquatic development.

Thus, metamorphosed individuals found in the water may result either after growth in a

terrestrial habitat, or by transforming at various stages of their aquatic life, possibly on

Source : MNHN, Paris

32 ALYTES 11 (1)

ALPINE NEWT

LIFE CYCLE

LAND WATER

DISTINCTIVE

FEATURES

LARVAL SKIN WITH

LEYDIG CELLS

UNDEVELOPED GONADS.

SEXUALLY INACTIVE

EXTERNAL GILLS

PAEDOMORPHIC

SUBADULTS

TERRESTRIAL

SUBADULTS

PAEDOGENETIC

ADULTS

SECONDARY

SEXUAL CHARACTERS

MULTILAYERED AND

CORNIFIED SKIN

EXO-EPYTHELIAL

GLANDS

METAMORPHOSED MATURE AND

ADULTS Le FUNCTIONAL GONADS

COURTSHIP BEHAVIOUR

Fig. 11. — Outline of the life cycle in Triturus alpestris apuanus. The left part summarizes the passages

between water and land throughout growth and metamorphosis. The black arrows indicate

habitat changes, but do not suggest any preference, depending on the characteristics and

constraints of each individual and population. The right part summarizes some typical features

of the extreme categories of aquatic newts, from an early larval stage to a final paedogenetic-

metamorphosed phase. Grey arrows indicate the correspondence of phases and biological

features. Between these two extremes several intergrades are present. The distinctive features of

metamorphosed newts, excluding the characteristics related with sexual activity, are common

both to adults and to juveniles.

Source : MNHN, Paris

ANDREONE, DORE, USAI & PARANINFO 33

the occasion of pond desiccation or of water pollution (see ANDREONE & DORE, 1991). The

reasons for this plasticity should be sought in adaptive explanations: paedomorphism —

as remarked by BREUIL (in press) — may lead to earlier sexual maturity, life in a more

stable aquatic habitat, or better utilization of trophic resources (KALEZIÉ, DZUKkIC &

TvRTKOVIÉ, 1990).

RÉSUMÉ

L’histologie de la peau du triton alpestre (Triturus alpestris apuanus) a été analysée et

comparée avec des données concernant l’histoire naturelle de l'espèce. La peau des jeunes

larves est composée d’un nombre réduit de couches cellulaires et contient des cellules de

Leydig, mais pas de glandes extra-épithéliales; celles-ci deviennent visibles plus tard, avant

la métamorphose. Chez les tritons métamorphosés elle est typiquement pluri-stratifiée et

cornifiée, avec beaucoup de glandes muqueuses et séreuses. L'organisation épithéliale des

individus pédomorphiques, qui ont des branchies et vivent dans l’eau, est assez variable:

chez certains exemplaires (correspondant aux larves géantes immatures), la peau présente

des caractéristiques typiquement larvaires (par exemple les cellules de Leydig sont encore

présentes), tandis que chez d’autres (habituellement sexuellement actifs bien que pédogé-

nétiques) elle est très similaire à celle des tritons métamorphosés. Ainsi, chez les tritons

pédomorphiques, la métamorphose n’est pas abolie, mais seulement retardée. Il en résulte

une mosaïque de caractéristiques larvaires et post-larvaires. Ces caractéristiques sont en

relation avec l'écologie plastique du triton alpestre.

ACKNOWLEDGEMENTS

Thanks are due to M. BreuiL for bibliographie support and for the exchange of information

about neoteny, and to two anonymous referees for critical advice. This work was supported by grants

from Ministero della Pubblica Istruzione and Ministero dell'Università e della Ricerca scientifica e

tecnologica (funds 40 % and 60 %).

LITERATURE CITED

ANDREONE, F., 1990. — Variabilità morfologica e riproduttiva in popolazioni di Triturus alpestris

(Laurenti, 1768) (Amphibia: Salamandridae). Doctoral research thesis, University of Bologna.

ANDREONE, F. & DORE, B., 1991. — New data on paedomorphism in Italian populations of the Alpine

newt, Triturus alpestris (Laurenti, 1768) (Caudata: Salamandridae). Herpetozoa, 4 (3/4):

149-156.

—— 1992. — Adaptation of the reproductive cycle in Triturus alpestris apuanus to an unpredictable

habitat. Amphibia-Reptilia, 13: 251-261.

Source : MNHN, Paris

34 ALYTES 11 (1)

ANDREONE, F., DORE, B. & Usai, P., 1991. — Histological and ecological aspects of neoteny in

Triturus alpestris apuanus (Bonaparte). In: G. GHiARA et al. (eds.), Symposium on the evolution

of terrestrial vertebrates, U.Z.I. Selected Monographs and Symposia, Modena, Mucchi Editore,

vol. 4: 413-420.

ANDREONE, F. & GiacomA, C., 1989. — Breeding dynamics of Triturus carnifex at a pond in

north-western Italy (Amphibia, Urodela, Salamandridae). Holaret. Ecol., 12 (3): 219-223.

ANDREONE, F., GlAcOMA, C., CAVALLOTTO, L. & FRANCILLON-VIEILLOT, H., in press. — Le cycle

reproducteur de Triturus alpestris: influence des facteurs externes. Actes du Colloque d'Ecologie

et de Biogéographie Alpines.

BovERo, S., 1991. — Analisi del comportamento riproduttivo in esemplari sintopici — metamorfosati e

pedogenetici — di Triturus alpestris apuanus (Amphibia: Salamandridae). Degree thesis,

University of Turin.

Breuil, M. in press. — La néoténie dans le genre Triturus: mythes et réalités. Bull. Soc. herp. Fr.

Dore, B. & Usai, P., 1986. — Glycol methacrylate embedding for enzyme preservation. Bas. appl.

Histochem., 30 (suppl): 82.

DouGHErTY, W., 1981. — Preparation of semi-thin sections of tissues embedded in water soluble

methacrylate for light microscopy. /n: G. CLARK (ed.), Staining procedures, Baltimore &

London, William & Wilkins, Biological stain commission.

Dumois, A., 1987. — Neoteny and associated terms. Alytes, 4 (4): 122-130.

DuELLMAN, W. E. & TRUEB, L., 1986. — The biology of amphibians. New York, McGraw Hill.

Dzuxié, G., KaLEzié, M. L., Tvrrirvoc, M. & DKPrPvoc, A., 1991. — An overview of the

occurrence of paedomorphosis in Yugoslav newt (Triturus, Salamandridae) populations. BHS

Bull., 34: 16-22.

Fox, H., 1977. — The anuran tadpole skin: changes occurring in it during metamorphosis and some

comparisons with that of the adult. /n: R. IL. C. SPEARMAN (ed.), Comparative biology of the skin,

Symp. zool. Soc. London, London & New York, vol. 39: 269-289.

me 1986a. — Changes in amphibian skin during larval development and metamorphosis. /n: G.

NaLLs & M. BOWNES (eds.), Metamorphosis. Symp. brit. Soc. dev. Biol., Oxford Univ. Press,

vol. 9-87.

ee 1986b. — The skin of Amphi In: J. BEREITER-HAHN, A. G. MATOLTSY & K. S. RICHARDS

(eds.), Biology of the integument, vol. I, Vertebrates, Heidelberg, Springer: 78-135.

ne 1988. — Riesenzellen, globlet cells, Leydig cells and the large cells of Xenopus, in the amphibian

larval epidermis: fine structure and a consideration of their homology. J. submicrosc. Cytol.

Pathol., 20 (2): 437-451.

GABRION, J. & SENTEIN, P., 1976. — Structure histologique de la peau et phénomènes de

dégénérescence chez Triturus helveticus Raz. au cours de la métamorphose. Bull. Soc. zool. Fr.

101, suppl. 5: 33-39.

Hay, E. D., 1961. — Fine structure of an unusual intracellular supporting network in the Leydig cells

of Amblystoma epidermis. J. fBiophys. Biochem. Cytol., 10: 457-463.

KaLezié, M. L., Dzukié, G. & TvrTkovié, N., 1990. — Newts (Triturus, Salamandridae, Urodela)

of the Bukovica and Ravni Kotari regions (Yugoslavia). Spixiana, 13 (3): 329-338.

KOLLMANN, J., 1884a. — Das Ueberwintern von europäischen Frosch und Tritonen Larven und die

Umwandlung des mexikanischen Axolotl. Verh. Naturf. Ges. Basel, (1883): 387-398.

en 1884b. — L'hivernage des larves dé grenouilles européennes et de tritons. La métamorphose de

l'axolotl mexicain. C. r. Assoc. fr. Avanc. Sci., (1883), 12: 567-570, 570-571 (LATASTE’s remarks).

= 1884c. — L'hivernage des larves de grenouilles et de tritons d'Europe et la métamorphose de

l’axolotl du Mexique. Rec. Zool. Suisse, 1: 75-89.

Lot, G., 1968. — Il tegumento del tritone crestato in condizioni sperimentali diverse. Boll. Zool., 35:

415.

LoDt, G., ANDREONE, F., DORE, B., PARANINFO, A., USsat, P. & BicioTri, M. in press. — Active

sodium transport and morpho-functional organization in the skin of the Alpine newt, Triturus

alpestris, during the life cycle. Boll. Zool.

Lot, G. & Bant, G., 1971. — Microscopic, submicroscopic and histoenzymologic features of the

-epidermis of the normal and hypophysectomized crested newt. Boll. Zool., 38: 111-125.

Loypa, Z., GossRAU, R. & SCHLEBER, T. H., 1979. — Enzyme histochemistry. A laboratory manual.

Springer-Verlag.

Source : MNHN, Paris

ANDREONE, DORE, UsAI & PARANINFO 35

Mazzi, V., 1977. — Manuale di tecniche istologiche ed istochimiche. Padova, Piccin.

MENNELLA, C., 1967. — 11 clima d'Italia nelle sue caratteristiche e varietà e quale fattore dinamico del

paesaggio. Napoli, Edart.

MoLa, L. & BERTOLANI, R., 1981. — Osservazioni istologiche e istochimiche sul differenziamento

della cute in esemplari metamorfosati e non di Triturus alpestris apuanus (Bonap.) (Amphibia

Urodela). Arch. ital. Anat. Embriol., 86 (3): 195-207.

RAFFAELLI, J., 1989. — Etat des données sur quelques traits de la biologie des Amphibiens Urodèles.

Bull. Soc. Herp. Fr., 48: 1-21.

VerreLz, P. À., 1985. — Return to water by juvenile amphibians at a pond in southern England.

Amphibia-Reptilia, 6: 93-96.

Wizsur, H. M. & CoLLINS, J. P., 1973. — Ecological aspects of amphibian metamorphosis. Science,

182: 1305-1314.

ZACCONE, G., FAsuLo, S., Lo Cascio, P. & LicaTA, A., 1986. — Enzyme cytochemical and

immunocytochemical studies of flask cells in the amphibian epidermis. Histocemistry, 84: 5-9.

Corresponding editor: Milos KALEZIC.

Source : MNHN, Paris

Alytes, 1993, 11 (1): 36.

Dates de publication du journal Alytes

(1992)

Alain Dugois

Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

Cette liste fait suite à celles que nous avons déjà publiées (Dugois, 1988, 1989, 1990,

1991, 1992) pour les années 1982-1991, et a été préparée de la même manière.

Volume Fascicule Pages Date figurant Date réelle

sur le fascicule de publication

9 4 89-144 December 1991 27 février 1992

10 1 1-36 May 1992 13 mai 1992

10 2 37-80 September 1992 30 septembre 1992

10 3 81-112 November 1992 27 novembre 1992

10 4 113-144 December 1992 23 décembre 1992

RÉFÉRENCES BIBLIOGRAPHIQUES

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

1989. — Dates de publication du journal Alytes (1988). Alytes, 7: 75.

1990. — Dates de publication du journal Alyres (1989). Alytes, 8: 22.

1991. — Dates de publication du journal Alyres (1990). Alytes,

1992. — Dates de publication du journal Alyres (1991). Alytes,

Source : MNHN, Paris

AINTTES

International Journal of. Batrachology

published by ISSCA

EDITORIAL BOARD FOR 1993

Chief Editor: Alain DuBois (Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire

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

Deputy Editor: Günter GOLLMANN (Institut für Zoologie, Universität Wien, Althanstr. 14, 1090 Wien,

Austria).

Other members of the Editorial Board: Ronald G. ALTIG (Mississippi State University, U.S.A.);

Emilio BALLETTO (Torino, Italy); Stephen D. Busack (Ashland, U.S.A.); Alain COLLENOT

(Paris, France); Tim HALLIDAY (Milton Keynes, United Kingdom); William R. HEYER

(Washington, U.S.A.); Walter HôDL (Wien, Austria); Pierre JOLY (Lyon, France); J. Dale

ROBERTS (Perth, Australia); Petr ROTH (Libechov, Czechoslovakia); Ulrich SINsCH (Bonn,

Germany); Marvalee H. Wake (Berkeley, U.S.A.).

Index Editor: Annemarie OHLER (Paris, France).

GUIDE FOR AUTHORS

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

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

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

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

beautiful or rare species, interesting behaviors, etc.

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

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

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

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legends of figures and tables should be assembled on a separate sheet. Each figure should be

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References in the text are to be written in capital letters (SOMEONE, 1948; So & So, 1987;

EveryBopy et al., 1882). References in the literature Cited section should be presented as follows:

BourReT, R., 1942. - Les batraciens de l'Indochine. Hanoi, Institut Océanographique de l’Indochine:

ix + 1-547, pl. IV.

GRAF, J.-D. & POLLS PELAZ, M., 1989. - Evolutionary genetics of the Rana esculenta complex. In:

R. M. DawLey & J. P. BOGART (eds.), Evolution and ecology of unisexual vertebrates, Albany, The

New York State Museum: 289-302.

INGER, R. F., Voris, H. K. & Voris, H. H., 1974. - Genetic variation and population ecology of some

Southeast Asian frogs of the genera Bufo and Rana. Biochem. Genet. 12: 121-145.

Manuscripts should be submitted in triplicate either to Alain DuRois (address above) if dealing

with amphibian morphology, systematics, biogeography, evolution, genetics or developmental

biology, or to Günter GOLLMANN (address above) if dealing with amphibian population genetics,

ecology, ethology or life history.

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

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

should be sent to the Chief Editor. We welcome the following formats of text processing: (1)

preferably, MS Word DOS and Windows, WordPerfect (4.1 to 5.1) or WordStar (3.3 to 5.5); (2) less

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

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

Charged. For each published paper, 25 free reprints are offered by A/ytes to the author(s). Additional

reprints may be purchased.

Published with the support of AALRAM

tion des Amis du Laboratoire des Reptiles et Amphibiens

du Muséum National d'Histoire naturelle, Paris, France).

Directeur de la Publication: Alain Dusois.

Numéro de Commission Paritaire: 64851.

Alytes, 1993, 11 (1): 1-36.

Contents

Jordi SERRA-CoBo

Descripcién de una nueva especie europea

de rana parda (Amphibia, Anura, Ranidae) ........................... 1

Leszek BERGER & Mariusz RYBACKI

Growth and maturity of brown frogs, Rana arvalis

and Rana temporaria, in central Poland ............................... 17

Franco ANDREONE, Bruno DORE, Pasquale Usa & Adriana PARANINFO

Skin morphology in larval, paedomorphic and

metamorphosed Alpine newts, Triturus alpestris apuanus ............... 25

Miscellanea

Application for membership of ISSCA

BOOM SUDSCTIPHONALO ALES EU En EE RENE nue 16

Alain Durois

Dates de publication du journal Alytes (1992) ......................... 26

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 1993.

Source : MNHN, Paris.