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AITES

INTERNATIONAL JOURNAL OF BATRACHOLOGY

May 1996 Volume 14, N° 1

Alytes, 1996, 14 (1): 1-26. 1

Two new Telmatobius species

(Leptodactylidae, Telmatobiinae

of Ancash, Peru

Antonio W. SALAS * & Ulrich SINSCH **

* Museo de Historia Natural, Universidad Ricardo Palma, Av. Benavides cdra. 54,

Santiago de Surco, Lima, Peru

** Institut für Biologie, Universität Koblenz-Landau, Rheinau 1, 56075 Koblenz, Germany

The taxonomic status of two populations of telmatobiine frogs in the

Peruvian department Ancash is evaluated using data from external morphol-

ogv. The intrapopulational variation of 18 morphometric measures is com-

pared with those of six telmatobiine species from adjacent regions: Batracho-

Phrynus brachydactylus, B. macrostomus, Telmatobius brevirostris, T. car-

rillae, T. jelskii and T. rimac. The frogs inhabiting the Laguna Conococha and

those of the Rio Sihuas are distinct from the already described species and

from each other. They represent two new species of the genus Telmatobius.

Diagnostic features of external morphology and skin histology are given to

distinguish among the central Peruvian Telmatobiinae.

Bibliothèque Centrale Muséum

INTRODUCTION 3 3001 00111592 1

The streams and lakes of the central and northern regions of the Peruvian Andes are

inhabited by leptodactylid frogs of the genera Telmatobius Wiegmann, 1835 and

Batrachophrynus Peters, 1873 (DUELLMAN, 1979; SinscH, 1990; WiEns, 1993). Batracho-

phrynus is endemic to central Peru, whereas Telmatobius has a widespread distribution

ranging from Ecuador in the north to Chile and Argentina in the south (Cet, 1986). The

taxonomic status of populations of telmatobiine frogs is difficult to evaluate, if only based

on external morphology, because shape and coloration are usually similar due to the

adaptation to the mostly riparian habitats. The original descriptions and diagnoses of most

species are inadequate and the taxonomic classification of populations often requires

comparison with type material. In many taxa, multivariate statistics such as discriminant

analyses are necessary to distinguish between intraspecific and interspecific variation of

Source : MNHN, Paris

2 ALYTES 14 (1)

morphometric measures (WIEns, 1993; SiNSCH et al., 1995). In others, genetic markers such

as allozyme loci are used to assess the specific status (WIENs, 1993). These approaches were

recently used to evaluate the status of several populations inhabiting the Andean regions

of the northern Peruvian departments of Amazonas, Cajamarca, La Libertad and Piura

CWiEns, 1993) and of the central Peruvian departments of Ancash, Ayacucho, Cerro de

Pasco, Huancavelica, Huanuco, Junin and Lima (SINsCH et al., 1995; SiNsSCH & JURASKE,

1995). The known species of the north-Peruvian departments are Telmatobius brevipes

Vellard, 1951, T. ignavus Barbour & Noble, 1920, T. latirostris Vellard, 1951, and the six

species recently described by WIEns (1993), T. atahualpai, T. colanensis, T. degener, T.

necopinus, T. thompsoni and T. truebae. The reassessment of the status of the central

Peruvian populations led to the recognition of Batrachophrynus brachydactylus Peters,

1873, B. macrostomus Peters, 1873, Telmatobius brevirostris Vellard, 1955, T. carrillae

Morales, 1988, T. jelskii (Peters, 1873) and T. rimac Schmidt, 1954.

Information on the Telmatobiinae inhabiting the department of Ancash are still

scarce. The few populations which have been treated taxonomically have been assigned to

three species of Telmatobius: T. rimac (VELLARD, 1955; MORALES, 19884), T. jelskii

(MORALES, 1988a) and T. carrillae (MORALES, 1988b; SALAS, 1990). The most recent

checklist of amphibian species of Peru (RODRIGUEZ et al., 1993) recognizes only the

occurrence of T. carrillae and T. rimac in Ancash. A thorough survey of the amphibians

of this region during eight years (1986-1994) by the senior author revealed the existence

of two populations of telmatobiine frogs which apparently differed in some characters of

external morphology from these species and others known to inhabit the adjacent central

Peruvian departments (SALAS, unpublished data). These observations motivated the senior

author to reevaluate the taxonomic assignment of telmatobiine frogs collected in Ancash

and preserved in local collections. The morphometric analysis (using the classification

criteria of SiNsCH et al., 1995) of the specimens collected in the Rio Huaylas and assigned

to T. jelskii by MORALES (1988a) showed that they had been confounded with T. rimac

(SALAS, in preparation). In contrast, the populations of telmatobiine frogs collected in the

Laguna Conococha and in the Rio Sihuas remained unidentified, though the first

superficially resembles Batrachophrynus brachydactylus, and the second Telmatobius

carrillae. Both populations differ in several aspects from all other species and also between

each other.

The aims of this paper are to: (1) establish the distinction of the populations

inhabiting the Laguna Conococha and the Rio Sihuas from the already described species

of this region; (2) describe two new species; and (3) justify their inclusion in the genus

Telmatobius.

MATERIAL AND METHODS

The material examined included adult frogs pertaining to six previously known species

(Batrachophrynus brachydactylus, N = 53; B. macrostomus, N = 13; Telmatobius

brevirostris, N = 5; T. carrillae, N = 43, T. jelski, N = 71; T. rimac, N = 42), and 25

unclassified specimens which were collected in the Laguna Conococha, Provincia Recuay,

Source : MNHN, Paris

SALAS & SINSCH 3

Department of Ancash, Peru (13 adults, 2 subadults), and in the Rio Sihuas, Provincia

Sihuas, Department of Ancash, Peru (10 adults), respectively. The geographical distribu-

tion of the collection sites are shown in figure 1. The detailed list of specimens of the 6

previously known species, with their localities and museum collections, was already

published by SiNsCH et al. (1995: 43-44, Appendix I). As for the 25 unclassified specimens,

their detailed list is given below under the two newly described species. Institutional

abbreviations are as follows: KU, Museum of Natural History, The University of Kansas,

USA; MHNSM, formerly MHNJP, Museo de Historia Natural, Universidad Nacional

Mayor de San Marcos, Lima, Peru; URP, Museo de Historia Natural, Universidad

Ricardo Palma, Lima, Peru; ZFMK, Zoologisches Forschungsmuseum Alexander Koenig,

Bonn, Germany.

Standard morphometric measurements were recorded from all adult specimens to the

nearest 0.1 mm with needle-tipped calipers, as in WiENS (1993) and SInsCH et al. (1995).

Note that measurements of limbs refer to the portions of body containing the bones: (1)

SVL, snout-vent length; (2) BH, height of body (at the pectoral girdle); (3) HWID,

maximum width of head; (4) EYE, eye diameter; (5) IOD, interorbital distance; (6)

ENOSE, eye-nostril distance (anterior margin of eye to posterior edge of naris); (7)

ESNOUT, distance between the eye and the tip of the snout; (8) HUML, humerus length

(upper forelimb); (9) RADL, radioulnar length (lower forelimb, elbow to distal edge of

outer palmar tubercle); (10) HNDL, hand length (proximal edge of outer palmar tubercle

to tip of third finger); (11) FG3L, length of the third finger; (12) FEML, femur length

(thigh); (13) TIBL, tibia length (shank, knee to heel);, (14) FOOTL, foot length (from

union with tibia to the tip of fourth toe); (15) TOEIL, length of first toe; (16) TOE4L,

length of fourth toe; (17) CIL, length of callus internus; (18) WEBL, maximum length of

toe web between III and IV (from the middle of the web, i.e. lowest part, to the union of

the toes).

Multivariate analyses were performed on log,ç-transformed data (BOOKSTEIN et al.,

1985) and morphometric ratios. The empiric measurements were transformed to ratios

(range: 0-1) by calculating measures relative to SVL (SINsCH et al., 1995). Moreover, two

indices were used for further analysis: CIL/TOEIL and FEML/TIBL.

The first step of classification was to calculate the discriminant scores for the adult

specimens from the Laguna Conococha and Rio Sihuas using the discriminant functions

published by SINsCH et al. (1995), which distinguish among Batrachophrynus brachydacty-

lus, B. macrostomus, Telmatobius brevirostris, T. carrillae, T. culeus, T. jelskii and T. rimac.

The second step consisted in subjecting sets of the log,,-transformed data to principal

component analysis to explore the morphometric variability independent of taxonomic

assignment. Data sets were: (1) Batrachophrynus brachydactylus, B. macrostomus, taxon

from Rio Sihuas and taxon from Laguna Conococha; (2) Telmatobius brevirostris, T.

carrillae, T. jelsküi, T. rimac, taxon from Rio Sihuas and taxon from Laguna Conococha.

Principal components (PC) are linear combinations of the measured variables, uncorre-

lated with each other and explaining the maximum amount of variation. The first principal

component (PCI) of morphometric data generally describes differences in size, but size

effects may be present in subsequent principal components (HUMPHRIES et al., 1981).

Techniques such as shearing have been developed to correct PC2 and PC3 for possible size

Source : MNHN, Paris

4 ALYTES 14 (1)

effects (BOOKSTEIN et al., 1985), but they are controversial and size effects may still persist

(RoLr & BookSTEIN, 1987). Therefore, we present the uncorrected PC2 and PC3. The

next step consisted in a stepwise canonical discriminant analysis to distinguish between the

taxonomic groups delimited a priori. We used stepwise forward selection of variables

(criterion to enter: F = 4.0) to minimize the number of variables needed for group

distinction. The resulting discriminant functions (CAN: canonical variables) are linear

combinations of those measured variables that maximize the differences between the

groups. Discriminant functions were derived from the log,,-transformed data. The final

step of analysis was to look for diagnostic morphometric ratios which differ significantly

among the known species and the taxa from Conococha and Sihuas. We applied a multiple

range test using the Least Square method and a significance level of 1 %. AI calculations

were performed on a Pentium PC using the program package STATGRAPHICS Plus,

version 1.4.

The descriptions of the new species follow the format of TRUEB (1979) and WIENS

(1993). The diagnosis only distinguishes among the species included in this paper. The

formulae for toe webbing follow SAVAGE & HEYER (1967) as modified by Myers &

DUELLMAN (1982).

RESULTS AND DISCUSSION

CLASSIFICATION WITH THE DISCRIMINANT FUNCTIONS WHICH DISTINGUISH AMONG THE

CENTRAL PERUVIAN SPECIES

The morphometric features of the adult frogs which were collected in the Laguna

Conococha and in the Rio Sihuas are listed in Tables I and II. The corresponding data

for Batrachophrynus brachydactylus, B. macrostomus, Telmatobius brevirostris, T. carrillae,

T. jelskii and T. rimac have been published by SiNsCH et al. (1995: Tables I-IT).

Eighteen log,,-transformed morphometric characters were used to obtain discriminant

functions which distinguish among the described telmatobiine species of central Peru

(SiNsCH et al., 1995: Tables III-IV). The first step of classification consisted in calculating

the scores for the adult individuals of the Conococha and Sihuas samples using these

discriminant functions. If the unclassified frogs are conspecific with any of the described

central Peruvian species, we expect that the discriminant scores are completely or at least

to a large amount within the known ranges of these species.

The discriminant scores based on the functions which distinguish among Batracho-

phrynus brachydactylus, B. macrostomus, Telmatobius brevirostris and T. carrillae are

shown in figure 2. AI scores of the specimens from Rio Sihuas are placed outside the

variation of any of the known species with respect to CANI and CAN2. In contrast, the

scores of the Conococha individuals completely overlap with the range of variation of T.

brevirostris. However, the scores obtained using CAN3 distinguish both Conococha and

Sihuas specimens from T. brevirostris. In a three-dimensional plot of these discriminant

functions there is no overlap of the distributions obtained for the samples from Laguna

Source : MNHN, Paris

SALAS & SINSCH 5

Conococha and from Rio Sihuas with that of Batrachophrynus brachydactylus, B.

macrostomus, Telmatobius brevirostris or T. carrillae. In conclusion, the telmatobiine frogs

of the unclassified populations remain unidentified and are probably not conspecific with

any of these species.

The same analysis was done applying the discriminant functions which distinguish

among T. culeus, T. jelskii and T. rimac. The distribution of scores obtained for the frogs

of the two unclassified populations does not overlap with the range of T. culeus, but some

scores are inside the ranges of T. jelskii and T. rimac (fig. 3). Nevertheless, most scores of

both populations are outside the ranges of either T. jelskii or T. rimac, especially those of

the frogs from Laguna Conococha. These results do not suggest that the unclassified frogs

pertain to either species, but due to the slight overlap conspecificity cannot entirely be

ruled out. However, the frogs of Rio Sihuas are not only morphometrically similar to T.

Jjelskii and T. rimac, but also share the presence of yellow-orange patches on the ventral

side of the thigh with these two species.

Only T. rimac is known to occur in Ancash, in three localities along the occidental

cordillera (SALAS, in preparation), whereas the nearest locality of a T. jelskii population is

situated more than 300 km south of the unclassified populations (VELLARD, 1955; SINSCH

et al., 1995). The centres of distribution of T. jelskii are clearly the more southern

departments of Ayacucho, Junin and Huancavelica. Considering our limited knowledge on

the distribution of most Peruvian Telmatobiinae, the biographical argument against the

conspecificity with T. jelskii is admittedly weak.

Finally, we have to consider the characters related to sexual maturity. A diagnostic

character for T. jelskii among the central Peruvian Telmatobiinae is the presence of horny

excrescences on the chest of reproductive males. This feature is not shared by the males

collected in the Laguna Conococha and in the Rio Sihuas. The minimum size of the

Conococha adults is about 67 mm SVL (Table I); two smaller individuals (54 mm and 57

mm SVL) were still sexually immature. At all localities and elevations so far known, T.

jelskii and T. rimac reach maturity at a considerably smaller size: 47 mm and 42 mm SVL,

respectively. In contrast, the size distribution of the Rio Sihuas frogs clearly falls within

the range of these species.

In conclusion, the morphometric data indicate that the taxon inhabiting the Laguna

Conococha is certainly not conspecific with any of the described central Peruvian species.

The taxon occurring in the Rio Sihuas is certainly distinct from Batrachophrynus

brachydactylus, B. macrostomus, Telmatobius brevirostris, T. carrillae, T. jelskii and T.

culeus, but some individuals cannot be morphometrically distinguished from T. rimac.

MORPHOMETRIC DISTINCTION OF THE UNCLASSIFIED TAXA FROM THE CENTRAL PERUVIAN

SPECIES

In the second step of classification, we applied principal component and discriminant

analyses to distinguish the unidentified populations from described central Peruvian

species. Analyses were performed on two data sets: (1) Batrachophrynus brachydactylus, B.

macrostomus and the samples from Rio Sihuas and Laguna Conococha; (2) Telmatobius

Source : MNHN, Paris

6 ALYTES 14 (1)

brevirostris, T. carrillae, T. jelskü, T. rimac and the samples from Rio Sihuas and Laguna

Conococha.

Generally, the interspecific differences in size (PC1) by far exceeded those in shape

(PC2, PC3). The size effects on PC2 and PC3 appeared to be small, because shearing

showed little effect. Discriminant analysis led to an optimal separation of species by

combining differences in size and shape.

In the data set used to distinguish the Conococha and Sihuas taxa from the

Batrachophrynus species, the first three principal components explained 95.4 % of the total

variance. PCI accounting for 88.9 % of total variance separates the large B. macrostomus

from the smaller B. brachydactylus and the unidentified taxa. The plot of PC2 (3.9 % of

total variance) and PC3 (2.6 % of total variance) scores shows that the scores of the

similar-shaped B. brachydactylus and B. macrostomus form one completely overlapping

group, and those of the Conococha and Sihuas taxa another group (fig. 4A). The slight

overlap between the two groups is due to scores of the Conococha taxon, whereas the

scores of Sihuas taxon vary outside the range of the Batrachophrynus species. A perfect

separation of the four taxa — 100 % of the specimens correctly classified — was obtained

by stepwise discriminant analysis (fig. 4B, Table III). The taxa are distinguished based on

only four out of 18 variables: FG3L, HUML, RADL and TOEAL, i.e. parameters of limb

morphology.

In the data set used to distinguish the Conococha and Sihuas taxa from the central

Peruvian Telmatobius species, the first three principal components accounted for 84.4 %

of the total variance. PCI accounting for 71.4 % of total variance separates the small T.

carrillae from the larger taxa. The plot of PC2 (8.1 % of total variance) and PC3 (4.9 %

of total variance) scores shows a complete separation of Conococha taxon from T.

brevirostris, T. jelskii and T. rimac, but a considerable overlap with T. carrillae and the

Sihuas taxon (fig. SA). The best separation of the six taxa was obtained by discriminant

functions based on a set of 13 out of 18 variables (Table IV). As five discriminant

functions are necessary to separate six taxa, a presentation in a single plot would require

five dimensions. Therefore, we present, as an example, a plot of CANI versus CAN2

which distinguishes T. carrillae and T. jelskii from all other species (fig. 5B). Based on five

discriminant functions, 94.3 % of all specimens were correctly classified. The erroneous

classifications were: 1 out of 53 T. carrillae which was confounded with T. rimac; 5 out

of 71 T. jelskii which were confounded with T. brevirostris, T. rimac and the Sihuas taxon,

respectively; 5 out of 42 T. rimac which were confounded with T. brevirostris and T. jelskii,

respectively. Thus, none of the unidentified specimens was confounded with a known taxon.

In conclusion, the analyses presented demonstrate that the two samples of unidenti-

fied telmatobiine frogs represent morphometrically well-defined taxa which can be

distinguished without erroneous classification from the six sympatric Batrachophrynus and

Telmatobius species, and from each other.

TAXONOMIC DECISIONS AND GENERIC ASSIGNMENT

The taxa inhabiting the Laguna Conococha and the Rio Sihuas, respectively, possess

unique characters that easily and consistantly separate them from the other central

Source : MNHN, Paris

SALAS & SINSCH 7

Peruvian Telmatobiinae (external morphology: figs. 6-7; skin histology: HEIN, 1994; HEIN

& SinscH, 1995; SnscH & HEIN, in preparation). Moreover, there is no indication that any

of the unidentified taxa in the department Ancash is conspecific with the north Peruvian

Telmatobius species which inhabit the Andes near the Huancabamba depression (WIENS,

1993; WiENs, personal communication; SALAS, unpublished observations). Therefore, we

conclude that the telmatobiine frogs of the populations inhabiting the Laguna Conococha

and the Rio Sihuas are members of new species.

The generic assignment of the new taxa to Telmatobius is based on the following

considerations. In central Peru, the Telmatobiinae are represented by the genera

Telmatobius and Batrachophrynus. There are two presumptive synapomorphies for the

monophyly of Telmatobius (WIENs, 1993): frontoparietals fused posteriorly, and nuptial

excrescences on finger I only. In contrast, evidence for the monophyly of Batrachophrynus

is based on allozymes, and on diagnostic features such as the absence of maxillary and

prevomerine teeth, and nuptial pads without horny excrescenses (PETERS, 1873; LYNCH,

1978; SINsCH & JURASKE, 1995). A/sodes is assumed to be the sister taxon of Te/matobius

(Lyc, 1978), though the only presumptive synapomorphy is the presence of an enlarged

crista medialis on the humerus in males (WIEns, 1993). However, allozymes and skin

morphology rather indicate that Telmatobius and Batrachophrynus are sister taxa (HEIN &

SiNscH, 1995; SiNsCH & JURASKE, 1995; SNsCH & HEIN, in preparation): (1) NErs genetic

distances between the species of these genera are low; (2) Telmatobius and Batrachophrynus

share the presence of granular glands with small granules which are absent in A/sodes (4.

montanus), (3) Telmatobius (except for T. carrillae) and Alsodes share the presence of

granular glands with large granules, but granules and gland structure are very different in

the two genera (SINSCH & HEIN, in preparation); (4) Telmatobius and Batrachophrynus

share the absence of nuptial excrescences on finger II which are present in A/sodes.

Analyzing the character states considered as diagnostic for the genera Alsodes, Batracho-

phrynus and Telmatobius in the two new taxa, we find: (1) horny nuptial excrescences are

present only on finger I; (2) maxillary and premaxillary teeth are present; (3) two types of

granular glands (small and large granules) are present. A conservative evaluation of these

character states suggests a provisional inclusion of the new taxa in the genus Telmatobius.

Further comparative studies on allozymes, osteological and histological characters are

needed and in work to test the validity of this assignment.

ACCOUNT OF THE NEW SPECIES

Telmatobius hockingi sp. nov.

(figs. 8-9)

Holotype. — URP 116, adult male, from Rio Sihuas 5 km from Sihuas, Provincia Sihuas,

Departamento Ancash, Peru, 2700 m altitude, 77°38‘14"W 08°30'00"S, collected on 19

december 1992 by Antonio W. SALAS.

Paratypes. — URP 112-115 and 117-119, 3 males and 4 females; ZFMK 57260, 1 male;

KU 220844, 1 female; all collected at the same site simultaneously with the holotype by

Antonio W. SALAS.

Source : MNHN, Paris

8 ALYTES 14 (1)

Diagnosis. — (1) Premaxillary teeth present; (2) tympanum absent; (3) nuptial spines

moderately small on the dorsal and ventral surfaces of the thumb; nuptial pads continuous

with inner palmar tubercle; (4) dorsum brownish grey (in preservative) with small patches;

(S) venter dark cream with diffuse grey; (6) forelimbs and hindlimbs always without

ornamentation or transverse bars; (7) dorsal skin smooth; (8) snout-vent length in males

to 52.5 mm, in females to 64.8 mm.

This species resembles in habitus the riparian Te/matobius (fig. 8). Confusion with the

sympatric Batrachophrynus species is impossible due to the difference in adult size, the

easily noticeable premaxillary teeth, and the presence of nuptial excrescences and of

granular glands with large granules in the dorsal skin. Moreover, the morphometric ratios

HWID/SVL and FG3L/SVL are diagnostic for the distinction of T. hockingi from

Batrachophrynus (fig. 6). T. hockingi differs from T. brevirostris, T. jelskii and T. rimac by

the ratio HUML/SVL, and from T. carrillae and the new species described below by the

ratio FG3L/SVL (fig. 7). The yellow-orange patches on the ventral side of the thigh

distinguish T. hockingi from T. brevirostris, T. carrillae and the new species described

below.

Description. — Head slightly narrower than body; head wider than long: HLEN 88.3 %

of HWID; head length 30.4 % of SVL; head width 34.4 % of SVL. Dorsal view of snout

rounded, in lateral profile gently sloping (fig. 9A). Nostrils not protuberant, located at the

extreme anterior terminus of snout, anterolaterally oriented. Canthus rostralis indistinct

dorsally, in lateral profile short and elevated; loreal region concave. Eyes protuberant on

top of head, eye diameter 29.3 % of head length. Tympanum absent, tympanic annulus

conspicuous. Supratympanic fold present and well developed, extending from posterior

corner of eyelid to the anteroventral insertion of forelimb. Maxillary and premaxillary

teeth embedded in labial mucosa, fanglike and protruding, but easily noticeable when

passing on top with finger tips. Dentigerous processes of vomer well developed, five times

closer to choanae than to each other, located anterior to choanae; choanae about the same

size and circular. Tongue rounded with slightly elevated lateral borders, posteriorly free.

Vocal slits absent.

Robust, stout forelimbs. Dermal wrist fold present, but inconspicuous. Fingers

uniform in diameter, long and slender; I and II separated due to well developed muscles

at the palmar region of insertion. Relative length of fingers: III > IV > I > II (fig. 9B),

tips of fingers round to spherical, palmar webbing absent. In males, large and raised

nuptial pad covering the dorsal and lateral surface of thumb; nuptial spines, moderately

large, conical, keratinized. Inner palmar tubercle oval, continuous with nuptial pad. Outer

palmar tubercle oval and large, but smaller than the inner, located proximally on fingers

IT and III. Conspicuous, supernumerary tubercles close to the base of fingers I and II.

Subarticular tubercle present proximally on each finger except III, smaller subarticular

tubercles in the middle of each finger and distal ones in III and IV.

Robust, but slender hind limbs. Hind limb length (foot plus tibia) 41.5 % of SVL.

Relative length of toes (fig. 9C): IV > II > V > I > I; webbing formula: I 1 — 2+

IH2— — 32/3 III 2+ — 3— IV 3- — 1 V; webbing diminishes gradually to form a

lateral fringe along the edge of toe IV. Tips of toes spherical and of the same size as finger

tips. Inner metatarsal tubercle small, oval and slightly raised; outer metatarsal tubercle

Source : MNHN, Paris

SALAS & SINsCH 9

round, 1/3 length of inner. Small, round subarticular tubercles distributed on toes as

follows: I(1), II(1), II(2), IV(3) and V(2). Tarsal fold extending to 1/3 length of tarsus,

confluent with lateral fringe of toe I.

Dorsal, ventral and lateral skin smooth. Ventral skin covered with few and isolated,

unconspicuous pustules. Cloacal opening dorsoventrally flattened.

Colour in life. — Dorsum yellowish orange with large irregular shaped black patches;

venter creamy yellow with large yellow-orange patches in the pubic region; iris yellow.

Colour in preservative. — Dorsum and dorsal surfaces of limbs blue grey with large dark

patches, venter and underside of limbs dull cream with scattered pale grey regions

distributed over the whole area, underside of thighs with isolated or connected light

patches.

Measurements (mm) of the holotype. — SVL 52.5; BH 14.2; HWID 18.1; EYE 4.7; IOD

12.2; ESNOUT 8.1; HUML 8.9; RADL 13.5; HNDL 12.3; FG3L 7.4; FEML 26.4; TIBL

24.9; FOOTL 40.9; TOEIL 5.6; TOE4L 27.0; CIL 2.6; WEBL 5.8.

Distribution. — Telmatobius hockingi is known only from the type locality and from

Piscobamba, Ancash.

Ecology. — Frogs of the type series were collected during the day under rocks in a stream

(Rio Sihuas) of strongly running water passing through an alder (A/nus jorullensis) forest.

The stream is used for the irrigation of the adjacent agricultural areas. Sometimes, the

stream dries, but small pools persist. These pools and moist soil below rocks are used by

the frogs to survive the dry period.

Etymology. — The specific name (a noun in the genetive case) is a patronym for Pedro

HockiNG of the Natural History Museum of the San Marcos University (MHNSM),

Lima, in recognition for his important contributions to the knowledge of biodiversity of

Peru.

Telmatobius mayoloi sp. nov.

(figs. 10-11)

Holotype. — URP 106, adult male, from the mouth of Rio Santa, 500 m from Lake

Conococha, Provincia Recuay, Departamento Ancash, Peru, 4050 m altitude, ca.

77°17:50"W 10°06’25"S, collected on 29 december 1992 by Eladio Turya CASTILLO.

Paratypes. — URP 103-105 and 107-111, 1 male, 6 females and 1 juvenile; MHNSM 7413

and 7419-7421, 1 male, 2 females and 1 juvenile; ZFMK 57259, 1 female; KU 220842, 1

female; all collected at the same site as the holotype by Antonio W. SALAS.

Diagnosis. — (1) Premaxillary teeth present, almost completely embedded in labial

mucosa; (2) tympanum absent; (3) nuptial spines minute, on dorsal and ventral surface of

the thumb; (4) dorsum blue grey (in preservative) with large dark blotches; (5) venter light

grey (in preservative) with small black spots; (6) forelimbs and hindlimbs with transverse

bars; (7) skin of dorsum smooth; (8) snout-vent length in males to 90.3 mm (MHNSM

7413), in females to 84.3 mm (ZFMK 57259).

Source : MNHN, Paris

ALYTES 14 (1)

TROPICAL

LOWLANDS

Er 7/|

2 à Li

rrare [QZS de LÉ

7h É\, “pe e EE

Co 77 x

fes Hô:

GS"

NORTH

£ 114 io Apuriae

EST

Oum.

Cu 100 ka

PACIFIC OCEAN

Fig. 1. — Distribution of northern and central Peruvian telmatobiine populations. Inverted triangle,

Batrachophrynus brachydactylus; triangle, B. macrostomus; 5, Telmatobius brevirostris; rhombus,

T. carrillae; open square, T. hockingi; circle, T. jelskif, filed square, T. mayoloi, dots, T. rimac.

Localities are approximated from distances by roads; multiple localities in close proximity are

represented by a single symbol. The main Andean river systems and lakes are indicated.

Source : MNHN, Paris

a Laguna Conococha e Rio Sihuas

A B.brachydactylus

brevirostris

AT. brevirostris

Canonical Variable 2

Canonical Variable 3

—2

—4 —4

—6 0 +6 +12 +18 6 0 +6 +12 +18

Canonical Variable 1 Canonical Variable 1

Fig. 2. — Plot of the discriminant function scores obtained for the populations from the Laguna Conococha and Rio Sihuas using the

functions which distinguish among the ranges of morphometric variation of Batrachophrynus brachydactylus, B. macrostomus,

Telmatobius brevirostris and T. carrillae (SiNsCH et al., 1995). (A) CANI versus CAN2. (B) CANI versus CAN3.

HOSNIS @ SVIVS

Source : MNHN, Paris

12 ALYTES 14 (1)

a Laguna Conococha

e Rio Sihuas

+ +

[e] vd 8

Canonical Variable 2

LŸ]

T. culeus

VTT. jelskii

—4 —2 0 +2 +4 +6

Canonical Variable 1

Fig. 3. — Plot of the discriminant function scores obtained for the populations from the Laguna

Conococha and Rio Sihuas using the functions which distinguish among the ranges of

morphometric variation of Telmatobius culeus, T. jelskii and T. rimac (SINSCH et al., 1995).

This species externally resembles Batrachophrynus brachydactylus, the only sympatric

telmatobiine species similar in size and coloration (fig. 10). The morphometric ratio

HWID/SVL is diagnostic for the distinction of T. mayoloi from Batrachophrynus

brachydactylus (fig. 6). Moreover, the presence of embedded premaxillary teeth and nuptial

excrescences as well as the rarely occurring granular glands with large granules in the

dorsal skin distinguish T. mayoloi from Batrachophrynus. T. mayoloi differs from T.

brevirostris, T. jelskii and T. rimac by the ratio HUML/SVL, and from T. carrillae and T.

hockingi by the ratio FG3L/SVL (fig. 7).

Description. — Head width almost equal to body width; head width and length almost

equal: HLEN 97 % of HWID; head length 34 % of SVL; head width 35 % of SVL. Dorsal

view of snout rounded, in lateral profile similar to T. atahualpai (fig. 11A). Nostrils not

Source : MNHN, Paris

e Batrachophrynus brachydactylus a Telmatobius mayoloi sp.n.

o Batrachophrynus macrostomus é o Telmatobius hockingi sp.n.

Ce]

+1 oi

ë co

= 4

a 0 É

£ cl 4

Q] de È

(e] LE ë

= 5 æ

g — 4 A

[on [=] 2

— © &

El $

pa Ù

Ês8

—3

—2 =1 0 #4 FR m6 -2 +1 +4 47 +10 +13

Principal Component 2 Canonical Variable 1

Fig. 4. — Plot of (A) principal component scores and (B) discriminant function scores of Batrachophrynus brachydactylus, B. macrostomus,

T. hockingi and T. mayoloi. Discriminant functions (1-3) and classification success are given in Table III. TR

Source : MNHN, Paris

o Telmatobius brevirostris + Telmatobius jelskii 5

a Telmatobius carrillae e Telmatobius mayoloi sp.n.

* Telmatobius hockingi sp.n. ao Telmatobius rimac

+2 M

ë 2

8 8

8 +1 É se

E o si

o — el

= 0 g ÉA

g : sn

2 ë &

2 É a

| ä

si [a]

& —1

—3 —-2 —1 O +1 +2 +3 —6 —-4 -2 0 +2 +4 +6

Principal Component 2 Canonical Variable 1

Fig. 5. — Plot of (A) principal component scores and (B) discriminant function scores of Telmatobius brevirostris, T. carrillae, T. hockingi,

T. jelsküi, T. mayoloi and T. rimac. Discriminant functions (1-5) and classification success are given in Table IV.

Source : MNHN, Paris

0.45

= = 0.16

@) in

® 0.35 a

A =)

= FE

n E0de e

0.25 0.08 9

D JS D ww D OÙ â

ss as . sŸ , s® PL as 4 s ù sŸ

Ÿ ct° 0 d° \Ÿ ct° x 9 d°

g «°° ÿ a «® X W)

ie Us Fe

®: { { @: g: { {

Fig. 6. — Box- and whisker-plot of morphometric ratios which permit the distinction among the Batrachophrynus species, T. hockingi and

T. mayoloi (multiple range test, LSD-method, P < 0.01). (A) HWID/SVL: B. macrostomus > T. mayoloi = T. hockingi > B.

brachydactylus. (B) FG3L/SVL: B. macrostomus > T. hockingi > B. brachydactylus = T. mayoloi.

Source : MNHN, Paris

16 ALYTES 14 (1)

0.35

Lo.25

S0.

pe)

0.15

Ê w & w

PS A \#" ET à sŸ . A

D 4. 4. nd”

2 . N 0

0.20

0.15

[a]

ro]

0.10

S av © v 4

PS 3° A#" FES à A s

nv co {- (9 4°

0 - XX pŸ

| 4.

Fig. 7. — Box- and whisker-plot of morphometric ratios which permit the distinction among the

known central Peruvian Telmatobius species, T. hockingi and T. mayoloi (multiple range test,

LSD-method, P < 0.01). (A) HUML/SVL: T. brevirostris = T. jelskii = T. rimac > T. carrillae

= T. mayoloi = T. hockingi. (B) FG3L/SVL: T. brevirostris = T. jelskit = T. rimac = T.

hockingi > T. carrillae = T. mayoloi.

Source : MNHN, Paris

SALAS & SINSCH 17

Fig, 8. — Photograph of a male Telmatobius hockingi.

protuberant, located at the anterior terminus of snout. Canthus rostralis indistinct

dorsally, in lateral profile short and elevated. Eyes not protuberant on top of head, eye

diameter 27 % of head length. Tympanum absent, tympanic annulus inconspicuous.

Supratympanic fold present, extending from posterior corner of eyelid to insertion of

forelimb. Maxillary and premaxillary teeth embedded in labial mucosa, slightly protrud-

ing, but almost unnoticeable when passing on top with finger tips. Well developed vertical

fold posterior to corner of jaw, extending below supratympanic fold to throat. Dentigerous

processes of vomer large and well developed, three times closer to choanae than to each

other, located slightly anterior to choanae; choanae small and oval. Tongue large and

rounded, attached through its complete length. Vocal slits absent.

Robust forelimbs, triangular shaped in cross section. Dermal wrist fold conspicuous,

but weakly developed. Relative length of fingers: II > IV > II > I (fig. 11B), tips of

fingers bluntly pointed, palmar webbing absent, lateral fringes absent. In males, inner

palmar tubercle large and oval, continuous with nuptial pad. Outer palmar tubercle

elliptical, about 2/3 of size of the inner. One large subarticular tubercle present proximally

on each finger, smaller subarticular tubercles present along the longitudinal axis of each

finger. In males, densely packed nuptial spines forming plush-like pads, extending on

dorsal, medial ventral surface of thumb.

Source : MNHN, Paris

18 ALYTES 14 (1)

Fig. 9. — Morphological details of male holotype URP 123 of Telmatobius hockingi. (A) Lateral view

of head. (B) Palmar view of right hand. (C) Plantar view of left foot. Scales = 5 mm.

Source : MNHN, Paris

SALAS & SINSCH 19

Fig. 10. — Photograph of a male Telmatobius mayoloi.

Stout hind limbs, dorsoventrally flattened; thighs with bagginess as in the lake-

dwelling B. macrostomus and T. culeus. Hind limb length (foot plus tibia) 47.9 % of SVL.

Relative length of toes (fig. 11C): IV > V > II > II > I; webbing formula: I 1 2/3 —

2+ 111 1/3 — 3— II 2+ — 3 1/3 IV 3 1/3 — 1 2/3 V; webbing diminishes gradually

to form lateral fringes along the edges of toes II, III, IV and V. Tips of toes spherical in

I, Il and III, more pointed in IV and V. Inner metatarsal tubercle ovally elongated, raised;

outer metatarsal tubercle equally shaped and elevated as inner, but only 2/3 in size. Small,

round subarticular tubercles distributed on toes as follows: I (1), II (1), HIT (2), IV (3) and

V (2). Tarsal fold extending to about 50 % of length of tarsus, confluent with lateral fringe

of toe I.

Dorsal, ventral and lateral skin usually smooth. Ventral skin covered with few and

isolated, inconspicuous pustules. Cloacal opening hidden due to the bagginess of skin.

Colour in life. — Dorsum pale brown with orange tone, frequently covered with irregular

shaped black blotches which often contain clear spots; forelimbs and hindlimbs with

transverse black bars and clear spots as on the dorsum; venter creamy yellow with orange

tone and black spots; iris orange with black reticulations.

Colour in preservative. — Dorsum grey with large, irregular shaped blotches; venter light

grey with isolated black dots, forelimbs and hindlimbs with transverse bars.

Source : MNHN, Paris

20 ALYTES 14 (1)

DCR.

Fig. 11. — Morphological details of male holotype URP 111 of Telmatobius mayoloi. (A) Lateral view

of head. (B) Palmar view of right hand. (C) Plantar view of left foot. Scales = 5 mm.

Source : MNHN, Paris

SALAS & SINSCH 21

Table I. - Morphometric data for Telmatobius mayoloi and T. hockingi. The first line is mean + 1 SD;

the second line is range. All values are in millimeters; see text for abbreviations of variables.

Telmatobius mayoloi Telmatobius hockingi

Character

Males Females Males Females

N=3 N = 10 N=5 N=5

SE 78.54 11.6 76.3 + 5.7 47.7 + 4.8 60.1 + 4.2

672 - 90.3 69.7 843 422-525 53.2: 64.8

Re 17.8 + 2.7 17.0 # 2.2 13.9 + 0.9 16.6 + 0.7

15.1: 20.4 146-213 12.9 : 15.2 15.5: 17.3

D 27.9 + 42 26.8 + 3.2 17.0 + 0.8 21.9 #14

236-320 2341318 16.2: 18.1 19.8 - 23.7

DE 6.1 + 1.0 5.7+04 4.4 + 0.6 49404

52-71 S1-65 3.9-53 4.6-5.6

16 17.5 + 24 17.6 + 12 12.1 +08 14.2 +07

15.3 - 20.0 15.8- 19.4 11.1-13.0 134-154

113 +12 10.6 + 0.6 7.6 + 0.5 8.8 +04

FNOSE 101-125 98-116 7.0-83 84-93

16.3 + 2.2 15.2 + 1.0 10.9 + 1.3 12.7 +05

FSNOUT 14.12 18.5 13.5 16.7 9.1-12.6 119-133

192 +58 172 43.1 9.9 + 0.6 115 +07

FUME 15.1 - 25.9 12.8-22.1 89-105 10.7 - 12.3

16.7 + 2.7 18.6 à 2.6 IL8+ 1.0 14.8 + 0.6

RADE 13.6: 18.9 147-231 112-135 13.8 - 15.5

Non 153441 15.6 + 2.6 IL8E 13 W2E17

113-194 11.7 19.7 10.5 - 13.8 12.7 : 16.9

FE 924 1.5 88412 82427 94H13

78-108 73-115 65-130 86-116

NL 35.8 + 4.6 38453 23.24 1.9 28.6 # 1.0

312-403 268-434 217-264 27.8 - 302

mu 35.8 + 4.8 34447 2.4 & 1.6 27.5 & 1.0

314-409 254-414 213-249 26.5 -290

57.3 & 7.8 54.1 + 62 35.743. 4.4 +24

ROOTL 499-655 44.2 - 66.0 33.0 - 40.9 416-471

BIEIA TB+O7 50405 65+04

ARE 6.6-9.3 6.3- 8.6 4.3-5.6 61-70

38.2 + 6.2 354 +41 23.2 #24 28.8 #15

TORE 73-46 28.2 - 42.6 211-170 26.9 - 30.9

me 34408 3.2 40.5 2.174 04 32404

2.6-4.2 2.3-3.9 21-30 2-37

112 #17 10.2 & 2.7 TTE28 8.04 1.8

APE 9.5-12.8 6.7-15.7 53-122 6.0 - 106

Source : MNHN, Paris

22 ALYTES 14 (1)

Table II. - Ratios of morphometric data for Telmatobius mayoloi and T. hockingi. Data are given as

mean + 1 SD. See text for abbreviations of variables.

Ratio Telmatobius mayoloi Telmatobius hockingi

N=13 N=10

BH/SVL 0.224 + 0.023 0.287 + 0.035

HWID/SVL 0.351 + 0.020 0.361 + 0.026

EYE/SVL 0.076 + 0.006 0.087 + 0.007

IOD/SVL 0.228 + 0.009 0.246 + 0.016

ENOSE/SVL 0.141 + 0.005 0.154 + 0.010

ESNOUT/SVL 0.201 + 0.012 0.221 + 0.014

HUML/SVL 0.228 + 0.033 0.201 + 0.020

RADL/SVL 0.237 + 0.035 0.248 + 0.016

HNDL/SVL 0.202 + 0.031 0.242 + 0.019

FG3L/SVL 0.116 + 0.010 0.164 + 0.034

FEML/SVL 0.456 + 0.044 0.484 + 0.039

TIBL/SVL 0.451 + 0.033 0.467 + 0.026

FOOTL/SVL 0.713 + 0.041 0.747 + 0.035

TOEIL/SVL 0.102 + 0.007 0.107 + 0.009

TOE4L/SVL 0.468 + 0.030 0.484 + 0.020

CIL/SVL 0.042 + 0.007 0.054 + 0.008

WEBL/SVL 0.135 + 0.027 0.147 + 0.040

FEM/TIBL 1.010 + 0.068 1.037 + 0.041

CIL/TOEIL 0.416 + 0.074 0.516 + 0.093

Measurements (mm) of the holotype. — SVL 67.2; BH 15.1; HWID 23.6; EYE 5.2; IOD

15.3; ESNOUT 14.1; HUML 15.1; RADL 17.5; HNDL 11.3; FG3L 7.8; FEML 31.2;

TIBL 31.4; FOOTL 49.9; TOEIL 6.6; TOEA4L 32.3; CIL 3.3; WEBL 9.5.

Distribution. — Telmatobius mayoloi is known only from the type locality.

Ecology. — During the day frogs were found under rocks and among submerged plants

Within the mouth of the Rio Santa. Between 11.00 and 12.00 h, some individuals were

observed swimming slowly in river parts with little current. Specimens were never seen

outside the water. This species occurs in the Puna. Tadpoles have been found over the year

in river pools and will be described in detail elsewhere.

Etymology. — The specific name (a noun in the genetive case) is a patronym for Antuñez

DE MAYOLO, a renowned engineer native from Ancash.

Remarks. — Four of the specimens examined (URP 103-104 and 109, KU 220842) are

large gravid females in an externally visible advanced state of egg development. The shape

of gravid females is almost ovoid, whereas the shape of non-gravid females and males

is slender and spindle-like. The head of the largest female is broad and similar-shaped

as in B. macrostomus. The thumbs of the reproductive males show well-developed

nuptial pads with minute, densely packed spines (fig. 11B). The two smallest individuals

Source : MNHN, Paris

SALAS & SINSCH

Table III. - Discriminant functions to distinguish among Batrachophrynus macrostomus, B.

brachydactylus, Telmatobius hockingi and T. mayoloi based on a stepwise discriminant analysis

(procedure: forward selection) using 18 log: transformed morphometric characters.

A. Statistical significance

Eigenvalue

quared

Degrees of

Freedom

19.00

2.14

0.71

B. Unstandardized discriminant function coefficients

0.643

Canonical Wilks

correlation Lambda

0.975 0.0093 392.5

0.825 0.1869 140.9

44.8

Character (loB10)

Coefficients of CANI

Coefficients of CAN2

Coefficients of CAN3

HUML

RADL

FG3L

TOE4L

Constant

4.80

8.93

9.70

-0.92

-23.83

C. Classification success

-0.87

-1.13

16.85

-21.28

17.92

D. Group centroids

Species

Predicted group

Actual group B. brachydactylus | B. macrostomus T. hockingi T. mayoloi

B. brachydactylus 53 (100%) =. F *

B. macrostomus L 13 (100%) : 4

T. hockingi = - 10 (100%) =

T. mayoloi - - - 13 (100%)

B. brachydactylus

B. macrostomus

T. hockingi

T. mayoloi

Source : MNHN, Paris

Table IV. - Discriminant functions to distinguish among Telmatobius brevirostris, T. carrillae, T.

hockingi, T. jelskii, T. mayoloi and T. rimac based on a stepwise discriminant analysis

ALYTES 14 (1)

(procedure: forward selection) using 18 logo transformed morphometric characters.

A. Statistical significance

EEE 1

6.98 0.935 0.0074 899.8 65 < 0.00001

223 0.831 0.0592 518.8 48 < 0.00001

124 0.744 0.1910 303.8 3 -< 0.00001

075 0.654 0.282 155.6 20 < 0.00001

033 0.501 0.7488 53.1 9 -< 0.00001

B. Unstandardized discriminant function coefficients

CA) of CANI of CAN2 of CAN3 of CAN4 of CANS

SvL 222.97 3.6 721 729 1431

BH -2.45 0.25 143 15.63 5.70

HWID 17.71 9.77 TT -2.42 -11.60

EYE -64.30 9.49 1.13 13.69 76.91

10D 103.47 24.03 9.64 2.6 113.66

ESNOUT 20.77 8.78 15.76 2.00

HUML 6.49 6175 223 12.12

RADL 237 0.50 1128 243

FG3L 8.77 5.51 6.90 3.93

TIBL 8.95 -23.58 228 “628

TOEIL -4.00 -15.26 4.33 -2.45

IL -3.06 3.05 2.30 3.68

WEBL 1.00 1.07 21.99 4.67

Constant 58.94 623 32.74 -80.43

C. Classification success

Predicted group

Acual group

T. brevirostris | 5 (100%) - 5 $ à :

T. carrillae 52 (08%) - 10%)

T. hockingi < Ë 10 (100%) - - ;

T. jelski 1%) - 26%) | 603%) - 26%)

T. mayoloi - g é : 13 (100%) -

T. rimac 2(5%) - 37%) - 37 (88%)

D. Group centroids

Species CAN 1 CAN2 CAN3 CAN4 CANS

T. brevirostris 1.65 “1.72 124 305 2.67

T. carrillae 4.05 0.63 -0.4 0.09 ou

T. hockingi 0.19 -0.86 -025 2.4 115

T. jelskit 237 137 0.14 0.10 0.01

T. mayoloi 0.76 -2.73 -3.46 0.43 0.24

T: rimac 0.72 -1.86 127 0.61 20.17

Source : MNHN, Paris

SALAS & SINSCH 25

(SVL 54.2 mm and 57.0 mm) without external sexual characters are considered as subadult

juveniles.

RESUMEN

Se evalüa la situacion taxonémica de dos poblaciones de ranas Telmatobiinae del

Departamento de Ancash, Perü, mediante la comparaciôn de la variaciôn intrapoblacional

de 18 de sus medidas morfométricas con las de seis especies de telmatobinidos de regiones

adyacentes: Batrachophrynus brachydactylus, B. macrostomus, Telmatobius brevirostris, T.

carrillae, T. jelskii y T. rimac. Las ranas, que habitan la Laguna Conococha y el Rio

Sihuas, no son miembros de las otras especies de la region. Las dos poblaciones

representan dos especies nuevas del género Telmatobius: T. hockingi y T. mayoloi. Se

presenta caracteres diagnosticos de la morfologia externa y de histologia de la piel para

distinguir entre los Telmatobiinae del Perû central.

ACKNOWLEDGEMENTS

We are grateful to Dra. N. CARRILLO and Lic. J. CORDOVA, ex and current curator of the

herpetological section of the MHNSM, Lima, permitting us access to the Telmatobiinae of the local

collection, especially to the type specimens assigned by J. VELLARD and V. MORALES. Likewise, to Dr.

M. Orriz, Director of the URP, Lima, and to Dr. W. BÔHME, curator of the herpetological section

of the ZFMK, Bonn, for the support of our research. The first hints revealing the existence of frogs

decribed in this paper as T. mayoloi were given to the first author by Lic. D. Mayo. M. ANTIGNANI,

C.S. Arias, J. ICOCHEA and E. TurYA CAsTiLLO helped us to collect frogs in the field. B. GLUMP

prepared the drawings of the new species. Finally, A.W.S. acknowledges the financial support given

by the Consejo Nacional de Ciencia y Tecnologia de Perü (CS-CONCYTEC N°0183-06-92-OAI) and

two anonymous donors. The paper benefited from the comments of W. R. HEvER, E. LAVILLA and

J. D. LyNCH.

LITERATURE CITED

BARBOUR, T. & NoOBie, G. K., 1920. — Some amphibians from northwestern Peru, with a revision

of the genera Phyllobates and Telmatobius. Bull. Mus. comp. Zool. Harvard, 63: 395-427.

BOokSTEIN, F. L., CHERNOFF, B. C., ELDER, R. L., HUMPHRIES, J. M., SmiTH, G. R. & STRAUSS, R.

E., 1985. — Morphometrics in evolutionary biology. Acad. nat. Sci. Philad. spec. Publ., 15:

1-277.

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HEN, K., 1994. — Lichimikroskopische Untersuchungen zur Histologie der Haut neotropischer Frôsche

(Gattungen Batrachophrynus und Telmatobius). Unpubl. Thesis, Institut für Biologie, Univ.

Koblenz: 1-163.

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52-66.

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Corresponding editor: W. Ronald HEYER.

Source : MNHN, Paris

Alytes, 1996, 14 (1): 27-41. 27

A contribution to the ecological genetics

of frogs: age structure and frequency

of striped specimens

in some Caucasian populations

of the Rana macrocnemis complex

David N. TARKHNISHVILI & Ramaz K. GOKHELASHVILI

Group of Population Ecology, Department of Biology, Tbilisi State University, 380030,

Chavchavadze avenue 1, Georgia

Four populations of Caucasian brown frogs (Rana macrocnemis) from

different elevations and different mountain systems (Great and Minor Cauca-

sus) were studied. In populations from the Minor Caucasus, the percentage of

striped frogs increases with elevation, but not in the Great Caucasus. At the

same time, age at sexual maturity in Caucasus Minor populations does not

differ between forest and subalpine populations. It is suggested that in this

region the increasing proportion of genetically striped frogs is the main

adaptation preventing a decrease of reproductive potential with elevation. In

the brown frogs metapopulation inhabiting the Great Caucasus, such a

mechanism is absent.

INTRODUCTION

Specimens with a light mid-dorsal stripe (phenotype “striata”) are found in

populations of many anuran species. The inheritance of this character has been studied in

several ranid species. Various authors have examined progeny produced as a result of

crossings between striped and unstriped frogs collected from populations with different

frequencies of the striped phenotype. A bright mid-dorsal stripe has been shown to be

determined by a simple dominant gene in Rana limnocharis (MorrWakt, 1953), R.

ridibunda (BERGER & SMIELOWSKI), R. sylvatica (BROWDER et al., 1966) and R. arvalis

(SHCHUPAK, 1977; SHCHUPAK & ISHCHENKO, 1981), though it has also been found that

isolated striped specimens may appear even among offspring of unstriped parents of R.

limnocharis (Morrwaki, 1953) and R. macrocnemis (TARKHNISHVILI & MAMRADZE, 1989;

TARKHNISHVILI, 1995). The most plausible reason for this appears to be the existence of

phenocopies.

At the same time, the proportions of striped specimens in populations of some frog

species display clinal variations, following climatic and landscape features. For example,

the proportion of striped R. sylvatica in North America generally increases towards the

west and north (FISHBECK & UNDERHILL, 1971; SCHUELLER & Cook, 1980). STUGREN

Source : MNHN, Paris

28 ALYTES 14 (1)

(1966) has shown that the proportion of striped specimens in populations of R. arvalis

increases in an eastern direction; at the same time, northern populations of R. limnocharis

in Japan are characterized by a reduced proportion of striped frogs (Morrwaki, 1953).

Previous authors have connected clinal changes in the proportion of different colour

morphs with genetic-stochastic processes (e.g. STUGREN, 1966) or their adaptive impor-

tance (e.g. MERRELL, 1969, 1973; NEVO, 1973; DaAPKkUS, 1976; ISHCHENKO, 1978). In

particular, for different colour morphs of the hylid Acris crepitans (the best studied species

in this respect), the hypothesis about the direct adaptive value of different colour morphs

(for escaping predation) competes with the hypothesis proposing a correlation of these

morphs with important physiological characteristics, i.e. thermotolerance and desiccation

resistance (NEVO, 1973); however, neither of these hypotheses has been supported

experimentally (GRAY, 1977, 1978). A similar situation is observed for a light mid-dorsal

stripe in brown frogs: e.g. for R. sylvatica, SCHUELLER & Cook (1980) suggest an

advantage of striped specimens in open areas with the cryptic character of this pattern.

Conversely, ISHCHENKO (1978) explains differences in the proportion of striped frogs

between different populations of R. arvalis on the basis of physiological differences

between different morphs. SCHWARZ & ISHCHENKO (1968), who compared striped and

unstriped froglets emerging from the same breeding site, have shown that striped froglets

have a relatively large liver, in comparison with unstriped ones, and that their weight

increases more rapidly. L. DoBriNskY (see ISHCHENKO, 1978), who used an optic-acoustic

gasoanalyser, demonstrated that metabolic exchange of striped froglets of R. arvalis is

especially high: they excrete up to twice as much CO, per gram of body mass than

unstriped ones. Tadpoles of striped R. arvalis need more time for completing metamor-

phosis (ISHCHENKO & SHCHUPAK, 1974) but, after metamorphosis, their growth is more

rapid than that of unstriped froglets, as shown by repeated measures of froglets with and

without stripes, after their emergence from the breeding pond (ISHCHENKO, 1978).

VERSHININ (1987) has shown that in demes of R. arvalis where striped frogs predominate,

froglets grow faster.

An interesting case of polymorphism is observed in populations of Caucasian brown

frogs (Rana macrocnemis complex). Different forms of brown frogs inhabit the Caucasian

Isthmus, Anatolia and mountain plateaus of the Middle East. The most widespread ones,

R. macrocnemis and R. camerani, represent closely related taxa included in the Rana

(Rana) temporaria group (Dusois, 1992). The taxonomic status of these forms is not very

clear. Some authors (e.g. MENsi et al., 1992) accept separate specific status of these frogs,

while demonstrating their close relations. BARAN and his co-authors (BARAN, 1969; BARAN

& ATATUR, 1986) demonstrated the presence of numerous populations with intermediate

characters, representing a probable hybrid zone between the two species. ISHCHENKO &

PYASTOLOVA (1973) obtained hybrids from parents caught in typical “macrocnemis” and

“camerani” populations; their viability, at least before and shortly after metamorphosis,

was not lower than in control groups. ISHCHENKO (1978, 1987) found no consistent

morphometric or coloration characters differentiating these two forms. He showed that the

multidimensional distance (based on 20 morphometric indices) between separate popula-

tions of “R. macrocnemis” and “R. camerani” in some cases exceeds the distance between

populations composed of the two different forms. He concluded that subdivision of

Caucasian brown frogs into two species is artificial.

Source : MNHN, Paris

TARKHNISHVILI & GOKHELASHVILI 29

At the same time, two forms of Caucasian brown frogs differ in the extent of

altitudinal variation of some characters, in particular the proportion of striped specimens.

Traditionally, one of the typical characters separating R. camerani from R. macrocnemis

is a light mid-dorsal stripe (TERENTYEV & CHERNOV, 1940; BARAN, 1969). In frogs

inhabiting Caucasus Minor, the proportion of striped specimens rapidly increases with

elevation and reaches 80 % in the subalpine belt. This is not observed in the Great

Caucasus, though some increase (up to 8 %) in the proportion of striped frogs with

elevation can be observed in North Caucasus; in these specimens the stripe is poorly

expressed (ISHCHENKO & PYASTOLOVA, 1973).

An increase in the percentage of specimens with a bright stripe at high elevations is

clearly expressed in the region of the Trialeti Mountain Ridge (Georgia), bordering the

north mountain plateaus of Caucasus Minor. Only a few striped specimens are found in

the lowlands and foothills, although they predominate in the subalpine belt, in spite of the

short distance between forest and subalpine populations and the very probable inter-

population migrations: specimens which had been marked in the forest populations during

the breeding period were sometimes caught later near the upper limit of the forest belt (our

data), and, thus, the distance between populations does not exceed ranges of individual

migrations. There are no barriers preventing interbreeding between frogs inhabiting

foothill and subalpine populations. According to our long-term observations, in any

population inhabiting the ridge a wide spectrum of phenotypes is found, from typical

“camerani” with a bright stripe, relatively short legs and sharp snout, to typical

“macrocnemis”, and pair formation among these two forms appears to be totally random.

In such conditions, the altitudinal differences we have described must presumably be the

result of strong selection favouring striped specimens in the mountains.

The reproductive success of an animal depends on its fecundity, mortality at different

stages of its life cycle and the period between successive generations (BEGON et al., 1986).

Fecundity, reflected in the number of eggs per clutch, depends directly on body size.

Differential mortality of different phenotypes can be estimated from changes in their

proportions in consecutive age classes (ISHCHENKO, 1978). The period between generations

can be estimated by studying the age distribution of adult animals. In connection with the

data on the different growth rates of striped and unstriped frogs, a comparative analysis

of the length of generation appears to be especially interesting. We studied the age

distribution of five populations of R. macrocnemis inhabiting localities at different

elevations and including different proportions of striped specimens.

MATERIAL AND METHODS

With the exception of frogs from Borjomi Canyon, animals were collected mainly

during the period April to July, 1993 and 1994, from the following localities (fig. 1, Table I).

(1) Borjomi Canyon, western foothills of the Trialeti Mountain Ridge, northern slope.

Forested canyon of the river Nedzura. Elevation 900-1100 m. Annual precipitation

1000-1200 mm (VLADIMIROV et al., 1991). Winter mild and wet. Period of activity for

amphibians about six months, from the beginning of April to the beginning of October.

Source : MNHN, Paris

30 ALYTES 14 (1)

CAUCASUS

Fig. 1. — Map of Georgia, with indication of studied localities. BC, Borjomi Canyon; GV, Gujareti

Village; LT, Lake Tabatskuri; MP, Mamisoni Pass; DU, Duruji Upstreams.

Total number of studied specimens, during two reproductive seasons (1992-1993), 138

adults: 106 males and 32 females.

(2) Gujareti Village. Western part of the Trialeti Ridge, northern slope. Subalpine

meadows. Elevation 1900 m. Annual precipitation 800-1000 mm. Winter cold and dry.

Period of activity for amphibians about four months, from the beginning of May to the

beginning of September. Total number of studied specimens 47: 13 adult males, 6 adult

female, 28 yearlings (body length 20-45 mm).

(3) Lake Tabatskuri, southern slopes of Trialeti Ridge, north-west of Javakheti

Plateau, Caucasus Minor. Mountain steppe. Elevation 2000 m. Annual precipitation

1000 mm. Winter cold and dry. Period of activity for amphibians four months, from early

May to early September. Total number of studied specimens 63: 27 adult males, 13 adult

females, 23 juveniles.

(4) Upstreams of the river Duruji, southern slopes of the eastern part of the Great

Caucasus Mountain Ridge, in Kvareli District of Georgia. Subalpine meadows on the

upper edge of the forest belt. Elevation 1950-2000 m. Annual precipitation 1500 mm.

Period of activity for amphibians about five months, from early May to late September.

Total number of studied specimens 22: 21 adult males, 1 adult female.

(5) Surroundings of the Mamisoni Mountain Pass in Racha Province, the central part

of the Great Caucasus, southern slopes. Alpine meadows. Elevation 2550 m. Annual

precipitation 1500 mm. Period of activity for amphibians about three months, from

mid-May to the end of August. Total number of studied specimens 22: 21 adult males,

1 adult female.

Source : MNHN, Paris

TARKHNISHVILI & GOKHELASHVILI 31

Table I. - Climatic conditions of studied locations. E, elevation above sea level

(meters); R, sum of sun radiation (ccal/cm?/year); TJA, mean temperature of

January (°C); TJU, mean temperature of July (°C); DWP, duration of the

period without freeze (days); SumT, sum of temperatures for the period with

stable mean temperature above 5°C; AP, median annual precipitation (mm);

DS, duration of the period of stable snow cover (days). BC, Borjomi Canyon;

GV, Gujareti Village, subalpine belt; LT, Lake Tabatskuri, subalpine belt;

DU, upstreams of the river Duruji, upper limit of forest belt, MP, Mamisoni

Pass, Racha. Most of the data are based on the Atlas of the Georgian SSR

(DrAVAKHISHVILI et al., 1964). Data on the annual precipitation are according

to VLADIMIROV et al. (1991). For E, DWP, AP and DS, median values

between minimal and maximal average estimations are given, for TJA,

maximal average estimations; for R, TJU and SumT, minimal average

estimations.

Climatic conditions at these localities (according to DJaAvAkHisHviLi et al., 1964) are

shown in Table I.

Body length (L: snout-urostyle length) of each specimen was measured by sliding

calipers with the distance between points 0.1 mm. The presence and brightness of the light

mid-dorsal stripe was recorded as clear, unclear or absent. Age was estimated by standard

skeletochronological methods (SMIRINA, 1989; CASTANET & SMIRINA, 1990).

For skeletochronology, femur (all frogs from Gujareti, Tabatskuri and Mamisoni

Pass and 20 specimens from Borjomi Canyon) as well as second phalange of fourth toe of

right foot (the remaining frogs from Borjomi Canyon) were used. Sections 25 u thick were

prepared with a cryostat, stained with Boemer hematoxylin and examined under a light

microscope. The line of the first hibernation is usually resorbed, as in other species of

brown frogs (LEDENTSOV, 1990). In most cases, age was estimated as N + 1, where N is

the number of fully visible lines of arrested growth (LAGs). In frogs collected during early

spring, the last LAG is invisible as well. In such cases, age was assumed as N + 2. The

numbers of visible LAGs in phalanges and femurs of a specimen were always equal.

Duplicated or additional LAGs, which can be observed sometimes on the sections of

tubular bones of brown frogs together with true ones (e.g. SMIRINA, 1989; LEDENTSOV,

1990) were rare.

Source : MNHN, Paris

32 ALYTES 14 (1)

Statistical analysis of differences between samples in body length of frogs was

conducted using the Student test (ordinary method and modified method for small

samples with different dispersions; ZAITsEv, 1984). Differences in age distribution were

tested with a nonparametric Kolmogorov-Smirnov à test. Differences in proportion of

striped specimens in samples were tested with Fisher’s angular method (ZaïTsev, 1984).

RESULTS

OCCURRENCE OF THE “STRIATA” PHENOTYPE

In Borjomi Canyon, only four of 138 examined frogs (2.9 %) had a bright mid-dorsal

stripe, and 89 % had not even an unclear stripe. In Gujareti, 35 frogs (74 %) had a bright

stripe and only 13 % were unstriped. Among frogs collected near Lake Tabatskuri, 77 %

had a bright stripe and 11 % were unstriped. No obviously striped frogs were found in

samples from Great Caucasus (Duruji Upstreams and Mamisoni Pass), though in each of

these samples a few frogs with very unclear light stripe in the middle part of the back were

found (Table II). Differences in the proportion of specimens with bright stripe are

significant, not only between “striped” populations from Gujareti and Tabatskuri and all

other populations (P < 0.001) but also between the population from Borjomi Canyon and

the sample from Mamisoni Pass (P < 0.01). Therefore, in populations from the Trialeti

Ridge an increase in the proportion of striped frogs with elevation was very clear, though

it was not observed in the Great Caucasus.

BODY LENGTH OF ADULT FROGS

Frogs from Tabatskuri were characterized by the smallest body size (62-63 mm on

average; Table II). Frogs from Duruji Upstreams were slightly larger: 62-66 mm. Body

length of specimens from other localities showed no significant differences (though frogs

from Mamisoni Pass were especially large). In Borjomi Canyon, females were significantly

(P < 0.01) larger, in comparison with males. Mean body length of specimens from

Borjomi Canyon, Gujareti and Mamisoni Pass varied from 67 to 73 mm. Differences

between most of samples are significant (Table III).

AGE DISTRIBUTION OF ADULT FROGS

Age distributions of adult frogs are shown in Table IV. In sections of tubular bones

of frogs from Trialeti Ridge (Borjomi Canyon, Gujareti, Tabatskuri), as well as from

Duruji Upstreams, one to six LAGs were observed (fig. 2) suggesting that the ages of the

animals are two to seven years. In the femur sections of frogs from Mamisoni Pass, from

four to ten LAGs were seen (i.e., ages five to eleven years). The “‘youngest” population

inhabits Lake Tabatskuri (mean age of adults 2.6-2.8 years). The mean age of frogs

Source : MNHN, Paris

TARKHNISHVILI & GOKHELASHVILI 33

Table II. - Morphological features of R. macrocnemis populations from different

localities. S, percentage of frogs with clear stripe; PS, percentage of frogs

with unclear stripe (“pseudostriata”); N, sample size; M, mean; SE, standard

error. For other abbreviations, see Table I. For Borjomi Canyon,

measurements of frogs collected during a five-year study (since 1989) are

given.

Body length (mm)

Coloration

N S(%) PS(%)|]

Table III. - Significance of inter-sample differences in body length of Rana

macrocnemis. Values of Student t are given as well as levels of significance: *,

P < 0.05; **, P < 0.01; ***, P < 0.001. For other abbreviations, see Table I.

Source : MNHN, Paris

(D) +I SHLATV

Fig. 2. — Femur sections of R. macrocnemis from different populations. A, Borjomi Canyon, female, three full

LAGs (four years of age); B, Gujareti Village, female, three full LAGs (four years of age); C, Mamisoni Pass,

male, eight full LAGs (nine years of age).

Source : MNHN, Paris

TARKHNISHVILI & GOKHELASHVILI 35

Table IV. - Age distribution of Rana macrocnemis from different localities. N, sample

size; MA, mean age in years; SE, standard error. For other abbreviations, see

Table I. Percentage of frogs of different age classes is indicated.

Age in years (%)

13 21 31 14

16 21 27 21

4 à |+ à |+ à | à |+ à

Table V. - Significance of inter-sample differences in age distribution of Rana

macrocnemis, using Kolmogorov-Smirnov À test. For abbreviations, see

Tables I and III.

Source : MNHN, Paris

36 ALYTES 14 (1)

Table VI. - Index of absolute growth rate (IG = L x MA!) and weighted index (TIG

=IGxXt! x 1000, where t is the sum of effective temperatures for the warm

period: average twenty-four-hour temperature + 5°C and more) for different

populations of Rana macrocnemis. For abbreviations, see Tables I and III.

Fe £

22.74 19.25 | 20.80 20.74 123.84 21.9 | 17.48 16.58 | 9.01 8.04

7.6 6.4 | 10.4 10.4 | 11.9 11.0 | 5.8 5.5 6.0 5.4

inhabiting Borjomi Canyon, Gujareti and Duruji Upstreams varied from 3.2 to 4.2 years,

but the mean age of frogs from Mamisoni was much higher and reached about eight years.

Differences in the age distributions of frogs from Mamisoni Pass, on the one hand, and

all other populations, on the other, are significant. Frogs from Duruji Upstreams are

significantly older than those from Borjomi Canyon (Table V). In all localities females

were slightly older than males, but differences are significant only for Borjomi Canyon (P

< 0.05).

POST-METAMORPHIC GROWTH RATES

The ratio IG = L x MA‘! of mean body size of adults (L, mm) to their mean age

(MA, years) can be assumed to be a good index of absolute growth rate. This index varied

between 19 and 24 in animals from Borjomi Canyon, Gujareti and Tabatskuri; it was less

(about 17) in Duruji Upstreams, and did not exceed 9 at Mamisoni Pass. No differences

in the growth rates of the sexes were detected.

Obviously, growth rates depend on the climatic conditions of the location. We cannot

detect genetic interpopulation differences in growth rates based only on observed growth

rates, but we have to take into account climatic differences between locations. Sum of

temperatures for the activity period of frogs (which more or less coincides with the period

when stable temperature exceeds 5°C), seems to be the most important quantitative

climatic variable affecting the growth rates of frogs. The modified index of growth, more

available for interpopulation comparisons than IG, was calculated in the following

manner: TIG = IG x xt' x 1000, where t represents the sum of temperatures for the

activity period. Calculated values are given in Table VI. Judging from estimated values of

TIG, similarity between different populations where unstriped frogs predominated (Duruji

Upstreams, Mamisoni Pass and Borjomi Canyon) was higher than between any of these

populations and populations with striped frogs (Gujareti and Tabatskuri).

Source : MNHN, Paris

TARKHNISHVILI & GOKHELASHVILI 37

DISCUSSION

Despite previous studies (see Introduction), the Rana macrocnemis complex remains

poorly known and more work is needed before a clear taxonomy of this group can be

proposed. Pending such studies, we here adopt a conservative attitude, and we use for the

Caucasian brown frogs the oldest available name for frogs of this complex, i.e. Rana

macrocnemis Boulenger, 1885.

Taking into consideration the great intra-population variability in all three localities

from Trialeti Ridge (Borjomi Canyon, Gujareti Village and Lake Tabatskuri), as well as

the free interbreeding that occurs between different phenotypes, we could unify them in the

same metapopulation system (“Trialeti”?). The most important question appears to be why

there is such marked morphological differences between different populations within this

system.

At the intraspecific level, growth rates of specimens are related to two main factors:

climatic conditions at the locality and genetically determined growth rates. Moreover,

actual growth rate of each individual depends on the attained body size: growth slows

down in animals reaching definitive species-specific size. Populations from Trialeti Ridge

differed one from another in each of these three characteristics. Borjomi Canyon, situated

at an elevation of about 1000 m in a forested gorge, is characterized by a relatively mild

climate (sum of effective temperatures about 3000, January temperature —4°C, etc.; see

Table I) in comparison with the other two localities. Attained body size is especially small

in the population at Lake Tabatskuri. At the same time, judging from the proportion of

striped frogs, the genetic composition of populations from Gujareti and Lake Tabatskuri

clearly differ from that of the population at Borjomi Canyon (though the distance between

populations from Borjomi Canyon and Gujareti is less than 18 km and the only natural

barrier between them is a small ridge of about 2000 m maximum elevation).

We considered the role of climatic conditions and weighted the indices of growth rates

according to the sum of effective temperatures (reflecting the period of activity of frogs)

at different elevations. Weighted growth index was especially high for frogs inhabiting the

vicinity of Lake Tabatskuri. This may have resulted in especially early maturation, at the

expense of decreased mean adult body size, in this population. However, frogs from

Gujareti, which mature at the same age and the same body size as frogs from Borjomi

Canyon, also grow much more rapidly than frogs from the latter locality. Thus, we

propose that post-metamorphic growth rates for Trialeti metapopulation are due to

genetic differences between local populations, reflected in the different frequencies of

striped specimens. The growth index of frogs from the populations where striped animals

predominate is about 1.5 times more than in the ‘“unstriped” population inhabiting

Borjomi Canyon. Accelerated growth in the mountain populations of Trialeti Ridge has

an adaptive value. If frogs from Gujareti had the same growth index as in the Borjomi

Canyon, they would mature 1.5 times later (taking into account differences in the period

of activity and sum of effective temperatures). If mean age of females from the Borjomi

Canyon reaches 3.6 years, in Gujareti it would reach about 5.4 years.

Source : MNHN, Paris

38 ALYTES 14 (1)

Intrinsic growth rate of a population, in accordance with well-known demographic

models (e.g. WiLLIAMSON, 1972), is described by the equation KT à b-d+1= HT

where b is the mean value of fecundity, d the mean mortality rate for adult animals and

+ the mean age of adult frogs. If the animals from two populations have the same fecundity

and mortality rates, the ratio of their productivities would be k = A(/! — 12, where +,

and r, are the mean ages of animals in populations where animals mature at a younger

and older age, respectively.

Genetically fixed rapid growth of frogs from Gujareti prevents displacement of local

genotypes by the genotypes predominating in Borjomi Canyon, in spite of the latter

breeding in more favourable climatic conditions.

Frogs from Lake Tabatskuri grow and mature even faster than in Gujareti. If

fecundity and mortality of these two populations were equal, the reproductive success of

frogs inhabiting the surroundings of Tabatskuri would be higher. However, the fecundity

of females with a body length of about six centimeters (mean size in Tabatskuri

population) is 1.5 times lower than that of females of the same species with a body length

of seven centimeters (TARKHNISHVILI, 1993). The small size of frogs inhabiting surround-

ings of Tabatskuri is probably the cost of advantages associated with rapid maturation.

The productivities of frogs from different populations on Trialeti Ridge appear to be

similar. This allows the stable coexistence of populations dominated by different morphs

without the displacement of morphological characters as a result of interbreeding.

The results presented in this study enable the high proportion of striped specimens in

some populations to be explained. However, the inverse situation, the very low proportion

of striped frogs in Borjomi Canyon, remains to be explained. The hypothesis that in

forested canyons selective pressure works against striped frogs cannot be excluded. In

particular, it may be connected with the very unstable breeding sites in this habitat (see

TARKHNISHVILI, 1993), taking into account the longer larval period of genetically striped

brown frogs, demonstrated in Rana arvalis (ISHCHENKO & SHCHUPAK, 1974). However, this

question requires further study.

In both populations from Great Caucasus, frogs with a bright stripe are absent,

independently of the elevation and the climatic conditions. Overall, the climate in the

Great Caucasus is more humid and mild, in comparison with Caucasus Minor localities

situated at the same elevations (Table 1): the sum of effective temperatures in the upper

reaches of streams in Duruïji (elevation 1900 m) is similar to that in Borjomi Canyon (1000

m). At the same time, the growth rates of frogs from this population are slightly lower than

those of unstriped frogs from Borjomi Canyon and markedly lower than in “striped”

populations from Gujareti Village and Lake Tabatskuri. The growth rates of frogs from

Mamisoni Pass are lowest; even the growth index (TIG), which takes into account the

coldest climate in this locality (Table I), shows a low value (Table IV). As a result, the

actual productivity of frogs from Great Caucasus clearly declines with elevation. The only

reason that can be hypothesized for this situation is an absence of a genotype, correlated

with rapid growth, in the gene pool of the metapopulation of brown frogs inhabiting Great

Caucasus.

An interesting conclusion can be outlined. In spite of the high external similarity

between populations from Borjomi Canyon and Great Caucasus, they belong to different

Source : MNHN, Paris

TARKHNISHVILI & GOKHELASHVILI 39

metapopulation systems. The first system (western part of the Trialeti Ridge) includes

genotypes connected with rapid growth. This facilitates the rapid redistribution of

genotypes when the population is exposed to new climatic conditions and the appearance

of specific “mountainous”’ populations, composed almost exclusively of striped frogs. In

the gene pool of the second system (southern slopes of Great Caucasus) such genotypes

are simply absent.

In À. macrocnemis populations from Armenia, inhabiting elevations of 1900-3000 m,

frogs of one to three years of age predominate (LEDENTSOV & MELKUMYAN, 1986). The

situation is similar to that in localities from Trialeti Ridge, which belong to the same

mountain system of Caucasus Minor.

This point of view requires further studies. Expected difficulties could be outlined.

For instance, the presence of striped frogs in the population does not necessarily prove the

presence of the genotype “striata” in its gene pool, because under changed developmental

conditions they can appear even in the descendants of genetically unstriped parents.

ACKNOWLEDGEMENTS

The authors are obligated to the Georgian-German Organization CUNA Georgica and Dr. Udo

HiscH, foundator of this organization, which supported our field work for 1993 and 1994. We are

greatly thankful to Dr. Alain Dumois, Dr. Tim HALLIDAY and Dr. Sergius KUZMIN for helpful

comments and remarks on the first draft of the manuscript. We greatly appreciated the efforts of Mr.

Boris ARDABIEVSKY and Mr. Alexander GAVASHELISHVILI who accompanied and helped us in the field

work.

LITERATURE CITED

BarAN, L, 1969. — Anadolu dag kurbagalari uzerindeki sistematik arastirma. [A study on the

taxonomy of the mountain frogs of Anatolia]. Sci. Rep. Fac. Sci. Ege Univ., 80 (54): 1-78. [In

Turkish, with English Summary].

BARAN, I. & ATATUR, M. K., 1986. — A taxonomical survey of the mountain frogs of Anatolia.

Amphibia-Reptilia, 7: 115-133.

BEGoN, M., HARPER, J. L. & TOWNSEND, C. R., 1986. — Ecology: individuals, populations and

communities. Part II. Blackwell Scientific Publications.

BERGER, L. & SMIELOWSKI, J., 1982. — Inheritance of vertebral stripe in Rana ridibunda Pall.

(Amphibia, Ranidae). Amphibia-Reptilia, 3: 145-151.

BROWDER, L. W., UNDERHILL, J. C. & MERRELL, D. J., 1966. — Mid-dorsal stripe in the wood frog.

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CASTANET, J. & SMIRINA, E., 1990. — Introduction to the skeletochronological method in amphibians

and reptiles. Ann. Sci. nat., Zool., (13), 11: 191-196.

Darkus, D., 1976. — Differential survival involving the Burnsi phenotype in the northern leopard

frog, Rana pipiens. Herpetologica, 32: 325-327.

DyAVAKHISHVILI, A. N., ASLANIKASHVILI, A. F., DZOTSENIDZE, G. S. et al., 1964. — Arles Gruzinskoj

Sovetskoj Sozialisticheskoj Respubliki. [Atlas of the Georgian Soviet Socialist Republic]. Tbilisi

and Moscow, Main Department of Geodesy and Cartography: 1-269.

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Lyon, 61 (10): 305-352.

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FisaBeCK, D. W. & UNDERHILL, J. C., 1971. — Distribution of stripe polymorphism in wood frogs,

Rana sylvatica LeConte, from Minnesota. Copeia, 1971 (2): 253-259.

Gray, R. H., 1977. — Lack of physiological differentiation in three color morphs of the cricket frog

(Acris crepitans) in Illinois. Trans. Ill. State Acad. Sci., 70 (1): 73-79.

Le 1978. — Nondifferential predation susceptibility and behavioral selection in three color morphs

of Illinois cricket frogs, Acris crepitans. Trans. III. State Acad. Sci., T1 (4): 356-360.

ISCHCHENKO, V. G., 1978. — Dinamicheskij polimorfizm burikh lyagushek fauni SSSR. [Dynamic

polymorphism of the brown frogs of USSR fauna]. Moscow, Nauka: 1-148. [In Russian].

LS 1987. — The level of morphological similarity between the populations of the Caucasian brown

frog, R. macrocnemis Blgr. In: L. J. BORKIN (ed.), Herpetological investigations in the Caucasus,

USSR Academy of Sciences, Proc. zool. Inst. Leningrad, 158: 100-104. [In Russian, with English

summary].

ISHCHENKO, V. G. & PYAsTOLOVA, O. A., 1973. — A contribution to the taxonomy of Caucasian

brown frogs. Zool. Zhurnal, 52 (11): 1733-1735. [In Russian, with English summary].

ISHCHENKO, V. G. & SHCHUPAK, E. L., 1974. — Ecological differences of individual genotypes in a

moor frog population. Soviet J. Ecol., 5 (4): 379-380.

LEDENTSOV, A. V., 1990. — Prostranstvennaya i vozrastnaya struktura reprodukyivnoj chasti populyatsii

ostromordoj lyagushki (Rana arvalis). [Spatial and age structure of the reproductive part of

population of moor frog (Rana arvalis)]. Ph.D. Thesis, Sverdlovsk: 1-24. [In Russian].

LEDENTSOV, A. V. & MELKUMYAN, L. S.,1987. — On longevity and growth rate in amphibians and

reptiles in Armenia. Proc. zool. Inst. Leningrad, 158: 105-110. [In Russian, with English

summary].

MENSI, P., LATTES, A., MACARIO, B., SALVIDIO, S., GIACOMA, C. & BALLETTO, E., 1992. — Taxonomy

and evolution of European brown frogs. Zool. J. linn. Soc., 104: 293-311.

MERRELL, D. J.,1969. — Limits on heterozygous advantage as an explanation of poymorphism. J.

Hered., 60: 180-182.

me 1973. — Ecological genetics of anurans as exemplified by Rana pipiens. In: J. L. ViaL (ed.),

Evolutionary biology of the anurans, Columbia, Missouri, University of Missouri Press: 329-335.

MoriWaki, T., 1953. — The inheritance of the dorso-median stripe in Rana limnocharis Wiegmann.

J. Sci. Hiroshima Univ., Sec. 3, Div. 1 (Zool.), 14: 159-164.

Nevo, E., 1973. — Adaptive color polymorphism in cricket frogs. Evolution, 27 (3): 353-367.

SCHUELLER, F. W. & Cook, F. R., 1980. — Distribution of the middorsal stripe dimorphism in the

wood frog, Rana sylvatica, in eastern North America. Canad. J. Zool., 58 (9): 1643-1651.

ScHWaARZ, S. S. & ISHCHENKO, V. G., 1968. — [Dynamics of genetical composition of populations of

moor frog]. Bulleten MOIP, Otdel biologichesky, 73 (3): 127-134. [In Russian].

SncHupak, E. L., 1977. — [Inheritance of the mid-dorsal stripe in moor frog]. Informacionnie

Materiali Instituta Ekologi Rastenij i Zhivotnikh, Sverdlovsk: 36-37. [In Russian].

ScHuPaK, E. L. & ISHCHENKO, V. G., 1981. — On the hereditary base of colour polymorphism in

moor frog (Rana arvalis Nilss). I. Light mid-dorsal stripe. In: Herpetological researches in

Siberia and Far East, Leningrad, Nauka: 128-132. [In Russian, with English summary].

SMIRINA, E. M., 1989. — [The method of determination of the age in amphibians and reptiles with

bone layers]. /n: SHCHERBAK, N. N. (ed), Rukovodstvo po izucheniyu zemnovodnikh i

presmikayushchikhsya, Kijev: 143-153. [In Russian].

STUGREN, B., 1966. — Geographic variation and distribution of the moor frog, Rana arvalis Nilss.

Ann. zool. fenn., 3 (1): 29-39.

TARKHNISHVILI, D. N., 1993. — Anurans of Borjomi Canyon: clutch parameters and guild structure.

Alytes, 11 (4): 140-154.

——. 1995. — On the inheritance of the mid-dorsal stripe in the Iranian wood frog (Rana

macrocnemis). Asiat. herpet. Res., 6: 120-131.

TARKHNISHVILI, D. N. & MAMRADZE, R. G., 1989. — Modification of the phenotype of Caucasian

brown frogs under the influence of high temperature. Bull. Acad. Sci. Georgia, 135 (2): 437-440.

[ln Russian, with English summary].

TERENTYEV, P. V. & CHERNOV, S. A., 1949. — Opredelitel zemnovodnikh i presmikayushchikhsya

SSSR. [Guide for amphibians and reptiles of the USSR]. Moscow, Sovetskaya Nauka. [In

Russian].

Source : MNHN, Paris

TARKHNISHVILI & GOKHELASHVILI 41

VERSHININ, V. L., 1987. — Some features of the phenetical structure of groupings of the moor frog

in the industrial city. In: Vliyanie sredi na dinamiku strukturi i chislennosti zhivotnikh,

Sverdlovsk: 74-79. [In Russian].

VLADIMIROV, L. A., GIGINEISHVILI, G. A., DJAVAKHISHVILI, À. I. & ZAKHARASHVILI, N. N., 1991. —

Vodny balans Kavkaza i ego geograficheskie zakonomernosti. [Water balance of Caucasus and

its geographic conformity to natural laws]. Tbilisi, Metsniereba. [In Russian].

WILLIAMSON, M., 1972. — The analysis of biological populations. London, Edward Arnold.

ZAITsEV, G. N., 1984. — Matematicheskaya statistika v eksperimentalnoy botanike. [The mathematical

statistics in experimental botany]. Moscow, Nauka: 1-424. [In Russian].

Corresponding editors: Alain Dugois & Tim R. HALLIDAY.

Source : MNHN, Paris

Alytes, 1996, 14 (1): 42-52.

Use of terrestrial habitats by amphibians

in the sandhill uplands

of north-central Florida

C. Kenneth Dopp, Jr.

National Biological Service, 1920 N.W. 71st Street, Gainesville, Florida 32653, U.S.A.

kdodd@nervm.nerde.ufl.edu

A total of 506 individuals of 12 amphibian species was captured during

sampling of two upland communities in north-central Florida, USA, in 1989

and 1990. Amphibians were found as far as 914 meters from the nearest water

body, although the actual breeding site could have been farther away. Of the

species dependent on water for breeding, three (Bufo terrestris, Gastrophryne

carolinensis, Scaphiopus holbrooki) accounted for 87 % of the amphibians

captured. No significant correlation was found between the total number of

amphibians captured per trap and trap distance to nearest water body. Most

amphibians (83 %) were caught less than 600 meters from the nearest water.

Upland communities appear to be used extensively by certain amphibians,

especially terrestrial burrow users. As such, management programs need to be

expanded to include surrounding uplands if amphibian declines are to be

prevented.

INTRODUCTION

For amphibians that rely on water for reproduction, the vast majority of field studies

center on activities at or near breeding sites (e.g., references in D'UELLMAN & TRUEB, 1985).

Amphibians are conspicuous at breeding locations as males call to attract females and

establish territories, amplectant pairs mate and deposit eggs, larvae grow and either

metamorphose or become neotenes, and adults and metamorphosed young begin to

disperse to uplands or other habitats used during non-reproductive times of the year.

The life history of wetland-breeding amphibians away from breeding sites is poorly

understood. It seems generally accepted that individuals may disperse some distance from

breeding sites, perhaps varying among species, life stages, or in response to quality

and availability of adjacent habitats. At least one text, however, terms distances moved

into adjacent habitats as “minor” (ZUG, 1993). Except for a few studies (e.g., PEARSON,

1955; WiccrAMs, 1973; SEMLITSCH, 1981), the presence of water-breeding amphibians in

uplands has been inadequately documented in the North American literature, and then

often on the basis of a single or relatively few observations on a few species (Table I). The

distances that most species in the southeastern United States can or normally disperse are

unknown.

Source : MNHN, Paris

Dopp 43

Table I. - Examples of distances that North American amphibians have been recorded moving overland

under natural conditions. Movements along watercourses and terrestrial movements associated

with displacement experiments are not included. M, mean.

Species Location Movement Reference

Salamanders

Ambystoma californiense California 120 m HOLLAND et al. (1990)

Ambystoma californiense Califomia 1600 m' AUSTIN & SHAFFER (1992)

Ambystoma jefersonianum Kentucky M=250m DOUGLAS & MONROE (1981)

Ambystoma jeffersonianum Indiana M=252m(20-625m) | WILLIAMS (1973)

Ambystoma jeffersonianum Indiana M=92m(3-247m) | WiLLiaMs (1973)

Ambystoma jeffersonianum Michigan 152m WACASEY (1961)

Ambystoma jeffersonianum New York 1610 m BisHoP (1941)

Ambystoma macrodactylum Califomia 30m STEBBINS (1951)

Ambystoma maculatum North Carolina 18-823 m GORDON (1968)

Ambystoma maculatum Michigan M=192m(157-249m) | KLEEBERGER & WERNER (1983)

Ambystoma maculatum Kentucky M=150m(6-220m) | DouGLas & MONROE (1981)

Ambystoma maculatum Missouri M=150m(to172m) | SexroNet al. (1986)

Ambystoma maculatum New York 75m WILSON (1976)

Ambystoma maculatum Indiana M = 64 m (0-125 m) WILLIAMS (1973)

Ambystoma opacum Indiana M=193m(0450m) | WiLLIAMS (1973)

Ambystoma talpoïdeum South Carolina 81-261 m SEMLITSCH (1981)

Ambystoma texanum Indiana M = 52 m (0-125 m) WILLIAMS (1973)

Ambystoma tigrinum South Carolina 162m SEMLITSCH (1983)

Notophthalmus viridescens Massachusetts 800 m HEALY (1975)

Frogs

Acris crepitans Texas 167 m PYBURN (1958)

Acris gryllus Florida 823m CARR (1940)

Acris gryllus Kansas 183m FITCH (1958)

Bufo americanus Minnesota 1000 m EWERT (1969)

Bufo americanus Ontario 594 m OLDHAM (1966)

Bufo cognatus Minnesota 300-1300 m EWERT (1969)

Bufo hemiophrys Minnesota 25m OLDFIELD & MORIARTY (1994)

Bufo hemiophrys Minnesota 61m BRECKENRIDGE & TESTER (1961)

Bufo woodhousei Kansas 579m FITCH (1958)

Gastrophryne olivacea Kansas t0183m FITCH (1956)

Pseudacris nigrita Kansas 183 m' FITCH (1958)

Pseudacris regilla Oregon 237 m! JAMESON (1956)

Pseudacris triseriata Indiana 100 m KRAMER (1974)

Rana capito Florida 1600 m CARR (1940)

Rana capito Florida 2000 m FRANZ et al. (1988)

Rana catesbeiana New York 76m INGRAM & RANEY (1943)

Rana catesbeiana New York 107 m RANEY (1940)

Rana palustris Minnesota 500 m OLDFIELD & MORIARTY (1994)

Rana pipiens Minnesota 1500 m OLDFIELD & MORIARTY (1994)

Scaphiopus bombifrons Kansas 94m FITCH (1958)

Scaphiopus holbrooki Florida 402m PEARSON (1955)

* Represents juvenile dispersion.

? Estimated from map.

Source : MNHN, Paris

44 ALYTES 14 (1)

In 1989 and 1990, Dopp & FRANZ (1995) conducted an inventory of the snake

community inhabiting upland sites on the Katharine Ordway Preserve in north-central

Florida. During the course of the survey, substantial numbers of amphibians were

captured in wire mesh funnel traps. Inasmuch as little information was available on the

presence of amphibians in these physically harsh environments, I tabulated capture results

to determine which species used upland habitats and how far they were from the nearest

potential breeding site. Although the original study was not designed to survey the

amphibian community, these data may be helpful in planning future research and in

directing attention to the importance of uplands in the conservation of amphibian

populations that depend upon isolated wetlands for breeding.

STUDY SITE AND METHODS

The Katharine Ordway Preserve-Swisher Memorial Sanctuary is a 3750-ha tract

located approximately 5 km SE of Melrose, Putnam County, Florida. This upland sandhill

region lies within the Interlachen Karstic Highland at the southern end of Trail Ridge. The

area represents a portion of a dune complex that probably formed in association with

active beach development during periods of higher sea levels (WHiTE, 1970). The dunes

have been secondarily modified by solution activities in the underlying limestone to form

sinkholes and karst basins. Many of these solution features hold water to form ponds,

lakes, and wetlands. More than 70 water bodies exist on the property. There are 27 species

of amphibians recorded from the Ordway Preserve (FRANZ, 1995), and at least 16 species

have been recorded in a single small temporary pond in upland habitat (Dopp, 1992).

Two of the eight vegetative communities known from the Ordway Preserve (FRANZ

& HALL, 1991) were sampled during this study. Both upland communities, high pine forest

and sand live oak hammock, have been influenced by human disturbance and past fire

histories. Also known as “sandhill”, high pine forest is dominated by longleaf pine (Pinus

palustris), turkey oak (Quercus laevis), and wiregrass (Aristida stricta). The community

occurs on deep sands associated with dune ridges. Sand live oak hammock occurs as

fringes around certain wetland types and on ruderal sites. Dominated by sand live oak (Q.

geminata) and occasionally by laurel oak (Q. hemisphaerica), sand live oak hammocks can

have dense understories composed of sapling oaks, blueberries (Vaccinium spp.), myrtle

oak (Q. myrtifolia), and other woody plants. Reindeer lichens (C/adonia spp. and Cladina

spp.) and herbaceous species are more prevalent in open hammocks without a dense

understory. General information and references on these and other Florida communities

are in MYERs & EWEL (1990).

Between 15 and 25 % of the property is believed to have been cleared for agriculture

and human habitation since 1850 (R. FRANZ, personal communication). Several of these

areas have undergone succession to xeric sand live oak hammocks. Regular prescribed

burning of high pine forests was established in 1983 as a part of the Ordway Preserve’s

management plan for reestablishing the native longleaf pine ecosystem. Summer air

temperatures in upland habitats routinely approach 36°C, and substrate temperatures of

50°C have been recorded. The porous sandy soils dry rapidly at and immediately below

Source : MNHN, Paris

Dopp 45

the surface. A combination of poor soil moisture retention and high temperatures at or

near the substrate surface make these upland sandhill habitats potentially harsh for small

amphibians.

In 1989, 100 individually numbered screen wire mesh double-opening funnel traps

(90 cm long by 18 to 25 cm diameter) were placed at six upland sites as follows: 31 traps

in closed xeric (sand live oak) hammock; 59 traps in sandhill (high pine) habitat; and 10

traps in open xeric (sand live oak) hammock. Exact locations of the traps and descriptions

of the habitats are presented elsewhere (Dopp & FRANZ, 1995).

Most traps were set along fallen trees and branches that formed natural drift fences.

At certain locations, traps were set along drift fences made of 10 m sections of galvanized

metal set in 4-pronged arrays (see figure 1 in CAMPBELL & CHRISTMAN, 1982, and figure

11A in CorN, 1994). AIl traps were covered with palmetto fronds to prevent captured

animals from overheating in the direct sun and to provide cover. In 1989, traps were

checked daily from April 4 through November 17 (23,800 trap nights) between 07.00 and

12.00 h. Species identifications were recorded and animals were released in cover within

several meters of the trap.

In 1990, the same areas were resampled using the same general techniques except that

all sites were not sampled simultaneously. In addition, 30 traps were set in closed xeric

hammock habitat in the vicinity of a temporary pond (Breezeway Pond). Traps were

placed in the same positions as in 1989. From 20 to 30 traps were checked daily from April

4 to September 27. The dates when individual sites were sampled are provided in DopD

& FRANZ (1995). This protocol resulted in a sampling period of 4,490 trap nights.

The location of each trap (excluding the Breezeway Pond traps) was plotted on aerial

photographs, and the distance to the nearest potential source of water for breeding by

amphibians was measured to the nearest meter. I examined possible effects of trap

placement on amphibian capture in relation to habitat (sandhill, live oak hammock with

open understory, live oak hammock with dense understory), type of water body (lake

versus pond), and specific water body. Ponds had surface areas less than 4 ha and usually

dried during droughts. Although Smith Lake dried during the intense drought of the late

1980’s to early 1990’s, the other lakes were permanent. Inasmuch as the data were not

normally distributed, most comparisons were made using the nonparametric Kruskal-

Wallis test (procedure NPARIWAY, ANONYMOUS, 1988). The effect of trap distance from

nearest water body on the total number of amphibians captured was examined using

Spearman rank correlation. Eleutherodactylus planirostris has terrestrial development and

therefore was excluded from analyses of the relationship between trap distance and nearest

water body. Statistical analyses were performed using the SAS program for microcomput-

ers (ANONYMOUS, 1988) and ABSTAT version 4 (ANONYMOUS, 1987). The level of

significance was set at « — 0.05.

RESULTS

A total of 506 amphibians comprising 12 species was captured during trapping for

snakes (0.2 amphibians/trap night in 1989; 0.1 amphibians/trap night in 1990). Amphib-

Source : MNHN, Paris

46 ALYTES 14 (1)

ians were found in funnel traps at distances from 42 m to 914 m from the nearest water

(Table IT). Individuals were found in 90 different traps; there was no significant difference

in mean distance (MD) to nearest water body between funnel traps in which amphibians

were caught (MD = 427.9 m) and those in which amphibians were not caught (MD =

334.5 m) (2 = 3.05, 1 DF, P = 0.08).

Trapping location was not random with respect to water bodies. The mean distance

from traps to the nearest water body varied significantly among different ponds and lakes

(Table IL; x? = 69.4, 5 DF, P = 0.0001) and in relation to water body type (lakes, MD

= 495 m, N = 57 traps; ponds, MD = 312 m, N = 33 traps; x? = 18.8, 1 DF, P =

0.0001). Perhaps because of these potential trap biases, there was no significant correlation

between the total number of amphibians captured per trap and the distance to nearest

water body (fig. 1; r, = 0.3084, P > 0.05, N = 100). Likewise, there was no significant

difference in the mean distance to nearest water body among the traps in different habitat

types (Table IV; Ê = 3.3, 2 DF, P = 0.19).

Only 28 % of the amphibians captured were in traps less than 400 m from the nearest

wetland, although 51 % of the traps were less than 400 m from the nearest water body.

As distance increased to 500 m (accounting for 77 % of the traps), the amphibian capture

percentage increased to 67.6 %, and at 600 m (accounting for 88 % of the traps) the

percentage increased to 82.9 %. Few specimens (11) were captured from 600 to 800 m (9 %

of the traps), or at distances greater than 900 m (14 amphibians and 2 % of the traps).

However, 11.6 % of all captures were recorded from 800 to 900 m; these traps accounted

for only 4 % of the trapping effort. Capture was not random with respect to habitat type.

More amphibians were captured in open xeric habitat, and less in closed xeric hammock,

than might be expected if the number of amphibians captured among habitats was in direct

proportion to trapping effort (x? = 10.73, 2 DF, P = 0.0047) (Table IV).

DISCUSSION

Trap biases exist in the survey protocol, and a rigorous assessment needs to be made

concerning factors that influence amphibian presence in upland communities. However,

these results suggest that the presence of amphibians in southeastern upland habitats may

be more significant than is usually recognized, especially by land and resource managers,

and that amphibians occupy habitats even at considerable distances from the nearest

potential breeding site. Amphibians captured during the inventory may have bred in more

distant wetlands than the nearest wetland to the trap in which they were captured.

Therefore, the maximum distances shown in Table II should not be confused with the

maximum distances that amphibians are capable of traveling. Likewise, the data in Table

IV should not be inferred to mean that amphibians prefer closed xeric hammock to the

other habitat types in Florida uplands. These data do suggest avenues for potential

research, however.

Although the data are not amenable to analysis of species’ preferences because of the

biased sampling protocol, it appears that burrow-using terrestrial frogs (toads, spadefoots,

narrow-mouthed toads) are more likely than the more arboreal and aquatic species (hylids

Source : MNHN, Paris

Dopp

Table II. - Species collected and distances (m) from nearest water body for amphibians

captured during funnel trapping in upland habitats of north-central Florida, 1989 -

1990. SD, standard deviation.

Species

Total number captured

Mean + SD (range)

Acris gryllus

Bufo quercicus

Bufo terrestris

Eleutherodactylus planirostris'

Gastrophryne carolinensis

Hyla cinerea

Hyla femoralis

Hyla squirella

Notophthalmus perstriatus

Pseudacris ocularis

Rana utricularia

Scaphiopus holbrooki

! Has terrestrial development.

383 + 81.4 (255-492)

574 + 216.8 (404-914)

515 + 202.2 (46-914)

478 + 136.7 (46-895)

420 + 216.8 (42-914)

545 + 181.1 (457-914)

266 + 317.5 (42-815)

594 + 188.3 (446-914)

225 + 180.2 (42-709)

434

539 + 211.2 (95-914)

Table III. - Trap distances (m) in relation to nearest water body on the Ordway Preserve. SD,

standard deviation.

| Name Wetland type Number of traps pres ? M pr

Blue Pond 8 FEU 156.5

Enslow Lake 20 DA 3172)

Goose Lake 10 Soin 76 (17.8)

One-Shot Pond 30 QE 7 91 (21.3)

Ross Lake 22 hé 180 (42.2)

Smith Lake 10 FRET 64 (15.0)

Source : MNHN, Paris

48 ALYTES 14 (1)

Table IV. - Amphibian captures in relation to habitat type and trap effort. Data for 1989

captures. SD, standard deviation.

Distance (m) to water: | Number of amphibians

Habitat Number of traps

mean + SD (range) (% of capture)

432. 229.

Sandhilis 59 4 248 (58 %)

Closed Xeric Hammock 10.021006 95 (22 %)

(243.8-579.1)

Open Xeric Hammock He En e 83 (19 %)

TOTAL NUMBER OF AMPHIBIANS IN TRAP

be]

00 100.0 2000 300.0 400.0 500.0 600.0 700.0 8000 900.0 1000

DISTANCE (m) TO NEAREST WATER

Fig. 1. — The relationship between the total number of amphibians captured in funnel traps and the

distance of the funnel trap to the nearest potential breeding site.

Source : MNHN, Paris

Dopp 49

and ranids) to be captured by randomly placed terrestrial traps. Arboreal species travel

well into uplands in dense oak hammocks surrounding lakes on the Ordway Preserve, but

they appear to travel through the tree canopy rather than on the ground (R. BOUGHTON,

personal communication). Ranids are also known to make extensive overland movements

in Florida uplands (e.g., FRANZ et al., 1988), but their travel routes, time and duration of

travel, and susceptibility to trapping are poorly understood.

In upland Florida habitats, amphibians are found in burrows of other animals such

as lizards (e.g., Gastrophryne carolinensis in the burrows of Cnemidophorus sexlineatus),

pocket gophers (Geomys sp.), and gopher tortoises (Gopherus polyphemus), under logs and

other surface debris, and in tree cavities (personal observation). Gopher tortoise burrows,

in particular, are excellent retreat sites, with nine amphibian species recorded from them

(JACKSON & MILsTREY, 1989). The extensive collection of amphibians in funnel traps

suggests that these animals are not sedentary but instead leave burrows and other cover

sites and move around.

Most North American amphibian field studies involving wetland-breeding species are

centered around the breeding site. Such a bias is akin to studying sea turtles only on a

nesting beach. Both amphibians and sea turtles spend a great majority of their lives away

from the habitats most easily studied by researchers. Just as sea turtle biologists have

gained new insights into the life histories of turtles by developing methodologies that allow

them to investigate activity away from nesting beaches, amphibian biologists must adopt

research methods that begin to probe an amphibian’s life away from the breeding pond

(DENTON & BEEBEE, 1992; HEYER et al., 1994). Few researchers have conducted field studies

of amphibians away from the breeding site (e.g., PEARSON, 1955; DENTON & BEEBEE, 1993;

PASANEN et al., 1993; LOMAN, 1994). However, such studies have allowed investigators to

take a more holistic view of the ecological requirements and activities of a species.

There has been great concern for the status of amphibian populations and species

throughout the world (Wake et al., 1991; BLAUSTEIN, 1994; BLAUSTEIN et al., 1994).

Declines have been reported in a variety of habitats and often have involved wetland-

breeding species. Few studies, however, have assessed habitat requirements away from

breeding sites. Biologists conducting inventories of upland communities should routinely

note the distances to nearest wetlands if wetland-breeding amphibians are found.

Management guidelines that promote wetland protection in order to conserve

amphibians yet ignore non-breeding upland habitats (e.g., WiLsON, 1994) are destined to

failure if resident animals move far from ponds and other wetlands. Buffer zones need to

be established around breeding ponds to ensure survival of the amphibian community. In

this regard, 82.9 % of the amphibians I captured were within 600 m of the nearest breeding

site, although I could not determine if this distance would be effective at protecting the

local amphibian community because of the study’s sampling biases. DuBois (1991: 396)

suggested that in tropical regions protection of a buffer zone of 100 to 500 m along each

side of watercourses would help conserving a large proportion of the batrachofauna. The

need for buffer zones to protect wetland-resident turtle populations has also been

recognized (BURKE & GiIBBONS, 1995; K. BUHLMANN, personal communication).

Source : MNHN, Paris

50 ALYTES 14 (1)

ACKNOWLEDGMENTS

1 thank the Board of the Katharine Ordway Preserve-Swisher Memorial Sanctuary, especially

John EISENBERG, for allowing me to carry out these observations on the Preserve. Joseph A. BUTLER,

Richard FRANZ, Joseph C. MITCHELL, and Joseph PECHMANN offered helpful comments on the

manuscrit.

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Source

AVTES

International Journal of Batrachology

published by ISSCA

EDITORIAL BOARD FOR 1996

Chief Editor: Alain Dupois (Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire

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

Deputy Editor: Janalee P. CALDWELL (Oklahoma Museum of Natural History, University of Oklahoma,

Norman, Oklahoma 73019, U.S.A.).

Editorial Board: Jean-Louis ALBARET (Paris, France), Ronald G. ALTIG (Mississippi State University,

U.S.A.); Emilio BALLETTO (Torino, Italy); Alain COLLENOT (Paris, France); Günter GOLLMANN (Wien,

Austria), Tim HALLIDAY (Milton Keynes, United Kingdom); W. Ronald Hever (Washington,

U.S.A.}; Walter HÔDL (Wien, Austria); Pierre JoLy (Lyon, France), Masafumi MATSUI (Kyoto,

Japan); Jaime E. Péraur (Mérida, Venezuela); J. Dale RoBerTs (Perth, Australia); Ulrich SINSCH

(Koblenz, Germany); Marvalee H. Wake (Berkeley, U.S.A.).

Technical Edirorial Team (Paris, France): Alain Dumois (texts); Roger Bour (tables); Annemarie

OuLer (figures).

Index Editors: Annemarie OHLER (Paris, France); Stephen J. RicHarDs (Townsville, Australia).

GUIDE FOR AUTHORS

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

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

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

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

species, interesting behaviors, etc.

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

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

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

Spanish abstract, acknowledgements, literature cited, appendix.

Figures and tables should be mentioned in the text as follows: fig. 4 or Table IV. Figures should not

exceed 16 X 24 cm. The size of the lettering should ensure its legibility after reduction. The legends of figures

and tables should be assembled on a separate sheet. Each figure should be numbered using a pencil.

References in the text are to be written in capital letters (BOURRET, 1942; GRar & POLLS PELAZ, 1989;

INGER et al., 1974). References in the literature cited section should be presented as follows:

BouRkeT, R., 1942. — Les Batraciens de l'Indochine. Hanoï, Institut Océanographique de l’Indochine: i-x

+°1:547, pl. I-IV.

Grar, J.-D. & POLLS PELAZ, M., 1989. — Evolutionary genetics of the Rana esculenta complex. In: R. M.

DawLey & J. P. BOGART (eds.), Evolution and ecology of unisexual vertebrates, Albany, The New York

State Museum: 289-302.

INGER, R. F., Vors, H. K. & Voris, H. H., 1974. — Genetic variation and population ecology of some

Southeast Asian frogs of the genera Bufo and Rana. Biochem. Genet., 12: 121-145.

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

amphibian morphology, systematics, biogeography, evolution, genetics or developmental biology, or to

Janalee P. CALDWELL (address above) if dealing with amphibian population genetics, ecology, ethology or

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

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du Muséum national d'Histoire naturelle, Paris, France).

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Numéro de Commission Paritaire: 64851.