ee Gun?

ISSN 0753-4973

AINTTES

INTERNATIONAL JOURNAL OF BATRACHOLOGY

January 1997 Volume 14, N° 4

Source : MNHN, Paris

International Society for the Study

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Source : MNHN, Paris

AINTES

INTERNATIONAL JOURNAL OF BATRACHOLOGY

January 1997 Volume 14, N° 4

Alytes, 1997, 14 (4): 129. 129

Editorial: 15 vears of Alytes

Founded in February 1982, Alytes is now reaching the term of its fifteenth year of existence.

Started as a modest national bulletin, its printing standard and the quality of its contents have been

regularly increasing over the years. The journal has now published a number of interesting and

important papers and is now recognized worldwide as a major scientific periodical dealing with all

aspects of the study of amphibians. This is also testified by the growing mean quality of manuscripts

submitted for publication in the journal over the years.

The experience accumulated by the editorial team of the journal over this fifteen-year period has

allowed us to make important progress in several respects, and, in particular, regarding the standards

of preparation and processing of manuscripts necessary to obtain a high quality journal. We think it

is useful to share this experience with our readers and with potential authors of papers intended for

publication in Alytes. For this reason, we present in this issue very detailed Jnstructions to Authors,

which will hopefully prove useful to colleagues to prepare high quality manuscripts and will

contribute to a further increase in the standards of the journal.

We feel that the experience of these first fifteen years has amply demonstrated that an

international journal of batrachology was really needed. Gathering together of papers once dispersed

in various journals, written in three major languages, and concerning very diverse aspects of the study

of a relatively poorly known group of vertebrates, has proved extremely stimulating, and will no

doubt become more and more productive as time goes on. The recent international interest in the

problems of conservation of amphibian populations and species has also clearly emphasized the need

for a particular effort of the international community, not only to understand the causes of the

observed declines and to try and stop or reduce them, but also to increase and speed up our efforts

for the mere work of inventory and description of amphibian species on our planet: it is now clear

that, if we delay considering this task as a major priority for amphibian specialists of our time, many

amphibian taxa will become extinct in the forthcoming years, without their having even been collected

and recognized by man. This is well exemplified in this issue of A/ytes, which contains the descriptions

of several new species, all of which are clearly threatened with extinction, and some of which may even

be already extinct as these descriptions are published. It is rather unusual to publish a paper dealing

with the contemporaneous fauna that describes four new species but at the same time states that they

were probably “already” extinct when these descriptions were written. Unfortunately, it is very likely

that in the coming future we will have to confront such situations more and more often, as a

consequence of ongoing massive destructions or extensive modifications of amphibians’ habitats

throughout the world. It results from this that basic alpha taxonomic work of inventory and

description of amphibian species is an urgent matter that should deserve all our attention, including

the creation of academic positions and financial support for field work, collection management and

descriptive activity. All well considered, this might be much more urgent than concentrating all efforts

on more “theoretical” issues of biology, however interesting and important the latter may be.

Understanding of these priorities is the reason for the clear choice made in Alytes to encourage

publication of “purely descriptive” papers of high quality, which has been characteristic of the journal

since its beginnings and which will continue to be so in the years to come.

Alain DuBois

Alytes Chief Editor

Bibliothèque Centrale Muséum KA 19 December 1996

3 3001 00037987 3

\

Source : MNHN, Paris

Alytes, 1997, 14 (4): 130-146.

Description of two new frogs of the genus

Mantidactylus from Madagascar,

with notes on Mantidactylus klemmeri

(Guibé, 1974) and Mantidactylus webbi

(Grandison, 1953)

(Amphibia, Ranidae, Mantellinae)

Miguel VENCES *, Frank GLAW * & Franco ANDREONE **

* Zoologisches Forschungsinstitut und Museum Alexander Koenig,

Adenauerallee 160, 53113 Bonn, Germany

** Museo Regionale di Scienze Naturali, Sezione di Zoologia, Via G. Giolitti 36, 10123 Torino, Italy

Two new species of the endemic Malagasy frog genus Mantidactylus were

discovered during recent fieldwork in northeastem Madagascar, and these are

formally named and described in this paper. The first one inhabits mossy stone

habitats in the Marojezy massif and is mainly characterized by its olive-

greenish coloration and a short snout. The second new species is similar but

larger and has a copper red iris. It is characterized by a very distinctly

protruding inner metatarsal tubercle which is less developed in other known

Mantidactylus. Both new species are tentatively included in the subgenus

Gephyromantis, but their relationships with other Mantidactylus remain

obscure. Morphologically they are most similar to M. klemmeri and M. webbi;

we therefore provide an updated diagnosis of these two species and a detailed

description of the call of M. klemmeri, as well as first data on its ecology and

coloration in life.

INTRODUCTION

The systematics of the frog genus Mantidactylus Boulenger, 1895, endemic to

Madagascar, has been the subject of several studies during the last 20 years. The

biosystematic investigations of BLOMMERS-SCHLÔSSER (1979) showed that the distinction

between the genera Gephyromantis and Mantidactylus, as made by GuIBé (1978) and

previous workers, was largely artificial. As a conclusion, BLOMMERS-SCHLÔSSER (1979)

considered Gephyromantis as a junior synonym of Mantidactylus. She also described two

new species of the genus and assigned 31 Mantidactylus species to 10 phenetic species

groups. BLOMMERS-SCHLÔSSER & BLANC (1991) assigned all Mantidactylus species to these

groups and analysed phylogenetic relationships within Mantidactylus. The cladogram

presented by these authors characterized the species groups by assumed synapomorphies,

Source : MNHN, Paris

VENCES, GLAW & ANDREONE 131

and thus suggested that each represents a monophyletic unit. DuBois (1992) elevated some

of these groups to subgeneric rank. Based on new biosystematic data, GLAW & VENCES

(1994) argued that the monophyly of the subgenera within Mantidactylus would be more

convincing recognizing four additional subgenera. They also noted that no phylogenetic

arguments remain to regard Laurentomantis Dubois, 1980 as a genus separate of

Mantidactylus, and thus considered it as a subgenus, transferring the three Laurentomantis

species to Mantidactylus. Following GLAW & VENCES (1994), 12 subgenera within

Mantidactylus are presently accepted: Mantidactylus Boulenger, 1895; Gephyromantis

Methuen, 1920; Hylobatrachus Laurent, 1943; Laurentomantis Dubois, 1980; Blommersia

Dubois, 1992; Brygoomantis Dubois, 1992; Guibemantis Dubois, 1992; Spinomantis

Dubois, 1992; Chonomantis Glaw & Vences, 1994; Ochthomantis Glaw & Vences, 1994;

Pandanusicola Glaw & Vences, 1994; Phylacomantis Glaw & Vences 1994.

Since 1990, eight new Mantidactylus species have been described (BLOMMERS-

SCHLÔSSER & BLANC, 1991; GLAW & VENCES, 1992b, 1994), one species was transferred to

this genus from the genus Boophis (GLAW & VENCES, 19924) and three species were

resurrected from synonymy (BLOMMERS-SCHLÔSSER & BLANC, 1991; RAXWORTHY &

NussBAUM, 1994; ANDREONE & GAVETTI, 1994). Despite this intensive work on the genus,

the species inventory of Mantidactylus is far from being complete. The continued discovery

of new species even in well studied areas shows that the genus is much more speciose than

presently recognized (59 named species according to GLAW & VENCES, 1994).

The subgenus Gephyromantis currently contains 10 species which are arranged in two

species groups according to GLAW & VENCES (1994): the M. asper group with four

scansorial primary forest species, which can be found during the day on the forest floor

and which call predominantly at night from leaves and branches; and the M. boulengeri

group with six rather small ground-dwelling frogs which call mainly during the day.

Probably closely related subgenera are Laurentomantis with three species and Phylacoman-

tis with six species. The monophyly of these taxonomic units (subgenera and species

groups) must still be verified since the attribution of species to subgenera within

Mantidactylus is often only based on phenetic similarity, the diagnostic synapomorphies of

the groups generally being derived from studies of only a few species.

Two species of the subgenus Gephyromantis differ from other species of this subgenus

mainly in terms of their coloration which is partly greenish, and their ecology which is

associated with mossy rocks along forest brooks. These species are M. webbi, and one

species which up to now was regarded as M. klemmeri (GLAW & VENCES, 1994).

Fieldwork conducted during 1994 and 1995 yielded new data on these two forms, and

resulted in the discovery of two additional species which are morphologically similar to M.

webbi and to the species previously regarded as M. klemmeri. After examination of the

types of Mantidactylus klemmeri and M. webbi in the context of the two similar, newly

discovered species, we conclude that the four forms studied by us are M. klemmeri, M.

webbi, and two unnamed forms which we will here describe as new species.

Source : MNHN, Paris

132 ALYTES 14 (4)

MATERIAL AND METHODS

Specimens were captured by hand both during the day and at night with the aid of

flashlights. They were fixed for a few minutes in 96 % ethyle alcohol, and stored in 70 %

ethyle alcohol. Live animals were photographed to document color and pattern variation.

Morphometric measurements were taken by the first author with a ruler to the nearest 0.5

mm or with a precision calliper to the nearest 0.1 mm. Abbreviations used are SVL

(snout-vent length) and HW (head width). Tympanum and eye diameter were measured

along a horizontal plane.

Calls of M. klemmeri were recorded with a Tensai portable tape recorder and an

external VIVANCO EM 238 microphone. These were analysed with the sound analyzing

system “MEDAV Spektro 3.2”.

Type material was examined both in the Natural History Museum, London, United

Kingdom (BM) and the Muséum National d'Histoire Naturelle, Paris, France (MNHN).

Other abbreviations used in this paper are as follow: MRSN, Museo Regionale di Scienze

Naturali, Torino, Italy; ZFMK, Zoologisches Forschungsinstitut und Museum Alexander

Koenig, Bonn, Germany.

RESULTS

THE IDENTITY OF MANTIDACTYLUS KLEMMERI

In 1994, GLaw & VENCES (color plate 101) showed a picture of a brook-dwelling frog

from the Marojezy mountains which they considered to be Mantidactylus klemmeri. The

pictured species occurred along brooks near the M. klemmeri type locality and was

morphologically similar to that species. Furthermore, its unique character combination

separated it clearly from any other described Mantidactylus. In 1995, however, F. GLAW

collected a second frog species in Marojezy which inhabited the forest floor and showed

some important morphological and ecological differences from the species previously

assigned to M. klemmeri. These differences are summarized in tab. 2, whereas tab. 1 shows

the absolute measurements of all specimens considered in the present publication. A direct

comparison with the types of Mantidactylus klemmeri showed that the species inhabiting

the forest-floor should be assigned to this taxon. An updated diagnosis of M. klemmeri

together with first data on its advertisement call is presented below. The brook-dwelling

form is consequently described as a new species.

Source : MNHN, Paris

VENCES, GLAW & ANDREONE 133

Tab. 1. - Measurements (all in mm) of the Mantidactylus specimens considered in the present study.

Number: abbreviation of museum where specimen is held, and its collection number. Holotypes

are marked with a * behind the number, paratypes with a + behind the number. Other

abbreviations used are: M, male; F, female; J, juvenile; SVL, snout-vent length; HW, head

width; Eye, horizontal eye diameter; Tym, horizontal tympanum diameter; E-N, distance

between anterior eye margin and nostril; N-ST, distance between nostril and snout tip; HL,

hand length: FLIT, foot length including tarsus; TibT, tibiotarsal articulation reaching (1) eye,

(2) beyond eye, (3) nostrils, (4) snout tip, (5) slightly beyond snout tip, (6) clearly beyond snout

tip; ToeL,, relative length of third toe compared with fifth toe; FGS, size of femoral glands,

given as length x width; FGD, distance between inner margins of glands on opposite femurs.

All specimens are from the type localities of the respective species, except for MNHN 1975.781

(M. klemmeri; female from the Chaînes Anosyennes) and MNHN 1975.951-952 (male and

female of M. webbi from Andohahela).

Species and number | Sex SVL HW Eye Tym E-N N-ST HL FLiT TibT ToeL FGS ro |

Mantidacrylus klemmeri

MNHN 1973.960+ M 208 73 30 12 20 1.2 69 165 (5 1515 27

MNHN 1973,959 + M 208 72 26 II 23 15 62 171 -— 114 26

MNHN 1973.957 + M 200 69 30 09 22 13 66 175 (4) 19x12 24

MNHN 1973.958+ M 212 66 28 09 22 13 GS 160 (S) - -

ZFMK 59944 M 20.5 70 27 09 22 1.5 70 162 (6) 20414 1.5

ZFMK 59943 M 195 65 25 09 22 15 62 156 (5 241.7 22

ZFMK 59942 M 20.5 66 25 10 22 1.5 64 173 (S 1813 27

MNHN 1973.962 + F 208 65 27 09 22 12 65 172 (5

MNHN 1973.961 + F 240 73 33 12 25 IS 7.7 184 (4)

MNEHN 1973.956+ F 240 72 3.1 10 25 14 70 185 -

MNHN 1973.955* F 240 75 28 II 26 14 8O 189 (5

MNHN 1973.963+ FL AGE ME NT EN Le Ou RE

MNHN 1975.781 F 26 VS 32 14 25) 22 82 &@ 3-5

Mantidacrylus rivicola

ZFMK 57428* M 225 75 26 10 20 14 78 16.7 (4) 1.513 0.8

ZFMK 59898+ M 243 88 34 17 25 17 80 180 (5

ZFMK 57429 + F 238 78 3.2 12 20 16 75 163 (2)

ZFMK 59946 + F 243 77 29 14 19 14 70 165 (4)

ZFMK 59945 + F xl

ZFMK 59947 + J 2

Mantidactylus silvanus

MRSN A1661* 30.5 10.5 39 22 29 20 95 3<5 2.2x1.5 ca. 1.0

Mantidactylus webbi

MNHN 1975.95

MNHN 1975.92

ZFMK 52725

ZFMK 52726

Source : MNHN, Paris

134 ALYTES 14 (4)

Tab. 2. - Comparison between the four Mantidactylus species treated in the present study.

Morphometric data from tab. 1. IMT: inner metatarsal tubercle. Other abbreviations as in tab.

1. Values are given as range; mean + standard deviation is given in brackets. Significant

differences (Mann-Whitney U test) were detected between value pairs marked with stars (* P <

0.05; ** P <0.005). Values for snout-vent length are given separately for males (M) and

females (F). For snout-vent length only specimens from the type localities were considered;

other values are based on all specimens listed in tab. 1.

M. klemmeri M. rivicola M. silvanus M. webbi

SL. (M) Imm 19.5-21.2 22.5-243 30.5 249

SVL (F) mm] 20.8-24.0 238-243 _ 30.5

HW:SVL 0.30 - 0.35 (0.324-0.02) | 0.30 - 0.36 (0.33 0.02) 0.34 0.34 - 0.36 (0.3540.01)

HL:SVL 0.29 - 0.34 (0.320.02) | 0.29 - 0.35 (0.31 +0.02) os 0:31 -0.35 (0.3340.02)

FLITSVL 0.40 - 0.88 (0.770.12) | 0.64 - 0.74 (0.71 +0.04) 0.69 0.71 - 0.80 (0.75 0.04)

Tym:Eye +*0.30 - 0.44 (0.37-0.05) | 0.36 - 0.50 (0.41 +0.06) 0.56 **0.45 - 0.60 (0.52:+-0.08)

{e-N + N-ST):SVL | *0.14 - 0.19 (0.170.01) | *0.14 - 0.17 (0.15+0.01) 0.16 0.16-0.17 (0.170.004)

Eÿe-N: NST **0.48 - 0.68 (0.60:+-0.07) | **0.68 - 0.80 (0.730.05) 0.69 0.49 - 0.75 (0.6140.1)

General habitat forest floor forest brooks forest brooks? forest brooks

Micro-habitat on leaf liter/roots on mossy rocks at night on leaves on mossy rocks

Habitus very slender slender less slender less slender

Dorsal skin finely granular some large granules finely granular coarsely granular

Snout long, pointed short long long

IMT small protruding strongly protruding reduced

Color brownish olive greenish olive greenisbrown | olive greenish

Tab. 3. - Parameters of six notes from calls of Mantidactylus klemmeri, recorded in the Marojezy Strict

Nature Reserve. Notes 1-3 from Camp 3 (26.02.1995; air temperature 23.8°C), notes 4-6 from

Camp 4 (28.02.1995; air temperature 22.5°C). All temporal measurements are given in

milliseconds (ms).

Note duration

Number of primary pulses

Duration of primary pulses (range)

Duration of primary pulses (mean)

Interval between primary pulses (range)

Interval between primary pulses (mean)

Number of double clicks

Duration of double clicks

Duration of secondary pulse series

Number of secondary pulses 17

Repetition rate of secondary pulses 29.6

Source : MNHN, Paris

ALYTES 14 (4) 135

Mantidactylus klemmeri (Guibé, 1974)

Gephyromantis klemmeri Guibé, 1974; GuIBé, 1978.

Mantidactylus klemmeri: BLOMMERS-SCHLÔSSER & BLANC, 1991; GLAW & VENCES, 1992b, 1994

(partim).

Mantidactylus (Gephyromantis) klemmeri: Dugois, 1992.

Diagnosis. — A small, extremely slender, brownish frog, belonging to the genus

Mantidactylus as is evident from the presence of femoral glands in males (not recognizable

in females). The absence of webbing between the toes, in combination with the small size,

connected lateral metatarsalia, and the presence of a subgular vocal sac (laterally blackish)

in males allow a distinction to be made from most other Mantidactylus. M. eiselti and M.

thelenae are morphologically very similar but their advertisement calls differ distinctly. M.

webbi and the two new species described below have a different coloration (partly

greenish). Additionally, M. webbi males can be recognized by the white vocal sac, and the

two new species have distinct, protruding inner metatarsal tubercles (small and not

protruding in M. klemmeri).

The tibiotarsal articulation of M. klemmeri reaches at least the tip of snout or beyond

as was noted by GUIBÉ (1974) in the original description, not between nostrils and tip of

snout as was first stated by BLOMMERS-SCHLÔSSER & BLANC (1991) and subsequently

quoted by GLAW & VENCES (1992, 1994).

Type material. — Holotype MNHN 1973.955, adult female, and paratypes MNHN

1973.956-963, four adult males, three adult females and one juvenile specimen, all collected

by C.-P. BLANC in December 1972 in the “Massif du Marojezy” at an altitude of 600 m.

Measurements of the types and of three additional ZFMK specimens from the type

locality are given in tab. 1.

Color in life. — Based on color slides of ZFMK specimens, the dorsum and flanks are

marbled grey or brownish without distinct transversal markings. No specimens with a light

median dorsal stripe were found at Marojezy. Indistinct dark crossbands are present on

fore- and hindlegs. The lower lip is dark with some rather regularly spaced light spots.

Sometimes this pattern also occurs on the upper lip, resulting in transverse dark and light

bands. In other specimens a light frenal stripe is faintly recognizable. The tympanic area

is brown. This brown patch runs from the eye to the insertion of forelegs. The thorax and

throat are marbled brown and white, the marbling on the venter and ventral limb surfaces

being lighter and less contrasting. A light median line is sometimes present on the throat.

Males have a distinct subgular vocal sac which is laterally blackish, indicating a possible

paired or bilobate shape of the inflated sac.

Distribution. — Apart from the type material and our specimens, which were all collected

in the Marojezy Strict Nature Reserve (Réserve Naturelle Intégrale) in northeastern

Madagascar, one additional specimen from the Anosy mountains (“Chaînes Anosyennes”)

in southeastern Madagascar is available in the MNHN. This female (MNHN 1975.781) is

somewbhat larger than the other specimens and has a light median dorsal stripe, but shows

no other morphological differences (tab. 1) from the types. We therefore consider it as

belonging to the species M. klemmeri pending the obtention of more data.

Source : MNHN, Paris

136 ALYTES 14 (4)

Fig. 1. — Specimen of Mantidactylus klemmeri from Marojezy Strict Nature Reserve, Camp 3,

26.02.1995.

Habitat and habits. — Specimens were found in 1995 at Marojezy between 600 and 1300

m elevation (up to above Camp 4). Males call during the day from the ground, mostly

from between tree roots where specimens hide when disturbed. Calling males occur in

small groups clustered in certain areas, but do not form dense choruses. They are not

confined to the vicinity of water bodies. No activity was observed at night.

Call. — The calls were recorded on 26.02.1995 and 28.02.1995 in the Marojezy reserve

(near Camp 3, about 700 m elevation, at 23.8°C, and at Camp 4, 1250 m, at 22.5°C). The

first locality is very near to the type locality of M. klemmeri: “Massif du Marojezy

(600 m)” (Gumé, 1974: 1175).

As in some other members of the M. boulengeri group (M. boulengeri, M. decaryi, M.

eiselti, M. thelenae), the general structure of a call can be described as a series of notes.

In M. klemmeri a call consists of up to 6 notes. Note duration ranges from 626 to 982 ms;

the duration of intervals between two notes of a series is about 2 s (measured minimum

1709 ms).

The general structure of a note is often quite complex and can be described as a series

of 1-5 primary pulses, followed by 0-2 double-clicks, followed by a series of 17-31

secondary pulses. The pulse repetition rate of the secondary pulses is 56.7-61.1/s at a

temperature of 23.8°C (recorded at Camp 3). The notes recorded at higher altitude (Camp

4) had a much lower pulse repetition rate (26.3-33.9/s) which may only partly be explained

Source : MNHN, Paris

VENCES, GLAW & ANDREONE 137

0 500 1000 ms

primary puises _ double click secondary puises

0 500 1000ms

Fig. 2. — Sonagram and oscillogram of the call of Mantidactylus klemmeri, recorded in the Marojezy

Strict Nature Reserve, Camp 3, on 26.02.1995 (air temperature 23.8°C).

by the slightly lower temperature (22.5°C). The frequency of all call components range

from 3700 to 5100 Hz. Detailed data of six notes are summarized in tab. 3.

The structure of the call of M. klemmeri is similar to that of Mantidactylus thelenae.

However, the call of the latter species is characterized by a shorter note duration (540-650

ms) and shorter intervals between notes (1140-1480 ms). A striking difference is the much

higher pulse repetition rate (about 250/s) in the call of M. thelenae.

Subgeneric attribution. — As indicated by its morphology (subgular vocal sac laterally

blackish, connected lateral metatarsalia, femoral glands present only in males) and

especially natural history (calling not concentrated around water, therefore probably direct

development), this species belongs to the subgenus Gephyromantis, M. boulengeri group. It

is probably closely related to M. eiselti and M. thelenae.

Source : MNHN, Paris

138 ALYTES 14 (4)

Mantidactylus rivicola sp. nov.

Diagnosis. — A rather small olive-greenish frog belonging to the genus Mantidactylus as

is evident from the presence of femoral glands in the males (absent in females). The

character combination of small size, connected lateral metatarsalia, scarcely webbed feet,

and presence of a protruding inner metatarsal tubercle is diagnostic for this species. Other

riparian Mantidactylus with partly greenish dorsal coloration are Mantidactylus lugubris

(subgenus Hylobatrachus) and young Mantidactylus microtympanum (subgenus Manti-

dactylus) which have strongly webbed feet and separated lateral metatarsalia, and M.

webbi which has only rudimentary metatarsal tubercles and a laterally white vocal sac.

Members of the subgenus Laurentomantis generally have a coarse granular skin and a very

broad head; however, the species Mantidactylus cf. malagasius which occurs sympatrically

with M. rivicola is morphologically rather similar to the latter. It can be distinguished by

the more granular skin, different coloration (not greenish), absence of recognizable vocal

sac in males, and slightly longer hindlimbs (tibiotarsal articulations reaching distinctly

beyond snout tip). :

M. rivicola is most similar to the second new species described below: for distinctive

characters see the diagnosis of that species.

Derivatio nominis. — From the Latin rivus (brook) and colere (to inhabit). This species

typically is an inhabitant of brook edges.

Holotype. — ZFMK 57428, adult male, from near Camp 1, Marojezy Strict Nature

Reserve (Réserve Naturelle Intégrale), northeastern Madagascar, altitude about 300 m

above sea level, collected by F. GLAW, N. RABIBIsOA and ©. RAMILISON on 27.03.1994.

Paratypes. — ZFMK 57429, adult female, same collection data as the holotype; ZFMK

59945-59946, two adult females, ZFMK 59898, adult male, and ZFMK 59947, juvenile,

from same locality as holotype, collected by F. GLAW and O. RAMILISON from 25 to

28.02.1995.

Description of the holotype. — See tab. 1 for measurements. Body slender. Widest part of

head slightly wider than widest part of body. Dorsal outline of head triangular, snout

rather short. Snout protruding over upper jaw in lateral view. Nostrils not distinctly

protruding, with lateral openings. Loreal region slightly concave. Tympanum distinct,

rather small, slightly larger than 1/3 of eye diameter. Distinct supratympanic fold above

the tympanum, which does not distinctly continue posterior to the tympanum. Forelegs

and hindlegs slender. Outer metacarpal tubercle present. Finger discs enlarged, semicir-

cular. No webbing between fingers. Comparative finger length 1 < 2 < 4 < 3. Toe discs

slightly enlarged, smaller than finger discs. Traces of webbing between toes. Comparative

toe length 1 < 2 < 3 < 5 < 4. Inner metatarsal tubercle rather large (length about 1 mm),

protruding forewards. Lateral metatarsalia largely connected. Dorsal skin rough, with

longitudinal tubercles in two rows bordering the median line. Some larger tubercles on the

flanks. Small, oblique femoral glands with rather indistinct borders. Vomerine teeth

rudimentary, hardly recognizable. Subgular vocal sac structure distinctly recognizable.

Greyish lateral color on the sac surface possibly indicates that the (subgular) vocal sac may

have a paired or bilobate shape when inflated.

Source : MNHN, Paris

VENCES, GLAW & ANDREONE 139

Fig. 3. — Paratype of Mantidactylus rivicola, ZFMK 57429, from Marojezy Strict Nature Reserve,

Camp 1, 27.03.1994.

Coloration of the holotype. — After 1.5 year in alcohol, the head, dorsum and flanks are

mossy olive greenish with two indistinct dark transversal markings. A light median line

runs from tip of snout to the anus. In the middle of the dorsum to the left and the right

of this line there are very thin dark and discontinuous folds; on the anterior dorsum these

folds run laterally towards the eyes. Fore- and hindlimbs as well as hands and feet have

distinct dark crossbands. The tips of the fingers are white. The eyes are completely white.

The lower lip is dark, broken by some alternating light spots, which do slightly correspond

to the markings on the upper lip. The tympanum is brownish and lighter than the skin

surrounding it. There is no distinct brown spot from the eye through the tympanum to the

insertion of the foreleg and no light stripe along the upper lip. Throat and venter are grey

with no distinct spots. A distinct white stripe extends from the tip of snout to the thorax.

In the thorax area this stripe is surrounded by brown. The ventral surface of the limbs is

greyish.

Description of paratypes. — Morphological features of the paratypes are consistent with

those of the holotype. See tab. 1 for measurements. The supratympanic folds are quite

irregular, often nearly unrecognizable behind the tympanum or fusing with large lateral

tubercles. Coloration of the paratypes is similar to that of the holotype. À median dorsal

stripe is absent in all paratypes except ZFMK 59947. The dark transverse markings on the

Source : MNHN, Paris

140 ALYTES 14 (4)

dorsum are more distinct in ZFMK 57429 and 59945, the dark bands on hands and feet

are very distinct in ZFMK 59898. The latter specimen also has some white tubercles on

the flanks which are less distinct in the other specimens. The light stripe on throat and

thorax is broken in most paratypes (complete only in the dorsally striped paratype ZFMK

59947), whereas it is reduced to a light spot on the thorax in ZFMK 59946. None of the

paratypes has a distinct brown spot in the tympanic region or a light stripe on the upper

lip.

ZFMK 57429 was pictured in life by GLAW & VENCES (1994) as color photo 101. The

iris was golden, the pupil probably of horizontal shape. Dorsum was olive greenish with

dark grey-brown markings. Patterns were similar to those in preservative. Distinct white

tubercles were present on the flanks. The fingertips were white.

Other specimens. — Additional specimens are kept in the herpetological collection of the

University of Antananarivo, Madagascar. Collection data are the same as for the type

material.

Distribution. — Only known from the type locality.

Habitat and habits. — Males and females were found during day and night along stony

brooks in the rain forest. No calling activity could be observed. Specimens were mostly

sitting on mossy stones in the brooks.

Subgeneric attribution. — Tentatively placed in the subgenus Gephyromantis, Mantidacty-

lus boulengeri group.

À NEW SPECIES FROM Nosy MANGABE

In 1953, A. GRANDISON described Rhacophorus webbi which had been collected by C.

S. Weg8 in the small Malagasy offshore island Nosy Mangabe. GUIBÉ (1978) transferred

the species to the genus Gephyromantis. BLOMMERS-SCHLÔSSER & BLANC (1991) included

the species in Mantidactylus. The presence of a paired subgular vocal sac in combination

with connected lateral metatarsalia were the main characters used by GLAW & VENCES

(1994) to group Mantidactylus webbi within the subgenus Gephyromantis. However, it

differs from all other members of this taxonomic group by the laterally white (not

blackish) vocal sac (GLAW & VENCES, 19924).

ANDREONE (1993) made reference to an unidentified Mantidactylus species of which

two specimens had been photographed at Nosy Mangabe. It was similar to M. webbi by

general habitus but differed by the more finely granular skin (coarsely granular in M.

webbi) and by body and iris coloration. During a second visit of F. ANDREONE to Nosy

Mangabe, a third specimen of this form was seen and collected. Further study has

confirmed that it belongs to an undescribed species of Mantidactylus which is superficially

similar to M. webbi. Before the formal description of the new species, we present here a

short diagnosis and a review of biology and distribution of M. webbi.

Source : MNHN, Paris

VENCES, GLAW & ANDREONE 141

Fig. 4. — Specimen of Mantidactylus webbi from Nosy Mangabe, January 1991 (not preserved).

Mantidactylus webbi (Grandison, 1953)

Rhacophorus webbi Grandison, 1953.

Gephyromantis webbi: GuBÉ, 1978.

Mantidactylus webbi: BLOMMERS-SCHLÔSSER & BLANC, 1991; GLAW & VENCES 1992a-b, 1994;

ANDREONE, 1993.

Mantidactylus (Gephyromantis) webbi: Dunois, 1992.

Diagnosis. — A small olive-greenish frog which belongs to the genus Mantidactylus as is

evident from the presence of femoral glands in the males. Metatarsal tubercles are

rudimentary. Nostrils are situated dorsolaterally. The paired subgular vocal sac of the

male is characteristic for most species of the subgenera Phylacomantis and Gephyromantis.

The connected lateral metatarsalia in combination with the small size and greenish

coloration allow a distinction from all Phylacomantis, whereas the presence of a laterally

white (instead of blackish) vocal sac in males is a unique feature of M. webbi and can serve

as an immediate recognition character.

Holotype. — BM 1953.1.2.34, adult female, SVL 33.3 mm. The original description

(GRANDISON, 1953) included a picture of this specimen.

Distribution. — The type locality is the small Malagasy offshore island Nosy Mangabe. On

the mainland near Nosy Mangabe we found this species at Voloina and Navana (GLAW &

VENCES, 1992a) and at Ambanizana (ANDREONE, unpublished). Specimens from Andohahela

Source : MNHN, Paris

142 ALYTES 14 (4)

in south-eastern Madagascar (MNHN 1975.951-952) were tentatively excluded from M.

webbi by GLAW & VENCES (1994); however, after detailed measurements of these specimens

(see tab. 1), we consider them as M. webbi pending further study.

Habitat and habits. — Activity and calls were noticed only during the day. The species lives

and calls on big mossy stones along brooks in primary forest and was never found at

distances of more than 20 m from the water (GLAW & VENCES, 1992a; ANDREONE, 1993).

Two egg clumps (one found in March, one in August), each with 7 yellowish eggs

(diameter 4 mm), were found on rocks in the brooks (GLAW & VENCES, 19924; ANDREONE,

1993). One of the egg clumps was guarded by a male during the night (ANDREONE, 1993).

Four additional clumps of similar size were observed by F. ANDREONE in August 1994 at

Nosy Mangabe.

Calls. — The advertisement call has been described by GLAW & VENCES (1992a). It is a

series of up to 10 unharmonious notes (note repetition rate 6.25/s). Note duration is 26-46

ms, duration of intervals between notes is 122-168 ms, and frequency ranges from 0.5 to

5 kHz.

Subgeneric attribution. — Presently placed in the subgenus Gephyromantis, Mantidactylus

boulengeri group (GLAW & VENCES, 1994).

Mantidactylus silvanus sp. nov.

Diagnosis. — A medium-sized member of the genus Mantidactylus as is evident from the

presence of distinct femoral glands and lack of nuptial pads in males (females unknown).

General color olive-greenish to light brown. The most distinctive character is the highly

protruding inner metatarsal tubercle which allows a distinction from all other Mantidacty-

lus thus far known. Additionally, M. silvanus can be distinguished from the superficially

similar M. lugubris, young M. microtympanum, and M. webbi by the same character

combinations specified above in the diagnosis of M. rivicola. A distinction from M. cf.

malagasius (from Marojezy) can easily be made by the smaller size and the granular skin

of that species.

M. silvanus is most similar and probably closely related to M. rivicola. Besides the size

differences between both species, the following distinctive characters can be used for

diagnosis: M. rivicola has a more granular skin on the dorsum, a larger distance between

the femoral glands, a less developed metatarsal tubercle, and a more granular tympanic

region with a more curved and rather discontinuous and irregular supratympanic fold

(smooth, with a regular fold which is curved only partly, in M. silvanus).

Derivatio nominis. — Classical Latin Silvanus (name of the God of forests). This species

lives in lowland forest.

Holotype. — MRSN A1661, adult male, from Nosy Mangabe Special Reserve, northeast-

ern Madagascar, collected by F. ANDREONE on 27.06.1995.

Description of the holotype. — See tab. 1 for measurements. Body more or less slender.

Widest part of head clearly wider than widest part of body. Dorsal outline of head

triangular, snout rather long. Snout protruding over upper jaw in lateral view. Nostrils

Source : MNHN, Paris

VENCES, GLAW & ANDREONE 143

Fig. 5. — Holotype of Mantidactylus silvanus, MRSN A1661, adult male, Nosy Mangabe Special

Reserve, 27.06.1995.

protruding with lateral openings. Loreal region slightly concave. Tympanum distinct, its

diameter accounting for about 3/5 of eye diameter. Tympanic region without granules.

Distinct and continuous supratympanic fold which runs rather straight and curves

downwards only at a certain point behind the tympanum, near the foreleg insertion.

Forelegs and hindlegs slender. Finger discs enlarged, semicircular to triangular. No

webbing between fingers. Comparative finger length: 1 < 2 < 4 < 3. Toe discs enlarged.

smaller than finger discs. Traces of webbing between toes. Comparative toe length:

1<2<3<5 < 4, Inner metatarsal tubercle large, protruding, resembling a “sixth toe”,

its size 1.8 X 0.5 mm. Outer metatarsal tubercle not recognizable. Foot length (not

including tarsus) 13.5 mm. Lateral metatarsalia connected. Dorsal skin rugged to finely

granular with few small dispersed tubercles. Oblique femoral glands, nearly touching each

other, their diameter 2.2 X 1.5 mm. Small but distinct vomerine teeth. No externally

recognizable vocal sac (neither distinct dark color in the angle of the lower jaw nor a skin

fold is visible).

Coloration of the holotype. — After one month in alcohol, the head, dorsum and flanks

are marbled brownish and light grey. Fore- and hindlimbs show distinct brown crossbands

which extend to the fingers and toes. The tips of the fingers are grey. The eyes are dark.

The lower lip is dark, broken by some alternating light spots, resulting in a pattern which

partly corresponds to the markings on the upper lip. The tympanum is light brown. There

is no distinct brown spot from the eye through tympanum to the insertion of foreleg and

no light stripe along the upper lip. Throat, venter and the anterior ventral

Source : MNHN, Paris

144 ALYTES 14 (4)

Fig. 6. — Specimen of Mantidactylus silvanus from Nosy Mangabe, August 1988 (not preserved).

region of the femur are light grey without distinct spots. The posterior region of the femur,

ie. between the femoral glands and anus, as well as the other ventral parts of the limbs,

are more dark grey. An indistinct and incomplete white stripe runs ventrally from the tip

of snout to the thorax.

Coloration in life was yellowish-beige marbled with olive grey and with two dark

transverse markings. The hindlegs were also beige with crossbands which were less distinct

than after preservation. The toes and fingers, especially the fingertips, were white. The

pupil was horizontal, the iris had copper-red coloration. The venter and ventral aspect of

femurs were pinkish translucent, with some silvery white color on the belly.

Other specimens. — Photographs of two specimens from Nosy Mangabe which were not

collected are available. Coloration patterns are in agreement with those of the holotype.

Distribution. — Only known from the type locality.

Habitat and habits. — The limited available data refer to the three specimens observed by

F. ANDREONE. The holotype was collected at night (about 21 h 00) whilst perched on a leaf

of a small tree, about 1 m above the water level of a very small brook flowing through

large stones in primary forest. In sympatry a larger group of calling males of

Mantidactylus redimitus were observed. The second and third specimens were photo-

graphed at night along a forest brook near a small cascade.

Subgeneric attribution. — We tentatively include this species in the subgenus Gephyro-

mantis, Mantidactylus boulengeri group.

Source : MNHN, Paris

VENCES, GLAW & ANDREONE 145

DISCUSSION

The divergence of the different Mantidactylus groups is evident in terms of the

synapomorphies in tadpole morphology, but is much less obvious in adult morphology

(GLAw & VENCES, 1994). Mantidactylus rivicola and M. silvanus show clear affinities to

members of the subgenus Gephyromantis sensu GLAW & VENCES (1994), especially to M.

webbi. However, reproductive biology of none of these three species is sufficiently

understood. The reproductive mode of only two species of the subgenus, M. eiselti (M.

boulengeri group) and M. asper (M. asper group), are known. Both undergo direct

development in terrestrial eggs (BLOMMERS-SCHLÔSSER, 1979; GLAW & VENCES, 1994).

Little doubt remains that M. boulengeri, the subgeneric type species, has a similar breeding

biology, and that direct development is typical for Gephyromantis.

Egg clutches attributed to M. webbi were found overhanging brooks, thus not

independent from water (GLAW & VENCES, 19924; ANDREONE, 1993). It can be assumed

that this species has a different breeding biology, with more or less developed tadpoles

dropping into the brook. A similar reproductive mode may exist in M. rivicola and M.

silvanus, which also live along brooks, but is very unlikely for M. klemmeri, which, like M.

eiselti, calls far from any water body.

The lack of available information points to the fact that new data on the subgenera

Gephyromantis, Laurentomantis and Phylacomantis are needed. The phylogenetic relation-

ships between the various species of these groups, some of which are probably

paraphyletic, need to be resolved as well as their status within the genus Mantidactylus.

Such a revision would also be useful for conservation purposes. Many species of the

subgenera Gephyromantis, Phylacomantis and Laurentomantis are confined to primary

rainforest habitats. Exceptions are M. eiselti and M. thelenae, which also inhabit

secondary fern scrub at the forest edge or in clearings, M. b. boulengeri, which also lives

in wood patches along brooks or in secondary eucalyptus forest, and M. granulatus, which

can live in degraded forest along brooks and small streams. Nevertheless, these frogs are

one of the Malagasy anuran groups which are mostly restricted to little disturbed habitat.

Therefore they may be useful as indicator taxa for the assessment of the status and degree

of forest habitat degradation. Since many of these species are active during the day and

may easily be located by means of their characteristic calls, their presence can be

established even by little experienced researchers. Detailed knowledge of the systematic

status, distribution range and habitat preferences of each species are basic requirements if

they are to be used as habitat indicators.

ACKNOWLEDGEMENTS

We thank Annemarie OHLER and Alain DuBois for their assistance during our visits to the Paris

Museum (MNHN). Nirhy RaBIBisoA and Olivier RAMILISON helped during fieldwork. The work of

Frank GLAW was made possible by cooperation between the Zoological Institute of Antananarivo

and the Zoologisches Forschungsinstitut und Museum A. Koenig (ZFMK), and financially supported

by the “Deutscher Akademischer Austauschdienst” (DAAD), that of Franco ANDREONE by coope-

Source : MNHN, Paris

146 ALYTES 14 (4)

ration between the Parc de Tsimbazaza (Antananarivo) and the Museo Regionale di Scienze Naturali

(Torino) and financially supported by Zoo-Project s.r.l. (Perugia). We are also indebted to two

anonymous referees, and especially to Marius BURGER and Catherine PRICE (Grahamstown), who

helped improving earlier manuscript versions.

LITERATURE CITED

ANDREONE, F., 1993. — Kommentierte Liste von Amphibienfunden auf Madagaskar. Salamandra, 29

(3/4): 200-211.

ANDREONE, F. & GAverri, E., 1994. — On the identity of Mantidactylus aerumnalis (Peracca, 1893)

(Anura: Mantellidae). Boll. Mus. reg. Sci. nat. Torino, 12 (1): 57-71.

BLOMMERS-SCHLÔSSER, R. M. A., 1979. — Biosystematics of the Malagasy frogs. I. Mantellinae

(Ranidae). Beaufortia, 29 (352): 1-71.

BLOMMERS-SCHLÔSSER, R. M. A. & BLANC, C. P., 1991. — Amphibiens (première partie). Faune de

Madagascar, 75 (1): 1-379, pl. 1-12.

Duois, A., 1992. — Notes sur la classification des Ranidae (Amphibiens Anoures). Bull. Soc. linn.

Lyon, 61 (10): 305-352.

Giaw, F. & Vences, M, 19924. — Zur Kenntnis der Gattungen Boophis, Aglyptodactylus und

Mantidactylus (Amphibia: Anura) aus Madagaskar, mit Beschreibung einer neuen Art. Bonn.

zool. Beitr., 43 (1): 45-77.

19926. — A ieldguide to the amphibians and reptiles of Madagascar. Këln, Vences & Glaw: 1-331.

1994. — A fieldguide to the amphibians and reptiles of Madagascar. Second edition, including

mammals and freshwater fish. Kôln, Vences & Glaw: 1-480.

GRANDISON, À. G. C., 1953. — A new species of rhacophorid frog from Madagascar. Ann. Mag. nat.

Hist., (12), 6: 855-856.

Guné, J., 1974. — Batraciens nouveaux de Madagascar. Bull. Mus. nan. Hist. nat., (3), 171 (Zoo!.

116): 1169-1192.

ee 1978. — Les batraciens de Madagascar. Bonner zool. Monogr., 11: 1-140.

RAXWORTHY, C. J. & NUSSBAUM, R. A., 1994. — A rainforest survey of amphibians, reptiles and small

mammals at Montagne d'Ambre, Madagascar. Biol. Conserv., 69: 65-73.

Corresponding editor: Alain Dumois.

© ISSCA 1997

Source : MNHN, Paris

Alytes, 1997, 14 (4): 147-174. 147

A review of the

Eleutherodactylus milesi-like frogs

(Anura, Leptodactylidae) from Honduras

with the description of four new species

James R. MCCRANIE * & Larry David WILSON **

* 10770 SW 164th Street, Miami, FL 33157, USA

** Department of Biology, Miami-Dade Community College, Kendall Campus, Miami, FL 33176, USA

Four new species of the Eleutherodactylus milesi group are described from

the Caribbean versant of Honduras. Eleutherodactylus stadelmani, recently

resurrected from the synonymy of E. milesi, is shown to represent a valid

species, but not in the recently suggested concept. A key to the identification

of the Honduran members of the E. milesi group is also provided.

INTRODUCTION

MCCRANIE et al. (1989) described two new species of streamside frogs of the

Eleutherodactylus milesi group (E. chrysozetetes and E. cruzi) from Honduras. These

authors also assigned frogs from seven localities across northern Honduras to the species

E. milesi Schmidt, 1933, although noting that variation occurred among some of these

populations in several characters previously used by SAVAGE (1975) to diagnose the species.

CAMPBELL (1994) described two new species of this group (E. adamastus and E.

trachydermus) from Guatemala and resurrected E. stadelmani Schmidt, 1936 (type locality:

Portillo Grande, Departamento de Yoro, Honduras) from the synonymy of E. milesi.

CAMPBELL (1994) examined two paratypes of E. stadelmani (sample 4 of MCCRANE et al.,

1989) and concluded that they were conspecific with a series of frogs from the Quebrada

de Oro region, Departamento de Atläntida, Honduras (sample 5 of MCCRANIE et al., 1989)

on the one hand, and were distinct from E. milesi (sample 6 of MCCRANIE et al., 1989)

from the Sierra de Omoa of northwestern Honduras on the other. CAMPBELL (1994) also

examined the frogs in samples 2 and 7 of MCCRANIE et al. (1989), but was undecided on

the specific identities of these specimens. Unfortunately, the two paratypes of E.

stadelmani examined by CAMPBELL (1994) are subadults. Had he examined the adults in the

type series of E. stadelmani, he might have reached different conclusions. Examination by

us of the entire type series of E. stadelmani, plus a series of frogs recently collected from

nearby localities, demonstrates that Æ. stadelmani is indeed a distinct species. Re-

examination of the Quebrada de Oro series demonstrates that this population also

Source : MNHN, Paris

148 ALYTES 14 (4)

represents a species distinct from E. milesi (as noted by CAMPBELL, 1994) and E. stadelmani

as well. In recent years, we have also collected a small series of E. milesi-like frogs from

forested hillsides well above streams, a previously unknown habitat for frogs of this group.

Additionally, a field party from the Museum of Vertebrate Zoology made an incidental

collection of E. milesi-like frogs while conducting salamander studies in the Sierra de

Omoa. Examination of the forest specimens and the MVZ specimens demonstrates that

each series also represents an undescribed species (a few additional specimens that are

conspecific with the MVZ series were also located in the FMNH and MCZ collections).

Finally, re-examination of the series of E. milesi-like frogs from the Sierra de Agalta

(samples 2 and 3 of MCCRANIE et al., 1989) demonstrated that these specimens represent

an undescribed species as well.

Early in this study, when it became apparent that several populations of E. milesi-like

frogs were morphologically distinct, a decision was made to attempt to collect tissues from

as many of these populations as possible. It was hoped that the tissues would substantiate

the taxonomic conclusions based on morphological data. Thus, in February 1995, the

senior author visited the still pristine Quebrada de Oro-Cerro Büfalo region where three

species of the milesi group were known to occur (E. chrysozetetes, E. cruzi, and E. sp.

nov.). However, in 10 days and nights of collecting, not a single streamside Eleuthero-

dactylus was seen, although three species of forest Eleutherodactylus were collected well

above the nearest streams (the forest E. milesi-like frog alluded to above, plus E. chac and

E. ridens). In July 1995, both authors returned to a stream in the Cordillera Nombre de

Dios in northern Yoro where E. stadelmani was common in July-August of 1991 and 1993.

In four daily and nightly visits to this stream by ourselves and/or other members of the

field party, no streamside Eleutherodactylus were seen, even though there was no change

in the streamside vegetation from previous years. Eleutherodactylus milesi also was

common along small streams that flow into the Rio Cusuco in Parque Nacional El

Cusuco, Departamento de Cortés, Honduras, in April 1979, May 1980, August 1982 and

July 1983. The senior author searched these same small streams during three days and four

nights in August 1992 without seeing a single Æ. milesi. Mario EsPINAL also collected in

Parque Nacional El Cusuco on two trips in July-August 1993. Although he collected

several specimens of the forest-occurring E. rostralis and two species of streamside hylids

(Plectrohyla dasypus and P. teuchestes), no E. milesi were collected. These recent collecting

efforts suggest that at least some populations of streamside frogs in the E. milesi group

may be experiencing population declines or have collapsed altogether. Several species of

moderate and intermediate elevation streamside frogs have been documented to have

disappeared from pristine habitats in Costa Rica and are now feared extinct (K. R. Lips,

J. M. SAVAGE, personal communication). Thus, the possibility exists that one or more of

the new species described below may have already vanished from at least their previously

known localities.

Source : MNHN, Paris

MCCRANIE & WILSON 149

MATERIALS AND METHODS

Museum abbreviations follow LEvITON et al. (1985); BMNH, Natural History

Museum, London, UK; FMNH, Field Museum of Natural History, Chicago, USA; KU,

Museum of Natural History, University of Kansas, Lawrence, USA; LACM, Los Angeles

County Museum of Natural History, Los Angeles, USA; MCZ, Museum of Comparative

Zoology, Harvard University, Cambridge, USA; MVZ, Museum of Vertebrate Zoology,

University of California, Berkeley, USA; ROM, Royal Ontario Museum, Toronto,

Canada; UIMNH, University of Illinois Museum of Natural History, Urbana, USA;

UMMZ, Museum of Zoology, University of Michigan, Ann Arbor, USA: USNM,

National Museum of Natural History, Washington, USA.

The numbered characters in each species diagnosis follow the standard established by

LyncH & DUELLMAN (1980). Discussion of some of these characters is expanded in the E.

milesi and E. stadelmani sections as no follow up description is provided for either species.

Additional to the specimens referred to the four new species described below (see species

descriptions), the following Honduran specimens of the E. milesi group were examined,

which include the holotypes and all extant paratypes of E. chrysozetetes, E. cruzi, E. milesi,

and E. stadelmani:

Eleutherodactylus chrysozetetes (30). — Atläntida: KU 209035 (holotype), 209036; USNM

497054-81.

Eleutherodactylus cruzi (2). — Atläntida: KU 209037 (holotype), 209038.

Eleutherodactylus milesi (89). — Copän: KU 209076-79, 209097; Cortés: FMNH 4699

(holotype), 4700, 4701 (24), 4702, 4704-11, 4713; KU 209040-57, 209060-75, 209107,

209141; LACM 137298-305;, MCZ 17435; UIMNH 39946; UMMZ 120388; USNM

118202.

Eleutherodactylus stadelmani (78). — Olancho: USNM 497120-68; Yoro: FMNH 21862-64;

MCZ 21290 (holotype), 21291-92; USNM 497169-91.

We have not recently examined the BMNH specimen of “E. milesi” (O’SHEA, 1989;

MCCRANIE et al., 1989) reported from the Departamento de Colôn, Honduras, and cannot

comment upon its specific status. We also follow the lead of CAMPBELL (1994) and do not

consider E. matudai in this discussion of the E. milesi group.

AII measurements were made to the nearest 0.1 mm with dial calipers and a dissecting

microscope. The following standardized abbreviations for measurements are used

(following CAMPBELL, 1994): SVL (snout-vent length), HL (head length: tip of snout to

angle of jaw), HW (greatest width of head), EL (eye length), E-N (distance between

anterior border of eye and posterior border of nostril), TM (tympanum length), E-T

(shortest distance between eye and tympanum), TL (tibia length, including covering

tissues), FL (distance from posteriormost portion of inner metatarsal tubercle to tip of

fourth toe), EW (greatest width of upper eyelid), IOD (shortest interorbital distance), and

F3 (disc width of third finger).

Because tympanum condition varies considerably among members of the milesi

group, we have adopted the following terminology and somewhat subjective criteria:

tympanum prominent (annulus distinctly raised and strongly evident throughout its

circumference; this condition does not occur in any milesi group member); tympanum

Source : MNHN, Paris

150 ALYTES 14 (4)

distinct (tympanum and annulus covered by thin skin, but annulus evident almost

throughout its circumference); tympanum indistinct (tympanum and annulus covered by

thicker skin, ca. one-half of annulus evident); tympanum very indistinct (tympanum and

annulus covered by thick skin, ca. one-third of annulus evident); tympanum hidden

(tympanum and annulus completely concealed by thick skin). The fact that the tympanum

may vary from one side of the head to the other in some individuals of those milesi group

species lacking distinct tympana further compounds the use of this feature. In addition, the

tympanum condition is sexually dimorphic. We follow CAMPBELL (1994) in the usage of the

terms lateral keels and lateral fringes for the ornamentation of the toes and fingers.

Comparative data for the Guatemalan species E. adamastus and E. trachydermus are

taken from CAMPBELL (1994). Snout shape terminology follows HEYER et al. (1990). Color

notes in each diagnosis are taken from preserved specimens. Each diagnosis is based upon

adults only, unless otherwise stated.

SYSTEMATIC ACCOUNTS

External characters defining the E. milesi group include inner tarsal fold absent,

dorsum with scattered to numerous tubercles, tympanum distinct to hidden in males and

indistinct to hidden in females, and pale para-anal bars or spots commonly present.

First we provide new diagnoses and distributional statements for E. milesi and E.

stadelmani, followed by the description of four new species of E. milesi-like frogs from

Honduras. Tables 1-2 compare selected characters among the Honduran members of the

E. milesi group.

Eleutherodactylus milesi Schmidt, 1933

Gig. 1)

Diagnosis. — (1) Skin of dorsum of head and body, flanks, and upper surfaces of limbs

not granular, but wrinkled in most specimens, weakly granular in others; numerous tiny

to small tubercles on top of head and anterior part of body; tubercles larger on flanks and

posterior part of dorsum of body than on rest of dorsum; margin of upper lip smooth to

rough; upper lip not flared; row of raised skin, usually discontinuous, with or without

tubercles, forming occipital fold from posterior edge of upper eyelid to scapular region;

shorter row of raised skin, with or without tubercles, extending from posterior edge of eye,

curving downward above tympanum; skin of venter slightly wrinkled, almost smooth; (2)

tympanic annulus distinct in males, indistinct in females, length about 47-68 % length of

eye in males, 27-47 % in females; tympanum separated from eye in males by distance

29-47 % length of tympanum, 71-118 % in females; (3) snout nearly rounded to rounded

in dorsal view, rounded to nearly vertical (with rounded upper end) in profile; canthus

rostralis well defined, angular; loreal region concave; (4) upper eyelid 100-118 %

interorbital distance, wrinkled, covered with tiny to large tubercles; no cranial crests; (5)

vomerine dentigerous processes round to oval in outline, larger than choanae, lateral edge

Source : MNHN, Paris

Tab. 1. - Comparison of selected characters of the Honduran members of the Eleutherodactylus milesi group. Abbreviations used are:

TM, tympanum length; EL, eye length.

Character

chrysozeretes

cruzi

epochthidius

fecundus

milesi

omoaensis

saltuarius

SVL (mm)

Males

Females

Male vocal slits

Tympanum

Males

Females

TM/EL

Males

Females

Upper lip

IToe lateral structure

1. Rarelÿ very indistinet

33.5-413

373-456

present

hidden

hidden

not flared”

fleshy fringes

27.0-332

present

hidden!

not flared

fleshy fringes

2. In one subadult female, adult females unknown.

3. Flared in large females

4. Occasional large females have weakly infolded fringes on toe IV.

20.9 - 26.9

33.1 - 36.7

present

indist. to hidden

indist, 10 very indist.

34-39%

29%

not flared

fleshy fringes

21.1-23.5

29.8 -37.3

present

indist. to hidden

indist. 10 hidden

36-49%

34-43%

not flared

keel

19.4-25.5

25.0-37.1

present

distinct

indistinct

47-68%

27-47%

not flared

keel

26.2 - 30.0

25.6-384

absent

distinct

indistinct

72-85%

39- 52%

flared

Keel

19.6-22.4

present

dist. to indist.

very indistinct?

35-46%

not flared

keel

stadelmani

273-331

332-474 Z

a

present 9

à 4

hidden Êl

ji &

£

Z

a

Source : MNHN, Paris

152 ALYTES 14 (4)

Tab. 2 . - Toe webbing formulae for the Honduran members of the Eleutherodactylus milesi

group.

Species Modal webbing formula

Eleutherodactylus chrysozetetes 111/3-21111/3-21/2112-31/31V31/3-2V

Eleutherodactylus cruzi 12-23/4112-31/2113 -41V4-21/2V

Eleutherodactylus epochthidius 12-23/4112-3 1/2 III 3 - 4 1/4 IV 4 1/4-23/4V

Eleutherodactylus fecundus 12-24/5112-3 3/4 III 3 - 4 1/4 IV 4 1/4-2 3/4 V

Eleutherodactylus milesi 12*-24/5112-33/4II13-41/21V41/2-3 V

Eleutherodactylus omoaensis 12-24/5112-3 3/4 III 3-4 1/2 IV 41/4-23/4V

Eleutherodactylus saltuarius 1-112*-3 4/51113* -41/21V41/2-3 V

Eleutherodactylus stadelmani 12-23/4112-31/2113 -41V4-21/2V

of dentigerous process not extending laterally to median portion of choanae; (6) vocal slits

and pale nuptial pads present in males; (7) first finger shorter than second or first and

second fingers about equal in length; fingers bearing moderately well developed pads; discs

on fingers III-IV slightly broader than long; disc width on finger III 14-36 % length of

tympanum in males, 25-56 % in females; (8) fingers bearing lateral keels; (9) no ulnar

tubercles on forearm in most specimens, ulnar tubercles on forearm occasionally arranged

in linear series, but not developed into fold in those specimens with weakly granular dorsa;

antebrachium smooth or with few tiny tubercles; (10) heel varying from nearly smooth

with few tiny tubercles to covered with about 30 small tubercles; no linear series of

tubercles or fold along outer edge of tarsus; no inner tarsal fold; (11) inner metatarsal

tubercle oval to elongate (length about 2-3 times width), 2-3 times size of small rounded

outer metatarsal tubercle; no plantar tubercles; (12) toes bearing well developed lateral

keel, webbing moderate for E. milesi group (modal formula 1 2* — 2 4/5 II 2 — 3 3/4 III

3—41/21V 41/2 — 3 V); discs on toes III-IV about as broad as or slightly broader than

those on fingers III-IV; (13) dorsum of head and body pale brown to dark brown; thin,

pale middorsal stripe rarely extending from level posterior to tympanum to vent; larger

tubercles with slightly paler colored tips; obscure pale interorbital bar frequently present;

dark bars present on upper lip; some individuals with conspicuous pale blotch below

canthus; darker crossbars present on dorsal surfaces of limbs; para-anal pale bars or spots

well defined, indistinct, or absent; posterior of thigh heavily flecked with brown, with paler

brown tiny to small spots or mottling; belly and venter of thigh cream colored, lightly to

moderately flecked with brown, belly flecking sometimes coalesced into scattered spots,

chin and throat more heavily flecked, especially in some males; (14) adults moderate-sized

relative to other members of E. milesi group, 25 males 19.4-25.5 (x = 21.8) mm SVL, 21

females 25.0-37.1 (x = 32.1) mm SVL.

Distribution. — Moderate and intermediate elevations (1050 to 1720 m) of the sierras

Omoa and Espiritu Santo of northwestern Honduras (fig. 2). See Materials and Methods

for a list of specimens examined. The habitat surrounding this species’ type locality

(mountains west of San Pedro [Sula], Honduras. Altitude 4500 feet.” ; SCHMIDT, 1933: 18)

Source : MNHN, Paris

MCCRANIE & WILSON 153

Fig. 1. — (A) Adult male (KU 209049) of E. milesi, SVL 22.5 mm. (B) Adult female (USNM 497172)

of E. stadelmani, SVL 46.8 mm. (C) Adult female (USNM 497098) of E. fecundus, SVL 33.3 mm.

(D) Adult male (USNM 497115) of E. saltuarius, SVL 22.4 mm. Photograph (A) by Jim

BRIDGES, remainder by senior author.

has been severely altered since SCHMIDT collected the type series. It is unlikely that this

species presently occurs at its type locality. Evidence also suggests that the El Cusuco

population is experiencing a population decline, or has collapsed altogether (see

Introduction).

Eleutherodactylus stadelmani Schmidt, 1936

Gig. 1)

Diagnosis. — (1) Skin of dorsum of head and body, flanks, and upper surfaces of limbs

not granular to moderately granular, frequently wrinkled; numerous tiny to moderate-

sized tubercles on top of head and anterior part of body; tubercles larger on flanks and

posterior portion of body than rest of dorsum; margin of upper lip smooth to rough; upper

lip usually not flared, sometimes slightly flared in large females; row of raised skin,

frequently tuberculate, continuous or discontinuous, forming occipital fold from posterior

Source : MNHN, Paris

154 ALYTES 14 (4)

edge of upper eyelid to scapular region; shorter row of raised skin, frequently tuberculate,

extending from posterior edge of eye, curving downward above tympanum; other short

rows of raised skin sometimes present dorsolaterally and on mid-back region; skin of

venter usually slightly wrinkled, almost smooth, although moderately wrinkled in some

specimens with more granular dorsa; (2) tympanic annulus usually hidden in males (rarely

very indistinct), hidden in females; (3) snout nearly rounded to rounded in dorsal view,

rounded to nearly vertical (with rounded upper end) in profile; canthus rostralis well

defined, angular; loreal region concave; (4) upper eyelid 102-124 % interorbital distance,

covered with tiny to large tubercles; no cranial crests; (5) vomerine dentigerous processes

round, oval, or somewhat triangular in outline, slightly larger than choanae in males,

much larger than choanae in females, lateral edge of dentigerous process not extending

laterally to median portion of choanae; (6) vocal slits and pale nuptial pads present in

males; (7) first finger shorter than second or first and second fingers about equal in length;

fingers bearing moderately well developed pads; discs on fingers II-IV slightly broader

than long; (8) fingers bearing lateral keels; (9) no ulnar tubercles on forearm in most

specimens, ulnar tubercles on forearm occasionally arranged in linear series, but not

developed into fold in those specimens with more granular dorsa; antebrachium smooth

with few small to moderate-sized tubercles; (10) heel covered with about 20-40 tiny to

moderate-sized tubercles; no linear series of tubercles along outer edge of tarsus; no inner

tarsal fold; (11) inner metatarsal tubercle oval to elongate (length about 2-3 times width),

3-5 times size of small rounded outer metatarsal tubercle; no plantar tubercles; (12) toes

bearing well developed lateral fleshy fringes that fold ventrally; webbing well developed for

E. milesi group (modal formula 12 — 2 3/4 II 2 — 3 1/2 III 3 — 4 IV 4 — 21/2 V); discs

of toes ITI-IV about as broad as or slightly broader than those on fingers III-IV; (13)

dorsum of head and body pale brown to dark brown, frequently with darker brown

mottling or spotting, some specimens with broad, pale dorsolateral longitudinal stripe,

others with broad, pale middorsal band beginning on snout and extending laterally to

cover upper eyelids and gradually narrowing to V-shape above vent; larger tubercles

frequently with slightly paler colored tips; obscure pale interorbital bar frequently present;

dark bars present on upper lip; some individuals with conspicuous pale blotch below

canthus; darker crossbars present on dorsal surfaces of limbs; para-anal pale bars or spots

well defined, indistinct, or absent; posterior of thigh heavily flecked with brown, with paler

brown tiny to small spots or mottling; belly and venter of thigh cream colored, lightly to

moderately flecked with brown, chin and throat more heavily flecked, especially in some

males; occasional specimens have dark flecks on belly, chest, chin, and throat concentrated

into mottled or vermiculate pattern; (14) adults large relative to other members of E. milesi

group, 18 males 27.3-33.1 (x = 30.2) mm SVL, 19 females 33.2-47.4 (x = 39.8) mm SVL.

Comparisons. — Eleutherodactylus stadelmani is easily distinguished from E. milesi by its

larger size (see above and tab. 1), tympanum condition (hidden or rarely very indistinct in

male and hidden in female stadelmani versus distinct in male and indistinct in female

milesi), toe margin structure (well developed lateral fleshy fringes that fold ventrally in

stadelmani versus lateral keels in milesi), and amount of toe webbing (see above and tab.

2). Eleutherodactylus stadelmani is apparently most closely related to E. cruzi (the latter

known only from 1520 m at a single locality in the central portion of the Cordillera

Nombre de Dios, Departamento de Atläntida, Honduras), differing from that species only

Source : MNHN, Paris

MCCRANIE & WILSON 155

in color pattern (no pale middorsal stripe in stadelmani versus thin, pale middorsal stripe

extending from tip of snout to just above vent in cruzi) and skin texture (row of raised

skin, that is frequently tuberculate, forming well developed occipital fold, numerous tiny

to moderate-sized tubercles on heel, and upper eyelid with many tiny to large tubercles in

stadelmani versus occipital fold indistinct to poorly developed, heel mostly smooth with

only tiny tubercles, and much of upper eyelid smooth or wrinkled in cruzi). Eleuthero-

dactylus chrysozetetes (known only from 880 to 1130 m in the central portion of the

Cordillera Nombre de Dios, Departamento de Atläntida, Honduras) is similar to E.

stadelmani in having well developed fleshy fringes on the toe margins and hidden tympana.

However, E. stadelmani differs from E. chrysozetetes in smaller male size (see tab. 1) and

by having less toe webbing (see tab. 2). Eleutherodactylus stadelmani is most easily

distinguished from £. adamastus of Guatemala by tympanum condition (annulus usually

hidden, rarely very indistinct in male and hidden in female stadelmani versus distinct in

male and indistinct in female adamastus) and toe margin structure (well developed lateral

fleshy fringes in stadelmani versus lateral keels in adamastus). Eleutherodactylus stadelmani

can be distinguished from E. trachydermus of Guatemala by skin texture (dorsal surfaces

not granular to moderately granular and ventral surfaces usually slightly wrinkled, almost

smooth in stadelmani versus dorsal surfaces strongly granular and ventral surfaces

coarseley wrinkled in trachydermus), male tympanum condition (annulus usually hidden,

rarely very indistinct in stadelmani versus indistinct in trachydermus) and chin and throat

coloration (lightly to heavily flecked with brown in stadelmani versus chin and throat with

heavy suffusion of dark gray melanophores to almost uniformly brown in trachydermus).

Distribution. — Isolated localities at moderate and intermediate elevations (1125 to 1900

m) of northern Honduras from the western portion of the Cordillera Nombre de Dios

southward to Montaña de Pijol and eastward to Parque Nacional La Muralla in

northwestern Olancho (fig. 2). See Materials and Methods for a list of specimens examined.

The habitat surrounding this species’ type locality (“Portillo Grande, Yoro, Honduras, at

4800 feet altitude”; SCHMIDT, 1936: 44) has been severely altered since the type series was

collected and it is unlikely that the species still occurs in the region. Evidence also suggests

that the population at the single known locality for the species in the western Cordillera

Nombre de Dios may also be declining (see Introduction).

Eleutherodactylus omoaensis sp. nov.

Holotype. — MVZ 115286, adult male, from about 10 airline km WSW San Pedro Sula

on road to Perü (15°28’N, 88°06'W), elevation 1150 m, Sierra de Omoa, Departamento de

Cortés, Honduras, 9 February 1974, James KEZER and James F. LyNCH.

Paratypes. — Sixteen specimens: MVZ 115283-84, 115287-88 and 115290, all adult males,

and MVZ 115281-82, 115285 and 128749-52, all adult females, all with same locality and

data as holotype; MCZ 21295-96, both adult males, and FMNH 21820 and 21829, both

adult females, all from Montaña Santa Ana W of San Pedro Sula, Departamento de

Cortés, Honduras.

Source : MNHN, Paris

I

(&) +I SALATV

L L 1 L L 1

89 88 87 86 85 84

Fig. 2. — Distribution of four species of the Eleutherodactylus milesi group in Honduras. Circles: E.

milesi; triangles: E. stadelmani; square: E. chrysozetetes and E. cruzi. À single symbol may

represent more than one locality.

Source : MNHN, Paris

MCCRANIE & WILSON 157

Referred specimens. — Seven specimens: MVZ 128754, subadult male, and MVZ 115289 and

128753, both subadult females, same locality and data as holotype; MCZ 21298, subadult

male, MCZ 21299, subadult female, and MCZ 21297, juvenile female, all from Montaña Santa

Ana W of San Pedro Sula, Departamento de Cortés, Honduras; FMNH 4677, juvenile,

from Cañon Santa Ana W of San Pedro Sula, Departamento de Cortés, Honduras.

Diagnosis. — (1) Skin of dorsum varying from not granular, but wrinkled, to strongly

granular, with numerous tiny to moderate-sized tubercles; skin of venter slightly wrinkled,

almost smooth in many specimens, coarsely wrinkled in those specimens with strongly

granular dorsa; (2) tympanic annulus distinct in males, indistinct in females; (3) snout

semicircular to nearly rounded in dorsal view, rounded in profile; (4) upper eyelid

100-125 % interorbital distance, usually covered with tiny to large tubercles, rarely

wrinkled with relatively few tubercles; no cranial crests; (5) vomerine dentigerous processes

oval to nearly triangular in outline; (6) vocal slits absent, pale nuptial pads present in

males; (7) first finger shorter than second or first and second fingers about equal in length;

fingers bearing moderately well developed pads; (8) fingers bearing lateral keels; (9) ulnar

tubercles indistinct; (10) heel bearing small to moderate-sized tubercles; no linear series of

tubercles along outer edge of tarsus; no inner tarsal fold; (11) inner metatarsal tubercle

elongate, 3-4 times size of small rounded outer metatarsal tubercle; no plantar tubercles;

(12) toes bearing well developed lateral keels, webbing moderate (fig. 3) for E. milesi group

(modal formula 1 2 — 2 4/5 112 — 3 3/4 III 3 — 4 1/2 IV 4 1/4 — 2 3/4 V); discs of toes

II-IV about as broad as or slightly broader than those on fingers III-IV; (13) dorsum of

head and body medium brown to dark brown, some specimens with broad, pale

dorsolateral longitudinal stripe, others with broad, pale middorsal band beginning on

snout and extending laterally to cover upper eyelids and gradually narrowing to V-shape

above vent; para-anal pale bars or spots indistinct, occasionally absent; belly and venter

of thigh cream colored, lightly to moderately flecked with brown, chin and throat more

heavily flecked, especially in some males; (14) adult males large and adult females

moderate-sized relative to other members of E. milesi group, 8 males 26.2-30.0 (x = 28.1)

mm SVL, 9 females 25.6-38.4 (x = 32.2) mm SVL.

Comparisons. — Eleutherodactylus omoaensis is apparently most closely related to E. milesi

(the latter also known from the Sierra de Omoa). These two species have similar

tympanum conditions, toe webbing, and toe margin structure. Eleutherodactylus omoaensis

differs from E. milesi by having flared upper lips (not flared in milesi), broader heads (HW

44-48 % of SVL in male and 43-49 % in female omoaensis versus 39-43 % in male and

39-42 % in female milesi), adult males lacking vocal slits (present in milesi), larger adult

male size (see above and tab. 1), and larger tympanum size in adult males (see tab. 1).

Eleutherodactylus omoaensis can be distinguished from all other members of the E. milesi

group by the following combination of characters: relatively large adult male size and

moderate adult female size; tympanic annulus distinct in males and indistinct in females;

males lacking vocal slits; moderate toe webbing; and lateral keels on toes.

Measurements of holotype (mm). — SVL 30.0; HL 13.6; HW 14.0; EL 4.0; E-N 3.0; TM

34; E-T 1.2; TL 15.5; FL 15.0; EW 3.7; IOD 3.4; F3 0.6.

Colors of holotype in preservative. — Dorsum of head and body dark brown, most of larger

tubercles with pale gray tips; dark crossbars on limbs not well marked; iris gray; pale inter-

Source : MNHN, Paris

8sT

(+) bI SALATV

Fig. 3. — Ventral view of right foot of (A) E. omoaensis (MVZ 115282), (B) E. fecundus (USNM

497099) and (C) E. saltuarius (USNM 497115). Each scale bar equals 5 mm. The foot structure

of E. epochthidius resembles that of E. fecundus except that fleshy fringes are present on toes

II-IV or toe IV.

Source : MNHN, Paris

MCCRANIE & WILSON 159

orbital bar absent; dark bars on upper lip very obscure; posterior of thigh heavily flecked

with brown, with small, pale brown spots; para-anal pale bar indistinct; belly and venter

of thigh cream colored, lightly to moderately flecked with brown, chin, throat, and chest

more heavily flecked; palmar and plantar surfaces brown with tiny, scattered pale spots.

Description. — The following measurements and proportions are based on the entire type

series of eight males and nine females. Head usually wider than long (width 98-109 % of

length); HW 43-49 % of SVL; HL 43-46 % of SVL in males, 41-47 % in females; snout

semicircular to nearly rounded in dorsal view, rounded in profile; E-N 9-11 % of SVL;

upper eyelid usually covered with tiny to large tubercles, upper eyelid rarely wrinkled with

relatively few tubercles, EW 100-125 % of IOD. Top of head flat in interorbital region,

skin varying from not granular, but wrinkled to strongly granular, with numerous tiny, or

numerous tiny plus a few to many moderate-sized tubercles; no cranial crests; canthus

rostralis angular; loreal region concave; margin of upper lip smooth to rough; upper lip

distinctly flared below and posterior to orbit in both sexes; internarial area slightly

concave; nostrils slitlike, protuberant, directed posterolaterally. Supratympanic fold well

developed, usually consisting of series of enlarged tubercles on raised skin, extending from

posterior edge of eye, but not reaching insertion of forelimb; row of raised skin, usually

tuberculate, forming usually discontinuous occipital fold from posterior edge of upper

eyelid to scapular region, although fold may be poorly defined in some specimens; shorter

row of raised skin, usually tuberculate, extending from posterior edge of eye, curving

downward above tympanum; tympanic annulus distinct and rounded in males, evident

almost throughout circumference; tympanic annulus indistinct in females, usually evident

anteriorly and ventrally; tympanum length about 72-85 % length of eye in males, 39-52 %

in females; tympanum separated from eye by distance 26-37 % of tympanum length in

males, 70-106 % in females. Choanae smaller than vomerine dentigerous processes, round,

teardrop-shaped, or oval in outline, anterior edge flat and formed by anterior lateral

process of vomer, not concealed by palatal shelf of maxillary arch; vomerine dentigerous

processes about as long as wide, or longer than wide, oval to nearly triangular in outline;

vomerine dentigerous processes posteromedial to choanae, lateral edge of dentigerous

process not extending laterally to median portion of choanae; vomerine dentigerous

processes separated by distance less than width of each dentigerous process; each

dentigerous process bearing three to four teeth along posterior border in males, three to

six teeth in females. Tongue moderate in size, longer than wide, not notched posteriorly,

free posteriorly for about 25 % of its length.

Skin on dorsum of anterior part of body and limbs varying from wrinkled and not

granular to strongly granular, skin with numerous tiny or numerous tiny plus a few to

many moderate-sized tubercles; skin on posterior part of dorsum and on flanks with larger

tubercles; skin of venter slightly wrinkled, almost smooth in many specimens, coarsely

wrinkled in those specimens with strongly granular dorsa; skin below vent tuberculate;

irregular series of ulnar tubercles or ulnar tubercles sometimes arranged in linear series,

but not developed into fold in those specimens with strongly granular dorsa; antebrachium

with numerous tiny to small tubercles; heel covered with about 20-30 small to

moderate-sized tubercles; no linear series of tubercles or fold along outer edge of tarsus.

Cloacal opening directed posteroventrally, slightly below upper level of thigh.

Source : MNHN, Paris

160 ALYTES 14 (4)

Forearm moderately slender in both sexes; fingers relatively long and slender, thumb

about equal to or shorter than second finger; fingers with lateral keels; fingers bearing

weakly dilated discs (finger III disc 1.4-2.0 times width of phalanx just proximal to disc),

all discs bearing pads, pads about as wide as long; discs on fingers III-IV as broad as or

slightly broader than long; disc width on finger III 16-31 % length of tympanum in males,

26-53 % in females; discs of fingers I-II ovoid apically, those of fingers III-IV rounded;

relative length of fingers in decreasing order III, IV, IL, I or IL, IV, II & I; subarticular

tubercles on fingers rounded to slightly elongate in ventral view, scarcely protuberant, flat

to rounded in lateral view; supernumerary tubercles on fingers absent; palmar tubercle

cordiform, about as large as or slightly larger than suboval thenar tubercle (thenar tubercle

less than twice as long as wide); several small accessory palmar tubercles frequently

present; males with pale nuptial pads, but without vocal slits. Hindlimbs short; heels not

in contact or barely overlapping when hindlimbs flexed at right angles to axis of body; TL

52-62 % of SVL; FL 50-57 % of SVL. Inner tarsal fold absent; two metatarsal tubercles,

inner about three times as long as wide, 3-4 times size of small rounded outer tubercle; no

supernumerary tubercles on toes; no plantar tubercles; toes with discs and pads, disc tips

broadly rounded; toes bearing well developed lateral keels and moderate webbing,

webbing formula 1 2 — (2 3/4 — 2 4/5) II (2 — 2*) — (3 1/2 — 3 4/5) II (3 — 3*) —

(4* — 41/2) IV (41/4 — 4 3/4) — (2 3/4 — 3*) V; disc on toe III about as broad as

or slightly broader than discs on fingers III-IV; subarticular tubercles on toes slightly

longer than wide, somewhat protuberant.

The color in preservative of the entire series essentially agrees with that of the

holotype. Some paratypes have a broad, pale middorsal longitudinal band; some have a

broad, pale dorsolateral longitudinal stripe on each side; an obscure pale interorbital bar

is frequently present; some specimens have a conspicuous pale blotch below the canthus.

One subadult male (MVZ 128754) has a thin, pale middorsal stripe extending from the

snout to just above the vent.

Etymology. — The name omoaensis refers to the Sierra de Omoa, the mountain range

where the species occurs.

Distribution and natural history notes. — Moderate elevations (760 to 1150 m) in the Sierra

de Omoa W of San Pedro Sula, Honduras (fig. 4). The MVZ specimens of E. omoaensis

were collected at 1150 m during the day (9 February 1974) from around a small stream

flowing through a cafetal (J. F. LyYNCH, in litt.). We have not visited this area and can add

nothing on the present status of this population. The specimens from Montaña Santa Ana

were collected from an unknown elevation in August 1931, whereas the juvenile from

Cañon Santa Ana was taken at about 760 m on 21 March 1923. Both of these Santa Ana

localities are in an area in which the vegetation has long since been severely altered.

Remark. — The paratypes MVZ 128749-52 of E. omouensis have been ventrally dissected

and have their left legs missing.

Source : MNHN, Paris

L 2 1 L 1

88 87 86 85 84

Fig. 4. — Distribution of four species of the Eleutherodactylus milesi group in Honduras. Circle: E.

omoaensis; square: E. fecundus and E. saltuarius; diamond: E. fecundus; inverted triangles: E.

epochthidius; triangle: E. saltuarius. À single symbol may represent more than one locality.

NOSIIM ®@ HINVHDON

191

Source : MNHN, Paris

162 ALYTES 14 (4)

Eleutherodactylus fecundus sp. nov.

(ig. 1)

Holotype. — LACM 137311, adult male, from Quebrada de Oro (15°38'N, 86°47'W),

elevation 880 m, tributary of Rio Viejo, south slope of Cerro Büfalo S of La Ceiba,

Cordillera Nombre de Dios, Departamento de Atläntida, Honduras, 16 August 1982,

James R. MCCRANE, Kenneth L. WILLIAMS, and Larry David WiLsoN. Original number

LDW 6322.

Paratypes. — Nineteen specimens: KU 209084-85, 209093, 209099, 209101 and 209142,

LACM 137308, USNM 497088 and 497092, all adult males, and USNM 497082-87,

497089-91 and 497093, all adult females, all from the same locality as the holotype,

elevation 780 to 1260 m.

Referred specimens. — Fifty-one specimens: KU 209080, 209094, 209143 and USNM

497102, all adult males, KU 209086-92, 209095, 209098, 209100, 209102 and 209104-06,

LACM 137306-07 and 137309-10, USNM 497094-96, 497098-101 and 497103-08, all adult

females, and KU 206750, 209058, 209081-83, 209096, 209103, 209108 (2) and USNM

497097, all juveniles or subadults, all from the type locality, elevation 780 to 1130 m; MCZ

21293-94, both adult females, from Cordillera Nombre de Dios S of La Ceiba,

Departamento de Atläntida, Honduras; USNM 497109, adult female, and USNM

497110-12, all juveniles, from Cerro Calentura, Cordillera Nombre de Dios S of Trujillo,

Departamento de Colôn, Honduras, elevation 460 m.

Diagnosis. — (1) Skin of dorsum not granular, but wrinkled, with tiny to small tubercles;

skin of venter slightly wrinkled, almost smooth; (2) male tympanic annulus indistinct (5

out of 10 specimens in type series), very indistinct (2 out of 10), or hidden (3 out of 10);

female tympanic annulus indistinct (4 out of 10 specimens in type series), very indistinct

(2 out of 10), or hidden (4 out of 10); (3) snout nearly rounded to rounded in dorsal view,

rounded to nearly vertical (with rounded upper end) in profile; (4) upper eyelid 103-133 %

interorbital distance, wrinkled, with numerous tiny and about 10-20 small to large

tubercles; no cranial crests; (5) vomerine dentigerous processes oval or somewhat

triangular in outline; (6) vocal slits and pale nuptial pads present in males; (7) first finger

shorter than second or first and second fingers about equal in length; fingers bearing

moderately well developed pads; (8) fingers bearing lateral keels; (9) ulnar tubercles absent

or indistinct; (10) heel bearing small tubercles; no linear series of tubercles along outer edge

of tarsus; no inner tarsal fold; (11) inner metatarsal tubercle elongate, 3-4 times size of

small rounded outer metatarsal tubercle; no plantar tubercles; (12) toes bearing well

developed lateral keel, weakly developed fringes occasionally present on toe IV of large

females; webbing moderate (fig. 3) for milesi group (modal formula 1 2 — 2 4/5 II 2 —

3 3/4 113 — 41/4 IV 4 1/4 — 2 3/4 V); discs of toes III-IV about as broad as or slightly

broader than those on fingers III-IV; (13) dorsum of head and body medium brown to

dark brown, some specimens with broad, pale dorsolateral longitudinal stripe on each side,

thin, pale middorsal stripe rarely present from level just posterior to eyes to above vent;

para-anal pale bars or spots well defined, indistinct, or absent; belly and venter of thigh

cream colored, lightly to moderately flecked with brown, chin and throat more heavily

flecked, especially in some males; (14) adults moderate-sized relative to other members of

Source : MNHN, Paris

MCCRANIE & WILSON 163

E. milesi group, 10 males 21.1-23.5 (x = 22.2) mm SVL, 10 females 29.8-37.3 (x = 33.5)

mm SVL.

Comparisons. — Eleutherodactylus fecundus is most similar to E. milesi, but differs in

tympanum condition (annulus indistinct, very indistinct, or hidden in male and female

fecundus versus distinct in male and indistinct in female milesi) and amount of webbing on

toes III-V (see above and tab. 2). Eleutherodactylus fecundus differs from E. omoaensis (the

apparent closest relative of E. milesi) in the same tympanic annulus condition as it does

from £. milesi. These two species also differ in upper lip condition (not flared in fecundus

versus flared in omoaensis), adult male size (see above and tab. 1), male tympanum size

(length about 36-49 % length of eye in fecundus versus 72-85 % in omoaensis), and male

vocal slit condition (present in fecundus versus absent in omouensis). Eleutherodactylus

fecundus can be distinguished from the remaining Æ. milesi group members by the

following combination of characters: moderate adult size; indistinct, very indistinct, or

hidden tympanic annulus condition in both males and females; moderate toe webbing; and

toes with well developed lateral keels.

Measurements of holotype (mm). — SVL 22.1; HL 9.4; HW 9.5; EL 3.2; E-N 1.9; TL 12.9;

FL 12.0; EW 2.9; IOD 2.6; F3 0.4.

Colors of holotype in preservative. — Dorsum of head and body medium brown, most

larger tubercles with pale gray tips; dark crossbars on limbs well defined; iris gray; pale

interorbital bar well defined; dark bars on upper lip well defined; posterior of thigh heavily

flecked with brown, with tiny to small pale brown spots; pale para-anal bars fairly well

defined; belly and venter of thigh cream colored, moderately flecked with brown, chin and

throat more heavily flecked; palmar surface cream colored, moderately flecked with

brown; plantar surface cream colored, more heavily flecked with brown than palmar

surface.

Description. — The following measurements and proportions are based on the entire type

series of 10 adult males and 10 adult females. Head nearly as wide as long to slightly wider

than long (width 85-105 % of length); HW 37-46 % of SVL; HL 40-46 % of SVL in males,

38-42 % in females; snout nearly rounded to rounded in dorsal view, rounded to nearly

vertical (with rounded upper end) in profile; E-N 7-12 % of SVL; upper eyelid wrinkled

with numerous tiny and about 10-20 small to large tubercles, EW 103-133 % of IOD. Top

of head flat in interorbital region, skin not granular, but wrinkled, with numerous tiny, or

numerous tiny plus scattered small tubercles; no cranial crests; canthus rostralis angular;

loreal region concave; margin of upper lip smooth, or smooth with scattered tiny tubercles;

upper lip not flared; internarial area slightly concave; nostrils slitlike, protuberant, directed

posterolaterally. Supratympanic fold usually well developed, consisting of raised skin, with

or without tubercles, extending from posterior edge of eye, but not reaching insertion of

forelimb; row of raised skin, with or without tubercles, forming discontinuous occipital

fold from posterior edge of upper eyelid to scapular region; shorter row of raised skin, with

or without tubercles, extending from posterior edge of eye, curving downward above

tympanum; tympanic annulus indistinct (usually evident anteriorly and ventrally), very

indistinct (usually evident anteroventrally), or hidden in males and females; tympanum

length about 36-49 % length of eye in males (n — 5), 34-43 % in females (n = 4);

tympanum separated from eye by distance 56-67 % of tympanum length in males, 81-85 %

Source : MNHN, Paris

164 ALYTES 14 (4)

in females. Choanae slightly smaller than vomerine dentigerous processes in males, much

smaller than them in females, round, teardrop-shaped, or oval in outline, anterior edge flat

and formed by anterior lateral process of vomer, not concealed by palatal shelf of

maxillary arch; vomerine dentigerous processes about as long as wide, or longer than wide,

oval to nearly triangular in outline; vomerine dentigerous processes posteromedial to

choanae, lateral edge of dentigerous process not extending laterally to median portion of

choanae; vomerine dentigerous processes separated by distance about equal to or less than

width of each dentigerous process; each dentigerous process bearing three to five teeth

along posterior border in males, five to eight teeth in females. Tongue moderate in size,

longer than wide, not notched posteriorly, free posteriorly for about 25-35 % of its length.

Skin on dorsum of anterior part of body and limbs not granular, but wrinkled, skin

with numerous tiny or numerous tiny plus scattered small tubercles; skin on posterior part

of dorsum and flanks with larger tubercles; skin of venter slightly wrinkled, almost

smooth; skin below vent tuberculate; ulnar tubercles absent, or irregular series of ulnar

tubercles present, or ulnar tubercles occasionally arranged in linear series, but not

developed into fold; antebrachium smooth or wrinkled, occasionally few small tubercles

may be present; heel covered with about 15-30 small tubercles; no linear series of tubercles

or fold along outer edge of tarsus. Cloacal opening directed posteroventrally, slightly

below upper level of thigh.

Forearm moderately slender in both sexes; fingers relatively long and slender, thumb

about equal to or slightly shorter than second finger; fingers with lateral keels; fingers

bearing weakly dilated discs (finger III disc 1.3-1.8 times width of phalanx just proximal

to disc), all discs bearing pads, pads about as wide as long; discs on fingers INI-IV slightly

broader than long; disc width on finger III 21-42 % length of tympanum in males (n =

5), 31-38 % in females (n = 4); discs of fingers I-II ovoid apically, those of fingers IHI-IV

rounded; relative length of fingers in decreasing order III, IV, IL, L or HI, IV, I & I;

subarticular tubercles on fingers rounded to slightly elongate in ventral view, scarcely

protuberant, flat to rounded in lateral view; supernumerary tubercles on fingers absent;

palmar tubercle cordiform or oval, about as large as or slightly larger than suboval thenar

tubercle (thenar tubercle less than twice as long as wide); several small accessory palmar

tubercles frequently present; males with pale nuptial pads and vocal slits. Hindlimbs short;

heels not in contact or barely overlapping when hindlimbs flexed at right angles to axis of

body; TL 49-61 % of SVL; FL 44-54 % of SVL. Inner tarsal fold absent; two metatarsal

tubercles, inner tubercle about three times as long as wide, 3-4 times size of small rounded

outer tubercle; no supernumerary tubercles on toes; no plantar tubercles; toes with discs

and pads, disc tips broadly rounded; toes bearing well developed lateral keels (large

females occasionally with weakly developed fringes on toe IV) and moderate webbing,

webbing formula 1 (2 — 2*) — 2 4/5 112 — (3 1/2 — 3 4/5) III 3 — (4 — 4 1/2) IV (4*

— 43/4) — (21/2 — 2 4/5) V; disc on toe III about as broad as or slightly broader than

discs on fingers III-IV; subarticular tubercles on toes slightly longer than wide, somewhat

protuberant.

Color in life of four adult females from the Quebrada de Oro region. — USNM

497095: dorsum dark chocolate brown; broad, irregular ocher dorsolateral stripe extending

from posterior of eye to groin; ocher blotch present below canthus; ocher blotch on upper

Source : MNHN, Paris

MCCRANIE & WILSON 165

arm continuous with dorsolateral stripe; dorsum of thigh ocher with brown crossbars;

dorsum of tibia banded brown and dark brown; para-anal spots pale orange; chin and

chest gray; belly and venter of thigh pale lemon yellow, flecked with brown; iris black with

dense gold flecking. USNM 497096: dorsum very dark chocolate brown; broad, dark red

dorsolateral stripe extending from posterior of eye to groin; dorsal surfaces of limbs very

dark chocolate brown; posterior of thigh dark brown; chin and chest dark chocolate

brown; belly and venter of thigh lemon yellow, flecked with brown; iris black with dense

gold flecking. KU 209100: dorsum dirty dark greenish gray; pale red to rust red spot on

rostrum, at forelimb insertion, and on heel; chin and chest gray; belly and venter of thigh

lemon yellow with some brown flecking; iris pale copper. USNM 497103: dorsum brown,

larger tubercles with slightly darker brown tips; thin, orange middorsal stripe extending

from level just posterior to eyes to vent; dorsal surfaces of limbs dark brown; toe and

finger discs white dorsally; flanks dark brown; chin and chest pale brown; belly and venter

of thigh yellow with brown suffusion.

The color in preservative of the entire series essentially agrees with that of the

holotype, except that many specimens have darker brown dorsal surfaces. Some specimens

have a broad, pale dorsolateral longitudinal stripe on each side; occasional specimens have

a thin, pale middorsal stripe extending from level just posterior to eye to vent; the pale

interorbital bar varies from well defined to obscure; some specimens have a pale blotch

below the canthus; dark bars are present on the upper lip in most specimens; the pale

para-anal bars or spots vary from well developed to indistinct or absent.

Etymology. — The name fecundus is Latin, meaning fertile or fruitful. The name is used

in reference to the Cordillera Nombre de Dios, the mountain range in which this species

occurs, being fertile or fruitful grounds for the discovery of new species of amphibians and

reptiles.

Distribution and natural history notes. — Low and moderate elevations (460 to 1260 m) of

the Cordillera Nombre de Dios south of La Ceiba and Trujillo (fig. 4), Honduras. The

series from the Quebrada de Oro region was collected during May to August alongside

streams between 780 to 1260 m in the Premontane Wet Forest formation of HOLDRIDGE

(1967). Adults were usually active at night, while juveniles and subadults were active both

at night and during the day. No specimens of E. fecundus were seen during 10 days and

nights of collecting in the Quebrada de Oro region in February 1995 (see Introduction),

although other Honduran members of the E. milesi group (E. omoaensis, E. stadelmani,

and E. sp. nov.) have been collected while active during this month. The specimens from

near Trujillo were collected in June alongside a stream at 460 m in the Lowland Moist

Forest formation of HOLDRIDGE (1967). Data accompanying the MCZ specimens indicate

that they were collected in April.

Eleutherodactylus saltuarius sp. nov.

Gig. 1)

Holotype. — USNM 497115, adult male, from south slope of Cerro Büfalo (15°39°N,

86°48'W), elevation 1550 m, Cordillera Nombre de Dios, Departamento de Atläntida,

Source : MNHN, Paris

166 ALYTES 14 (4)

Honduras, 18 February 1995, James R. MCCRANIE and John C. RINDFLEISH. Original

number LDW 10432.

Paratypes. — Four specimens: USNM 497117, an adult male, USNM 497119, a subadult

male, and USNM 497118, a subadult female, all from 2.5 airline km NNE La Fortuna,

Cordillera Nombre de Dios, Departamento de Yoro, Honduras; USNM 497116, a

subadult male from the type locality.

Diagnosis. — (1) Skin of dorsum not granular, but wrinkled, with tiny to small tubercles;

skin of venter slightly wrinkled, almost smooth; (2) male tympanic annulus distinct or

indistinct; tympanic annulus in one subadult female very indistinct; (3) snout nearly

rounded in dorsal view, nearly vertical (with rounded upper end) in profile; (4) upper

eyelid 100-120 % interorbital distance (including subadult female), wrinkled, with

scattered small to moderate-sized tubercles; no cranial crests; (5) vomerine dentigerous

processes oval to nearly triangular in outline; (6) vocal slits and pale nuptial pads present

in males; (7) first finger shorter than second; fingers bearing moderately well developed

pads; (8) fingers bearing lateral keels; (9) ulnar tubercles absent; (10) heel mostly smooth

or bearing small tubercles; no linear series of tubercles along outer edge of tarsus; no inner

tarsal fold; (11) inner metatarsal tubercle oval, 3-4 times size of small rounded outer

metatarsal tubercle; no plantar tubercles; (12) toes bearing well developed lateral keel,

webbing minimal (fig. 3) for E. milesi group (modal formula 1 — II 2* — 3 4/5 III 3*

— 41/21V 41/2 — 3 V); discs of toes III-IV about as broad as those on fingers III-IV;

(13) (also includes all subadults) dorsum of head and body dark brown or grayish brown,

usually with darker colored spotting on back; para-anal pale bars well defined to

indistinct; belly and venter of thigh cream colored, moderately flecked with brown, with

some flecking on belly usually coalesced into scattered spots, chin and throat more heavily

flecked, becoming uniformly brown in some males; (14) adults small relative to other

members of E. milesi group, two males 19.6-22.4 (x = 21.0) mm SVL, one subadult female

17.8 mm SVL.

Comparisons. — Adult males of E. saltuarius are somewhat similar in size to those of E.

fecundus and E. milesi. Eleutherodactylus saltuarius differs from E. fecundus in amount of

toe webbing (see above and tab. 2), ventral coloration in preservative (some flecking on

belly usually coalesced into spots in saltuarius versus belly flecking more or less evenly

distributed in fecundus) and in life (belly and venter of thigh Clay Color [color 26 in

SMITHE, 1975] with numerous spots that were either dark brown or white in saltuarius

versus yellow with brown flecking in fecundus), and habitat (forest floor well away from

streams in saltuarius versus streamside in fecundus). Eleutherodactylus saltuarius can be

distinguished from E. milesi by tympanum condition (annulus distinct or indistinct in male

and very indistinct in female saltuarius versus distinct in male and indistinct in female

milesi), male tympanum size (length 35-46 % length of eye in saltuarius versus 47-68 % in

milesi), ventral coloration in life (belly and venter of thigh Clay Color with numerous spots

that were either dark brown or white in saltuarius versus belly yellow with brown flecking

and venter of thigh yellowish orange with brown flecks in milesi), and habitat (forest floor

well away from streams in saltuarius versus streamside in milesi). Streamside habitats at the

two known localities for E. saltuarius are occupied by either E. cruzi (Cerro Büfalo) or E.

stadelmani (NNE of La Fortuna). Both of these species have considerably larger males (to

Source : MNHN, Paris

MCCRANIE & WILSON 167

33.2 mm SVL in cruzi, 33.1 in stadelmani, 22.4 in saltuarius), fleshy toe fringes (lateral keels

in saltuarius), more toe webbing (see above and tab. 2), and usually hidden tympana in

males (distinct or indistinct in male saltuarius). Juvenile or subadult E. stadelmani (USNM

497178: 20.3 mm SVL; USNM 497179: 20.4 mm SVL; USNM 497186: 15.2 mm SVL)

from streamside habitats at the La Fortuna locality are easily distinguished from E.

saltuarius by having well developed toe webbing and fringes. Juvenile or subadult E. cruzi

are unknown, but presumably have toe fringes and webbing similar to the conditions

found in immature E. stadelmani.

Measurements of holotype (mm). — SVL 22.4; HL 9.0; HW 8.8; EL 2.9; E-N 2.0; TM 1.0;

E-T 0.6; TL 13.7; FL 12.7, EW 2.3; IOD 23; F3 0.3.

Colors of holotype in life. — Dorsum Clay Color with larger tubercles on back reddish

brown; dorsal and anterior surfaces of limbs Clay Color with darker brown crossbars;

posterior of thigh Clay Color with dirty white mottling; belly and venter of thigh Clay

Color with numerous spots that were either dark brown or white; chin and throat dark

brown; iris pale brown with dark brown reticulations.

Colors of holotype in preservative. — Dorsum of head and body grayish brown with darker

brown spotting on back; darker crossbars on limbs well defined; iris gray; pale interorbital

bar indistinct; dark bars on upper lip distinct; posterior of thigh heavily flecked with

brown, with cream colored spots and mottling; para-anal pale bars well defined; belly and

venter of thigh cream colored, moderately flecked with brown, some flecking on belly

coalesced into scattered spots; chin and throat uniformly brown.

Description. — The following measurements and proportions are based on two adult males

and one subadult female. Head longer than wide (width 95-98 % of length); HW 39-43 %

of SVL; HL 40-41 % of SVL in males, 45 % in subadult female; snout nearly rounded in

dorsal view, nearly vertical (with rounded upper end) in profile; E-N 9 % of SVL; upper

eyelid wrinkled, with scattered small to moderate-sized tubercles, EW 100-120 % of IOD.

Top of head flat in interorbital region, skin not granular, but wrinkled, with numerous

tiny, or numerous tiny plus a few small tubercles; no cranial crests; canthus rostralis

angular; loreal region concave; margin of upper lip smooth to rough; upper lip not flared;

internarial area slightly concave; nostrils slitlike, protuberant, directed posterolaterally.

Supratympanic fold not well developed, consisting of raised skin, with or without

tubercles, extending from posterior edge of eye, but not reaching insertion of forelimb; row

of raised skin, with or without tubercles, forming obscure, discontinuous occipital fold

from posterior edge of upper eyelid to scapular region; shorter row of raised skin, with or

without tubercles, extending from posterior edge of eyelid, curving downward above

tympanum; tympanic annulus distinct and rounded or indistinct (annulus evident

anteriorly and ventrally) in males and very indistinct (annulus evident anteriorly) in one

subadult female; tympanum length about 35-46 % length of eye in males; tympanum

separated from eye by distance 46-60 % of tympanum length in males. Choanae smaller

than vomerine dentigerous processes, round, teardrop-shaped, or oval in outline, anterior

edge flat and formed by anterior lateral process of vomer, not concealed by palatal shelf

of maxillary arch; vomerine dentigerous processes about as long as wide, or longer than

wide, oval to nearly triangular in outline; vomerine dentigerous processes posteromedial

to choanae, lateral edge of dentigerous process not extending laterally to median portion

Source : MNHN, Paris

168 ALYTES 14 (4)

of choanae; vomerine dentigerous processes separated by distance less than width of each

dentigerous process; each dentigerous process bearing three to four teeth along posterior

border in males. Tongue moderate in size, longer than wide, not notched posteriorly, free

posteriorly for nearly 50 % of its length.

Skin on dorsum of anterior part of body and limbs not granular, but wrinkled, skin

with numerous tiny or numerous tiny plus a few small tubercles; skin on posterior part of

dorsum and flanks with a few enlarged tubercles; skin of venter slightly wrinkled, almost

smooth; skin below vent tuberculate; no ulnar tubercles; antebrachium smooth; heel

mostly smooth or with up to about 10-15 small tubercles; no linear series of tubercles or

fold along outer edge of tarsus. Cloacal opening directed posteroventrally, slightly below

upper level of thigh.

Forearm moderately slender in both sexes; fingers relatively long and slender, thumb

shorter than second finger; fingers with lateral keels; fingers bearing weakly dilated discs

(finger III disc 1.5 times width of phalanx just proximal to disc), all discs bearing pads,

pads almost as wide as long; discs on fingers III-IV about as broad as long; disc width on

finger II 23-30 % length of tympanum in males; discs of fingers I-II ovoid apically, those

of fingers III-IV rounded; relative length of fingers in decreasing order III, IV, I, I;

subarticular tubercles on fingers rounded to slightly elongate in ventral view, scarcely

protuberant, flat to rounded in lateral view; supernumerary tubercles on fingers absent;

palmar tubercle cordiform, about as large as oval thenar tubercle (thenar tubercle roughly

twice as long as wide); several small accessory palmar tubercles present; males with pale

nuptial pads and vocal slits. Hindlimbs short; heels slightly overlapping when hindlimbs

flexed at right angles to axis of body; TL 61-64 % of SVL; FL 55-57 % of SVL. Inner

tarsal fold absent; two metatarsal tubercles, inner tubercle about two times as long as wide,

3-4 times size of small rounded outer tubercle; no supernumerary tubercles on toes; no

plantar tubercles; toes with discs and pads, disc tips broadly rounded; toes bearing well

developed lateral keels and minimal webbing, webbing formula I (absent or very basal) II

2* — 34/51 (3 — 3*) — (41/2 — 4 3/4) IV (41/2 — 4 3/4) — (3° — 3) V; disc on

toe IIT about as broad as discs on fingers II-IV; subarticular tubercles on toes slightly

longer than wide, somewhat protuberant.

Color in life of USNM 497116 was recorded as being very similar to that of the

holotype. The color in preservative of the paratypes is similar to that of the holotype,

except some details: some specimens have dark brown dorsa, the pale interorbital bar is

sometimes well defined, a large pale spot occurs posteromedially to the eyes in two

specimens, the pale para-anal bars may be indistinct, and one specimen lacks the brown

spots on the belly.

Etymology. — The name saltuarius is a Latin word meaning “keeper of the sacred forest

of Virtue”. The name is used in reference to the occurrence of this species in forested

habitats well away from streams.

Distribution and natural history notes. — Intermediate elevations (1550 to 1800 m) from

two localities in the Lower Montane Wet Forest formation of HOLDRIDGE (1967) in the

Cordillera Nombre de Dios of north-central Honduras (fig. 4). Specimens were collected

while active during the day on the forest floor in February and August. The forest at both

localities for the species was in a primary state with a closed canopy when the specimens

Source : MNHN, Paris

MCCRANIE & WILSON 169

were collected. However, in August 1995, four years after we collected this species NNE

of La Fortuna, we found these forests to be in a less than pristine condition. Tracts of

forest had been clear cut and burned to make way for crops. Probably not coincidently,

no specimens of E. saltuarius were seen at that time, although a concerted effort was made

to collect more specimens.

Remarks. — A subadult male E. saltuarius (USNM 497116), with a SVL of 14.9 mm, has

a single vocal slit on the right side and the nuptial pads beginning to develop.

Eleutherodactylus epochthidius sp. nov.

Holotype. — ROM 18109, adult female, from Rio Seco (14°55'N, 85°56'W), elevation 1050

m, tributary of Rio Guayape N of Catacamas, Sierra de Agalta, Departamento de

Olancho, Honduras, 8 August 1986, James R. MCCRANE, Kenneth L. WILLIAMS and

Larry David WiLson. Original number LDW 8297.

Paratypes. — Ten specimens: ROM 18110 and 18113, both adult males, and ROM 18112,

adult female, all from the same locality as the holotype, elevation 1000 to 1060 m; ROM

18100-01 and 18103-05, all adult males, and ROM 18099 and 18102, both adult females,

all from between rios Catacamas and Seco, Sierra de Agalta N of Catacamas,

Departamento de Olancho, Honduras, elevation 1450 m.

Referred specimens. — Five specimens: ROM 18108, subadult male, and ROM 18111,

juvenile, both from the type locality, elevation 1000-1050 m; ROM 18106-07, both

juveniles, from between rios Catacamas and Seco, Sierra de Agalta N of Catacamas,

Departamento de Olancho, Honduras, elevation 1450 m; KU 209059, subadult female,

from Montaña de Malacate ca. 10 airline km NW Dulce Nombre de Culmi, Sierra de

Agalta, Departamento de Olancho, Honduras, elevation 760 m.

Diagnosis. — (1) Skin of dorsum not granular, but wrinkled, with tiny to small tubercles;

skin of venter slightly wrinkled, almost smooth; (2) male tympanic annulus indistinct (2

out of 7 specimens in type series), very indistinct (4 out of 7), or hidden (1 out of 7); female

tympanic annulus indistinct (1 out of 4 specimens in type series) or very indistinct (3 out

of 4); (3) snout nearly rounded to rounded in dorsal view, rounded to nearly vertical (with

rounded upper end) in profile; (4) upper eyelid 100-121 % interorbital distance, wrinkled,

with scattered small to moderate-sized tubercles; no cranial crests; (5) vomerine

dentigerous processes oval or somewhat triangular in outline; (6) vocal slits and pale

nuptial pads present in males; (7) first finger shorter than second or first and second

fingers about equal in length; fingers bearing moderately well developed pads; (8) fingers

bearing lateral keels; (9) ulnar tubercles absent or indistinct; (10) heel bearing small

tubercles; no linear series of tubercles along outer edge of tarsus; no inner tarsal fold; (11)

inner metatarsal tubercle elongate, 3-4 times size of small rounded outer metatarsal

tubercle; no plantar tubercles; (12) toes III-IV or toe IV of males bearing distinct or weak

lateral fleshy fringes that fold ventrally, toes III-IV of females bearing distinct lateral fleshy

fringes that fold ventrally, remaining toes bearing well developed lateral keels; webbing

moderate for milesi group (modal formula 1 2 — 2 3/4 II 2 — 3 1/2 III 3 — 41/4 IV 4

Source : MNHN, Paris

170 ALYTES 14 (4)

1/4 — 2 3/4 V); disc of toes III-IV about as broad as or slightly broader than those on

fingers III-IV; (13) dorsum of head and body pale brown to dark brown, frequently with

darker brown blotches or hourglass figure on back; para-anal pale bars or spots well

defined, indistinct, or absent; belly and venter of thigh cream colored, heavily flecked with

brown; chin, throat, and chest with dense brown flecking surrounding pale “spots”; (14)

adults moderate-sized relative to other members of E. milesi group, seven males 20.9-26.9

(x = 24.4) mm SVL, four females 33.1-36.7 (x = 35.1) mm SVL.

Comparisons. — Eleutherodactylus epochthidius approaches E. chrysozetetes, E. stadelmani

and E. cruzi by having fleshy fringes on toes III-IV (females and some males) or on toe

IV only (remaining males) (fleshy fringes present on toes II-IV in chrysozetetes, stadelmani,

and cruzi). However, E. epochthidius differs from these three species by having less

toe webbing (see above and tab. 2), smaller male size (see above and tab. 1), and

tympanum condition (usually very indistinct, occasionally indistinct, or rarely hidden

versus hidden in chrysozetetes and usually hidden or rarely very indistinct in stadelmani

and cruzi). Eleutherodactylus epochthidius further differs from E. cruzi by lacking a pale

middorsal stripe (present in cruzi). Eleutherodactylus epochthidius is similar to E. fecundus

in adult size, amount of toe webbing and tympanum condition, but differs from

that species by having distinct to rather weak fleshy toe fringes that fold ventrally on

toes III-IV (females and some males) or on toe IV only (remaining males), whereas lateral

keels are almost always present on toes III-IV in fecundus (occasional large females of

fecundus have weakly infolded fringes on toe IV). Eleutherodactylus epochthidius further

differs from E. fecundus in ventral coloration (usually dense brown flecking on the

chin, throat and chest surrounding pale “spots” versus usually moderate flecking that is

more or less evenly distributed in fecundus). Eleutherodactylus epochthidius can be

distinguished from the remaining E. milesi group members by the following combina-

tion of characters: moderate adult size; usually very indistinct, occasionally indistinct, or

rarely hidden tympanic annulus condition; moderate toe webbing; toes III-IV or toe IV

with distinct to rather weak fleshy fringes; upper lip not flared; and male vocal slits

present.

Measurements of holotype (mm). — SVL 35.4; HL 15.3; HW 15.4; EL 5.2; E-N 3.1; TL

22.0; FL 20.5; EW 3.1; IOD 3.1; F3 0.8.

Colors of holotype in life. — Dorsum of body with dark chocolate brown middorsal

hourglass figure, rest of dorsum of body brick red; dorsal surfaces of limbs mottled and

banded with dark chocolate brown and brick red; chin and throat dark brown with several

white spots; chest and anterior portion of belly mottled dark brown and white; posterior

portion of belly and ventral surface of thighs pale yellow, rather heavily flecked with

brown; palms and soles dark brown; iris dark metallic green.

Colors of holotype in preservative. — Dorsum of head and body pale brown with darker

brown hourglass figure on back, most larger tubercles with pale gray tips; dark crossbars

on limbs fairly well defined; iris gray; dark bars on upper lip well defined; posterior of

thigh heavily flecked with brown, with tiny to small pale brown spots; pale para-anal bars

fairly well defined; belly and venter of thigh cream colored, heavily flecked with dark

brown; chin, throat, and chest with dense dark brown flecking surrounding pale “spots”;

palmar and plantar surfaces densely flecked with dark brown.

Source : MNHN, Paris

MCCRANIE & WILSON 171

Description. — The following measurements and proportions are based on the entire type

series of seven adult males and four adult females. Head nearly as wide as long in males

(width 95-98 % of length) and slightly wider than long in females (width 101-102 % of

length); HW 40-46 % of SVL; HL 41-48 % of SVL; snout nearly rounded to rounded in

dorsal view, rounded to nearly vertical (with rounded upper end) in profile; E-N 8-10 %

of SVL; upper eyelid wrinkled with numerous tiny and a few small to moderate-sized

tubercles, EW 100-121 % of IOD. Top of head flat in interorbital region, skin not

granular, but wrinkled, with numerous tiny, or numerous tiny plus scattered small

tubercles; no cranial crests; canthus rostralis angular; loreal region concave; margin of

upper lip smooth, or smooth with scattered tiny tubercles; upper lip not flared; internarial

area slightly concave; nostrils slitlike, protuberant, directed posterolaterally. Supratym-

panic fold usually well developed, consisting of raised skin, with or without tubercles,

extending from posterior edge of eye, but not reaching insertion of forelimb; row of raised

skin, with or without tubercles, forming discontinuous occipital fold from posterior edge

of upper eyelid to scapular region; tympanic annulus usually very indistinct (annulus

usually evident anteroventrally), occasionally indistinct (annulus usually evident anteriorly

and ventrally), or rarely hidden; tympanum length about 34-39 % length of eye in males

(n = 2), 29% in one female; tympanum separated from eye by distance 79-92 % of

tympanum length in two males, 160 % in one female. Choanae slightly smaller than

vomerine dentigerous processes in males, much smaller than vomerine dentigerous

processes in females, round, teardrop-shaped, or oval in outline, anterior edge flat

and formed by anterior lateral process of vomer, not concealed by palatal shelf of

maxillary arch; vomerine dentigerous processes about as long as wide, or longer than wide,

oval to nearly triangular in outline; vomerine dentigerous processes posteromedial to

choanae, lateral edge of dentigerous process not extending laterally to median portion

of choanae; vomerine dentigerous processes separated by distance about equal to or less

than width of each dentigerous process; each dentigerous process bearing three to four

teeth along posterior border in males, five to seven teeth in females. Tongue moderate in

size, longer than wide, not notched posteriorly, free posteriorly for about 25-35% of its

length.

Skin on dorsum of anterior part of body and limbs not granular, but wrinkled, skin

with numerous tiny or numerous tiny plus scattered small tubercles; skin on posterior part

of dorsum and flanks with larger tubercles; skin of venter slightly wrinkled, almost

smooth; skin below vent tuberculate; ulnar tubercles absent, or irregular series of ulnar

tubercles present, or ulnar tubercles occasionally arranged in linear series, but not

developed into fold; antebrachium smooth or wrinkled, occasionally with few small

tubercles; heel covered with about 10-30 small tubercles; no linear series of tubercles or

fold along outer edge of tarsus. Cloacal opening directed posteroventrally, slightly below

upper level of thigh.

Forearm moderately slender in both sexes; fingers relatively long and slender, thumb

about equal to or slightly shorter than second finger; fingers with lateral keels; fingers

bearing weakly dilated discs (finger III disc 1.6-2.0 times width of phalanx just proximal

to disc), all discs bearing pads, pads about as wide as long; discs on fingers III-IV slightly

broader than long; disc width on finger III 46-57 % length of tympanum in males (n =

2), 53 % in one female; discs of fingers I-II ovoid apically, those of fingers III-IV rounded;

Source : MNHN, Paris

172 ALYTES 14 (4)

relative length of fingers in decreasing order III, IV, IL, I or IL, IV, II & I; subarticular

tubercles on fingers rounded to slightly elongate in ventral view, scarcely protuberant, flat

to rounded in lateral view; supernumerary tubercles on fingers absent; palmar tubercle

cordiform or oval, about as large as or slightly larger than suboval thenar tubercle (thenar

tubercle less than twice as long as wide); several small accessory palmar tubercles

frequently present; males with pale nuptial pads and vocal slits. Hindlimbs short; heels not

in contact or barely overlapping when hindlimbs flexed at right angles to axis of body; TL

54-62 % of SVL; FL 51-58 % of SVL. Inner tarsal fold absent; two metatarsal tubercles,

inner tubercle about three times as long as wide, 3-4 times size of small rounded outer

tubercle; no supernumerary tubercles on toes; no plantar tubercles; toes with discs and

pads, disc tips broadly rounded; toes III-IV or toe IV of males bearing distinct to rather

weak lateral fleshy fringes that fold ventrally, toes III-IV of females bearing distinct lateral

fleshy fringes that fold ventrally, remaining toes bearing well developed lateral keels; toes

with moderate webbing, webbing formula 1 2 — (2 1/2 — 2 3/4) II 2 — (3 1/2 — 3 3/4)

I 3 — 41/4 IV (4* — 41/4) — (2* — 2 4/5) V; disc on toe III about as broad as or

slightly broader than discs on fingers III-IV; subarticular tubercles on toes slightly longer

than wide, somewhat protuberant.

Color in life of an adult female (ROM 18112) was as follows: dorsum of body

and head brown with dark outlined pale interorbital bar, bar preceded by and followed

by tan blotches; dorsal surfaces of limbs brown with dark brown crossbars; groin golden

yellow; chin brown with white punctations; chest mottled brown and white; belly and

venter of thigh dirty yellow; palms and soles brown; iris bronze with rust copper band in

pupil.

The color in preservative of the type series may be described as follows: dorsum of

head and body pale brown to dark brown, frequently with darker brown blotches or

hourglass figure on back; pale interorbital bar varying from well defined to obscure; dark

bars present on upper lip in most specimens; para-anal pale bars or spots well defined,

indistinct, or absent; belly and venter of thigh cream colored, heavily flecked with dark

brown; chin, throat, and chest densely flecked with dark brown, flecking surrounding pale

“spots” on many specimens, the pale “spots” extending onto belly in occasional

specimens. À single subadult female (KU 209059) has much paler ventral surfaces than the

remaining specimens, including the other subadult specimen and the juveniles.

Etymology. — The specific name epochthidius is formed from the Greek word epochthidios,

which means on or of the mountains. The name alludes to the montane habitat of this

species.

Distribution and natural history notes. — Moderate elevations (760 to 1450 m) in the

Premontane Wet Forest formation of HOLDRIDGE (1967) of the Sierra de Agalta north of

Catacamas and northwest of Dulce Nombre de Culmi (fig. 4), Honduras. The series from

between the rios Catacamas and Seco was taken in about 30 min of collecting along a

small stream just before midday on 5 August 1986. The Rio Seco series was collected both

during the day and at night along a large stream on 8-9 August 1986. The Montaña de

Malacate specimen was collected along a small stream just before midday on 11 June 1980.

Source : MNHN, Paris

MCCRANIE & WILSON 173

KEY TO THE HONDURAN SPECIES IN THE E. MILESI GROUP

+ Tocs-wWithilateral fleshy fringes sms isds hs cndiaeseensetines sucette 2

Toes with well developed lateral keel, but without fleshy fringes (toe IV with weak

fleshy fringes in some large female E. fecundus) ............................. 5

. Toes extensively webbed, modal formula I 1 1/3 — 2 11 1 1/3 — 2 1/2 III 2 — 3 1/3

IV 31/3 — 2 V; males to 41 mm SVL (moderate elevations of central part of

Cordillera Nombre de Dios).................................. E. chrysozetetes

Toes moderately webbed, modal formula 1 2 — 2 3/4 II 2 — 3 1/2 III 3 — 4 IV 4 —

21/2Vor12— 23/4112 — 31/2 III 3 — 41/4 IV 4 1/4 — 2 3/4 V; males less

than mm S VE tee nes sentir te eos else 3

. Toes III-IV or only toe IV (some males) with lateral fleshy fringes; webbing on toes

HI-V slightly reduced, modal formula III 3 — 4 1/4 IV 4 1/4 — 2 3/4 V; males

20.9-26.9 mm SVL (moderate elevations of Sierra de Agalta)... E. epochthidius

Toes II-IV with lateral fleshy fringes; increased webbing on toes III-V, modal formula

II 3 — 4IV 4 — 21/2 V; males 27.0-33.2 mm SVL....................... 4

. Thin, pale middorsal stripe extending from tip of snout to above vent (intermediate

elevations of central part of Cordillera Nombre de Dios) ............. E. cruzi

No pale middorsal stripe (moderate and intermediate elevations of western part of

Cordillera Nombre de Dios and west-central Yoro to northwestern Olancho) ...

£. stadelmani

. Upper lip distinctly flared; vocal slits absent in adult males; males 26-30 mm SVL; male

tympanum length 72-85 % of eye length (moderate elevations of Sierra de Omoa)

D Sptnie eve Miatomiepute vien ee monte des pulse tes SHUIMÉE oreLtre PHARE ENTRE ee Æ. omoaensis

Upper lip not flared; vocal slits present in adult males; males less than 26 mm SVL;

male tympanum length 35-68 % length of eye, or tympanum very indistinct to

hidden and unmeasureable.… 6

. Modal webbing formula 1 2 — 2 4/5 II 2 — 3 3/4 III 3 — 4 1/4 IV 4 1/4 — 2 3/4 V;

male tympanic annulus indistinct, very indistinct, or hidden (low and moderate

elevations of central and eastern portions of Cordillera Nombre de Dios).......

E. fecundus

Modal webbing formula 1 — I12* — 3 4/5 111 3* — 41/2 IV 41/2 — 3 Vor12*

— 24/5112 — 33/4113 — 41/21V 41/2 — 3 V; male tympanic annulus distinct

OF IRAISHNCE EL 5 ue ta ete Ben 6 RU ect da ee RME ae ve E 4É

. Male tympanic annulus distinct or indistinct, that of female very indistinct; male

tympanum length 35-46 % length of eye; occurs in forest well away from streams

(intermediate elevations of western and central portions of Cordillera Nombre de

Dios) E. saltuarius

Male tympanic annulus distinct, that of female indistinct; male tympanum length

47-68 % length of eye; occurs alongside streams (moderate and intermediate

elevations of sierras Omoa and Espiritu Santo) ...................... E. milesi

Source : MNHN, Paris

174 ALYTES 14 (4)

ACKNOWLEDGMENTS

Field assistance was provided by D. ALMENDAREZ, M. R. EsPiNAL, G. A. FLORES, J. C.

RINDFLEISH, and K. L. WiLLiams. Collecting and exportation permits have been provided over the

years by the personnel of the Departamento de Recursos Naturales Renovables, Tegucigalpa, and

those of Corporacién Hondureña de Desarrollo Forestal, Tegucigalpa. Assistance with recent permits

was provided by M. R. EsiNaL. Specimen loans were provided by the following colleagues: A.

ReseraR (FMNH); W. E. DUELLMAN and J. R. MENDELSON (KU); R. L. BEzy (LACM); J. E. CADLE

and M. L. E. STeINER (MCZ); D. B. Wake (MVZ); R. W. MurPHY (ROM); S. D. SRokA (UIMNH);

A. G. KLUGE and G. SCHNEIDER (UMMZ); R. W. MCDiaRMID and J. A. POINDEXTER (USNM).

Additionally, M. E. EsPINAL made available his collection of Eleutherodactylus, J. F. LYNCH and D.

B. Wake provided collecting information for MVZ specimens, J. M. SAVAGE reviewed an early draft

of the manuscript, and R. NUTT prepared fig. 3.

LITERATURE CITED

CAMPBELL, J. A., 1994. — New species of Eleutherodactylus (Anura: Leptodactylidae) of the milesi

group from Guatemala. Herpetologica, 50: 398-411.

HEYER, W. R., RAND, A. S., GONÇALVES DA CRUZ, C. AÀ., PEIXOTO, O. L. & NELSON, C. E., 1990. —

Frogs of Boracëia. 4rg. Zool., 31 (4): 231-410.

HOLDRIDGE, L. R., 1967. — Life zone ecology. Revised edition. San José, Costa Rica, Tropical Science

Center: 1-206.

LEviTON, A. E., Giss, R. H., Jr., HEAL, E. & DAWsON, C. E., 1985. — Standards in herpetology and

ichthyology. Part I. Standard symbolic codes for institutional resource collections in

herpetology and ichthyology. Copeia, 1985: 802-832.

LyncH, J. D. & DuELLMAN, W. E., 1980. — The Eleutherodactylus of the Amazonian slopes of the

Ecuadorian Andes (Anura: Leptodactylidae). Univ. Kansas Mus. nat. Hist. misc. Pub., 69: 1-86.

MCCRANIE, J. R., SAVAGE, J. M. & WiLsoN, L. D., 1989. — Description of two new species of the

Eleutherodactylus milesi group (Amphibia: Anura: Leptodactylidae) from northern Honduras.

Proc. biol. Soc. Washington, 102: 483-490.

O'SHEA, M. T., 1989. — New departmental records for northeastern Honduran herpetofauna. Herp.

Rev, 20: 16.

SAVAGE, J. M., 1975. — Systematics and distribution of the Mexican and Central American stream

frogs related to Eleutherodactylus rugulosus. Copeia, 1975: 254-306.

SCHMIDT, K. P., 1933. — New reptiles and amphibians from Honduras. Zool. Ser. Field Mus. nat.

Hist., 20: 15-22.

- 1936. — New amphibians and reptiles from Honduras in the Museum of Comparative Zoology.

Proc. biol. Soc. Washington, 49: 43-50.

SMITHE, F. B., 1975. — Naturalist's color guide. Part I. Color guide. New York, The American

Museum of Natural History: 182 color swatches.

Corresponding editors: W. Ronald HEYER & Alain DuBois.

© ISSCA 1997

Source : MNHN, Paris

Alytes, 1997, 14 (4): 175-200. 175

Instructions to authors

of papers submitted to Alytes

Alain Dugois

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Source : MNHN, Paris

176 ALYTES 14 (4)

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Source : MNHN, Paris

Dugois 177

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Source : MNHN, Paris

178 ALYTES 14 (4)

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bibliographic references or of figures and tables in the manuscript.

Names of new taxa described in the paper should not be mentioned in the abstract (see

below).

At the end of the paper, a summary should be given in one or several other languages,

including either French or Spanish. In papers written in English, this summary may be

slightly longer than the English abstract.

Source : MNHN, Paris

Dugois 179

SECTIONS OF THE MANUSCRIPT

The manuscript should preferably be organized according to the following sections:

abstract, introduction, material(s) and methods, abbreviations, results, discussion, conclu-

sion, French or Spanish summary, acknowledgements, literature cited, appendices.

Sections (as listed above) and possible subsections should not be numbered. The hierarchy

of subsections within sections should be limited to a low number of levels, preferably no

more than three. Section headings should be centered, whereas those of subsections should

be indented. All headings should be typewritten in lower case, except for initials that must

be printed in capitals for grammatical or nomenclatural reasons (see below).

ABBREVIATIONS, UNITS AND NUMERALS

Tables 1-3 provide a list of standard abbreviations and symbols that should be used

in Alytes. Please note that some abbreviations are italicized, while others are not, and that

none includes dots. All other abbreviations should follow if possible usual standards in

scientific literature, and should be defined in a special section or table of the manuscript.

As acknowledged by all major dictionaries (e.g., PROCTER, 1995: 12; THOMPSON, 1995: 13),

the term “‘acronym” designates an abbreviation formed of the initial letters of several

words and pronounced as a word, such as NATO, AIDS or ISSCA: therefore this term

should not be used to designate abbreviations that must be spelled out and cannot be

pronounced as a single word, such as most Museum or measurement abbreviations. In

such abbreviations, letters should not be separated by dots (eg, “MNHN”, not

“M.N.H.N.”). Signs, symbols or characters that cannot be typed with the printing

equipment used by author(s) should be added by hand on every occurrence in each copy

of the manuscript.

In text or in legends, a sentence should never start with an abbreviation.

Weights and measures should follow the International System of Units (SI). Table 4

gives the standard abbreviations for the most common of these units, which should be used

in text, tables and figures. Please note that none is italicized and that none includes dots.

Unusual units should be defined before their first use. Measurements should be given in

figures which are consistent with the degree of accuracy obtainable.

Numbers one to nine should be spelled out, unless they precede units or are part of

a series of numbers: e.g., “3 mm’, “3 to 4 days”, “3-4 days”. Numbers of 10 or larger

should be given as Arabic numerals except at the beginning of a sentence. Numbers up to

four digits should be written in one block: e.g., “1948”, “6500”. In numbers with five or

more digits, a space should be used to separate thousands from hundreds, in order to

avoid confusion between the English style on one hand (where comma is used) and the

French and Spanish styles on the other (where dot is used): e.g., “35 637”, “350 000”,

“12 000 000”. Dates should be given by day, month and year: e.g., “11 February 1960”,

“11 Feb. 1960” or “11.11.1960”. Times of day (24-hour clock) can be written either

as “17 h 00” or as “17.00 h”. Decimals should not be given naked: e.g., “0.5”,

not “.5”.

Source : MNHN, Paris

180

ALYTES 14 (4)

Tab. 1. - Various standard abbreviations and symbols to be used in text, plates and figure

legends in Alytes.

Abbreviation

or symbol

Meaning

Abbreviation

or symbol

Meaning

adult

Mahalanobis distance

degrees of freedom

desoxyribonucleic acid

Doctor

cast

first generation offspring

standard value of

Kruskal-Wallis test

null hypothesis

head length!

head width

imago?

individual

juvenile

natural logarithm

base 10 logarithm

median

maximum

minimum

sample size

population size

north

number

northeast

nord-ouest

not significant (P > 0.05)

northwest

ouest

probability of wrongly

rejecting the null

hypothesis

(significance level)

hydrogen ion activity

Doctor of Philosophy

ppm

1. Preferably measured on side of head (see e.g. LIU, 1950: 19).

2. International term for “just metamorphosed amphibian” (see DUuOIS, 1978).

parts per million

Professor

coefficient of correlation

ribonucleic acid

south

standard deviation

standard error of the mean

southeast

scanning electron microscopy

sud-ouest

subadult

snout-vent length

southwest

standard value of

Student 1 test

tadpole

tibia length

standard value of

Mann-Whitney U test

variation coefficient

mean

standard value of

chi-square test

significant

(P<0.05)

highly significant

(P<0.01)

very highly significant

(P<0.001)

per cent

per thousand

male

female

Source : MNHN, Paris

Dugois 181

Tab. 2. - Standard abbreviations for 20 associations, organizations and countries, to be used in

text, plates and figure legends in Alytes.

Abbreviation Meaning

Asociacién Herpetolégica Argentina

Asociacién Herpetolégica Española

American Society of Ichthyologists and Herpetologists

British Herpetological Society

Declining Amphibian Populations Task Force

Deutsche Gesellschaft für Herpetologie und Terrarienkunde

European Union

International Commission on Zoological Nomenclature

International Society for the Study and Conservation of Amphibians

World Conservation Union

North Atlantic Treaty Organization

Ôsterreichische Gesellschaft für Herpetologie

Societas Europaea Herpetologica

Société Herpétologique de France

Society for the Study of Amphibians and Reptiles

United Kingdom

United Nations Organization

United States of America

World Congress of Herpetology

World Wide Fund for Nature

CAPITALS

In the headings, subheadings, text, tables, and figure captions, the use of capital letters

as initials should be limited to situations where this is strictly compulsory for nomenclatural

or grammatical reasons: Latin scientific names above the species level, names of months

and days in English (e.g, “September”, “Saturday”), names of persons, countries,

standard or official geographic names, nouns in titles of books, periodicals, papers or

chapters when cited in the text (but not in the list of references: see below), nouns and

adjectives in titles of institutions when cited in the text (e.g., “Muséum National d'Histoire

Naturelle, Paris”), etc. Descriptive geographic adjectives such as “south”, “southern”,

“central”, “lower”, etc., should not start with a capital except when they are included in

standard or official administrative names of countries or regions: e.g., “New South

Wales”, “South Africa”, “Northern Frontier Province”, “Central Province”, “Hautes

Alpes”. Vernacular or common names of taxa (e.g., ‘‘anurans”, “brown frogs”, “ranids”),

names of anatomical structures (e.g., ‘“keratodont”, “canthus rostralis””) or of scientific

disciplines (e.g., “zoology”, “herpetology”, “genetics”) should not start with a capital.

Source : MNHN, Paris

182 ALYTES 14 (4)

Tab. 3. - Standard abbreviations for 30 herpetological Museum collections, to be used in text,

plates and figure legends in A/ytes.

Abbreviation

American Museum of Natural History, New York, New York, USA

Academy of Natural Sciences, Philadelphia, Pennsylvania, USA

Natural History Museum, London, United Kingdom

California Academy of Sciences, San Francisco, California, USA

Chengdu Institute of Biology, Academia Sinica, Chengdu, Sichuan,

una

Coleccién de Vertebrados de la Universidad de Los Andes, Mérida,

Venezuela

Fundaciôn Miguel Lillo, Tucumän, Argentina

Field Museum of Natural History, Chicago, Illinois, USA

Museum of Natural History, The University of Kansas, Lawrence,

Kansas, USA

Los Angeles County Museum of Natural History, Los Angeles,

California, USA

Museo Argentino de Ciencias Naturales Bernardino Rivadavia,

Buenos Aires, Argentina

Museum of Comparative Zoology, Cambridge, Massachusetts, USA

Muséum d'Histoire Naturelle, Genève, Switzerland

Museo Nacional de Ciencias Naturales, Madrid, Spain

Muséum National d'Histoire Naturelle, Paris, France

Museu Nacional do Rio de Janeiro, Rio de Janeiro, Brazil

Museo Civico di Storia Naturale Giacomo Doria, Genova, Italy

Museum of Vertebrate Zoology, University of California, Berkeley,

California, USA

Museu de Zoologia, Universidade de Säo Paulo, Säo Paulo, Brazil

Naturhistorisches Museum, Wien, Austria

Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands

Royal Ontario Museum, Toronto, Canada

Senckenberg Museum, Frankfurt-am-Main, Germany

Museum of Natural History, University of Illinois, Urbana, Illinois,

USA

Museum of Zoology, University of Michigan, Ann Arbor, Michigan,

USA

National Museum of Natural History, Washington, D.C., USA

Zoologisches Forschungsinstitut und Museum Alexander Koenig,

Bonn, Germany

Zoological Institute, Academy of Sciences, Saint-Petersburg, Russia

Zoologisches Museum an der Humboldt-Universität, Berlin, Germany

Zoological Survey of India, Calcutta, India

Source : MNHN, Paris

Dugois 183

Tab. 4. - Standard abbreviations for measurement units to be used in text, plates and figure

legends in Alytes.

Abbreviation

Meaning

Abbreviation

Meaning

cm

centimetre

day

gramme

hour

hectare

hertz

joule

kilogramme

kilohertz

kilometre

litre

metre

milligramme

minute (time)

millilitre

millimetre

month

millisecond (time)

million years

chromosome number

nanometre (10° m)

second (time)

volt

watt

year

micrometre (10 m)

degree Celsius

degree (angle)

minute (angle)

second (angle)

Names of authors of cited literature, as well as names of persons mentioned in the

acknowledgements, should be typed in capital letters (preferably small capitals), both in

text and in figure and table legends, and in the list of references. On the other hand,

authors’ names following Latin scientific names, in parentheses if necessary, should be

typed in lower-case letters, followed by a comma and the date: e.g., “Rana temporaria

Linnaeus, 1758”; “Bufo bufo (Linnaeus, 1758)”. The same rule should be followed for

persons” names that are included in standard experimental or statistical equipments,

procedures or methods: e.g., “Petri dish”, “Ringer solution”, “Feulgen reaction”,

“Lincoln-Petersen index”, “Mahalanobis distance”, “Mann-Whitney U test”.

ITALICS

Parts of the text that are meant to be finally printed in italics should be typed

underlined or printed in italics. Italics should be used for scientific names of the genus and

species groups, for new or newly defined scientific terms, for some anatomical names (e.g.,

names of muscles), for titles of books, periodicals, papers or chapters, and for emphasis

on particular terms or sentences. They should not be used for Latin or other foreign terms

or abbreviations of common use or “that are now considered standard English in science”

(HurH et al., 1994: 170), such as “e.g.”, “i.e.”, “etc.”, ‘et al.”, “s. str.”, “a priori”, “vice

versa”, “hoc loco”, “in vivo”, “canthus rostralis” or “sp. nov.”.

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184 ALYTES 14 (4)

FOOTNOTES

The use of footnotes is not encouraged in A/ytes. They may however be used to clarify

tables, and in the text when this is clearly the best solution to briefly present information

that does not belong in the text but is nevertheless useful to the understanding of the latter.

Footnotes should be numbered in Arabic numbers (1, 2, 3...) from the beginning to the end

of the paper.

SPECIMENS, TAXONOMIC AND GEOGRAPHIC INFORMATION

SPECIMENS

Papers based on animals killed for the purpose of research should indicate the exact

location and catalogue numbers of voucher specimens and, when appropriate, information

on collecting permits and licenses.

Holotypes of newly described taxa, and if possible all other studied specimens, should

be deposited in internationally recognized scientific institutions, preferably in Museums, as

these institutions have the duty to provide specimens with permanent storage and care,

and to make them available for study to the international scientific community. Table 3

provides a list of abbreviations that should be used in A/ytes to designate some major

herpetological Museum collections in the world; abbreviations used for other collections

should be defined in the paper (see above).

TAXONOMIC ACCOUNTS

To be acceptable in Alytes, any paper proposing description of new taxa should

include clear evidence that a thorough investigation of related taxa has been carried

out.

Species or subspecies taxonomic accounts should be organized following subsections

such as: synonymy, holotype, paratypes, type locality, etymology of name, diagnosis,

description, variation, measurements, coloration, comparisons, phylogeny, natural history,

distribution, comments. Genus or subgenus taxonomic accounts should be organized

following subsections such as: synonymy, type species (with its mode of designation),

etymology of name, species content, diagnosis, variation, comparisons, phylogeny, natural

history, distribution, comments. Mention of type species of genera should always be

accompanied by precise information on their mode of designation.

Synonymies presented, when justified, at the head of taxonomic accounts, should

preferably be limited to lists of genuine synonyms (sensu SmiTH & SmiTH, 1973) and of new

combinations and subsequent spellings. They should preferably not include chresonyms

(Suit & SmirH, 1973), or, if these are given, they should be presented in such a way as

Source : MNHN, Paris

Dusois 185

to be clearly identified and distinguished from true synonyms, e.g., by use of a colon

between the scientific name and the name of the (first) author of reference. The

mathematical symbol “=” is not appropriate to designate synonymies and should never

be used in this context. Instead, sentences like “Hyla viridis Laurenti, 1768 (subjective

junior synonym of Rana arborea Linnaeus, 1758)” should be used.

Descriptions of specimens should mention their sex and age class (e.g., tadpole,

imago, juvenile, subadult, adult). In descriptions, it is strongly recommended to include

measurements, at least the four following standard ones: SVL, HW, HL and TL (see

tab. 1). Other measurements not mentioned in tab. 1 should be defined in the paper.

Descriptions should be written in a homogeneous manner. In a given paper, a choice

should be made by the author(s) between telegraphic style (“Head wider than long”) or

complete sentences with verbs (“The head is wider than long”). If possible, telegraphic

style should be chosen. If several taxa are described in the same paper, their descriptions

should follow exactly the same format, possibly with numbered subdivisions.

In order to facilitate comparisons between different works, standard descriptive

methods, terms, formulae or stage tables, when available, should be used: e.g., DuBoIs’s

(1978) major developmental stages for amphibians, Gosner’s (1960) developmental table

for tadpoles, Duois’s (1995) format for the presentation of tadpole keratodont formulae,

or Myers & DUELLMAN'S (1982) format for the presentation of webbing formulae.

NOMENCLATURE

For all nomenclatural matters, the rules of the most recent edition of the International

Code of Zoological Nomenclature (ANONYMOUS, 1985; cited below as “the Code”) should

be strictly followed. Particular attention will be paid by the Editors to the fact that

scientific names used in papers published in Alytes are the valid ones according to the

Code, even if they are not the “usual” ones. In particular, family-group names, as well as

species-group and genus-group names, should follow the Rule of Priority, except for

specific rulings by the International Commission on Zoological Nomenclature.

The inclusion of the author’s name in citing the name of a taxon is primarily intended

for the purpose of helping the search for literature and information: “It is not, as many

detractors would have it, merely to promote the reputation of a scientist or simply an

egotistical exercise. The value of knowing the author’s name so as to be able to track down

the original reference (...) is much more important than merely knowing who the person

responsible for the name is. There is, as yet, no other effective way to keep track of

authors, publication dates and to trace the original references.” (NG, 1994: 509). With the

help of the Zoological Record, knowledge of the author’s name and of the date is usually

sufficient to find the reference of the original description of a taxon. However such a

search is much more difficult when only the author’s name, but not the date, is given.

Therefore, in all parts of manuscripts submitted to Alytes, scientific names of taxa should

be mentioned either followed by their complete authorship information, i.e. their complete

(not abbreviated) author’s name and their date (either in parentheses or not, according to

the Code), or without both, but never with the author’s name without the date: e.g., “Rana

Source : MNHN, Paris

186 ALYTES 14 (4)

catesbeiana Shaw, 1802” or “Rana catesbeiana”, not “Rana catesbeiana Shaw”; “Hyla

arborea (Linnaeus, 1758)” or “Hyla arborea”, not “Hyla arborea (L.)”; “Eleutherodactylus

Duméril & Bibron, 1841” or ‘“Eleutherodactylus”, not “Eleutherodactylus D. B.”;

“Ranidae Rafinesque-Schmaltz, 1814” or “Ranidae”, not “Ranidae Rafinesque”.

The Latin name of any species-group taxon or of any new combination should be

written in full at least once, on the occasion of its first occurrence in the paper. If

subsequently the same taxon is mentioned a high number of times, the generic name

should be replaced by its initial; the same rule may apply to the specific name of

subspecies. Such a use of shortened names is not necessary in papers where scientific names

are mentioned only a small number of times.

When proposing new scientific names, author(s) should pay close attention to

checking that these names are not preoccupied. New names should be formed according

to the rules and recommendations of the Code, and should, as far as possible, be simple,

short and euphonious. Authors are encouraged to use their imagination to propose new,

original names, rather than always using common epithets which have already been

applied to many animal species (e.g., “viridis”, ‘“rugosus”, “major”, “vulgaris”, “monta-

nus”, “‘occidentalis”, “sinensis”). For species-group names based on geographic terms,

attention of authors is drawn to the possibility to use local names in any language placed

in apposition to the generic name, which is often shorter, more euphonious and more

original than constant use of the suffixes “-ensis” or “-icus”: e.g., “Rana diuata Brown

& Alcala, 1977”; “Ranixalus gundia Dubois, 1986”; “Aubria masako Ohler & Kazadi,

1990”.

The name of a new taxon described in the paper should not appear before the heading

of the section devoted to the description of the taxon, so that there can be no doubt or

disagreement among subsequent authors as to the page to be cited as first page of appearance

of the new name.

GEOGRAPHIC INFORMATION

Both in the paper and in the addresses of authors in the title page, geographic names

of countries should be given in the language of the paper: e.g., “Germany” in a paper in

English, “Allemagne” in French, ‘“Alemania” in Spanish, never “Deutschland”. The same

is true for international geographic units, such as seas, lakes, rivers, mountains: e.g., in a

paper in English, “Amazon Basin”, “Indian Ocean”, “Rhine”, ‘“Alps”. In contrast, names

of provinces, districts, towns, rivers, lakes, mountains, etc., entirely included within a

country, should be given in the (or one of the) official language(s) of this country, not in

the language of the paper: e.g., “Wien”, not “Vienna”; “Massif Central”, not “Central

Massif”; “Castilla”, not “Castile”. However, generic geographic terms such as ‘“moun-

tain”, “river”, “lake”, should be given in the language of the paper: e.g., in a paper in

English, “Borjomi canyon”, “lake Dreibrüdersee”, “Samothraki island”, “Thévenon

pond”, “Masako forest”. Special care should be taken to ensure that geographic words are

correctly spelled in their original languages, including accents (e.g., “é”, “”, “à”, “é”) and

other diacritic marks such as tilde (‘ñ”'), diaeresis (‘“‘ü”') or cedilla (“£”).

Source : MNHN, Paris

Duois 187

Localities of study and/or collection of specimens should be referred to as precisely

as possible, preferably giving their latitude and longitude (in relation to Greenwich

meridian) and their altitude above sea level (in metres). Distances should be given in

metres or kilometres: e.g., “Masako forest, near Batibongena village, 15 km from centre

of Kisangani on the ancient road to Buta, Zaire”. As different atlases and maps often give

slightly different latitudes, longitudes or altitudes for the same locality, it is recommended

to provide, in the Literature Cited, references of the documents used to find the

coordinates of localities given in the paper. If altitude and/or position was/were

determined directly in the field, e.g. by use of an altimeter or a Global Positioning System

(GPS), rather than on a map, this should be stated in the Material(s) and Methods section

of the paper.

FIGURES, TABLES AND APPENDICES

FIGURES

Alytes encourages the publication of high quality figures (photographs, drawings,

graphs, etc.) illustrating the major points of a paper. A good figure may often be more

informative and convincing than long descriptions or discussions. To be acceptable in

Alytes, figures should be of good quality and should clearly show the character to be

depicted or the results of experimental work to be illustrated.

Each figure (“figure” in English and French; “figura” in Spanish) should be planned

to accommodate reduction and to occupy, together with its legend, no more than the

maximum height and width of the printed portion of a page in Alytes (13 x 18.5 cm).

Originals of figures submitted should not exceed 21 x 29.7 cm. The size of the lettering

should ensure its legibility after reduction. Optimum reproduction is achieved by reducing

slightly the size of the figures during the process of publication.

Figures should include a scale of magnification, dimension or distance where

appropriate. This should appear as a scale bar on the figure itself, not only in the legend.

AI axes of graphs should be labelled. If letters or numbers are used in the figure, these

should not be handwritten, but printed or transferred. Abbreviations in figures should

follow the conventions given above and should be defined either in the text or in the figure

legend.

Several figures or subfigures can be labelled and mounted on Bristol board to produce

a single plate that will occupy, with its legend, a full page of the journal. Subfigures of a

composite figure should be lettered in lower case (a, b, c...). It is suggested to protect

figures, in particular photographs mounted as plates, by a transparent overlay, e.g., tracing

paper; any areas of particular interest or importance on a photograph may be outlined in

pencil on the overlay for the guidance of the printer.

Figure legends should be intelligible without reference to the text. Legends of figures

showing specimens or parts of specimens should include information about the actual

Source : MNHN, Paris

188 ALYTES 14 (4)

specimen shown (collection number of specimen, date of collecting or of photograph, etc.).

Legends of figures should not appear on the figures themselves. They should be grouped

and presented double-spaced on separate sheet(s).

Figures should be numbered in Arabic numerals in the order of their first mention in

the text. In the text, figures should be referred to in an abbreviated form (e.g., “fig. 2”),

except at the beginning of a sentence in English (e.g., “Figure 4 shows...”). Several figures

should be cited using “fig.”, not “figs.”: e.g., “fig. 4-5”. If several figures or subfigures are

assembled on a plate (see above), they will always be referred to by their individual

numbers (e.g., “fig. 3”, “fig. 7a”; not “pl. IL, fig. 3”), and plates will not be numbered

as such (in A/ytes, pages occupied by plates are numbered just like standard pages of the

journal).

The top, bottom or back of each figure or plate should be labelled in pencil with the

(first) author’s name, the figure number and its date of submission.

TABLES

Alytes encourages the publication of complete tables of original data, that can be used

by subsequent authors for further analysis or critical reevaluation, rather than simply

providing results of statistical tests, phylogenetic analyses, etc.

Tables should be as clear and simple as possible, and should avoid the unnecessary

multiplication of columns and lines. During the preparation of tables in text processing,

tabulations, rather than simple spaces, should be used to separate columns.

Each table (“table” in English; “tableau” in French; “tabla”’ in Spanish) should be

typed or printed, double-spaced, on a separate sheet, together with its legend. Table

legends should be intelligible without reference to the text. Abbreviations in tables should

follow the conventions given above and should be defined in the text or in the table legend.

Tables should be numbered in Arabic numerals in the order of their first citation in

the text. In the text, tables should be referred to in an abbreviated form (e.g., “tab. 2”),

except at the beginning of a sentence in English (e.g., “Table 4 shows...”). Several tables

should be cited using “tab.”, not “tabs.”: e.g., “tab. 4-5”.

APPENDICES

AS far as possible, contributors should avoid the use of appendices in manuscripts

submitted to Alytes. In many cases, data that are sometimes provided in appendices can

be presented as in-text tables. Only very long tables (more than two pages, e.g. giving

details of original data), or very specific discussions of technical or other questions that

would unnecessarily make the text unwieldy, should be placed in an appendix (“appendix”

in English; “appendice” in French; “apéndice” in Spanish).

Appendices should be labelled in Arabic numbers in the order of their first citation

in text. In the text, appendices should be referred to in an abbreviated form (e.g., “app.

Source : MNHN, Paris

DuBois 189

2” in English and French; “ap. 2” in Spanish), except at the beginning of a sentence in

English (e.g., “Appendix 4 shows...”). Several appendices should be cited using “app.”,

not “apps.”: e.g., “app. 4-5”.

BIBLIOGRAPHIC REFERENCES

Authors are particularly urged to ensure that bibliographical details are accurate.

Correct references are the responsibility of the author(s). This implies that, as far as

possible, all cited references should have been checked personally by the author(s), not

copied from other sources. When this is impossible, the reference should be followed by

the mention “[Not seen]”, possibly followed by reference to the source used (see example

below).

CITATION IN TEXT

In the text (including synonymies of systematic accounts, figure legends and tables),

bibliographic references should be referred to by the name of the author(s), written in

capital letters, followed by the actual date of publication of the work: e.g., “GÜNTHER,

1859”; “GRAF & PoLLs PELAZ, 1989”; “INGER et al., 1984b”; ‘ANONYMOUS, 1993”.

References such as “GÜNTHER, ‘1858 (1859)” should not be used: if the date that is printed

in the publication itself is not the actual date of publication, this should simply appear in

the full reference given in the list of references (see below). Papers with two authors should

be referred to in the text by their names, connected by the ampersand (sign “&”,

international for “and”, “et”, “y”, “und”, etc.). Papers with more than two authors

should always be referred to by the first author’s surname followed by “et al.”. Strings of

cited references dealing with the same question should be given in chronological order of

publication, and not in alphabetical order of first authors. Special care should be taken to

ensure that all references mentioned in the text are included in the list of references, and vice

versa. Dates should match.

Two or more references by the same author(s), or having the same first author in case

of more than two authors, and published in the same year, should be distinguished by

adding, after the date, lower case, italicized letters (a, b, c...); if possible, these letters

should be given in chronological order of publication. These lower case letters may be

either connected by a hyphen (which points to all references from first to last including

those not listed between them) or separated by commas (which points to those references

only): e.g., “DUMÉRIL et al., 1854a-c” points to references appearing in the list of references

under the letters a, b and c, while “DUMÉRIL et al., 1854a,c” points only to references

appearing in the list of references under the letters a and c.

Source : MNHN, Paris

190 ALYTES 14 (4)

LIST OF REFERENCES

In the Literature Cited, references should be listed in alphabetical order of first author

and presented as follows (see also the list of references at the end of this paper):

(1) For a paper in a periodical:

ANONYMOUS, 1986. — [Footnote]. Bull. Mus. nain. Hist. nat., “1985”, (4), 7 (A) (4): 937.

BOULENGER, G. A., 1892. — A synopsis of the tadpoles of the European batrachians. Proc. zool. Soc.

London, 1891: 593-627, pl. 45-47. [Publication date according to DUNCAN, 1937].

CLARKE, B. T., 1981. — Comparative osteology and evolutionary relationships in the African Raninae

(Anura Ranidae). Monit. zool. ital., (n.s.), 15 (suppl.): 285-331.

DE L'IsLE, A., 1872. — De l’hybridation chez les amphibies. Ann. Sci. nat., (5), 17 (3): 1-24.

DELAUGERRE, M. & Dumois, A., 1986. — La variation géographique et la variabilité intrapopula-

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

nat., “1985”, (4), 7 (A) (3): 709-736. [Publication date according to ANONYMOUS, 1986].

Duéril, A., 1856. — Note sur les reptiles du Gabon. Rev. Mag. Zool. pure appl., (2), 8 (8): 369-377;

8 (9): 417-424; 8 (10): 460-470; 8 (12): 553-562.

Duncan, F. M., 1937. — On the dates of publication of the Society’s Proceedings, 1859-1926. Proc.

zool. Soc. London, 107 (A): 71-84.

HARPER, F., 1940. — Some works of Bartram, Daudin, Latreille, and Sonnini, and their bearing upon

North American herpetological nomenclature. Am. Mid. Natur., 23 (3): 692-723.

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

amphibians and reptiles from the Ponmudi, Kerala, South India. J. Bombay nat. Hist. Soc., 81

(2): 406-427, pl. 1-3.

——— 1984b. — A report on a collection of amphibians and reptiles from the Ponmudi, Kerala, South

India. (Continued). J. Bombay nat. Hist. Soc., 81 (3): 551-570, pl. 4-5.

Maïsui, M. & HikiDA, T., 1985. — Tomopterna porosa Cope, 1868, a senior synonym of Rana

brevipoda Ho, 1941 (Ranidae). J. Herp., 19 (3): 423-425.

SALVADORI, T., 1888. — Le date della pubblicazione della /conografia della fauna italica del

Bonaparte ed indice delle specie illustrate in detta opera. Bol. Mus. Zool. Anat. comp. Univ.

Torino, 3 (48): 1-25.

SHergorN, C. D., 1934. — Dates of publication of Catalogues of Natural History (post 1850) issued

by the British Museum. Ann. Mag. nat. Hist., (10), 13 (74): 308-312.

WILSON, A. C., MaxsoN, L. R. & SariCH, V. M., 1974a. — Two types of molecular evolution.

Evidence from studies of interspecific hybridization. Proc. nain. Acad. Sci. USA, 71 (7):

2843-2847.

WILSON, A. C., SARICH, V. M. & MaxsON, L. R., 1974b. — The importance of gene rearrangement

in evolution: evidence from studies on rates of chromosomal, protein, and anatomical

evolution. Proc. natn. Acad. Sci. USA, T1 (8): 3028-3030.

(2) For a book:

ANONYMOUS, 1977. — Zhonghua Renmin Gongheguo Fen Sheng Dituji. (Hanyua Pinyinban).

Zhongguo Beijing, Ditu Chubanshe: [i-vii] + [i-ii] + 1-169, pl. 1-51.

ANDERSON, J., 1879. — Anatomical and zoological researches, comprising an account of the zoological

results of the two expeditions 10 western Yunnan in 1868 and 1875 and monograph of the two

cetacean genera Platanista and Orcella. London, Quaritch, “1878”. Vol. 1: i-xxv + 1-985; vol.

2: pl. 1-81. [Publication date according to ZHAo & ADLER, 1993: 350].

BLaiR, W. F., (ed.), 1972. — Evolution in the genus Bufo. Austin & London, Univ. Texas Press: i-viii

+ 1:459.

MAN, W. E. & TRUEB, L., 1985. — Biology of amphibians. 1st edition. New York, MeGraw-Hill,

“1986”: i-xix + 1-670. [Publication date according to ZHao & ADLER, 1993: 188].

Duel

Source : MNHN, Paris

Dugois 191

FRosT, D. R., (ed.), 1985. — Amphibian species of the world. Lawrence, Allen Press & Ass. Syst. Coll.:

HV] + iv + 1.732.

GünTHER, A., 1859. — Catalogue of the Batrachia Salientia in the collection of the British Museum.

London, Taylor & Francis, “1858”: i-xvi + 1-160, pl. 1-12. [Publication date according to

SHErBORN, 1934].

NAKAMURA, K. & UÉNO, S.-I, 1963. — [Japanese reptiles and amphibians in colour]. Osaka,

Hoiku-sha: 1-214, pl. 1-42. [In Japanese]. [Not seen, cited according to MaTsuI & HikIDA,

1985].

THorN, R., 1969. — Les salamandres d'Europe, d'Asie et d'Afrique du Nord. Paris, Lechevalier,

“1968”: 1-376. (Publication date according to printing date mentioned in p. 377 of the book].

Ye, C., Fi, L., & Hu, S., 1993. — [Rare and economic amphibians of China]. Chengdu, Sichuan

Publishing House of Science and Technology: [i-ii] + 1:2 + 1-2 + 1-7 + 1-412. [In Chinese].

ZHao, E.-M. & ADLER, K., 1993. — Herpetology of China. Oxford, Ohio, USA, SSAR: 1-522 + [i-v],

pl. 1-48 + 1.

(3) For a chapter or a part of a book:

BONAPARTE, C. L., 1838. — Rana esculenta. Ranocchia verde. In: C. L. BONAPARTE, Jconografia della

Sauna italica per le quattro classi degli animali vertebrati, fasc. 22, Roma, Salviucci: puntata 117

Ip. 199-202], pl. 79. [Publication date according to SALVADORI, 1888].

Daunin, F. M. 1801. — La raine à verrues, Hyla verrucosa. In: C. S. SONNINI, XIV® genre. Raine,

Hyla. In: C. S. SONNINI & P. A. LATREILLE, Histoire naturelle des reptiles, tome 2, Paris,

Deterville: 186. [Publication date and authorship according to HARPER, 1940].

GRAF, J.-D. & PoLs PeLaz, M, 1989. — Evolutionary genetics of the Rana esculenta complex. In:

R. M. DawWLey & J. P. BOGART (ed.), Evolution and ecology of unisexual vertebrates, Albany,

The New York State Museum: 289-302.

GÜNTHER, R., 1985. — Ordnung Anura, Froschlurche. Jn: W.-E. ENGELMANN, J. FRITZSCHE, R.

Günraer & F. J. Onsr, Lurche und Kriechtiere Europas, Leipzig & Radebeul, Neumann:

113-184.

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

Grassé & M. DELsoL (ed.), Traité de zoologie, tome 14, Batraciens, fasc. 1 B, Paris, Masson:

594-797.

(4) For a volume of a series:

BOcqQuET, C., GÉNERMONT, J. & LAMOTTE, M., (ed.), 1977. — Les problèmes de l'espèce dans le règne

animal. Tome 2. Mém. Soc. zool. Fr., 39: 1-381.

DUCROTAY DE BLAINVILLE, H. M., 1822. — De l'organisation des animaux, ou principes d'anatomie

comparée. Tome premier, contenant la morphologie et l'aistésologie. Paris, Levrault: [i-üi] + i-lix

+ 1-574, 10 tab.

Dumérir, A.-M.-C., BIBRON, G. & DUMÉRIL, A., 1854a. — Erpétologie générale ou histoire naturelle

complète des reptiles. Tome 7 (1). Paris, Roret: i-vii + [i-iv] + i-xvi + 1-780.

. — Erpétologie générale ou histoire naturelle complète des reptiles. Tome 7 (2). Paris, Roret:

+ 781-1536.

Lis 1854c. — Erpétologie générale ou histoire naturelle complète des reptiles. Tome 9. Paris, Roret: i-xx

+ 1-440.

GorHam, S. W., 1966. — Liste der rezenten Amphibien und Reptilien. Ascaphidae, Leiopelmatidea

[sic], Pipidae, Discoglossidae, Pelobatidae, Leptodactylidae, Rhinophrynidae. Das Tierreich, 85:

i-xvi + 1-222.

HOFFMANN, C. K., 1878. — Klassen und Ordnungen der Amphibien wissenschaftlich dargestelidt in

Wort und Bild. In: H. G. BRONN, Die Klassen und Ordnungen des Thier-Reichs wissenschaftlich

dargestelldt in Wort und Bild, Leipzig & Heidelberg, Winter, 6 (2): 1-726, pl. 1-42.

LINNAEUS, C. A, 1767. — Systema naturae per regna tria naturae, secundum classes, ordines, genera,

species, cum characteribus, differentiis, synonymis, locis. Editio duodecima, reformata. Tomus 1,

pars 2. Holmiae, Laurentii Salvii: 533-1327 + [i-xxxvii].

Source : MNHN, Paris

192 ALYTES 14 (4)

(5) For a report or a thesis:

ANONYMOUS, 1987. — Essais des eaux. Détection en milieu aquatique de la génotoxicité d'une substance

vis-d-vis de larves de batraciens (Pleurodeles waltl er Ambystoma mexicanum). Essai des

micronoyaux. Fascicule de Documentation AFNOR T90-325, Paris, Association Française de

Normalisation: 1-12.

LyncH, J. D., 1969a. — Program of the final public examination for the degree of Doctor of Philosophy.

Lawrence, The University of Kansas: 1-4.

is 19696. — Evolutionary relationships and osteology of the frog family Leptodactylidae. PhD Thesis,

Lawrence, The University of Kansas: 1-800.

OHLER, À. & PIERRON, E., 1991. — Le peuplement de batraciens du glacis du Fort de Noisy (Bufo

calamita, Triturus vulgaris, Triturus helveticus): composition, effectifs, activités. Rapport

préliminaire. Etampes, AALRAM: 1-11.

(6) For a meeting abstract:

Dunoïs, A., 1988. — Taxa should have names. Program and abstracts, Combined Meetings of the

Herpetologists” League, American Elasmobranch Society, Early Life History Section, AFS,

SSAR, ASIH, celebrating the 75th anniversary of Copeia, The University of Michigan, Ann

Arbor, Michigan, USA, 24-29 June 1988: 86.

_ 1990. — Declining amphibian populations — a global phenomenon? Some preliminary comments.

Report distributed to participants of the workshop Declining amphibian populations, sponsored

by the Board on Biology of the National Research Council, Irvine, California, USA, 19-

20 February 1990: 1-7.

DETAILED FORMAT, CONVENTIONS AND ABBREVIATIONS

Only publications with a finite number of identified authors should be cited under

their names. Other publications, even if published under the name, acronym or

abbreviation of a committee, an institute or any other collective body, should be cited,

both in the text and in the references, as “ANONYMOUS” (see examples above and below).

Except when the contrary is expressly required by their language, authors’ surnames with

a separated prefix (e.g., “DE”, “LE”, VAN”) should be cited, both in the text and in the

references, with the prefix preceding the name (see example above).

Special care should be taken to ensure that all names and words in references are

correctly spelled in their original languages, including accents (e.g., 6”, “4”, “à”, 6”) and

other diacritic marks such as tilde (“n°”), diaeresis (“ü”’) or cedilla (“£”'). Titles of papers

and books should be quoted exactly, even when they contain mistakes: in such cases, the

mention “[sic]” may be added after the identified mistake (see example above).

Dates given for references should be the actual date of publication, i.e. of public

distribution, not of writing or printing, of the book or periodical. This is of particular

importance for publications containing new scientific names or nomenclatural acts. Actual

publication dates may be ascertained using various kinds of evidence, including specialized

publications, and this may be indicated at the end of the reference (see examples above and

below).

When several editions of a book have been published, the reference should indicate

which one was used by the author(s) of the manuscript. For books, reports and theses, city

of publication should precede the name of publisher and complete pagination should be

Source : MNHN, Paris

Duois 193

given, including possible additional unnumbered pages at the beginning or end of the

volume, which should be written in small Roman numbers and between square brackets:

e.g., “[i-xix]”. In-text figures and plates included in the book pagination should not be

mentioned. Plates having their own numbers should be cited using Arabic numbers (e.g.,

“pl. 1-88”), even when in fact they bear Roman numbers (e.g., “pl. I-LXXXVIIT”) in the

actual publication. Table 5 provides standard abbreviations that should be used in

references for terms like “fascicule”, “figure”, or “plate”. Please note that they are not

italicized, that they end with a dot, and that plural should not be indicated (e.g. by adding

“s” at their end or by doubling the abbreviation, e.g. “pp.”).

Titles of periodicals composed of one single word or two words linked by a hyphen

should be given in full: e.g., “Herpetologica”, “ Amphibia-Reptilia”. Abbreviations can be

used for titles of periodicals composed of several separated words. In order to avoid

having several distinct abbreviations for the same term (e.g., “herp.”, “herpet.” or

“herpetol.” for “herpetological”), a list of common abbreviations that should be used for

terms in titles of periodicals is provided in tab. 5. Please note that all are italicized and end

with a dot. These abbreviations should not be used for other terms than those given in tab.

5. Other, less usual, abbreviations, should be clear, self-evident and non-ambiguous; in

case of doubt or of possible confusion, words should not be abbreviated. Except those in

tab. 5, very short terms should be given in full: e.g., “Acta”, “New”. Names of cities

should not be abbreviated: e.g., “London”, “San Francisco”. In titles of periodicals

including several words, nouns (or their abbreviations) should start with a capital letter,

and adjectives (or their abbreviations) with a lower case letter, except for first word of title

(e.g., “Occ. Pap. Mus. nat. Hist. Univ. Kansas”). Number of volume should always be

given as a bold Arabic number (e.g., “88”), even when in fact it appears as a Roman

number (e.g. “LXXXVIII”) in the actual publication. For a given periodical, series,

volume, section, and number, should be referred to respectively in parentheses followed by

a comma, in bold followed by a space, in parentheses followed by a space, and in

parentheses followed by a colon: e.g., “(2), 14 (B) (3):”.

Works referred to in the Literature Cited should be either published or accepted for

publication in an identified journal: in the latter case, they should be cited as “in press”

both in the text and in the list of references. Papers either “in preparation” or “submitted”

(but not yet accepted) should not appear in the Literature Cited: the corresponding work

may be mentioned, if this is necessary, in the text as “unpublished” or “in preparation”.

These terms should not be abbreviated, nor should phrases like “personal communica-

tion”, “unpublished results/data”, or other undefined terms. Personal observations or

papers by other authors which have not yet appeared in print may be cited only when

written authorization from the communicator is submitted with the manuscript.

Source : MNHN, Paris

194

ALYTES 14 (4)

Tab. 5. - Standard abbreviations to be used in lists of references in Alytes. Most of these

abbreviations can be used both for a noun (starting with a capital letter) and for an

adjective (starting with a lower-case letter, except if first term of title). Besides,

most of them can correspond to slightly different words according to the language

of the publication and the word being in the masculine or feminine, in the singular

or plural: e.g., "Herp." can mean "Herpetology", "Herpétologie", "Herpetologia",

etc.; "herp" can mean "herpetological", “herpétologique”, “herpétologiques",

"herpetolôgico", "herpetolégica", "herpetolégicos", "herpetolôgicas", etc. In the

table below, only one spelling is given (usually the English noun) as an example.

Proper terms can easily be deduced from the context (language of the publication,

place in a series of abbreviations). Some of these abbreviations can be combined

with other terms to produce longer abbreviatio: eg, "Biochem" for

"Biochemistry", "Hydrobiol" for "Hydrobiology", "Naturwiss" for "Natur-

wissenschaft", "neotrop." for "neotropical", "southwest." for "southwestern".

Abbreviation Meaning Abbreviation Meaning

ap. apéndice(s) Bras. / bras. Brasil

app. appendix/ce(s) Brit. british

ed. editor(s) Bull. Bulletin

fasc. fascicule(s) Cr. Compte rendu

fig. figure(s) Cah. Cahier

lim. lämina(s) Cal. / cal. California

ns. new series Can. / can. Canada

P. page(s) caribb. caribbean

pl plate(s) Cat. Catalogue

réd. rédacteur(s) cent. central

ser. series Chem. / chem Chemistry

suppl. supplement(s) Gi. Ciencia

tab. table(s) Co. County

vol. volume(s) Col. Coleoptera

Coll. Collection

Abh. Abhandlung comp. comparative

Abst. Abstract Conserv. Conservation

Acad. / acad. Academy Contr. Contribution

Afr. / afr. Africa Crust. Crustacea

Akad. Akademie Cuad. Cuaderno

Am. / am. America Fe /cyt. Cytology

An. Anales Department

Anat. / anat. Anatomy De. / dev. Development

Anim. / anim. Animal Dip. Diptera

Ann. Annals Diss. Dissertation

An. Anzeiger Disch. / disch. Deutschland

appl. applied east. eastern

Ara. Arachnid Ecol. / ecol. Ecology

Arch. Archive Embr. / embr. Embryology

Arg. /arg. Argentina Endocr. / endocr. Endocrinology

Arth. Arthropode Ent. /'ent. Entomology

As. / as. Asia Entw,. Entwicklung

Asoc. Asociaciôn Env. / env. Environment

Ass. Association Esp. / esp. España

Aust. / aust. Australia Evol. / evol. Evolution

Behav. / behav. Behaviour exp. experimental

Beitr. Beitrag Fac. Faculté

Belg. / belg. Belgique Forsch. Forschung

Ber. Bericht Fortschr. Fortschritt

Biol. / biol. Biology Fr. fr. France

Bol. Boletin gen. general

Boll. Bolletino Genet. / genet. Genetics

Bot. / bot. Botany Geogr. / geogr. Geography

Source : MNHN, Paris

Dugois 195

Abbreviation Meaning Abbreviation Meaning

Geol. / geol. Geology occid. occidental

Ges. Gesellschaft Dis. Oiseau

Gesch. Geschichte Okol. / kol. Okologie

Giorn. Giornale ori. oriental

hebd. hebdomadaire Orn. / orn. Ornithology

Helm. / helm. Helminthology Orth. Orthoptera

Hem. Hemiptera Qst. /'ost. Osteology

Herp. / herp. Herpetology Ost. / st. Osterreich

Hist. / hist. History Pap. Paper

Histol. / histol. Histology Paras. / paras. Parasitology

Hym. Hymenoptera Path. / path. Pathology

Ich. /'ich. Ichtyology Per. Periodical

imp. imperial Phil. / phil. Philosophy

Ind. / ind. India philom. philomatique

Inf. Information Phys. / phys. Physics

Ins. Insecta Physiol. / physiol. Physiology

Inst. Institut Pol. Poland

int. international Pop. / pop. Population

I lit. Jtalia preuss. preussich

J. Journal Proc. Proceedings

Jap. / jap. Japan Prot. / prot. Protistology

Jb. Jahrbuch Prov. / prov. Province

kaïs. kaiserlich Psych. / psych. Psychology

kônigl. kôniglich Publ. Publication

Lab. Laboratory q. quarterly

Lep. Lepidoptera Qld. Queensland

Limn. / limn. Limnology r. royal

linn. linnaean Rec. Record

Mag. Magazine reg. regionale

Mai Malacology Rep. Report

Mamm. Mammalogy Repr. / repr. Reproduction

Math. / math. Mathematics Res. Research

Med. / med. Medicine Rev. Review

Meded. Mededelingen Riv. Rivista

Mem. Memoirs Sber. Sitzungsbericht

mens. mensuel Sci. / sci. Science

Micr. / micr. Microscopy Serv. Service

Midl. Midlands sin. sinica

misc. miscellaneous Soc. Society

Mitt. Mitteilungen south. southern

Moll. Mollusca spec. special

Mon. Monograph LA Storia

Monatsb. Monatsberichte Stud. Study

Monit. Monitore Symp. Symposium

Morph. / morph. Morphology Syst. / syst. Systematics

Mus. Museum Tech. / tech. Technique

Mpyol. / myol. Myology Technol. / technol. | Technology

Myr. Myriapoda theor. theoretical

nac. nacional Trans. Transactions

Nat. / nat. Nature Trav. Travaux

nain. national trop. tropical

Natur. Naturalist Univ. University

Naturf. Naturforscher Ver. Verein

Neth. Netherlands Verh. Verhandlung

north. northern west. western

Novit. Novitates Wiss. / wiss. Wissenschaft

Nutr. / nutr. Nutrition Z. Zeitschrift

occ. occasional Zool. / zool. Zoology

Source : MNHN, Paris

196 ALYTES 14 (4)

SUBMISSION AND REVIEW OF MANUSCRIPTS

Manuscripts should be submitted in triplicate to either the Chief Editor or the Deputy

Editor of Alytes. Please consult the last published issue of the journal for their names and

addresses, and the distribution of scientific fields within batrachology between them.

Submission of a manuscript to Alytes implies recognition of the present Instructions to

Authors and consequently of possible modifications of the format of the manuscript by the

Editors if these instructions are not followed. A strong envelope should be used to dispatch

the manuscript. All three copies of the manuscript should be complete, including tables

and figures. For each figure, the original and two good copies should be provided, except

for photographs: photocopies of photographs are not acceptable and three original prints

should be provided.

Address for correspondence should clearly be indicated, particularly in the case of

multiple authorship.

Each manuscript will be assigned by the Editors to an appropriate Corresponding

Editor (CE), chosen among the members of the Editorial Board of Alytes. The work of

the CE will include submission of the manuscript to at least two referees, correspondence

with the referees and then with the author(s), and final acceptance (possibly after slight or

important modifications) or rejection of the paper. Manuscripts will be judged on the basis

of their scientific merit (methodological rigour, originality, novelty and importance of

results, quality of discussion) and of the quality of their writing and illustration. As

emphasized by DuBois & GOLLMANN (1993), the Editors of Alytes view their work as a

positive, constructive one: rather than automatically reject all papers which are not

“perfect” at once, they are ready to work, in good collaboration with the authors, to

improve the standards of papers, especially when the basic data and results are interesting

and new. On the other hand, they recognize that the author(s) of a paper is/are the

person(s) who sign(s) it, not the reviewers or the Editors. Suggested modifications

transmitted by the CE to author(s) are therefore of two kinds, similar to the categories of

objective and subjective criticisms defined by Bour & DuBois (1994): they include

compulsory changes, which must be followed if the paper is to be accepted for publication

in Alytes, and suggestions for improvement of the paper, which the author(s) is/are free

to follow or not.

FINAL MANUSCRIPT AND PROOFS

After acceptance, a paper copy of the final manuscript should be sent to the Chief

Editor. The author(s) should keep one exact copy of this final manuscript, as it will not

be returned with the page proofs. If possible, a copy of the final manuscript on a floppy

disk (3 7 or 5 4) should also be sent to the Chief Editor with the paper copy: this will

save much time and avoid misprints in the proofs. We welcome the following applications

for text processing: (1) preferably, MS Word (1.1 to 6.0, DOS or Windows), WordPerfect

Source : MNHN, Paris

Dusois 197

(4.1 to 5.1, DOS or Windows) or WordStar (3.3 to 7.0); (2) less preferably, formated DOS

(ASCII) or DOS-formated MS Word for the Macintosh (on a 3 2 high density 1.44 MB

floppy disk only).

Page proofs will be sent to the author(s) for correction. Jt is the responsibility of the

author(s) to ensure that the corrected proofs contain no error. Particular attention should

be paid to checking titles, scientific names, symbols, figures and tables. Corrected proofs

should be returned to the Chief Editor as soon as possible to prevent a delay in

publication. Revisions should not be made in proofs. Changes, relative to the final

manuscript, made in proofs (in text, figures or tables), will be charged to the author(s).

Manuscripts, including illustrations, will be discarded by the Chief Editor one month

after publication, unless specific instructions were received from the author(s) for their

return.

Payment of printing costs will be requested only from authors who have access to

institutional funds for this purpose. However, publication of colour photographs will

always have to be charged to the authors.

For each published paper, 25 free reprints will be offered by ISSCA to the author(s).

Additional reprints may be ordered, using the form provided, at the time corrected page

proofs are returned to the Chief Editor; subsequent orders will not be possible.

AUTHOR CHECKLIST

The present Instructions to Authors are meant to help authors to prepare manuscripts

for Alytes, and Editors and referees to review them. They should ensure that manuscripts

received are in the good format, make for more rapid processing, acceptance and

publication of papers, and contribute to a final high quality journal. For additional help,

tab. 6 presents an Author Checklist which summarizes 50 major points that should be

checked by author(s) before submitting a manuscript to Alytes. It is recommended to make

a photocopy of this checklist, to mark it with crosses and to send it with the manuscript

to the Editors. The same should be done when dispatching the final manuscript to the

Chief Editor at the end of the review process. AI items of the checklist should be marked

with a cross in the column “Yes” after checking, except those which should be marked as

“Irrelevant” for a given manuscript: e.g., items (26)-(29) are not relevant for a paper

including no description of a new taxon, or items (40)-(46) are not relevant for a paper

including no figure.

ACKNOWLEDGEMENTS

The editorial rules presented above are the result of a continuous evolution through a fifteen-year

experience (1982-1996) of publication of Alyres, but no doubt they will keep on improving in the

future: constructive suggestions in this respect will be welcome. Establishment of these standards has

greatly benefited from the work of all the colleagues who served as Editors and Corresponding

Source : MNHN, Paris

198 ALYTES 14 (4)

Tab. 6. - Author checklist: major points that should be checked by any author(s) before

submitting a manuscript to Alytes. Irr.: irrelevant.

Item

(1) The manuscript submitted deals with amphibians as such and brings original

information concerning these animals.

C2) This manuseript has not been published previously and is not under

consideration for publication elsewhere.

(3) Observation or experimental intervention on animals, their capture or killing,

were strictly justified for the purpose of advancing scientific knowledge and

were made according to ethics of respect for all living beings.

(à) Information on collecting permits and licenses for the specimens used in the

study is provided in the manuscript.

(5) All pages of the manuscript are numbered, dated, and marked with the (first)

author's name.

(6) All authors have signed letter accompanying manuscript.

(7) Written authorization from communicators for citing personal observations or

unpublished manuscripts is provided.

(8) Title page includes title, authorship, address(es) of author(s), address for

correspondence, number of pages, of plates and of figures.

(9) The title follows the format of Alytes.

(10) The manuscript starts with an English abstract (a single paragraph) and ends

with a summary in one or several other languages, including either French or

Spanish [please circle language chosen].

(11) If the paper is written in English, consistent use of either the British or the

American standard [please circle standard chosen] has been checked throughout

the manuscript

(12) The text of the manuscript is not justified and words are not divided at the

right margin.

(13) Sections and subsections of the manuscript follow the format of 4/ytes.

(14) References in text and in list of references have been checked to correspond

exactly.

(15) Al references cited have been checked with the original publications; in case

some have not been seen by author(s), this is stated in the list of references.

(16) Dates given for references, especially those of nomenclatural importance, are

their actual publication dates.

(17) Presentation of references in text and in Literature Cited follows the format of

Alytes.

(18) In text, tables, figure legends and list of references, all names and words are

correctly spelled in their original languages, including accents and other diacritic

marks

(19) Abbreviations, symbols, measurement units, numerals, use of capitals and

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(20) Abbreviations used in the text, in particular those for Museum collections and

for measurements, are defined in a special section or table ofthe manuscript.

(21) Signs, symbols or characters that cannot be typed with the printing equipment

used by author(s) have been added by hand on every occurrence in cach copy of

the manuscript.

(2) The zoological nomenclature used in the paper strictly follows the

International Code of Zoological Nomenclature

Source : MNHN, Paris

Dusois

Item

(23) All scientific names of taxa are given either followed by their author name and

their date, or without both.

(24) Use of parentheses for inclusion of authors names and dates of names of taxa

is made according to the Code.

(25) AI scientific names of taxa, including new combinations, are written in full at

least once, on the occasion of their first occurrence in the paper.

(26) If the paper proposes the description of new taxa, it provides clear evidence

that a thorough investigation of related taxa has been carried out.

(27) New scientific names have been thoroughly checked not to be preoccupied.

(28) The name of any new taxon described in the paper does not appear, either in

the text, or in tables and figure legends, before the heading of the section

devoted to the description of the taxon.

(29) Holotypes of new species-group taxa have been deposited in a permanent and

intenationally recognized scientific institution.

(30) The exact location and catalogue numbers of specimens mentioned in the

paper are given.

(31) Taxonomic accounts follow the format of Alytes.

(32) Synonymies provided in taxonomic accounts follow the format of Ares.

(33) Mention in the paper of type species of genera is always accompanied by

paper ol sofg

precise information on their mode of designation.

(34) Geographic information is provided following the format of Alytes.

(35) Each table is presented on a separate sheet, with legend above.

(36) If the manuscript was prepared in text processing, columns in tables are

separated by tabulations, not by simple spaces.

(37) Figures and tables are numbered in Arabic numerals in the order of their first

mention in text.

(38) AI abbreviations in tables and figures are defined in their legends or elsewhere

in the manuscript.

(39) Legends of tables and figures are intelligible without reference to text.

(40) Legends of figures showing specimens or parts of specimens include

information about the actual specimens shown.

(41) Subfigures of a composite figure mounted as a plate are lettered in lower

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(42) Letters or numbers used in figures are printed or transferred, not handwritten.

(43) Size of original figures is conform to the requirements of 4/ytes.

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bars, not in legends.

(46) If colour photographs are submitted, it is understood by author(s) that their

colour publication will be at his/her/their cost.

(47) Name of (first) author, date of submission and number of figure, plate or table

appear on top, bottom or back of all figures, plates and tables.

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are sent to A/ytes, one exact copy kept by author(s).

(50) The manuscript is dispatched in a strong envelope.

Source : MNHN, Paris

200 ALYTES 14 (4)

Editors of the journal since its creation. For their comments on earlier drafts of this paper, I am much

indebted to Philippe BOUCHET (Paris, France), Janalee P. CALDWELL (Norman, USA), Thierry DEUVE

(Paris, France), Pierre Dumois (Souvigny, France), Serge Goras (Paris, France), Günter GOLLMANN

(Wien, Austria), W. Ronald Heyer (Washington, USA), Jean-Lou JUSTINE (Paris, France) and Jon

LOMAN (Lund, Sweden). I am grateful to Roger BOUR and Annemarie OHLER (Paris, France) for their

help in the preparation of the tables of this paper.

LITERATURE CITED

ANONYMOUS, 1985. — /nternational code of zoological nomenclature. 3rd edition. London, Interna-

tional Trust for zoological Nomenclature: i-xx + 1-338.

= 1993, — Lexique des règles typographiques en usage à l'Imprimerie Nationale. Paris, Imprimerie

Nationale: 1-197.

= 1994, — Statutes of ISSCA. Circalyres, 6 (6): 62-71.

Bour, R. & DuBois, A., 1994. — Dumerilia: présentation d’un nouveau journal herpétologique.

Dumerilia, 1: 1-4.

Dusois, A., 1978. — Les principaux stades de développement significatifs en écologie et en génétique

des populations des amphibiens anoures. Terre & Vie, 32 (3): 453-459.

ee 1995. — Keratodont formulae in anuran tadpoles: proposals for a standardization. J. zool. Syst.

Evol. Res., 32 (4): 297-318; 33 (1): I-XV.

Duois, A. & GOLLMANN, G., 1993. — Report of the Editors of Alytes for 1993. Circalytes, 6 (4):

35-36.

GoLLMANN, G., 1992. — The relationship of batrachology to herpetology. Copeia, 1992 (3): 887.

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

identification. Herpetologica, 16 (3): 183-190.

HuTH, E. J., BROGAN, M., DANCIK, B. P., KOMMEDAHL, T., NADZIEIKA, D. E., ROBINSON, P. &

Swanson, W., 1994. — Scientific style and format. 6th edition. Cambridge, Cambridge Univ.

Press, Council of Biology Editors: i-xv + 1-825.

Liu, C.-C., 1950. — Amphibians of western China. Fieldiana: Zool. Mem., 2: 1-400, pl. 1-10.

Myers, C. W. & DUELLMAN, W. E., 1982. — A new species of Hyla from Cerro Colorado, and other

tree frog records and geographical notes from western Panama. 4m. Mus. Novit., 2752: 1-32.

NG, P. K. L. 1994. — The citation of species names and the role of the author’s name. Raffles Bull.

Zool., 42 (3): 509-513.

ProcrER, P., (ed.), 1995. — Cambridge international dictionary of English. Cambridge, Cambridge

Univ. Press: i-xviii + 1-1774.

QuiRk, R., GREENBAUM, S., LEECH, G. & SVARTVIK, J., 1980. — À grammar of contemporary English.

9th edition. Harlow, Longman: i-xii + 1-1120.

Suit, H. M. & Smirn, R. B, 1973. — Chresonymy ex synonymy. Sys. Zool., “1972”, 21 (4): 445.

[Publication date according to front cover of issue].

Summers, D., (ed.), 1995. — Longman dictionary of contemporary English. 3rd edition. Harlow,

Longman: i-xxii + 1-1668 + B1-B22.

THowpson, D. (ed.), 1995. — The concise Oxford dictionary of current English. 9th edition. Oxford,

Clarendon Press: i-xxi + 1-1673.

Paris, 4 December 1996

© ISSCA 1997

Source : MNHN, Paris

AINTTES

International Journal of Batrachology

published by ISSCA

EDITORIAL BOARD FOR 1996

Chief Editor: Alain Dusois (Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire

naturelle, 25 rue Cuvier, 7005 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 HEyER (Washington,

U.S.A.); Walter HôDL (Wien, Austria); Pierre JoLy (Lyon, France); Masafumi Marsu1 (Kyoto,

Japan); Jaime E. Péraur (Mérida, Venezuela); J. Dale RomerTs (Perth, Australia); Ulrich SINSCH

(Koblenz, Germany); Marvalee H. Wake (Berkeley, U.S.A.).

Technical Editorial Team (Paris, France): Alain Dunois (texts); Roger Bour (tables); Annemarie

Ouer (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 tab. 4. Figures should not

exceed 16 x 24 cm. The size of the lettering should ensure its legibility after reduction. The legends of figures

and tables should be assembled on a separate sheet. Each figure should be numbered using a pencil.

References in the text are to be written in capital letters (BOURRET, 1942; GRAF & POLLS PELAZ, 1989;

INGER et al., 1974). References in the literature cited section should be presented as follows:

BOURRET, R., 1942. — Les Batraciens de l'Indochine. Hanoï, Institut Océanographique de l'Indochine:

+°1-547, pl. 14.

GR4r, J.-D. & POLLS PELAZ, M., 1989. — Evolutionary genetics of the Rana esculenta complex. In: R. M.

DawLey & J. P. BOGART (eds.), Evolution and ecology of unisexual vertebrates, Albany, The New York

State Museum: 289-302.

INGER, R. F., Voris, H. K. & Voris, H. H., 1974. — Genetic variation and population ecology of some

Southeast Asian frogs of the genera Bufo and Rana. Biochem. Genet., 12: 121-145.

Manuscripts should be submitted in triplicate either to Alain DuBois (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

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

referces.

If possible, after acceptance, a copy of the final manuscript on a floppy disk (3 % or 5 %4) should

be sent to the Chief Editor. We welcome the following formats of text processing: (1) preferably, MS Word

(1 to 6.0, DOS or Windows), WordPerfect (4.1 to 5.1, DOS or Windows) or WordStar (3.3 to 7.0); (2) less

preferably, formated DOS (ASCII) or DOS-formated MS Word for the Macintosh (on a 3 % high density

1.44 Mo floppy disk only).

No page charges are requested from the author(s), but the publication of color photographs is

charged. For each published paper, 25 free reprints are offered by Alytes to the author(s). Additional reprints

may be purchased.

-x

Published with the support of AALRAM

(Association des Amis du Laboratoire des Reptiles et Amphibiens

du Muséum national d'Histoire naturelle, Paris, France).

Directeur de la Publication: Alain DuBois.

Numéro de Commission Paritaire: 64851.

SRCR D Source : MNHN, Paris

Alytes, 1997, 14 (4): 129-200.

Contents

Alain DuBois

Editorial : 15 years of Alytes 129

Miguel VENCES, Frank GLAW & Franco ANDREONE

Description of two new frogs of the genus Mantidactylus

from Madagascar, with notes on Mantidactylus klemmeri

(Guibé, 1974) and Mantidactylus webbi (Grandison, 1953)

(AMD RANCE MAntElnAe) EE 130-146

James R. MCCRANIE & Larry David WiLsON

A review of the Eleutherodactylus milesi-like frogs

(Anura, Leptodactylidae) from Honduras with the

(dESCTIDHONIOMÉOUT NEWASPECIES +... serrer e 147-174

Alain Dupois

Instructions to authors of papers submitted to Alytes .............. 175-200

Alytes is printed on acid-free paper.

Alytes is indexed in Biosis, Cambridge Scientific Abstracts, Current Awareness in Biological

Sciences, Pascal, Referativny Zhurnal and The Zoological Record.

Imprimerie F. Paillart, Abbeville, France.

Dépôt légal: 1° trimestre 1997.

© ISSCA 1997

Source : MNHIN, Paris