D 6445 a

ALYTES

Décembre 1986 Volume 5, fascicule 4

Source : MNHN, Paris

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ALVTES

Bulletin trimestriel Volume 5

Décembre 1986 Fascicule 4

Alytes, 1986, 5 (4): 153-164. 153

Diets of tadpoles living in a Bornean rain forest

Bibliothèque Centrale Muséum

MON

3

3001 00110039 4

Field Museum of Natural History,

Roosevelt Road at Lake Shore Drive,

Chicago, Illinois 60605-2496, U.S.A.

Diets of 16 larval forms of Bornean anurans are generally similar to those

of tadpoles from other regions : gut contents consist mainly of small algae and

other protists. Despite broad overlap, there appear to be differences between diets

of some co-occurring species in size and type of food ingested. Five feeding types

are represented by these 16 kinds of tadpoles : obligate benthic, macrophagous,

midwater suspension, surface film, and bottom suspension feeders. All except

the last are associated with morphological specializations that appear to be func-

tionally related to the size or kind of food particles ingested. Modes of feeding

are related to differences in microhabitat distribution and to some of the diffe-

rences in composition of the diets. It is this relationship that lends significance

to food resource partitioning as an element in the organization of this Bornean

tadpole community.

INTRODUCTION

Most of the recent spurt in non-taxonomic study of larval anurans, aside from

developmental and neurobiology, has been focused on morphology and function of the

buccopharynx (e.g., KENNY, 1969 ; SEALE & WASSERSUG, 1979 ; VIERTEL, 1985 ;

WASSERSUG, 1980 ; WASSERSUG & HOFF, 1979) or on population ecology (e.g., HEYER,

1979 ; MORIN, 1983 ; WILBUR, 1984). Studies on the diets of free-living tadpoles are

less numerous and usually concerned with one or two species (e.g., COSTA & BALASU-

BRAMANIAN, 1965 ; JENSSEN, 1967 ; SAVAGE, 1952 ; SEALE, 1980). HEYER (1973), who

examined the gut contents of 17 larval forms in Thailand, found much overlap in the

type of food ingested by tadpoles having keratinous beaks and denticles and a broader,

though still overlapping, spectrum of food types in larvae lacking keratinous mouth

Source : MNHN, Paris

154 ALYTES 5 (4)

parts (Microhylidae). DIAZ-PANIAGUA (1985) related the modest differences among the

diets of five species of tadpoles in Spain to differences in their distribution within the

water column of ponds.

In this paper, I report observations on the diets of 16 species of tadpoles living

in a Bornean rain forest at Nanga Tekalit, Sarawak. Twelve of these larval forms live

in small streams where they occupy a variety of microhabitats : torrents around large boul-

ders and rocks, shallow riffles over gravel, quiet open pools and areas of shingle rock,

areas of leaf drifts trapped by eddies, shallow side pools cut off from the main current,

and potholes in rocky banks. The remaining four larval forms live in pools distant from

streams : either rain filled floor depressions made by forest pigs or tanks formed by anas-

tomosing buttresses of three trunks. These 16 kinds of tadpoles are a subset of the 36

now known from Nanga Tekalit (INGER, 1985), occupy almost the entire array of micro-

habitats known to be used by tadpoles there, and represent 8 of the 14 genera.

This report has the nature of a preliminary survey. Sample sizes (see below) are

not large enough to provide definitive descriptions of the larval diets of the individual

species. However, even these limited samples reveal relationships among diet, morpho-

logy, and microhabitat distributions and the possibilities for food resource partitioning

within a complex assemblage of tadpoles.

One of the difficulties in some studies of tadpole diets (and in the present one as

well) is that objects identified as food may not be the actual sources of nourishment. Some

protists, e.g., blue-green algae, other algae, Volvox, etc., are known to pass through the

gut of tadpoles undamaged (COSTA & BALASUBRAMANIAN, 1965 ; SAVAGE, 1952). Con-

ceivably, the true food may be bacteria or viruses, unseen and unrecorded, as suggested

by HEYER (1973). Nonetheless, any systematic differences between species in either taxo-

nomic category or size of items in the gut indicate at least differences in feeding habits.

MATERIALS AND METHODS

Processing of larvae in the field is described in INGER, VORIS & FROGNER (1986).

To obtain samples of food, I cut half a centimeter of the foregut of a tadpole close to

the esophagus and teased its contents on to a glass slide. I removed the gut wall and visi-

ble portions ofits lining and added several drops of Melzer’s solution (iodine and chloral

hydrate). After spreading the gut contents as thinly as possible, I placed a cover slip over

it and sealed the edges. I scanned the entirety of each smear within a few days of prepara-

tion, using a compound microscope equipped with an ocular grid.

Every item that was identifiably organic and had reasonably intact cell boundaries

was measured (maximum diameter), recorded, and in the great majority of cases identi-

fied to major category (e.g., diatom, blue-green alga, fragment of tracheoid plant, etc.).

It was not possible to identify food items to genus or species. About half of the cases

in which identification was not possible involved small rod-like or spherical organic bodies

that I classified as “protists.”

Foregut smears were made from a total of 32 individuals. To minimize the risk

of confounding differences between days or microhabitat sites with differences between

species, wherever possible gut samples were taken from tadpoles of several species col-

Source : MNHN, Paris

INGER 155

Table I. — Microhabitat distribution, size, and stage of development of Bornean tadpoles used as

sources of gut smears.

Head-body

Species Microhabitat Stage* length (mm)

Tadpoles from stream microhabitats

Leptobrachium gracilis riffle 35 23

Leptobrachium montanum shingle 26 19

open pool 26 22

Megophrys nasuta riffle 26 10

shingle 34 11

Bufo divergens side pool 37,37,39 7

Ansonia longidigita leaf drift 36 5

Microhyla petrigena pothole 29,35 4,7

Amolops phacomerus torrent 35,37 12,14

Rana blythi leaf drift 26,39 7,10

side pool 28 8

pothole 26 8

Rana ibanorum side pool 36 9

pothole 28,33 8,10

Rana chalconota side pool 26,28,40 8,14,15

Rana signata leaf drift 38 13

Rhacophorus bimaculatus riffle 36 9

Tadpoles from microhabitats away from streams

Microhyla borneensis pig wallow 37 6

Rhacophorus dulitensis pig wallow 27,36 16,17

Rhacophorus nigropalmatus pig wallow 26 14

Rhacophorus harrissoni buttress tank 27 11

“Stages according to GOSNER (1960).

lected on the same day at the same site. The following constituted such “multispecies”?

samples of foregut smears from cooccurring tadpoles :

— Ansonia longidigita + Rana blythi, 2 samples of each from leaf drifts ;

— Bufo divergens + Rana chalconota, 2 samples of each from side pools ;

— Bufo divergens + Rana blythi + R. chalconota + R. ibanorum, 1 sample of each

from a side pool ;

— Rana blythi + R. ibanorum, 1 sample of each from a pothole ;

— Microhyla borneensis + Rhacophorus dulitensis, 2 samples of each from pig wallows.

Stages, sizes, and microhabitat sources of the tadpoles from which the food sam-

ples came are given in Table I.

Description of the general environment, full definitions of stream microhabitats,

and microhabitat distributions of stream tadpoles appear in INGER, VORIS & FROGNER

(1986) and descriptions of all larvae and definitions of non-riparian microhabitats in INGER

(1985).

Source : MNHN, Paris

156 ALYTES 5 (4)

Table II. — Frequency distribution of food particles of various sizes in smears from foreguts of

Bornean tadpoles. Number of smears per species given in Table I.

Species Food particle size (mm)

<:03 .03-05 06-10 .11-15 .16-20 .21-30 .31-40 >.4 mean*

Tadpoles from stream microhabitats

Leptobrachium gracilis 2 : Re 3 1 6 2 2 GT

Leptobrachium montanum 13 13 12 6 7 8 33 .188

Megophrys nasuta 36 8 10 4 LI ln 4 .050

Bufo divergens 121 40 86 8 4 9 041

Ansonia longidigita 3% 30 13 1 1 1 035

Microhyla petrigena 190 17 24 à 1 À 1 .026

Amolops phaeomerus 456 386 21 3 1 7 029

Rana blythi 30 52 88 38 21 23 5 6 .080

Rana ibanorum 43 48 55 32 16 13 # 5 .069

Rana chalconota 17 51 36 14 8 il 4 7.037

Rana signata Su. 12 17 2 4 7 1 075

Rhacophorus bimaculatus 40 34 1 028

Tadpoles from microhabitats away from streams

Microhyla borneensis 2 15 5 2 1 1.063

Rhacophorus dulitensis 2 73 48 28 8 7 1 10 .071

Rhacophorus harrissoni 29 F 52 8 10 20 3 4 081

Rhacophorus nigropalmatus 10 10 73 1 6 6 1 1 083

*Means calculated from class mid-points converted to logs. Assumed mid-point of smallest class = .02.

Differences in food size-frequency distributions were analyzed by means of the

Kolmogorov-Smirnov test. The G test was used for comparing types of food, with uniden-

tified items omitted.

RESULTS

Most of the gut samples include a wide spectrum of food types and sizes (Tables

Il and III). The community as a whole appears to be supported by a diet of very small

organisms, mainly single-celled protists and short strands (usually <16 cells) of algae and

fungi. Fragments of higher plants form the next most frequent category. There were also

miscellaneous fragments of invertebrate cuticle, several butterfly scales, and pieces of

arthropod exoskeleton.

The multispecies samples (see Merhods) yield the following :

— Ansonia longidigita x Rana blythi : both sets show the same trend, i.e., food size

smaller in À. longidigita (P <.01), and more tracheoid plant fragments eaten by R. blythi

(P<.001).

— Bufo divergens x Rana blythi : food size smaller in B. divergens (P = .005) ; more

tracheoid plant fragments eaten by R. blychi (P<.001).

Source : MNHN, Paris

INGER 157

Table III. — Frequency of food types in smears from foreguts of Bornean tadpoles. Number of smears

per species given in Table I.

Species Food types*

AL DI FN CI EU AM PR TP RO MS æ

Tadpoles from stream microhabitats

Leptobrachium gracilis 1 3 13 7

Leptobrachium montanum 18 8 54 4 8

Megophrys nasuta 35 7 2 20 4 25

Bufo divergens 35 10 2 4 64 100 9 1 3

Ansonia longidigita 47 4 3 5 ul

Microhyla perigena 5 I 6 8 sl

Amolops phacomerus 807 12 2 3 38

Rana blythi AB CA TEE T e 8 LI 65 14 74

Rana ibanorum 15-MNTS 397 No MN 2 24 7 48

Rana chalconota 90 14 16 1 T7 2 108 23 3 2 44

Rana signata DS MANS (ES; 1 6 2 1

Rhacophorus bimaculatus 43 32

Tadpoles from microhabitats away from streams

Microhyla borneensis 10 1 6 3, À 3

Rhacophorus dulitensis 68 39 4 3 I 1 19 12 1 4 14

Rhacophorus harrissoni 1e 20202 4 2 5 0.21

Rhacophorus nigropalmatus 16» “6 3 OL 07 3 1 6 À" 6° 25

* AL = algae, mainly blue-green ; DI = diatoms ; FN = fungi ; CI = ciliates ; EU = euglenoids ; AM = amoe-

bae ; PR = miscellaenous protists ; TP = tracheoid plant fragments ; RO = rotifers ; MS = miscellany, includ-

ing insect and crustacean exoskeleton fragments ; ? = unknown.

— Bufo divergens x Rana chalconota : food size smaller in R. chalconota in 2 sets

(P<.01) but not the third ; no significant difference in type of food eaten.

— Bufo divergens x Rana ibanorum : food size smaller in B. divergens (P < .001) ;

more tracheoid plant fragments eaten by R. ibanorum (P <.001).

— Rana blythi x R. ibanorum : food size smaller in R. ibanorum in one set (P < .03),

but not in the other ; no significant difference in type of food eaten.

— Rana blythi x R. chalconota : no significant difference in size or type of food.

— Rana ibanorum x R. chalconota : no significant difference in size or type of food.

— Rhacophorus dulitensis x Microhyla borneensis : no significant difference in size

or type of food eaten.

The 12 species from stream microhabitats fall into three groups on the basis of food

particle size (Table IT) : 2 (Leptobrachium gracilis and L. montanum) that fed on relatively

large objects (mean particle size > .12 mm), with heavy emphasis on fragments of tracheoid

plants ; 3 (Rana blythi, R. ibanorum, and R. signata) that had ingested a high proportion

of medium-sized items (means .069-.08 mm) ; and 7 that contained a high proportion of

very-Ssmall items (means .026-.05 mm). For convenience I term these three groups macro

meso-, and microphagous types, respectively. Différences within the microphagous group

Source : MNHN, Paris

158 ALYTES 5 (4)

are not significant (P >.10, Friedman 2- way ANOVA), but each of them differed signifi-

cantly from each of the mesophages in pair-wise comparisons of food size-frequency dis-

tributions (P<.01, Kolmogorov-Smirnov test). However, I doubt the position of Rana

chalconota with the microphagous species and the reality of the differences between it and

R. blythi and R. ibanorum because of the results with multispecies samples (see above).

The size-frequency distributions of the two species of Leprobrachium differ significantly

(P<.05) in pair-wise tests with all others except for the L. gracilis x R. signata pair. The

three mesophagous forms do not differ among themselves in size of food.

The two larval Leprobrachium are among the largest Bornean tadpoles (Table I and

data in INGER, 1985) and have large beaks. They occur mainly in open pools (L. monta-

num) and riffles (L. gracilis). The three larval Rana constituting the mesophagous group

are smaller (Table I) and have weaker beaks. One of them, R. ibanorum, lives primarily

in side pools and potholes and the other two, blyrhi and signata, mainly in leaf drift.

The microphagous stream larvae are heterogeneous in size, phylogenetic relations,

and ecological distribution. They include a small microhylid (Microhyla petrigena)

lacking beaks and labial denticles, a medium-sized, funnel-mouth pelobatid (Megophrys

nasuta), two small, generalized bufonids (Ansonia longidigita and Bufo divergens), a large,

heavy-beaked ranid (4molops phaeomerus), and a moderate-sized, heavy-beaked rhacopho-

rid (Rhacophorus bimaculatus). The diversity of microhabitat distribution within this group

is evident in Table I.

The four larval types collected away from streams do not differ significantly

among themselves in size or general type of particles ingested. The rain-filled pig wallows

in which three of them occurred had fine, silty bottoms and relatively few dead leaves.

Microhyla borneensis, which we saw occasionally near the surface of the turbid water, is

much smaller than the two rhacophorids (Table I) and lacked their horny beaks and

denticles. We did not see the two rhacophorids near the surface unless we disturbed the

water with nets. The buttress tank from which the sampled tadpole of Rhacophorus

harrissoni came had a deep layer of dead leaves at the bottom.

DISCUSSION

MODES OF FEEDING

Direct observations on the behavior of free-living tadpoles provide information on

where and roughly how seven forms feed : Megophrys nasuta, Bufo divergens, Microhyla

petrigena, M. borneensis, Amolops phaeomerus, Rana ibanorum, and R. chalconota. In

addition, we have reliable information on where in the water column nine additional forms

spend most of their time and, presumably, where they feed : Leprobrachium gracilis,

L. montanum, Ansonia longidigita, Rana blythi, R. signata, Rhacophorus bimaculatus,

R. dulitensis, and R. nigropalmatus.

Five of the six modes of larval feeding defined by SATEL & WASSERSUG (1981) are

recognizable in this community : (1) obligate benthic feeding, (2) creation of suspensions

over the bottom (=“generalist” of SATEL & WASSERSUG), (3) macrophagous, (4) midwater

suspension feeding, and (5) particulate surface film feeding.

Source : MNHN, Paris

INGER 159

(1) Larval Amolops phaeomerus cling to rocks by means of an abdominal sucker and

graze on the epilithic film of protists. The tadpoles are large enough to watch as they slowly

move across rocks in clear water. Rhacophorus bimaculatus, which has a cup-like, suctorial

oral disk, lives in the interstices of bottom rocks (INGER, 1985) and belongs in this category.

(2) Larval Rana chalconota, R. ibanorum, and Bufo divergens move slowly and irre-

gularly over bottom debris in shallow side pools and potholes. Often an individual pauses

in a snout-down, tail-elevated position, presumably creating and ingesting suspensions imme-

diately above the interface of water and substrate. Larval Rana blyrhi, R. signata, and Ansonia

longidigita live mainly within the layers of dead leaves that constitute leaf drifts and, given

their gut contents, appear to feed by creating bottom suspensions. Larval Rhacophorus

harrissoni, though living in tree buttress tanks, are like the preceding four species in living

on and within mats of dead leaves. The broad spectrum of food types and sizes found

in the sample from this species (Table II and III) suggests a similar mode of feeding. The

two rhacophorids from pig wallows, Rhacophorus dulitensis and R. nigropalmatus, also appear

to fit this feeding category.

(3) Larval Leprobrachium montanum and L. gracilis clearly ingest significant amounts

of relatively large fragments of tracheoid plants (Table II and III). They apparently obtain

much of their food by snipping off pieces of decaying vegetation. When larval L. monta-

num being reared in a field laboratory were offered dead leaves, they attacked the leaves

around the margins, not on a broad surface.

(4) Only two larval types, Microhyla borneensis and M. petrigena, are midwater sus-

pension feeders, a mode they share with other Asian microhylids (HEYER, 1973). Microhyla

petrigena was easily and frequently observed in small, clear potholes with individuals dis-

tributed throughout the water column and remaining fixed in position unless disturbed.

Although it was more difficult to see M. borneensis in the silty water of pig wallows, enough

individuals were seen near the surface to suggest behavior similar to that of M. perrigena.

(5) Megophrys nasuta is the only larval form in this series that feeds at the surface

film. Tadpoles of species of Megophrys have long been known to feed in this manner (e.g.,

SMITH, 1926 ; POPE, 1931 ; LIU, 1950).

MORPHOLOGICAL RELATIONS

Although this study did not involve critical investigation of functional relations,

there appear to be some associations of oral and buccopharyngeal morphology with diets

and mode of feeding for all except one group, those larvae that create and ingest bottom

suspensions. The obligate benthic feeding Amolops phaeomerus and Rhacophorus bimacu-

latus share a number of features : (a) a suctorial device — an abdominal sucker in the

former and the oral disk in the latter ; (b) heavy beaks with thick, coarse, marginal serra-

tions and ribbed outer surfaces ; (c) long, scoop-shaped denticles sharply angled towards

the mouth and having many marginal cusps (INGER, 1985 : fig. 33) ; (d) regular, pro-

nounced decrease in size of denticles from inner to outer rows of both lips ; (e) modifica-

tion of the anterior walls of the internal nares to form forwardly projecting flaps (INGER,

1985 : fig. 34) ; (f) no lingual papillae ; (g) few or no pustules in the interiors of buccal

roof and floor arenas. Characters (a), (b), (d), (e), and (g) are unique to these two larval forms

Source : MNHN, Paris

160 ALYTES 5 (4)

among those dealt with in this paper. The form of their beaks and the length, angulation,

and cusp wear pattern of their denticles suggest both these tadpoles use beaks and den-

ticles to scrape rocks, which is consistent with their gut contents (Table III) and observed

behavior of À. phaeomerus.

The macrophagous larval Leptobrachium have many rows of laterally compressed,

sharply pointed denticles (INGER, 1985 : fig. 1) and very heavy, sharp beaks that seem

suited to snipping bits of decaying vegetation. The array of large papillae in the buccal

cavities of both L. gracilis and L. montanum (INGER, 1983) may serve to shunt large food

particles away from the branchial baskets and glottis. Surface feeding Megophrys larvae

hang from the surface film by means of their upturned funnel mouth (SMITH, 1926). LIU

(1950) has described how the large palps and flaps just inside the mouth of M. minor act

to block entrance of large objects. Similar structures occur in the buccal cavity of larval

M. nasuta (INGER, 1985 : figs. 5-6), which has a much more slender beak than do tad-

poles of other genera of Oriental pelobatids (see illustrations in POPE, 1931, and Liu, 1950).

Mean food particle size is relatively small in M. nasuta (Table II). In common with other

larval Microhylidae, the midwater suspension feeding tadpoles of Microhyla borneensis and

M. petrigena lack beaks and denticles. Both have very large branchial baskets with dense

filter ruffles (INGER, 1985 : fig. 16), suggesting filtration of small particles, though the

food sample of only perrigena substantiates this suggestion (Table II). HEYER (1973) sug-

gested that the method of feeding used by microhylid tadpoles is less discriminating both

with respect to taxonomy and size of food items, but this idea is not borne out by the

Bornean data (Tables II and III).

The tadpoles feeding on bottom suspensions are a mixture taxonomically and mor-

phologically. Their morphological variation has no obvious functional or ecological cor-

relation. The one feature they share — beaks of moderate thickness having finely serrated,

sharp margins — is their only morphological distinction from all the other feeding types

in this assemblage. Other morphological characters either vary widely within this group

or overlap with one or several of the other feeding modes. For example, although all the

bottom suspension feeders have many pustules (20-100) in the interior of buccal roof arena,

so do the macrophagous larvae of Leptobrachium. Denticles of 5 of the 9 bottom feeders

are set with many (> 12)triangular marginal cusps and have the end and sides of the shaft

curved towards the mouth, features also found in the two obligate benthic feeders.

MICROHABITAT DISTRIBUTION

Modes of feeding and microhabitat distribution (Table I ; see also INGER, VORIS

& FROGNER, 1986) are clearly related. The obligate benthic feeders are excluded from

microhabitats, such as leaf drifts and silty side pools, where bottom cover would prevent

development of the epilithic flora these tadpoles graze on. Bottom suspensions would be

lost to tadpoles creating them in moderate or strong current, limiting distribution of this

mode of feeding to standing water (e.g., pig wallows used by Rhacophorus dulitensis) or

areas of weak current (e.g., side pools used by Bufo divergens). Similar physical constraints

limit midwater suspension feeders to potholes along stream banks (Microhyla petrigena)

or pools of standing water on the forest floor (M. borneensis). In contrast to the pre-

Source : MNHN, Paris

INGER 161

ceding types, the macrophagous Leprobrachium larvae are not restricted by either their

basic food source, dead leaves, or their mode of feeding. However, only one of these lar-

vae, L. montanum, actually has a wide microhabitat range (INGER, VORIS & FROGNER,

1986). Surface film feeding is apparently possible anywhere except in the most turbulent

areas ; larval Megophrys nasuta used almost the entire range of stream microhabitats from

riffles to potholes (INGER, VORIS & FROGNER, 1986).

TYPE OF FOOD

The dominant kinds of food in the diets of these Bornean tadpoles, as a group, resem-

ble those of other assemblages. The main food source consists of small algae and other

protists for larval communities investigated by SAVAGE (1952) in England, by HEYER (1973)

in Thailand, by SEALE (1980) in Central United States, and DIAZ-PANIAGUA (1985) in

Spain, as well as in the Bornean one. Interspecific variation within this framework can

be described in only broad terms because of difficulties associated with specific

identifications of food and with measurements and counts of food items. Hyla meridio-

nalis and Rana perezi ingested much more Cyanophyta than the other three larvae from

Spain (DIAZ-PANIAGUA, 1985). Diatoms were an important element in the food of all Thai

larvae except those of Kaloula pulchra (HEYER, 1973). Cyanophyta were more

important in gut contents of larval Amolops phaeomerus than in the other Bornean

tadpoles (Table III).

Fragments of tracheoid plants were relatively common in both the Bornean (Table

III) and Spanish assemblages (DIAZ-PANIAGUA, 1985), but were not reported for the Thai

samples (HEYER, 1973). SAVAGE (1952) said that “higher plants appear to be almost use-

less as food for tadpoles..?” because though “..aten in large quantities by starving ani-

mals. [they]... do not support growth?” SAVAGE’S statement is probably related to the short

time food is in the digestive tract of tadpoles (4-8 hrs in ones he studied) and the inability

of tadpoles to break down cellulose (SAVAGE, 1952). Nonetheless, apparently healthy Bor-

nean tadpoles ingest significant amounts of tracheoid plant matter, though they may be

digesting microorganisms growing on those fragments.

Animal matter appeared only sporadically in food remains in all these samples,

although SAVAGE thought that ingestion of micro-crustaceans had a significant positive

effect on growth rates. SAVAGE interpreted “fairly common” appearance of tadpole den-

ticles in the gut contents as evidence of feeding on dead larvae. Single denticles found

in foregut smears of four Bornean tadpoles were clearly from conspecifics and indicate

that tadpoles may sometimes swallow their own worn, shed denticles. This interpretation

seems particularly apt for one of these four, a larval Amolops phaeomerus, for it is difficult

to visualize how a tadpole of this species could feed on an object having the shape of a

dead tadpole.

CONCLUSION

The role of diet in organizing tadpole communities has been minimized (HEYER,

1976 ; TOFT, 1985 ; DIAZ-PANIAGUA, 1985), partly because attention has centered on taxo-

nomic composition of the diet. Given the overwhelming importance of protists as a food

source for all these communities and the coarse level of food identification in most

Source : MNHN, Paris

162 ALYTES 5 (4) -

studies, the observed high overlap between species in composition of the diet is expected.

To be sure, the weak indications of specific differentiation (see above) might be strength-

ened by improved identifications of algae. However, that advance would be offset by the

complications of temporal and microgeographic variation in algal blooms and microhabi-

tat and temporal distributions of tadpoles. Differences among diets, in terms of size of

food particles, exist (HEYER, 1973 ; this study, p. 156), although there is much overlap

between species (Table IT) and little relation to microhabitat distribution or larval size.

With improvements in measurement and identification of food and expansion of

studies to other larval assemblages, composition of diets may ultimately help us under-

stand organization of these communities. However, even with the present limitations of

our data, it is evident that mode of feeding is an important factor in mediating the struc-

ture of tadpole communities (INGER, VORIS & FROGNER, 1986). HEYER’s (1973) obser-

vation of three modes of feeding — bottom suspension feeders, midwater filter feeders,

and surface film feeders — accounts for most of the variation in positions in the water

column of the Thai tadpoles. The five feeding modes of the Bornean community (see

p. 158) are related to differences in microhabitat distributions and to some of the diffe-

rences in diet composition.

Those observations, however, leave an unanswered question : is the community

structure that is revealed by diets, feeding modes, and microhabitat distributions main-

tained by ecological forces such as competition ? Differences among species within com-

munities in modes of feeding and associated morphological specializations are correlated

with taxonomic boundaries, at least in the Thai and Bornean samples, which have the

largest arrays of species and genera. In Southeast Asia an abdominal sucker associated

with obligatory benthic habits is confined to tadpoles ofthe genus Amolops and expanded

suctorial lips limited to benthic feeding tadpoles of the Rhacophorus bimaculatus species

group. Sharp beaks and compressed, knife-like denticles are found only in macrophagous

pelobatid larvae (those of Leptobrachium, in this case) among Asian tadpoles. Surface fee-

ding by means of “funnel” mouths is restricted to larval Megophrys and certain species

of Microhyla in Southeast Asia, though these two groups have radically different bucco-

pharyngeal structures (WASSERSUG, 1980) and presumably very different ways of pro-

cessing food particles. Larvae of species groups (or subgenera) of Asian Rana show limi-

ted within-group morphological and behavioral variation (cf., HEYER, 1973 ; INGER, 1985 ;

POPE, 1931). Given this broad correspondence between taxonomic boundaries and modes

of feeding and morphology, the relation of feeding biology to organization of tadpole com-

munities appears to owe more to phylogenetic events than to contemporary ecological

forces.

ACKNOWLEDGEMENTS

Jam grateful to Richard J. WASSERSUG for many stimulating discussions on the biology of

tadpoles. Both he and Harold K. VORIS made helpfül critical suggestions on the manuscript. Four

colleagues, J. P. BACON, K. J. FROGNER, W. HOSMER, and F. W. KING, participated in the collec-

tion of tadpoles at various times. Field and laboratory work were partially supported by National

Science Foundation grants G20867 and GB7845X and a grant from the Allen-Heath Memorial

Foundation.

Source : MNHN, Paris

INGER 163

RÉSUMÉ

Les régimes alimentaires des têtards de 16 espèces d’Anoures vivant en forêt dense

humide, à Bornéo, sont étudiés. D’une manière générale, ils s’avèrent similaires à ceux

des têtards d’autres régions : les contenus digestifs sont principalement composés de peti-

tes algues et autres protistes. Malgré un large chevauchement, des différences sont consta-

tées entre les régimes alimentaires de certaines espèces qui se rencontrent ensemble ; ces

différences portent sur la taille et le type d’aliments ingérés. Cinq modes d’alimentation

sont représentés parmi ces 16 types de têtards : l'alimentation benthique stricte, la macro-

phagie, l'alimentation à partir de particules en suspension en pleine eau, l’alimentation

à partir du film en surface de l’eau et l’alimentation à partir de suspensions créées au-

dessus du fond. Tous ces modes d’alimentation, sauf le dernier, sont associés à des modifi-

cations morphologiques qui sont en rapport fonctionnel avec la taille ou le type des parti-

cules alimentaires ingérées. À divers modes d’alimentation correspondent également des

différences dans les microhabitats fréquentés par les têtards et certaines des différences

observées dans la composition des régimes alimentaires. Ces corrélations indiquent que

le partage des ressources alimentaires joue un rôle dans l’organisation de cette commu-

nauté de têtards de Bornéo.

(Résumé rédigé par 3.7. MORÈRE)

LITERATURE CITED

Cosra, H. H. & BALASUBRAMANIAN, S., 1965. — The food of the tadpoles of Rhacophorus cruciger

cruciger (Blyth). Ceylon J. Sci., 5 : 107-109.

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

cies in the Biological Reserve Donana (Huelva, Spain). Amph.-Repr., 6 : 307-322.

HEYER, W. R., 1973. — Ecological interactions of frog larvae at a seasonal tropical location in Thai-

land. 7. Herper., 7 : 337-361.

—— 1976. — Studies in larval amphibian habitat partitioning. Smithsonian Contr. Zool., 242 : iii

+ 1-27.

— 1979. — Annual variation in larval amphibian populations within a temperate pond. 3. Was-

hingion Acad. Sci., 69 : 65-74.

INGER, R. EF, 1983 — Larvae of Southeast Asian species of Leprobrachium and Leprobrachella (Anura :

Pelobatidae). In : RHODIN, A. & MIYATA, K. (eds.), Advances in herpetology and evolutionary

biology, Cambridge, Museum of Comparative Zoology : 13-22.

es 1985. — Tadpoles of forested regions of Borneo. Fieldiana : Zool., (n.s.), 26 : i-v + 1-89.

INGER, R. EF, VORIS, H. K. & FROGNER, K. J., 1986. — Organization of a community of tadpoles in

rain forest streams in Borneo. %. Tropical Ecol., 2 : 193-205.

JENSSEN, T. A., 1967. — Food habits of the green frog, Rana clamitans, before and during meta-

morphosis. Copeia, 1967 : 214-218.

KENNY, J. S., 1969. — Feeding mechanisms in anuran larvae. . Zoo!. London, 157 : 225-246.

Liu, C. C., 1950. — Amphibians of Western China. Fieldiana : Zool. Mem., 2 : 1-400.

MORIN, P. J., 1983. — Predation, competition, and the composition of larval anuran guilds. Ecol.

Monogr., 53 : 119-138.

Source : MNHN, Paris

164 ALYTES 5 (4)

Pope, C. H., 1931. — Notes on amphibians from Fukien, Hainan, and other parts of China. Bull.

Amer. Mus. Nat. Hist., 61: 397-611.

SATEL,S. L. & WASSERSUG, R.J., 1981. — On the relative sizes of buccal floor depressor and eleva-

tor musculature in tadpoles. Copeia, 1981 : 129-137.

SAVAGE, R. M., 1952. — Ecological, physiological and anatomical observations on some species of

anuran tadpoles. Proc. Zool. Soc. London, 122 : 467-514.

SEALE, D. B. & WASSERSUG, R. J., 1979. — Suspension feeding dynamics of anuran larvae related

to their functional morphology. Oecologia, 39 : 259-272.

SMITH, M. A., 1926. — The function of the “funnel” mouth of the tadpoles of Megalophrys, with

a note on M. aceras Boulenger. Proc. Zool. Soc. London, 1926 : 983-988.

Torr, C. A., 1985. — Resource partitioning in amphibians and reptiles. Copeia, 1985 : 1-21.

VIERTEL, B., 1985. — The filter apparatus of Rana temporaria and Bufo bufo larvae (Amphibia,

Anura). Zoomorph., 105 : 345-355.

WASSERSUG, R. J., 1980. — Internal oral features of larvae from eight anuran families : functional,

systematic, evolutionary, and ecological considerations. Misc. Publ. Mus. Nat. Hist. Univ.

Kansas, 68 : i-iv + 1-146.

WASSERSUG, R. J. & HOFE, K., 1979. — A comparative study of the buccal pumping mechanism

of tadpoles. Biol. J. Linnean Soc, 12 : 225-259.

WILBUR, H. M., 1984. — Complex life cycles and community organization in amphibians. Jn :

PRICE, P. W., SLOBODCHIKOFF, C. N. & GAUD, W. S. (eds), À new ecology : novel approaches

10 interactive systems, New York, Wiley : 195-224.

Source : MNHN, Paris

Alytes, 1986, 5 (4): 165-172. 165

Growth and metamorphosis of anuran larvae :

effect of diet and temperature

Ashok K. HOTA* & Madhab C. DASH**

* P. G. Department of Zoology, G. M. College,

Sambalpur — 768004, Orissa, India

#* School of Life Science, Sambalpur University,

Jyoti Vihar, Burla — 768017, Orissa, India

A diet that favours tadpole growth also quickens the onset of metamor-

phosis and promotes a greater transformation size. Within the range of 27°. 37°C

metamorphosis is accelerated by increased temperature more than growth is

and the larvae tend to transform at a lower size limit. At 15°C, growth was favou-

red over metamorphosis and the larvae grew beyond the normal upper limit sho-

wing a tendency toward facultative neoteny.

INTRODUCTION

Growth rate at each stage of development is an important part of a species life his-

tory strategy (COLE, 1954 ; GADGIL & BASSERT, 1970). In poikilothermic animals body

size and growth rate are controlled by environmental conditions. Among aquatic animals

body size and rate of growth are functions of the volume of water in which the animals

are raised (HOGG, 1854 ; ALLEE, 1931) as well as of the more usual analyzed environ-

mental factors of food supply and temperature. In general ectotherms grown at lower

temperature are larger than those grown at higher temperature (HOWE, 1967 ; LOCK &

MCLAREN, 1970).

Anuran metamorphosis, as a developmental process, involves both growth and dif-

ferentiation and it is very difficult to separate the two processes. The concept of a meta-

morphic threshold size (WILBUR & COLLINS, 1973 ; SALTHE & MECHAM, 1974 ; DASH

& HOTA, 1980 ; HOTA, 1984) indicates that conditions that affect the growth rate of lar-

val anurans also affect the time of and body size at metamorphosis. In a study on the

influence of food quality and quantity on early larval growth of two anuran species, STEIN-

WASCHER & TRAVIS (1983) reported that growth was greatest at the highest ratio of pro-

tein to carbohydrate offered in the diet but not at the highest food level in Hyla chrysosce-

lis. However larval growth of Rana clamitans was unrelated to the specific protein / car-

bohydrate ratio in the diet but responded proportionately to change in protein content

and food level. According to SMITH-GILL & BERVEN (1979), “environmental tempera-

ture is a major proximal factor in the growth, differentiation and overall life history pat-

terns observed in amphibians”. Given these considerations, in this study, we attempt to

evaluate the effect of diet and temperature on growth and metamorphosis of Rana tige-

rina (Daudin) and Bufo melanostictus Schneider larvae.

Source : MNHN, Paris

166 ALYTES 5 (4)

METHODS

The methods of spawn collections were the same as described by HOTA & DASH

(1981). The eggs were allowed to hatch in laboratory conditions and the hatchlings were

mixed to assure uniformity in initial genetic and developmental conditions before being

assigned at random to experimental treatment.

Experiments were started soon after the hatchlings begin to feed. Tap water, con-

ditioned with sodium thiosulphate at a concentration of 8 mg/4.5 1 (NACE & RICHARDS,

1972) and filtered, was used as the culture medium. According to GROMKO, MASON &

SMITH-GILL (1973) the cube root of tadpole volume is the best estimator of larval size.

MCNAB (1970) and BARTHOLOMEW (1977) argued in favour of weight as the best size

measurement. In this study body mass has been used to estimate larval growth. Body

mass was determined in a chemical balance sensitive to 0.001 g precision, by weighting

one or more individuals (after blotted on a cloth towel) in a preweighed beaker containing

10 ml of clear distilled water. All weighings were done in duplicate.

The larvae were selected at random from the homogeneous population of both spe-

cies and were assigned to the following experimental treatments.

(1) R. tigerina larvae in group of 5 and B. melanostictus larvae in group of 20, in

triplicate sets were reared in different diets : (i) boiled Amaranthus tricolor leaves, (ii) boi-

led Basella alba leaves, (iii) cooked minced goat meat, (iv) boiled chicken egg yolk and

(v) boiled Amaranthus leaves, boiled chicken egg yolk and cooked minced goat meat mixed

in the proportion of 5 : 1 : 1. The larvae were fed ad libitum. Such food qualities were

chosen with an aim to develop suitable culture method of anuran larvae and to produce

healthy froglets and toadlets for dispersal as part of a frog farming programme.

(2) Groups of 5 and 20 of R. rigerina and B. melanostictus larvae, respectively, were

reared at temperature 15°, 27°, 33° (room temperature in June and July) and 37°C with

sufficient food (above mentioned diet type v).

The tadpoles in the experiments were allowed to progress to metamorphic climax

stage. The emergence of first forelimb was taken as the criterion of onset of metamorpho-

sis and complete resorption of tail was taken to indicate termination of metamorphic events.

Metamorphosing individuals (froglets and toadlets with emergent forelimbs) were remo-

ved from the cultures into amphibious environments where they were allowed to com-

plete metamorphosis to emerge as juveniles. The cultures were examined daily to deter-

mine the survival. The dead individuals were removed from the cultures and were exclu-

ded from analysis. Thrice weekly the culture pots were cleaned, water renewed and new

food was added. Twice a week the masses of all larvae were determined to ascertain the

mean growth rates. All statistical analyses were done according to SOKAL & ROHLF (1969).

RESULTS

GROWTH

A simple F, max test of the body mass of R. rigerina and B. melanostictus shows that

the variances among groups reared in different diet are not significantly different (Tables

J'and Il). But oneway analysis of variance indicates that the means are significantly diffe-

rent from one another (Tables I and Il). À posteriori test suggests that each diet types

are significant in their effects on growth of R. rigerina larvae (Table I), whereas the effects

Source : MNHN, Paris

HOTA & DASH 167

Table I. — Test of variances among R. rigerina larvae after 15th days growth as a function of diffe-

rent diet.

: Average body

Nos. Diet weight (8) SD.

1. Boiled Amaranthus tricolor leaves 0.1690 0.0042

2. Boiled Basella alba leaves 0.2138 0.0093

3. Cooked minced goat meat 0.2746 0.0296

4. Boiled egg yolk 0.3600 0.0147

5. Mixed diet 0.4412 0.0151

F, max = 75.53 (ns)

Anova table

Variation df S.S. MS. F:

Treatments 4 0.30415 0.07604

267.45 **

Error 10 0.00284 0.00028

Total 14 0.30699

Coeff. det. = R2 = 0.98

#* Significant at 0.001 level

A posteriori test (S.N.K. test)

3 4 5

Q 3.15 3.88 433 4.66

L.SR. 0.0304 0.0374 0.0418 0.045

1<2<3<4<5

of each pair of diet types on growth of B. melanostictus larvae are similar (Table Il). It

is evident that the final mass of larvae of both the species increased with the supplemen-

ted carnivorous diet. After 35th day of rearing with a strictly plant diet, heavy mortality

and deformities were observed amongst R. tigerina larvae.

At lower temperature the larvae of both species were larger at any given stage

(fig. 1). The apparent decline in weight at 15°C in both species was due to heavy morta-

lity of larger animals causing a drop in the average weight.

METAMORPHOSIS

Table III summarises the effect of diet on size and time of metamorphosis. In exclu-

sively plant diet, not a single R. rigerina larva from the replicate pots developed fore-

limbs. In contrast B. melanostictus larvae reached metamorphic climax at around the 15th

day after hatching with all diets tested in this experiment. But there was distinct varia-

Source : MNHN, Paris

168 ALYTES 5 (4)

@ R.tigerina at 17th day

© B. melanostictus at 10th day

0.6

O5

[e)

=

z 04

©

ui

3 03

>

Q

O

2 02

0.1

5 10 15 20 25 30 35 40

TEMP. °C

Fig. 1. — Temperature dependent growth of individual R. tigerina and B. melanostictus larvae.

Source : MNHN, Paris

HOTA & DASH 169

Table II. — Test of variances among B. melanostictus larvae after 10 days growth as a function

of different diet.

Nos. Diet Average body S.D.

mass (2)

1. Boiled Amaranthus tricolor leaves 0.086 0.017

2. Boiled Basella alba leaves 0.096 0.007

3. Cooked minced goat meat 0.109 0.009

4. Boiled chicken egg yolk 0.120 0.005

5. Mixed diet 0.137 0.008

F, max = 75.53 (ns)

Anova table

Variation df S.S. MS. F.

Treatments 4 0.00485 0.001213

LLTO**

Error 10 0.00103 0.000103

Total 14 0.00588

Coeff. det. = R2 = 0.83

#* Significant at 0.01 level

À posteriori test (S.N.K. test)

3 4 5

Q 3.15 3.88 4.33 4.66

L.S.R. 0.0184 0.0227 0.0253 0.0273

1<2<3<4<5

tion in metamorphic size. À comparison of Tables I, II and III suggests that the diet

that favours growth also favours quickening of the onset of metamorphic events and pro-

motes a greater size at transformation.

Table IV enumerates the effect of temperature on the time of metamorphosis and

metamorphic size. Within the range of 27° to 37°C lower temperature favoured greater

transformation size in both species. But the larvae reared at 15°C grew for a long time

into giant larvae and did not metamorphose, developing neotenic tendency. Ultimately

they could not adapt to permanent neoteny and died.

DISCUSSION

In frogs and toads there is usually a drastic change from aquatic herbivory to ter-

restrial carnivory, demanding reorganization of the gut during metamorphosis. Rana tige-

rina undergoes a “first metamorphosis” in the middle of its larval life when it changes

from being a herbivore to being a carnivore (VARUTE, 1970). Probably because of this

first metamorphosis in the. middle of larval life, the growth of the R. rigerina larvae with

Source : MNHN, Paris

170 ALYTES 5 (4)

Table III. — Metamorphic size and the time of metamorphosis as a function of diet.

Diet .. R. tigerina LE B. melanostictus

X(g) + S.D. Timein days X (8) + S.D. Time in days

Boiled À. tricolor leaves Did not metamorphose 0.129 + 0.007 15

Boiled B. alba leaves -do- 0.144 + 0.006 15

Cooked minced goat meat 0.751 + 0.016 34 0.164 + 0.009 15

Boiled egg yolk 0.755 + 0.026 28 0.180 + 0.008 15

Mixed diet 0.855 + 0.025 28 0.215 + 0.011 15

Table IV. — Metamorphic size and time of metamorphosis as a function of temperature.

Temperature es R. tigerina Fa B. melanostictus

°C X (g) + SD. Time in days X (g) + S.D. Time in days

37 0.750 + 0.035 28 0.130 + 0.009 16

33 0.858 + 0.016 28 0.137 + 0.002 15

27 0.050 + 0.048 29 0.169 + 0.012 15

15 Did not metamorphose.After Did not metamorphose. After

64th day high rate of mortality 48th day high rate of mortality

was observed and experiment was observed and experiment

was discontinued. was discontinued.

plant diets is checked and they cannot attain the minimum threshold size for completing

the second metamorphic step. In nature, this does not happen. R. rigerina larvae can satisfy

their carnivorous habits with a variety of insect larvae or other microinvertebrate prey

during this period. So in this study the mixed diet gave best results. In contrast, the gut

reorganization in the larvae of R. clamitans occurs after the emergence of the forelimbs,

when they stop feeding (JENSSEN, 1967). In this case the first and second metamorphic

processes are not distinguishable. In our experiment B. melanostictus larvae behaved like

R. clamitans, so that the plant diet did not affect the transformation as the gut reorganiza-

tion which occurs after metamorphosis is initiated.

In this study the results with controlled temperature follow the trend reported for

R. pipiens by SMITH-GILL & BERVEN (1979). Here also, the results show that the envi-

ronmental temperature is a major proximal factor in the growth of R. rigerina and B.

melanostictus larvae. As in the case of R. pipiens (SMITH-GILL & BERVEN, 1979) either

direct temperature effects on the developing tissue or indirect temperature effect media-

ted by thyroxine, are sufficient to explain temperature dependence of growth in these

two species.

It has been shown that metan:orphosis and the effects of thyroid hormones on other

species tadpoles are completely inhibited below temperature 5°C (HUXLEY, 1929 ; LYNN

& WACHOWSKI, 1951 ; FRIEDEN, WAHI.BORG & HOWARD, 1965 ; ASHLEY, KATTI &

Source : MNHN, Paris

HOTA & DASH 171

FRIEDEN, 1968). Similarly, in this study metamorphosis is inhibited at 15°C. The tro-

pical climate of Sambalpur provides a mean daily temperature of 20°C in winter, 30°C

in summer (maximum being 42°- 45°C), and also 30°C in the rainy season. So this rise

in the lower temperature tolerance of metamorphic process of R. tigerina and B. melanos-

tictus might be a compensatory adaptation to tropical climate. The length of the larval

stages of the American bull frog in nature increases with the length and severity of the

winters (WILLIS, MOYLE & BASKETT, 1956). ETKIN (1964) has observed that within the

range of 15°- 30°C metamorphosis is accelerated by increased temperature and the ani-

mals metamorphose at a small size. This appears to be partly true also with R. rigerina

and B. melanostictus. In this study, within the range of 27°- 37°C the lower temperature

favoured transformation at a greater size and higher temperature hastens the initiation

of metamorphic events and favours the transformation at a lower size. However, in this

investigation unlike that of the American bull frog, there is no significant increase in the

larval period at low temperature.

ACKNOWLEDGEMENT

The authors wish to thank Dr. Ilan MCLAREN for his critical comments. Finan-

cial support for this study from University Grants Commission, New Delhi (Grant No.

057/Bio-Scs/76) is greatefully acknowledged.

RÉSUMÉ

L'influence de l’alimentation et celle de la température sur la croissance et la méta-

morphose des larves d’Anoures sont étudiées à partir d'expériences effectuées sur Rana

tigerina et Bufo melanostictus. Cinq types de régimes alimentaires (composés uniquement

de végétaux, de viande, de jaune d’œuf, ou mixte) sont testés. Les meilleurs résultats sont

obtenus avec un régime mixte. Une alimentation qui favorise la croissance des têtards

a pour effet également d’accélérer le déclenchement de leur métamorphose et d’entraîner

une plus grande taille des métamorphosés. Entre 27° et 37°C un accroissement de la tem-

pérature accélère l’apparition de la métamorphose plus que la croissance larvaire de sorte

que les têtards ont tendance à se transformer à une taille inférieure. À 15°C la croissance

est favorisée aux dépens de la métamorphose de sorte que les larves grandissent au-delà

des valeurs limites habituelles et qu’elles manifestent une tendance vers la néoténie

facultative.

(Résumé rédigé par J.-J. MORÈRE)

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WieuR, H. M. & COLLINS, J. P., 1973. — Ecological aspects of amphibian metamorphosis. Science,

182 : 1305-1314.

Wizuis, Y. L., MOYLE, D. L. & BASKETT, T. S., 1956. — Emergence, breeding, hibernation, move-

ments and transformation fo the bullfrog, Rana catesbeiana, in Missouri. Copeia, 1956 : 30-41.

Source : MNHN, Paris

Alÿtes, 1986, 5 (4) : 173-174. 173

Miscellanea nomenclatorica batrachologica (XIV)

Alain DUBOIS

Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

The family-group name “Oreolalaxinae” Tian & Hu, 1985, which should

be emended to Oreolalaginae, is a strict synonym of Leptobrachiinae Dubois,

1980. This subfamily includes the genera and subgenera Leptobrachium, Lepto-

lalax, Scutiger, Oreolalax, Aelurolalax, and also Leptobrachella, which was erro-

neously placed by Tian & Hu (1985) in the Megophryinae.

TiAN & HU (1985) ont proposé la création d’une nouvelle sous-famille des Peloba-

tidae, regroupant les genres (ou sous-genres) Leptobrachium, Leptolalax, Scutiger, Oreola-

lax. Pour cette sous-famille, ils ont créé le nom “Oreolalaxinae”, fondé sur le nom géné-

rique Oreolalax.

11 faut tout d’abord noter que le nouveau nom de sous-famille est mal formé, et

doit être émendé en “Oreolalaginae”. En effet le nom Oreo/alax est fondé sur le mot grec

AGAGË, dont le génitif est A&\wyos, et le radical de ce nom générique est donc Oreolalag-.

De toute manière, ce nom du groupe-famille n’aura pas lieu d’être utilisé, car il

s’agit d’un strict synonyme plus récent du nom Leptobrachiinae Dubois, 1980. Ce der-

nier nom fut d’abord proposé (DUBOIS, 1980 : 471) sous la forme Leptobrachiini, pour

la tribu regroupant les genres à tétards ‘‘généralisés”” des Megophryinae, opposée à la

tribu des Megophryini, dont les têtards ont une bouche en entonnoir. Ce taxon fut élevé

au rang de sous-famille Leptobrachiinae par DUBOIS (1983 a : 272), et mentionné à diverses

reprises par la suite (DUBOIS, 1983 b : 147-148, 1984 : 29, 1985 : 74, 1987 : 13 ; FROST,

1985 : 409).

Notons enfin que c’est à tort que TIAN & HU (1985) incluent le genre Leptobra-

chella dans la sous-famille des Megophryinae : le têtard de Leprobrachella mjobergi décrit

en détail par INGER (1983) s’avère très proche des têtards de Leprolalax, ce qui indique

que la place de Leprobrachella est au sein des Leptobrachiinae.

Les Leptobrachiinae comportent donc les genres et sous-genres suivants (DUBOIS,

1987) : Leptobrachium Tschudi, 1838 (dont Vibrissaphora Liu, 1945 est synonyme) ; Lep-

tolalax Dubois, 1980; Leptobrachella Smith, 1925 ; Scuriger Theobald, 1868 ; Oreolalax

Myers & Leviton, 1962 ; et Aelurolalax Dubois, 1987.

Source : MNHN, Paris

174 ALYTES 5 (4)

RÉFÉRENCES BIBLIOGRAPHIQUES

Dugois, A., 1980. — Notes sur la systématique et la répartition des Amphibiens Anoures de Chine

et des régions avoisinantes. IV. Classification générique et subgénérique des Pelobatidae

Megophryinae. Bull. Soc. linn. Lyon, 49 : 469-482.

1983 a. — Classification et nomenclature supragénérique des Amphibiens Anoures. Bull. Soc.

linn. Lyon, 52 : 270-276.

x 1983 b. — Note préliminaire sur le genre Leprolalax Dubois, 1980 (Amphibiens, Anoures), avec

diagnose d’une espèce nouvelle du Vietnam. Alyres, 2 : 147-153.

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

nat., (A), 131 : 1-64.

1985. — Miscellanea nomenclatorica batrachologica (VII). Alyres, 4 : 61-78.

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

FROST, D. R., 1985. — Amphibian species of the world. À taxinomic and geographical reference. Law-

rence, Kansas, Allen Press & A.S.C. : [i-iv] + i-v + 1-732.

INGER, R. F., 1983. — Larvae of Southeast Asian species of Leprobrachium and Leptobrachella

(Anura : Pelobatidae). In : A. RHODIN & K. MIYATA (éds.), Advances in herpetology and evo-

lutionary biology, Cambridge, Mass. : 13-32.

TiaN, W. & HU Q., 1985. — Taxonomical studies on the primitive Anurans of the Hengduan

Mountains, with descriptions of a new subfamily and subdivision of Bombina. Acta herpet.

sin., 4: 219-224.

Source : MNHN, Paris

Alytes, 1987, 5 (4) : 175-176. 175

Miscellanea nomenclatorica batrachologica (XV)

Alain DUBOIS

Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

The generic name Ranixalus Dubois, 1986 is a synonym of Indirana Lau-

rent, 1986, which was proposed independently and published a few months ear-

lier. The name Ranixalini Dubois, 1987 remains the valid name for the tribe which

includes Indirana, Nyctibatrachus and Nannophrys.

Nous avons récemment (DUBOIS, 1986) décrit sous le nom Ranixalus gundia une

nouvelle espèce de Ranoïidea du Karnataka, dont nous avons ensuite (DUBOIS, 1987) mon-

tré qu’elle appartenait à un genre de Ranidae endémique du sud de l’Inde, comprenant

Polypedates beddomii Günther, 1876 et les espèces voisines. Pour ce genre, nous avons

initialement (DUBOIS, 1987) retenu le nom Ranixalus Dubois, 1986, mais le nom généri-

que /ndirana Laurent, 1986, proposé indépendamment et publié quelques mois plus tôt

par LAURENT (1986 : 761) pour Polypedates beddomiï et les espèces voisines, s’avère avoir

priorité. Toutes les espèces placées par DUBOIS (1987) dans le genre Ranixalus doivent

donc être rapportées au genre Indirana. Notons toutefois que quelques-unes des espèces

rapportées par LAURENT (1986) à ce genre appartiennent en fait à d’autres genres (pour

plus de détails, voir DUBOIS, 1987).

A l’examen des descriptions et figures que GÜNTHER (1876) et INGER et al. (1984)

ont donné de /ndirana brachytarsus (Günther, 1876) il nous paraît que cette espèce pour-

rait être la même que /ndirana gundia (Dubois, 1986), mais jusqu’à présent nous n'avons

pas eu la possibilité d'examiner les types de Polypedates brachytarsus, ou d’autres spéci-

mens rapportés à cette espèce.

En ce qui concerne enfin la nomenclature supragénérique, la mise en synonymie

de Ranixalus n'implique nullement la nécessité d'abandonner le nom Ranixalini Dubois,

1987, qui reste le nom valide de la tribu comportant les genres Zndirana, Nyctibatrachus

et Nannophrys (voir DUBOIS, 1987).

Source : MNHN, Paris

176 ALYTES 5 (4)

RÉFÉRENCES BIBLIOGRAPHIQUES

Dugois, À., 1986. — Diagnose préliminaire d’un nouveau genre de Ranoidea (Amphibiens, Anou-

res) du sud de l’Inde. A/yres, 4 : 113-118.

_— 1987. — Miscellanea taxinomica batrachologica (I). Ayres, 5 : 7-95.

GÜNTHER, À., 1876. — Third report on collections of Indian Reptiles obtained by the British

Museum. Proc. zool. Soc. Lond., 1875 : 567-577, pl. LXIII-LXVI.

INGER, R. F., SHAFFER, H. B., KOSHY, M. & BAKDE, R., 1984. — A report on a collection of

Amphibians and Reptiles from the Ponmudi, Kerala, South India. 7. Bombay nat. Hist. Soc.,

81 : 406-427.

LAURENT, R. F., 1986. — Sous-classe des Lissamphibiens (Lissamphibia). Systématique. /n : P.-

P. GRASSÉ & M. DELSOL (éds.), Traité de zoologie, Tome XIV, Batraciens, Fasc. 1-B, Paris,

Masson : 594-797.

Source : MNHN, Paris

Veste

ALYTES

Volume 5

("1986")

INDEX

Annemarie OHLER et Alain DUBOIS

Contents

Dates of publication of issues

Authors and titles index

Systematic index ..…

Index of new taxons

Subjects index

Geographic index

Referees

Source : MNHN, Paris

ü ALYTES

DATES OF PUBLICATION OF ISSUES

N° 1-2, "Mars-juin 1986" (pages 1-96):

13 May 1987.

N° 3, "Septembre 1986" (pages 97-152):

15 September 1987.

N° 4, “Décembre 1986" (pages 153-176):

1st October 1987.

AUTHORS AND TITLES INDEX

Dugois, A. - Miscellanea taxinomica

batrachologica (I)... 7-95

Dugois, A. - Miscellanea nomenclatorica

batrachologica (XII) 97-98

Dugois, A. - Living amphibians of the

world: a first step towards a

comprehensive checklist 99-149

Dugois, A. - Miscellanea nomenclatorica

batrachologica (XII) ............. 150

Dugois, A. - Miscellanea nomenclatorica

batrachologica (XIV) 173-174

Dugois, A. - Miscellanea nomenclatorica

batrachologica (XV) 175-176

HorA, A. K. & DaAsH, M. C. - Growth

and metamorphosis of anuran larvae:

effect of diet and temperature

. 165-172

INGER, R. F. - Diets of tadpoles living in

a Bornean rain forest .…... 153-164

LAURENT, R. F. - The systematic

position of the genus Afrixalus

Laurent (Hyperoliidae) 1-6

BOOK REVIEW

AMET, J.-L. - Un livre sur

les Amphibiens d'Australie

occidentale …................. 151-152

ANNOUNCEMENT

First World Congress of Herpetology . 96

Source : MNHN, Paris

Abrana: 130

cotti: 130

Acanthixalus: 4

Acris: 135-136

Adenomera: 128

marmorata: 128

ADENOMINAE: 24-25

Aelurolalax: 14, 173

Aelurophryne

brevipes: 22

gigas: 21-22

glandulata: 22

maculata: 16, 18

mammata: 17, 19, 21-22

tainingensis: 17

Afrixalus: 1-5, 35, 133

laevis: 3

Allophryne: 25, 122

ALLOPHRYNIDAE: 118, 122, 124

ALLOPHRYNINAE: 25, 122

Alsodes: 128

monticola: 128

ALSODINI: 118, 124

Altiphrynoides: 27

malcolmi: 27

Altirana: 39, 114

ALYTAE: 11

Alytes: 7, 12, 128

cisternasii: 12

muletensis: 12

obstetricans: 12

talaioticus: 12

Alytes (Alytes)

obstetricans: 12

obstetricans boscaï: 12

obstetricans maurus: 12

INDEX VOL. 5 ji

SYSTEMATIC INDEX

obstetricans obstetricans: 12

Alytes (Ammoryctis)

cisternasii: 12

Alytes (Baleaphryne)

muletensis: 12

talaioticus: 12

ALYTIDAE: 11, 124

ALYTINAE: 12

ALYTINI: 12

Ambystoma: 128, 136, 142

kl. platineum: 142

kl. tremblayi: 142

platineum: 142

AMBYSTOMATIDAE: 118, 124, 134, 138,

142

AMBYSTOMATOIDEA: 118, 120

AMBYSTOMATOIDEI: 120

Amietia: 49-50

Ammoryctis: 12

Amolops: 39-40, 51, 53, 57, 64, 76, 141,

162

afghanus: 141

chapaensis: 51

kaulbacki: 141

longimanus: 51

monticola: 142

phaeomerus: 155-161

torrentis: 53

AMPHIBIA: 7, 101-106, 108-109, 114,

117-118, 120-121, 126, 134, 139-

140, 142, 144

AMPHIGNATHODONTINAE: 29, 124

AMPHIUMIDAE: 124, 134, 138

AMPHIUMOIDEA: 118, 120

AMPHIUMOIDEI: 120

Ansonia: 25

longidigita: 155-159

ornata: 130

Source : MNHN, Paris

ANURA: 11, 105, 132, 134, 138, 140

APODA: 118

ARTHROLEPTIDAE: 1, 34-35, 123-124,

134, 138

ARTHROLEPTINA: 123

ARTHROLEPTINAE: 34, 123

Arthroleptis: 2, 100, 123

ASCAPHIDAE: 118, 124

ASTEROPHRYINAE: 34, 118, 124

Asterophrys: 128

ASTYLOSTERNINAE: 2, 34, 124

Atelognathus: 141

ATELOPODIDAE: 118, 124

ATELOPODINAE: 25

Atelopus: 25

Atympanophrys: 22-23, 114-115

Aubria: 39, 66, 114

subsigillata: 66

Babina: 42, 130

Baleaphryne: 12, 115, 128

muletensis: 12

Barbourula: 12, 130

busangensis: 130

busuangensis: 130

BATRACHIA: 117

Batrachophrynus: 2

patagonicus: 141

BATRACHYLINI: 118, 124

Batrachylodes: 39-40, 101

Bolitoglossa: 115

BOLITOGLOSSINI: 118, 124

Bombina: 12, 97-98

Bombina (Bombina)

bombina: 97

orientalis: 97

variegata: 97

Bombina (Grobina)

fortinuptialis: 98

maxima: 98

microdeladigitora: 98

ALYTES

Bombinator

australis: 133, 135

maximus: 97

sikimmensis: 19-20

BOMBINATORIDAE: 11, 118, 124

BOMBINATORINA: 11

BOMBINATORINAE: 11-12, 120

BOMBINATORINI: 12

BOMBINIDAE: 118, 124

BOMBININAE: 120

BOMBITATORES: 11

Boophis: 127

Bourretia: 61-62

BRACHYCEPHALIDAE: 100, 118, 124,

134, 138

Brachycephalus: 2

Brachymerus

bifasciatus: 136

Brachytarsophrys: 22-23, 114-115

BREVICIPINA: 126

BREVICIPITINA: 126

BREVICIPITINAE: 34, 123-124, 126

Buergeria: 127

Bufo: 25, 29, 115, 141

andersoni: 19

anotis: 27

arabicus: 130

beddomii: 130

bufo formosus: 140

divergens: 155-160

glaberrimus: 130

himalayanus: 140

hololius: 130

Jjaponicus: 140

Kelaartii: 141

koynayensis: 130

mammatus: 21-22

melanostictus: 165-171

orientalis: 130

osgoodi: 26

sikkimmensis: 20

stomaticus: 141

sulphureus: 130

vertebralis group: 29

Source : MNHN, Paris

INDEX VOL. 5

vulgaris: 104

Bufoides : 25

BUFONIDAE: 2, 7, 24, 28, 49, 122, 128,

130, 134, 138, 140

BUFONINAE: 25

C

CACOSTERNINAE: 122, 124

Caecilia

bivittata: 133

bivittatum: 133

glutinosa: 133, 135

hypocyanea: 133

CAECILNDAE: 117, 122, 124

CAECILUNAE: 117, 124

Calamita

tinctorius: 130, 133

Callixalus: 4

Callula

triangularis: 131

CALYPTOCEPHALELLINI: 118, 124

Capensibufo: 25-26

Cardioglossa: 2

Carpophrys: 129

Cassina: 37

obscura: 35, 38

wealii: 35-36

Cassiniopsis: 37

CAUDATA: 117-118

Caudiverbera: 135-136

peruviana: 135-136

CECILNDAE: 117, 122, 134, 139

CECILINNAE: 117

CENTROLENIDAE: 107, 124, 134, 138

CERATOBATRACHIDAE: 58

Ceratobatrachus: 2, 57, 67, 101

CERATOPHRYDES: 122, 127

CERATOPHRYIDAE: 122, 124

CERATOPHRYINAE: 124

Ceratophrys: 2, 130

boiei: 131

Chiromantis: 129

Chrysobatrachus: 4, 136

cupreonitens: 136

Clinotarsus: 42

COLODACTYLI: 11

COLODACTYLIDAE: 11

Colodactylus: 11

Colostethus: 136

Conraua: 2, 39, 50, 57

Cophophryne

alticola: 17, 18

sikkimensis: 19, 20

COPHYLIDAE: 127

COPHYLINAE: 34, 127

Cornufer: 52, 141

baluensis: 52-54

xizangensis: 53-54, 65

CORNUFERINAE: 38, 57-58

Crepidophryne: 25

Crinia: 127

bilingua: 141

deserticola: 141

remota: 141

signifera: 141

CRYPTOBRANCHIDAE: 124, 134, 138

CRYPTOBRANCHOIDEA: 118, 120

CRYPTOBRANCHOIDEI: 120

Cryptothylax: 4

CYCLORAMPHINI: 117, 120

Cycloramphus: 127, 131

Julginosus: 131

fuliginosus: 131

CYCLORANINAE: 118, 124

Cynops: 128

CYSTIGNATHI: 122

CYSTIGNATHIDAE: 124

Cystignathus

nodosus: 128

rhodonotus: 131

senegalensis: 37

D

DACTYLETHRIDAE: 121

Source : MNHN, Paris

vi ALYTES

DACTYLETHRINAE: 117, 120, 122, 124

Dendrobates: 2, 130, 133, 141

DENDROBATIDAE: 118, 121, 124, 127-

128, 130, 134, 138, 141

Dendrophryniscus: 25

DERMOPHIIDAE: 117, 120, 124

DERMOPHIINAE: 117-118, 124

DESMOGNATHIDAE: 118

DESMOGNATHINAE: 118, 124

DICAMPTODONTIDAE: 123, 134, 138

DICAMPTODONTINAE: 127

DICROGLOSSIDAE: 58

DICROGLOSSINI: 57, 64, 66-67

Dicroglossus: 57-58

adolfi: 57

Didynamipus: 25

Discodeles: 52-53, 57, 67, 101

ventricosus: 131

vogti: 131

DISCOGLOSSIDAE: 7, 11-12, 118, 121,

124, 128, 130, 132, 134, 138

DISCOGLOSSINAE: 12

DISCOGLOSSOIDEI: 11

Discoglossus: 11-12

Duellmania: 32

DYSCOPHIDAE: 127

DYSCOPHINAE: 34, 124

Dyscophus: 136

insularis: 136

E

Echinotriton: 11, 103, 129, 142

Elachistocleis: 128

Elachyglossa: 57

ELEUTHERODACTYLINI: 7, 23, 118, 124

Eleutherodactylus: 23-24, 115

auriculatus group: 23-24

coqui: 24

gaigei: 131

gaigeae: 131

jasperi: 23

lineatus: 131

orcutti: 24

ELOSHIDAE: 122

ELOSIINAE: 118, 120, 124

Engystoma: 128

marmoratum: 131

rugosum: 136

ENGYSTOMATINAE: 123

Eodiscoglossus: 12

EPICRNDAE: 117, 120, 134, 139

EPICRINAE: 117, 120

Eremiophilus: 37, 128

EUBAPHIDAE: 141

Euchnemis

fornasinii: 133

Euphlyctis: 2, 39, 57-60, 130, 134, 141

Euphlyctis line: 39-40, 54, 56-57, 66

Eupsophus: 134

Exaeretus: 133

caucasicus: 133

F

Fejervarya: 60-62, 142

G

GASTROPHRYNAE: 123

Gastrophryne: 133, 136

Gastrotheca: 7, 29-30, 32-33, 133, 136

argenteovirens: 33

argenteovirens group: 33

aureomaculata: 33

ernestoi: 31

marsupiata group: 32

medemi: 31

plumbea group: 32-33

Gastrotheca (Duellmania)

argenteovirens: 33

argenteovirens group: 33

aureomaculata: 33

cavia: 33

lojana: 33

Source : MNHN, Paris

INDEX VOL. 5

monticola: 33

psychrophila: 33

riobambae: 33

riobambae group: 33

Gastrotheca (Gastrotheca)

chrysosticta: 32

gracilis: 32

marsupiata: 32

peruana: 32

Gastrotheca (Opisthodelphys)

andaquiensis: 31

angustifrons: 31

bufona: 31

christiani: 31

cornuta: 31

cornuta subgroup: 31

dendronastes: 31

ernestoi: 31

excubitor: 31

Jissipes: 31

galeata: 31

griswoldi: 31

griswoldi group: 31-32

helenae: 31

humbertoi: 31

longipes: 31

longipes subgroup: 31

medemi: 31

microdiscus: 31

nicefori: 31

ochoaï: 31

orophylax: 32

ovifera: 31

ovifera group: 30

ovifera subgroup: 31

plumbea: 32

plumbea group: 32-33

testudinea: 31

viridis: 31

walkeri: 31

weinlandii: 31

williamsoni: 31

yacambuensis: 31

GENYOPHRYNINAE: 34, 120-121, 124,

vii

127

Glandula: 97

Glyphoglossus: 129

molossus: 131

Gorhixalus: 72

GRIPISCINI: 118

Grobina: 97

GRYPISCINI: 117, 120, 124

GYMNOPHIONA: 118, 132, 134, 139-140

H

Hammatodactylus: 128

HELEOPHRYNIDAE: 124, 132, 134, 138

HEMIDACTYLHNI: 120, 124

HEMIDACTYLINI: 117-118, 120

Hemidactylium: 127

HEMIMANTIDAE: 121

HEMIMANTINAE: 121

Hemimantis: 121, 141

HEMIPHRACTINAE: 29, 127

Hemiphractus: 2, 128, 136

spixii: 136

HEMISIDAE: 34, 124, 134, 138

Hemisus: 2

obscurus: 136

HERPELINAE: 118, 124

Heterixalus: 4

Hildebrandtia: 39, 55-56, 114

Hildebrandtia (Hildebrandtia)

macrotympanum: 56

ornata: 56

ornatissima: 56

Hildebrandtia (Lanzarana)

largeni: 56

Hoplobatrachus: 60-61

ceylanicus: 60

HOPLOPHRYNINAE: 123, 125

Hydromantes: 129

Hydromantoides: 129

Hydrophylax: 42

Hyla: 9, 109, 115, 141

albovitrata: 130

Source : MNHN, Paris

vii ALYTES

argenteovirens: 32

aurifasciata: 71

boans: 139

chinensis: 139

chrysoscelis: 165

cinerea: 130

erythraea: 130

iris: 143

leucotaenia: 130

marsupiata: 32, 136

meridionali.

reinwardi

strigilata: 136

zonata: 136

Hyla (Pseudacris)

nigrita floridensis: 141

HYLAEDACTYLI: 123

Hylambates: 1, 3-4, 36-37

leonardi: 38

viridis: 130

Hylaplesia: 133, 141

borbonica: 130

Hylarana: 39-40, 42, 50, 54, 57, 65, 73,

113, 130, 142

mindanensis: 63

Hylarthroleptis: 141

HYLIDAE: 7, 29, 107, 118, 120, 125,

128, 130, 134, 138, 141

HYLINAE: 118, 125

Hylodes: 122

lineatus: 130-131, 133

HYLODIDAE: 122

HYLODINAE: 119-120, 122, 125

HYLOIDEA: 23

Hylorana

longipes: 80-81

Hymenochirus: 136

HYNOBIIDAE: 119-120, 125, 134, 138

Hynobius: 128

HYPEROLNDAE: 1-5, 34, 127-128, 130,

134, 138

HYPEROLIINAE: 2, 4-5, 34-35

HYPEROLINI: 3-4, 34, 125

Hyperolius: 1-5, 109, 152

houyi: 143

Hysaplesia: 133, 141

I

ICHTHYOPHIIDAE: 117, 120, 125, 127

ICHTHYOPHNNAE: 117, 125

Ichthyophis: 133, 135

hasselti: 133

hasseltii: 135

Indirana: 175

brachytarsus: 175

gundia: 175

Ingerana: 64-65

baluensis: 64-65

liui: 64-65

mariae: 64

tasanae: 64-65

tenasserimensis: 64-65

xizangensis: 64

Ingerana (Ingerana):

baluensis: 65

mariae: 65

sariba: 65

tasanae: 65

tenasserimensis: 65

Ingerana (Liurana)

liui: 65

xizangensis: 65

Ixalus

beddomii: 132

chalazodes: 132

diplostictus: 132

fuscus: 51, 53-54

jerdonii: 73

nasutus: 132

opisthorhodus: 51, 53, 132

sarasinorum: 52-54

silvaticus: 51, 53-54

Source : MNHNI, Paris

Kalophrynus: 128

Kaloula

pulchra: 141, 161

KALOULIDAE: 122

Kassina: 1-5, 35-38, 128

arboricola: 37

cochranae: 37

lamottei: 37

senegalensis: 35

wealei: 36

Kassina (Kassina)

arboricola: 37

cassinoides: 37

cochranae: 37

decorata: 37

fusca: 37

kuvangensis: 37

lamottei: 37

maculata: 37

maculosa: 37

mertensi: 37

parkeri: 37

senegalensis: 37

wealii: 37

wittei: 37

Kassina (Paracassina)

Kkounhiensis: 38

obscura: 38

Kassina (Phlyctimantis)

keithae: 38

leonardi: 38

verrucosa: 38

KASSININAE: 1, 125

KASSININI: 1, 3-4, 7, 34-37

Kassinula: 3-4, 35-37, 115, 136

wittei: 136

Kirtixalus: 72

Lacerta

INDEX VOL. 5 ix

caudiverbera: 135

salamandra: 137

subviolacea: 136

Ladailadne: 23-24

jasperi: 24

Lanzarana: 39, 55-56, 115, 141

Latonia: 12

Laurentomantis: 136, 141

ventrimaculata: 139

Laurentophryne: 27

LEIOPELMATIDAE: 132, 134, 138

Leptobrachella: 13, 129, 173

mjobergi: 173

LEPTOBRACHIINAE: 13, 119, 125, 173

LEPTOBRACHINI: 173

Leptobrachium: 13, 18, 127, 158, 160-

162, 173

boringii: 131

boulengeri: 16-17

gracilis: 155-160

montanum: 155-161

pullum: 131

pullus: 131

LEPTODACTYLIDAE: 7, 23, 120, 122,

126-128, 130, 134, 138, 141

LEPTODACTYLINAE: 125

Leptodactylus: 109, 133

Leptolalax: 7, 13, 18, 173

dringi: 13-14

heteropus: 13-14

oshanensis: 129

pelodytoides: 14

Leptomantis: 75-16

bimaculata: 75

LEPTOPELINAE: 125, 127

LEPTOPELINI: 4, 34, 38

Leptopelis: 2, 4, 152

brevirostris: 38

Leptophryne: 25, 134

Limnodynastes: 133

peronii: 139

LIMNODYNASTINAE: 119, 125

Limnodytes

phyllophila: 51, 53-54

Source : MNHN, Paris

x ALYTES

Limnomedusa: 134 rugulosus: 60

Limnonectes: 43, 57-58, 60, 62-64, 130, tigerinus: 60

134, 141 tigerinus group: 60-61

cancrivorus: 61 verruculosus: 60

doriae: 61-62 Limnonectes (Limnonectes)

finchi: 62 acanthi: 63

grunniens: 62 arathooni: 63

grunniens group: 62 blythii: 63

hascheanus: 64 corrugatus: 63

kohchangae: 61 dammermani: 63

Kuhlii group: 62 diuata: 63

limborgii: 63 Jinchi: 63

macrodon: 62 fragilis: 63

microdiscus group: 62 grunniens: 63

tigerinus group: 60 grunniens group: 63

Limnonectes (Bourretia) heinrichi: 63

dabanus: 62 ibanorum: 63

doriae: 62 ingeri: 63

kohchangae: 62 Kenepaiensis: 63

macrognathus: 62 Khammonensis: 63

mawphlangensis: 62 Khasiensis: 63

pileatus: 62 kuhliï: 63

plicatellus: 62 kuhlii group: 63

toumanoffi: 62 laticeps: 63

Limnonectes (Fejervarya) leytensis: 63

andamanensis: 61 macrocephalus: 63

greenii: 61 macrodon: 63

Kkeralensis: 61 magnus: 63

limnocharis: 61 malesianus: 63

murthii: 61 micrixalus: 63

nepalensis: 61 microdiscus: 63

nilagirica: 61 microdiscus group: 63

pierrei: 61 microtympanum: 63

rufescens: 61 modestus: 63

syhadrensis: 61 namiyei: 63

teraiensis: 61 nitidus: 63

vittiger: 61 palavanensis: 63

Limnonectes (Hoplobatrachus) paramacrodon: 63

cancrivorus: 60 parvus: 63

crassus: 60 timorensis: 63

demarchii: 61 tweediei: 63

occipitalis: 61 visayanus: 63

occipitalis group: 61 woodworthi: 63

raja: 60 Limnonectes (Taylorana)

Source : MNHN, Paris

INDEX VOL. 5

hascheanus: 64

limborgii: 64

LISSAMPHIBIA: 117-118

Lithodytes: 130, 133-134

Litoria: 115, 127, 152

bicolor: 152

microbelos: 152

nasuta: 152

rothii: 152

tornieri: 152

Liurana: 65

Lynchophrys: 2

M

MANTELLINAE: 34, 67, 114, 125

Mantidactylus: 115

MEANTES: 118

Megalixalus: 1, 128

fornasinii congicus: 133

MEGALOPHREIDINA: 123

Megalophrys: 15

boulengeri: 15, 17

weigoldi: 14

MEGOPHRYIDAE: 123

MEGOPHRYINAE: 13, 22-23, 102, 114,

119-120, 125, 129, 173

MEGOPHRYINI: 173

Megophrys: 13, 22-23, 114, 130, 159-

160, 162

minor: 160

montana: 131

monticola: 23, 131

nasuta: 155-161

oshanensis: 129

pachyprocta: 131

pachyproctus: 131

parva: 131, 141

Megophrys (Atympanophrys)

shapingensis: 23

Megophrys (Brachytarsophrys)

carinensis: 23

Megophrys (Megophrys)

xi

montana: 23

pachyproctus: 23

MELANOBATRACHINAE: 34, 123, 125

Melanophryniscus: 25

Mertensiella: 133

Mertensophryne: 28-29, 141

micranotis: 28-29, 49

rondoensis: 141

Micrarthroleptis: 141

MICRHYLIDAE: 125

Micrixalus: 39-40, 51-54, 57, 64, 141

borealis: 52-53, 59

diminutivus: 53, 59

Juscus: 52, 55

magnapustulosus: 53

mariae: 53-54

nudis: 52-53, 55

opisthorhodus: 54

phyllophilus: 54-55

saxicola: 55

silvaticus: 55

thampii: 52-53, 55

torrentis: 53

Microhyla: 128, 162

borneensis: 155-160

ornata: 141

petrigena: 155-160

MICROHYLIDAE: 2, 10, 34, 119-120,

122, 127-128, 131, 134, 138, 141,

154, 160

MICROHYLINAE: 34, 119, 127

MICROHYLOIDEA: 33

Microphryne: 136, 141

malagasia: 136

Mocquardia: 35, 38

MOLGINAE: 120

MYCETOGLOSSINI: 117, 120

MYOBATRACHIDAE: 125, 127, 134, 138,

141

MYOBATRACHINAE: 127

Source : MNHN, Paris

xii ALYTES

N Nyctimantis papua: 136

Nyctixalus margaritifer: 139

Nannobatrachus: 67-68 Nyctymystes: 136

anamallaiensis: 68

Nannophrys: 66-68, 175

ceylonensis: 68 oO

guentheri: 68

marmorata: 68 Occidozyga: 53, 57-60, 64

Nanorana: 39, 114 cyanophlyctis: 59

Nectophryne: 27 hexadactyla: 59

afra: 27 lima: 58-59

tornieri: 26 Occidozyga (Euphlyctis)

Nectophryne line: 25, 27 cornii: 59

NECTOPHRYNINI: 27, 29 cyanophlyctis: 59

Nectophrynoides: 25-27, 141 ehrenbergii: 59

cryptus: 26 hexadactyla: 59

malcolmi: 27 Occidozyga (Occidozyga)

minutus: 26 lima: 58

occidentalis: 9, 27 ODONTOPHRYNINI: 119, 125

osgoodi: 9 Ololygon: 133, 136

tornieri: 26 Onychodactylus: 127

viviparus: 26 Ooeidozyga: 53, 58

Necturus: 133 Ophryophryne: 13, 22-23, 114, 130

lateralis: 133 microstoma: 23, 130

Nimbaphrynoides: 27 poilani: 23

liberiensis: 27 Opisthodelphys: 30

occidentalis: 27, 49 Opisthothylax: 2, 4

Notaden: 2 Orchestes: 71, 142

Notodelphys: 30 OREOLALAGINAE: 173

ovifera: 30 Oreolalax: 13-15, 18-19, 113, 130, 173

Notokassina: 4, 35, 37 OREOLALAXINAE: 173

Nyctibatrachus: 66-69, 175 Oreophrynella: 25

aliciae: 68 Osornophryne: 25

beddomii: 68 Osteocephalus: 136

deccanensis: 68 taurinus: 136

humayuni: 68 Osteopilus: 134

Khempholeyensis: 68 Oxydozyga: 58

major: 68 Oxyglossus

minor: 68 laevis: 58

modestus: 68

sanctipalustris: 68

sanctipalustris var. modestus: 68

sinensis: 63

sylvaticus: 68

Source : MNHN, Paris

INDEX VOL. 5 xii

P

Paa: 40, 43-45, 60, 130

Palmatorappia: 57, 101

Paracassina: 35, 37-38

kounhiensis: 35

obscura: 35

Pararthroleptis: 141

nanus: 141

Parkerana: 55-56, 130

Pedostibes: 25, 128

tuberculosus: 130

Pelobates: 13

PELOBATIDAE: 7, 13, 22, 114, 123, 125-

126, 129, 131, 134, 138, 141, 173

PELOBATINA: 126

PELOBATINAE: 13

PELOBATOIDEA: 13

PELODRYADIDAE: 119, 125

PELODRY ADINAE: 119, 125

PELODYTIDAE: 125, 132, 134, 138

PELODYTINAE: 125

Pelophryne: 25

Peltophryne: 25, 134

Petropedetes: 121

PETROPEDETINAE: 117, 121-122, 125

PHILAUTINAE: 125, 127

PHILAUTINI: 69

Philautus: 68-72, 142

annandalii: 142

aurifasciatus: 69-70, 72

dubius: 73

hosii: 70

lissobrachius: 72

microtympanum: 73

pleurostictus: 73

schmackeri: 72

Philautus (Gorhixalus)

hosii: 72

Philautus (Kirtixalus): 74, 76

dubius: 73

Jerdonii: 73

microdiscus: 73

microtympanum: 73

pleurostictus: 73

Philautus (Philautus)

acutirostris: T2

aurifasciatus: 72

emembranatus: 72

leucorhinus: 72

lissobrachius: 72

nasutus: 72

parvulus: 72

schmackeri: 72

surdus: 72

Phlyctimantis: 1-4, 35-38

Phryniscus

australis: 135

PHRYNOBATRACHINAE: 34, 117, 121, 125

Phrynobatrachus: 2, 121, 141

Phrynoglossus: 52-53, 57-59

baluensis: 59

borealis: 52, 59

borealis group: 59

celebensis: 59

diminutivus: 52, 59

floresianus: 59

laevis: 59

laevis group: 59

magnapustulosus: 59

martensii: 59

semipalmatus: 59

vittatus: 59

Phrynohyas: 133, 136

PHRYNOMERIDAE: 125

PHRYNOMERINAE: 34, 125

Phrynomerus: 136

Phyllobates: 128

bicolor: 130

latinasus: 136

PHYLLOBATIDAE: 125

PHYLLOMEDUSIDAE: 119, 125

PHYLLOMEDUSINAE: 119, 125

Physalaemus

biligonigerus: 141

biligonigerus group: 141

cuvieri group: 141

nattereri: 141

Source : MNHN, Paris

xiv ALYTES

Pipa: 2

PIPIDAE: 120, 122, 125, 131, 134, 138

PIPINAE: 125

PIPOIDEI: 13

PIPRINA: 127

PLATYMANTINAE: 58, 125

PLATYMANTINI: 39, 57

Platymantis: 40, 52, 57, 64, 101, 141

liui: 53-54

Plethodon: 128, 135

serratus: 143

PLETHODONTIDAE: 125, 127, 134, 138

PLETHODONTINAE: 125

PLETHODONTINI: 125

PLETHODONTOIDEA: 118, 120

PLETHODONTOIDEI: 120

Pleurodeles: 7, 10-11, 142

walil: 10

Pleurodeles (Echinotriton)

andersoni: 11

asperrimus: 11

chinhaiensis: 11

hainanensis: 11

Pleurodeles (Pleurodeles)

poireti: 11

waltl: 11

Pleurodeles (Tylototriton)

kweichowensis: 11

taliangensis: 11

verrucosus: 11

PLEURODELINAE: 10

Pleurodema: 127

Polypedates: 74-76, 127, 135

appendiculatus: 70, 74-75

beddomii: 132, 175

biscutiger: 84

brachytarsus: 132, 175

cavirostris: 132

formosus: 51, 132

hascheanus: 52, 63

Jjerdonii: 73, 132

leucomystax: 131, 142

maculatus: 142

microtympanum: 52, 69-70, 72, 74

nasutus: 132

rufescens: 132

saxicola: 51, 53

POLYPEDATIDAE: 119, 125

PROTEIDAE: 125, 134, 139

PROTEOIDEA: 118, 120

PROTEOIDEI: 120

Proteus: 136

anguinus: 136

Pseudacris: 9, 133, 136, 141

nigrita verrucosa: 141

Pseudarthroleptis: 141

PSEUDIDAE: 107, 125, 132, 134, 138

Pseudobufo: 25, 128

Pseudohemisus: 136

Pseudophryne: 2, 133, 135

australis: 133

semimarmorata: 135

vivipara: 25

Pseudotriton: 127, 135-136

nigra: 135

Ptychadena: 39, 55-56, 114, 130, 152

PTYCHADENINI: 55

PYXICEPHALINI: 66

Pyxicephalus: 39, 66, 114, 127

adspersus: 66

delalandii: 56

fodiens: 57

frithi: 57, 61

pluvialis: 57

Ramanella: 131

simbiotica: 131

symbioitica: 131

symbiotica: 131

Rana: 38-42, 44, 50, 52-53, 55, 57, 60,

64-65, 67, 76, 100-101, 113-115,

130-131, 135, 141-142, 158, 162

aenea: 43

agricola: 60-61

aliilabris: 61

Source : MNHN, Paris

areolata group: 41

assimilis: 61

beddomii: 52

beddomii group: 67

berlandieri group: 41

bicolor: 130

blanfordii: 142

blythi: 155-159

bombina: 97

bonaspei: 131

boulengeri: 44

bourreti: 47

boylii group: 41

bragantina: 60

breviceps group: 40, 142

brevipalmata: 61

burkilli: 60

catesbeiana group: 41

chalconota: 155-159

clamitans: 165, 170

clamitans group: 41

conspicillata: 63

cornuta: 130

crassa: 142

dalmatina: 131

dauchina: 131

daunchina: 131

delacouri: 48, 139

dobsonii: 57

doriae group: 61

esculenta: 142

esculenta group: 41, 142

esculenta synklepton: 105

fasciata: 133

fasciculispina: 48

feae: 45-47

Jusca: 63, 104, 133, 142

Jfuscigula: 139

Juscigula group: 42, 50

gibbosa: 128

gracilis: 61

gracilis var. nicobariensis: 61

gracilis var. pulla: 60-61, 132

grunniens group: 142

INDEX VOL. 5

gryllus: 135-136

holsti: 130

hubeiensis: 143

humeralis: 142

hydraletis: 60

hydromedusa: 63

ibanorum: 155-159

kl. esculenta: 142

kuhlii: 60, 62, 130, 139

leptodactyla: 52, 68-69

leschenaultii: 59, 130

leucorhynchus: 57, 142

liebigit: 46, 130

liebigii group: 43-44

lima: 58

limnocharis: 61, 131

limnocharis group: 40

limnocharis mysorensis: 61

lineata: 130, 133

longimanus: 51

macrodon complex: 142

macrodon var. leporina: 63

mascareniensis: 55

mediolineata: 48

microdisca: 62

microdisca superspecies: 62

microlineata: 48

moluccana: 139

montezumae group: 41

moodiei: 60

mortenseni: 131

muta: 150

nantaiwuensis: 60

nigrita: 136

nigrovittata: 131, 142

occipitalis subgroup: 60

ovalis: 128

palavanensis: 42

Palmipes group: 41-42

paradoxa: 63

parambikulamana: 61

perezi: 161

phrynoides: 45, 47

picta: 60

Source : MNHN, Paris

bPipiens: 170

Pipiens complex: 41, 142

Pipiens group: 41, 142

polunini: 142

pullus: 52, 132

pygmaea: 68, 132

quadrana: 113, 131

quadranus: 48, 131

rostandi: 46

rufescens: 40

rufescens group: 40, 142

sanguinea: 139

sariba: 52-54

sauriceps: 61, 142

schlueteri: 60

scutigera: 81

semipalmata: 68-69

shini: 142

sichuanensis: 47

signata: 155-159

sikimensis: 142

spinosa: 47-48, 60, 142

spinosa group: 44

spinosa supergroup: 48

spinosa verrucospinosa: 44

subsaltans: 63

swani: 57

taipehensis: 142

tarahumarae group: 41

tasanae: 52-54

temporaria: 39-40, 150

temporaria group: 41

tenasserimensis: 52-54, 64-65

tibetana: 44

tigerina: 165-171

tigerina subgroup: 60

tigrina var. angustopalmata: 60

tigrina var. pantherina: 60

tinctoria: 130, 133

toumanoffi: 61

travancorica: 68

typhonia: 133

umbraculata: 42

ALYTES

unculuana: 113, 131

unculuanus: 131

variegata: 57

ventricosa: 131

ventricosus: 131, 143

verrucosa: 132

vertebralis: 42, 49-50

wasl: 61

yunnanensis: 45-47

yunnanensis group: 44

Rana line: 39-40, 54-56

Rana (Amietia)

umbraculata: 49

vertebralis: 49

Rana (Chaparana)

fansipani: 131

Rana (Euphlyctis)

cyanophlyctis group: 58

Keralensis: 131

tigerina group: 60

Rana (Hylarana): 73, 76

Rana (Paa)

aenea: 43

annandalii: 43

arnoldi: 43, 131

blanfordii: 43

boulengeri: 43-44

bourreti: 46

conaensis: 43, 49

delacouri: 44, 47-49

delacouri group: 44, 47

ercepeae: 43

exilispinosa: 43

fasciculispina: 44, 47

feae: 43, 46

hazarensis: 43, 131

liebigii: 43

liebigii group: 43-44

liebigii supergroup: 43

liui: 150

maculosa: 43

maculosa group: 43

minica: 43, 131

polunini: 43

Source : MNHN, Paris

quadranus: 44, 47, 49

rara: 43

rostandi: 43

shini: 43

sichuanensis: 47

sikimensis: 43, 49

sikimensis group: 43

spinosa: 43

spinosa group: 43-44

spinosa supergroup: 43-44, 47

sternosignata: 43-45

vicina: 43

yunnanensis: 43, 45-46

yunnanensis group: 43-44, 150

Rana (Rana)

amieti: 42

angolensis: 42

areolata group: 42

aurora group: 41

berlandieri subgroup: 42

boylii group: 41

catesbeiana group: 41

clamitans group: 41

desaegeri: 42

dracomontana: 42

Juscigula: 42

Juscigula group: 42

Jjohnstoni: 42

kl. esculenta group: 41

lateralis: 42

lateralis group: 42

montezumae subgroup: 42

nigrolineata: 42

Palmipes group: 42

palustris subgroup: 42

Pipiens group: 41

Pipiens subgroup: 42

pleuraden: 41

pustulosa group: 42

rugosa: 42

rugosa group: 42

ruwenzorica: 42

shuchinae: 42

sylvatica: 41

INDEX VOL. 5 xvii

tarahumarae group: 42

temporaria group: 41

tientaiensis: 42

unculuanus: 42, 49

wittei: 42

Rana (Strongylopus)

bonaespei: 50, 131

fasciata: 50

grayii: 50

hymenopus: 50

wageri: 50

Ranae

beddomianae: 66

Ranae chalconotae: 51

Ranae grunnientes: 61-62

Ranae hexadactylae: 58

Ranae kuhlianae: 62

Ranae tigrinae: 58

RANIDAE: 7, 34, 38, 67, 69, 102, 119-

120, 125, 129, 131, 134, 138, 141,

175

Ranidella: 141, 152

RANINAE: 7, 34, 38-40, 42, 54-57, 64,

66-67, 73, 76, 101-102, 113-114,

119, 125

RANINI: 39-40, 50, 54-55

RANIXALINAE: 67

RANIXALINI: 66-67, 175

Ranixalus: 64, 66-67, 69, 175

beddomii: 67, 69

brachytarsus: 69

diplostictus: 69

gundia: 66-67, 69, 175

leithii: 69

leptodactylus: 69

phrynoderma: 69

semipalmatus: 67, 69

tenuilingua: 69

RANOIDEA: 7, 33-34, 66, 175

RANOIDEI: 23

RHACOPHORIDAE: 34, 51-52, 69, 114,

119-120, 125, 129, 132, 134, 138,

142

Source : MNHN, Paris

xvii ALYTES

RHACOPHORINAE: 7, 10, 34, 69, 73-74,

119

RHACOPHORINI: 74

Rhacophorus: 19, 51, 70, 74-78

appendiculatus: 76

bimaculatus: 75, 155-159

bimaculatus group: 74-75, 162

buergeri chapaensis: 51

dubius: 73

dulitensis: 155-160

everetti: 70, 74

gauni: 75

harrissoni: 155-159

hosei: 70

hosii: 69-70, 72

Jjavanus: 76

Kajau: 70, 74-75

leucomystax: 18-79, 81

leucomystax megacephalus: 85

macrotis: 80

maculatus: 78-80, 84-86

maculatus himalayensis: 78, 81, 85

maculatus maculatus: 78-79, 84, 86

maximus: 142

microdiscus: 73

microtympanum: 10

moschatus: 76, 132

mutus: 80

naso: 75

nigropalmatus: 155-159

oxycephalus: 75

pleurostictus: 73

taeniatus: 79

translineatus: 716

verrucosus: 75

zed: 86

Rhacophorus (Leptomantis): 76

bimaculatus: 76

gauni: 76

oxycephalus: 76

sp. KB: 76

sp. NT: 76

Rhacophorus (Rhacophorus)

annamensis: 77

appendiculatus: 77

appendiculatus group: 77

arboreus: 77

bipunctatus: 77

bisacculus: 77

calcadensis: 77

chenfui: 77

chenfui group: 77

colletti: 77

cruciger: 17

dennysii: 77

dennysi group: 77

dugritei: 77

dugritei group: 77

dulitensis: 77

eques: 17

fasciatus: 77

fasciatus group: 77

feae: 77

georgii: 77

harrissoni: 17

hungfuensis: 77

kajau: 77

leucomystax: 77, 19

leucomystax group: 77-78

leucomystax leucomystax: 80

leucomystax megacephalus: 81

leucomystax teraiensis: 81

longinasus: 77

macrotis: 77

maculatus: 77, 82

maculatus biscutiger: 84

maculatus himalayensis: 84

maculatus maculatus: 83

malabaricus: 77

malabaricus group: 77

maximus: 77-78

moltrechti: 77

mutus: 77

nigropalmatus: 77

notater: 77

omeimontis: 77

otilophus: 77

owstoni: 717

Source : MNHN, Paris

INDEX VOL. 5 xix

pardalis: 77

pardalis group: 77

prasinatus: 77

prominanus: 49, 77

reinwardtii: 71

reinwardtii group: 77

rhodopus: 77

schlegelii group: 77

taeniatus: 77, 79

taipeianus: 77

verrucopus: T7

viridis: 77

yaoshanensis: 77

zed: 86

Rhamphophryne: 25

Rhinatrema: 133

RHINATREMATIDAE: 125, 127, 134, 139

Rhinoderma: 2

RHINODERMATIDAE: 125, 134, 138

RHINOPHRYNIDAE: 119, 125, 134, 138

Rhinophrynus: 2

dorsalis: 119

RHYACOTRITONIDAE: 123

RHYACOTRITONINAE: 125, 127

Rothschildia: 35, 38

Kkounhiensis: 35

S

Salamandra: 136-137

genei: 129

maculosa: 136

subfusca: 135-136

SALAMANDRIDAE: 7, 10, 119, 125, 134,

139, 142

SALAMANDROIDEA: 118, 120

SALAMANDROIDEI: 10, 120

Scaphiophryne: 136

marmorata: 136

SCAPHIOPHRYNIDAE: 34

SCAPHIOPHRYNINAE: 126

SCAPHIOPODIDAE: 119, 126

SCAPHIOPODINAE: 13, 119, 126

Scaphiopus: 13, 130

bombifrons: 130

Schismaderma: 28-29

carens: 28

Schoutedenella: 2

SCOLECOMORPHIDAE: 119, 126, 134, 139

Scutiger: 2, 7, 13-16, 19-20, 113, 130,

173

adungensis: 15, 18

alticola: 17-18

boulengeri: 15-16, 18

glandulatus: 15

mammatus: 15, 18, 22

mammatus group: 16

nepalensis: 15-16

nyingchiensis: 15-16, 19

occidentalis: 19

pingit: 15, 130

15-16, 18, 21

sikimmensis group: 15-16

sikkimensis: 19

weigoldi: 15

Scutiger (Aelurolalax)

weigoldi: 15

Scutiger (Oreolalax)

popei: 15

Scutiger (Scutiger)

boulengeri: 16

glandulatus: 22

mammatus: 21

mammatus group: 21

nepalensis: 21

nyingchiensis: 19

sikimmensis: 19

sikimmensis group: 16

Semnodactylus: 35, 37

thabanchuensis: 35

wealii: 35

SIPHONOPIDAE: 117, 120

SIPHONOPINAE: 117

Source : MNHN, Paris

xx ALYTES

Sirena

maculosa: 133

SIRENIDAE: 126, 134, 139

Sminthillus: 2

Somuncuria: 136

SOOGLOSSIDAE: 134, 138

SOOGLOSSINAE: 127

Spea: 130

Sphaenorhynchus: 128

Sphaeroteca: 56-57

strigata: 56-57

SPHENOPHRYNINAE: 120-121, 126

Spinophrynoides: 26-27

osgoodi: 26

Staurois: 39-40, 57, 64, 141

Stenorhynchus: 141

Stephopaedes: 27-29

anotis: 28

STEPHOPAEDINAE: 29

STEPHOPAEDINI: 27

Stombus: 130

Strongylopus: 39-40, 50, 101, 114-115,

127, 133, 135

Synapturanus: 135

Systoma: 129

T

Tachycnemis: 1-2, 4, 133

Taylorana: 63-64

TELMATOBII: 127

TELMATOBIIDAE: 126

TELMATOBIINAE: 23, 38, 127

TELMATOBINI: 126

Telmatobius

somuncurensis: 136

Theloderma: 127

Tomopterna: 39-40, 56, 101, 113-114,

141-142

breviceps: 56

rolandae: 56

strachani: 57

Tomopterna line: 39-40, 56

Tomopterna (Sphaeroteca)

breviceps: 57

labrosa: 57

rolandae: 57

Tomopterna (Tomopterna)

cryptotis: 56

delalandii: 56

krugerensis: 56

marmorata: 57

natalensis: 57

tuberculosa: 57

TOMOPTERNINI: 56

Tornierella: 3-4, 35-38, 115, 137

pulchra: 137

Tornieriobates: 25

TORNIERIOBATINAE: 7, 25

TORNIERIOBATINI: 25, 29

Trachycephalus: 128

Trachymantis: 129, 141

TRACHYSTOMATA: 118

Triton: 137

cristatus: 137

major: 135

palmatus: 104

TRITURINAE: 120

Triturus: 129, 137

Tylototriton: 11, 142

verrucosus: 10

TYPHLONECTIDAE: 126-127, 134, 139

U

Uperodon: 131

globulosus: 141

Uperoleia: 2

URAEOTYPHLINAE: 126

URODELA: 10, 117, 119, 132, 134, 138

140

Vibrissaphora: 173

Source : MNHN, Paris

INDEX VOL. 5 xxi

Werneria: 27

Wolterstorffina: 27

XENOPINAE: 122

XENOPODA: 121

XENOPODINAE: 117, 120, 122, 126

Xenopus: 105

boettgeri: 136

bunyoniensis: 131

(laevis) bunyoniensis: 131

wittei: 131

INDEX OF NEW TAXONS

Aelurolalax: 14

Altiphrynoides: 27

Amietia: 49

Bourretia: 61

Duellmania: 32

Gastrotheca (Duellmania)

group: 33

Gastrotheca (Opisthodelphys) griswoldi

group: 31

Gorhixalus: 72

Grobina: 97

Ingerana: 64

Kirtixalus: 63

Ladailadne: 23

Leptolalax dringi: 13

Liurana: 65

Nimbaphrynoides: 27

PTYCHADENINI: 55

Rana (Paa) bourreti: 46

Rana (Paa) delacouri group: 44

Rana (Paa) liui: 150

Rana (Paa) maculosa group: 43

Rana (Paa) sichuanensis: 47

Rana (Paa) spinosa group: 43

Rana (Paa) yunnanensis group: 43

Rana (Rana) fuscigula group: 42

Rana (Rana) lateralis group: 42

Rana (Rana) rugosa group: 42

riobambae

RANIXALINI: 66

Rhacophorus (Rhacophorus) appen-

diculatus group: 77

Rhacophorus (Rhacophorus) chenfui

group: 77

Rhacophorus (Rhacophorus) dennysii

group: 77

Rhacophorus (Rhacophorus) dugritei

group: 77

Rhacophorus (Rhacophorus) fasciatus

group: 77

Rhacophorus (Rhacophorus) leucomystax

group: 77

Rhacophorus (Rhacophorus) leucomystax

teraiensis: 81

Rhacophorus (Rhacophorus) malabaricus

group: 77

Rhacophorus

group: 77

Rhacophorus (Rhacophorus) reinwardtii

group: 77

Rhacophorus

group: 77

Rhacophorus (Rhacophorus) zed: 86

Spinophrynoïdes: 26

STEPHOPAEDINI: 27

Taylorana: 63

TOMOPTERNINI: 56

(Rhacophorus) … pardalis

(Rhacophorus) schlegelii

Source : MNHN, Paris

xxii ALYTES

SUBJECTS INDEX

Altitude: 13, 17-21, 30-33, 81-86

Amplexus: 5, 29, 49

Aneuchrony: 8, 30, 58

Anus: 48, 49

Body mass: 166-171

Buccopharingeal morphology: 13, 30-32,

69, 71-73, 153, 159, 160

Call: 71

Catalogue: 99-145

Chromosomes: 5

Classification: 7-86

Co-ossification: 78-82, 86

Colour: 5, 15, 18, 46, 78-86

Development: 8, 9, 23-27, 29, 30, 32,

33, 54, 62-65, 69, 70-74

Diet: 5, 37, 38, 153-163, 165-171

Distribution: 13, 15-21, 31-33, 38, 41,

42, 44-48, 50-57, 66, 67, 78-86

Ecology: 8, 12, 18, 34, 37, 50, 56, 75,

84, 153

Eggs: 4, 25-27, 54, 55, 62-65, 69, 71-74

Evolution: 25, 71

Eye: 5, 50

Faunistics: 151-152

Feeding behavior: 158-159

Fertilization: 24, 29, 30, 49

Food particle size: 156-158

Forelimbs: 3, 5, 46, 55, 64, 71-73

Fossil: 12

Genetics: 8

Glands: 2, 4, 5, 14, 15, 55, 64, 66, 67,

73

Growth rate: 165, 166

Habitat: 50, 154, 155, 160-162

Head: 16, 21, 46-48, 50, 61, 78-86

Hindlimbs: 5, 15, 45-48, 50, 58, 64, 71-

73, 78-86

Hybridizability: 9

Hybridization: 10, 12

Incubation pouch: 29-32

Intersexuality: 44, 45, 59

Larvae: 13, 22, 25-28, 32, 33, 50, 55,

58, 59-61, 66, 67-70, 74-76, 79,

152-163, 165-171

Larval anatomy: 13, 28

Larval feeding modes: 158, 159

Larval teeth: 4, 5, 26, 27, 50, 75, 76

Lateral line: 58, 60

Life history: 8, 165

Limbs: 5, 29

Metamorphosis: 165-171

Morphological variation: 18, 20, 33

Morphology: 8, 12, 23, 34

Musculature: 2, 4, 5, 25, 27, 71-74

Nomenclature: 7-86, 97-98, 150, 173,

175

Nomenclature family group taxons: 9, 10

Oral morphology: 28, 55, 58, 68, 69,

153, 159, 160

Osteology: 1, 3, 4

Paedomorphosis: 2-4

Pectoral girdle: 3, 5, 25, 27, 39, 40, 50,

53-55, 57, 59, 60, 64, 66, 71, 72

Pelvic girdle: 55, 56

Phylogeny: 1-5,8, 35, 39, 40, 56, 57, 66,

67

Reproduction: 8, 9, 23-27, 29, 30, 32,

33, 54, 63, 65, 70, 71

Reproductive behavior: 62

Review: 99-145, 151-152

Sampling: 154, 155

Secondary sexual characters: 14, 15, 16,

26, 29, 44, 45, 47-49, 55, 59-61,

63, 64, 67, 71-73, 78-86

Size: 1, 5, 13-16, 21, 22, 30, 32, 44-48,

50-52, 54, 58, 64, 65, 73, 74, 78-

86, 165

Skin: 5, 16, 18, 46-48, 54, 55, 64

Source : MNHN, Paris

INDEX VOL. 5 xxiii

Skull: 2-5, 32, 39, 50, 52, 55, 56, 57,

59, 60, 63, 66, 71, 72

Species recognition: 80-86

Systematics: 1

Taxinomy: 7-86

Teeth: 2, 5, 14, 15, 18 54, 59, 64, 71, 72

Temperature: 165-171

Toe dilatations: 3, 5, 14, 34, 36, 38, 40,

46-48, 54, 58, 59, 61, 64, 65

Tongue: 15, 54, 58, 59, 64, 65

Tympanum: 14, 15, 47, 54, 59, 64

Vertebral column: 2, 3, 5, 55, 71-73

Vocal sacs: 1-5, 14, 16

Webbing: 3, 5, 14-16, 18, 36, 46, 47,

50, 54, 58, 59, 64, 65

GEOGRAPHIC INDEX

Afghanistan: 44, 45

Africa: 25, 29, 35, 38-42, 55, 56, 57,

66, 152

America: 41

Argentina: 30, 32

Asia: 38, 40, 42, 56, 57, 102, 162

Australia: 151

Bangla Desh: 82

Bhutan: 20

Borneo: 13, 52, 53, 69, 70, 74, 75, 80,

81, 153-163

Brasil: 31

Burma: 46, 51, 53, 82

Cameroon: 66

Central Africa: 152

Central America: 56

Ceylon: see Sri Lanka

China: 15-19, 21, 22, 41, 45-47, 53, 75,

80, 81, 85, 100

Colombia: 30, 31, 33

Congo: 66

East Asia: 100

Ecuador: 30, 31, 33

England: 161

Ethiopia: 27

India: 19, 20, 51-54, 56, 66, 70, 73, 78,

79, 82-85, 171, 175

Indochina: 80, 85

Java: 80

Liberia: 66

Madagascar: 56, 67

Majorca: 12

Malaya: 13, 70, 80, 81

Nepal: 18-21, 78, 79, 81, 83-86

Nimba Mountain: 27

North America: 41

Pacific: 42

Pakistan: 44

Panama: 31

Peru: 30-33

Philippines: 53, 69, 70, 72, 80

Puerto Rico: 23

South Africa: 29, 50, 151

South East Asia: 24, 75, 162

Spain: 154, 161

Sri Lanka: 52, 67, 70, 84

Sumatra: 80

Tanzania: 26

Thailand: 43, 53, 153, 161

Tropical Asia: 57, 100

United States: 161

Venezuela: 31

Vietnam: 45-48, 80, 81

Western Australia: 151, 152

Source : MNHN, Paris

xxiv ALYTES

REFEREES

The editors of Alytes thank warmly the following colleagues, who accepted to

read, study and comment the papers submitted for publication in Alytes from 1st July

1985 to 30 June 1987:

Edouard-Raoul BRYGO0 (Paris) Pierre JOLY (Lyon)

Stephen D. BUSACK (Urbana) Raymond F. LAURENT (Tucumän)

Ronald I. CROMBIE (Washington) Jan MCLAREN (Halifax)

William E. DUELLMAN (Lawrence) Annemarie OHLER (Wien)

Carl GANS (Ann Arbor) Georges PASTEUR (Montpeller)

Jacqueline GAVAUD (Paris) Manuel POLLS PELAZ (Paris)

Jean-Daniel GRAF (Genève) Jean-Paul RisCH (Luxembourg)

Claude P. GUILLAUME (Montpellier) Josef Friedrich SCHMIDTLER (München)

W. Ronald HEYER (Washington) Richard WASSERSUG (Halifax)

Marinus S. HOOGMOED (Leiden) Earl WERNER (Ann Arbor)

© ISSCA 1993

Source : MNHN, Paris

AINTES

Journal International de Batrachologie

International Journal of Batrachology

édité par la Société Batrachologique de France

Rédacteurs : Alain DUBOIS et Jean-Jacques MORÈRE.

Adresse : Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire naturelle, 25 rue Cuvier,

75005 Paris, France.

Comité de rédaction : Jean-Louis AMIET (Yaoundé), Stephan D. BUSACK (Urbana), Benedetto LANZA

(Firenze), Raymond F. LAURENT (Tucumän), Michael J.TYLER (Adelaide), Richard J.

WASSERSUG (Halifax).

Recommandations aux auteurs. — Alytes publie des articles originaux en français ou en anglais, consa-

crés aux Amphibiens. Les manuscrits doivent être dactylographiés et accompagnés d’un résumé

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çais (pour ceux qui le souhaiteraient, les rédacteurs acceptent de revoir les résumés en fran-

çais à partir d’un texte en anglais). Tableaux et figures doivent comporter un titre. Les figu-

res, exécutées à l’encre noire, ne devront pas dépasser le format 16 x 24 cm. Indiquer leur

numéro au crayon ; légendes sur feuille séparée. Présenter les références bibliographiques

conformément au dernier numéro d” A/yres paru (les références de livres doivent comporter

la pagination). Adresser les manuscrits en trois exemplaires aux rédacteurs. L’acceptation

d’un article pour publication est décidée par les rédacteurs après lecture critique de celui-ci

par deux lecteurs ou plus.

Instructions to authors. — Alytes publishes original papers in English or in French, dealing with Am-

phibians. Manuscrits should be typewritten, and preceded by an English abstract. Papers in

English should be followed by a detailed French summary (for those who may wish so, the

editors accept to revise such French summaries on the basis of an English text). Tables and

figures should possess titles. Figures should be drawn in black ink and should not exceed

16 x 24 cm in size. Their numbers should be written in pencil. Figure captions should be

assembled on a separate sheet. Bibliographic references should be presented as in recent issues

of Alytes (book references should include the pagination). Send the manuscripts in triplicate

to the editors (address above). Acceptance for publication will be decided by the editors fol-

lowing review by two referees or more.

Tirés à part. — 25 exemplaires gratuits par article. Au-delà, les tirés à part seront facturés par tran-

ches de 25 exemplaires.

Publié avec le concours du Muséum national d'Histoire naturelle.

Directeur de la Publication : Alain DUBOIS.

Numéro de Commission Paritaire : 64851.

Source : MNHIN, Paris:

6 OCT. 1987

Alytes, 1986, 5 (4) : 153-176.

SOMMAIRE

Robert F. INGER

Diets of tadpoles living in a Bornean rain forest ..…

Ashok K. HOTA & Madhab C. DASH

Growth and metamorphosis of anuran larvae : effect of diet and

temperature

Alain DUBOIS

Miscellanea nomenclatorica batrachologica (XIV) ............

Alain DUBOIS

Miscellanea nomenclatorica batrachologica (XV) .............

Photocomposition/Photogravure : Alexandre, Paris. 42 46 17 57.

Imprimé aux Ateliers de la Couronnerie, 45750 Saint-Pryvé Saint-Mesmin, France.

Dépôt légal : 3° trimestre 1987.

153

165

173

175

Source : MNHN, Paris: