ISSCA

International Society for the Study

and Conservation of Amphibians

(International Society of Batrachology)

SEAT

Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire naturelle, 25 rue Cuvier, 75005

Paris, France. — Tel.: (33).(0)1.40.79.34.87. — Fax: (33).(0)1.40.79.34.88. — E-mail: dubois@mnhn.fr.

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Councillors: C. Kenneth Dopp, Jr. (Gainesville, USA); Britta GriLurrscH (Wien, Austria); Stéphane

GROSIEAN (Paris, France) ; Julio Mario Hoyos (Bogotä, Colombia); Thierry LODÉ (Angers,

France): Jon LOMAN (Lund, Sweden); Alain PAGANO (Angers, France).

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jource : | Paris

Bibliothèque Centrale Muséum

LUN

3 3001 00189260 2

AIVTES

NTERNATIONAL JOURNAL OF BATRACHOLOGY

October 2003 Volume 21, N° 1-2

Alytes, 2003, 21 (1-2): 1-2. Editorial

Should internet sites be mentioned

in the bibliographies

of scientific publications?

Alain DUBOIS

Vertébrés: Reptiles & Amphibiens,

USM 0602 Taxonomie & Collections,

Département de Systématique & Evolution,

+ Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

<dubois@mnhn.fr>

In the 19 century, when a long-gone optimistic attitude towards the future of our civilization was prevailing,

science was seen as a permanent process of increase of our positive, rational knowledge, not only in the aim of being able

to act more efliciently, ie. to improve our technical mastership, but also in the hope of getting a more comprehensive

basic understanding of all aspects of the world we live in, by pure intellectual curiosity. Ât that time, it was considered

of utmost importance to maintain a permanent link between all historical periods of development of science: any new

finding, any new theory, was placed in this historical context and was viewed as the addition of a new stone to an edifice

to the building of which all scientists of the past had contributed. Times have changed, and many scientists have now

adopted a much more limited scope in their activity, either by being only interested in the development of scientific

knowledge having predictable, and often immediate, practical applications, or by feeling only concerned with recent

findings and theories, without including them in a more general apprehension of the evolution of scientific knowledge.

The fact that, quite recently, a paper using (in a specific context, that of zoological nomenclature) a formula like “tyranny

of the past” to qualify this permanent link between current and previous science, has been accepted for publication by

different scientific journals (SAVAGE, 1990a-b, 1991), is an interesting illustration of this trend.

The laboratory in the Paris Museum where I have been working for thirty years is on the other side of a street

bordered by two major French Universities, and in the last decades 1 have not unfrequently found, in the dustbins of the

latter, piles of journals, especially those, like Nature or Science, which, being published weekly, represent a large volume

of paper each year: questioned about this, researchers or librarians of these Universities would reply that their

laboratories or libraries are 100 small and lack space to store important amounts of scientific literature.

furthermore, after à certain time has elapsed, these publications have become “obsolete” and are no more

ing research: for this latter reason, they did not even think useful to query, before throwing these publications away,

if other scientific laboratories or libraries would be interested in recovering them. Clearly, for a number of scientists

nowad. tific publications have become a short-term-use product, like many other products in our society, That this

tude hasstrong influences on the course of scientific research itself is hardly to de demonstrated: never before has scien-

search more slavishly followed fashions, with some dominant ideas, techniques, methods or research subjects being

supported orexplored by many researchers and teams duringa decade or two, and then completely abandoned when other

more recent ideas or subjects supplant them in the fund-raising systems of science, and, by waÿ of consequence, in the

minds of many.

Such a way of functioning of science may be very efMicient whenever science is viewed merely as a way to produce

more ellicient or cheaper pharmaceuticals, pesticides, cosmetics, Computers, cars, satellites or weapons, Le. as à basic

Support to technical improvement, but it may not be so if science is also understood as aimed at a better understanding

of the world we live in, irrespective of any aim at acting upon it or at modifying it. If science is to be a progressive

construction of a complex corpus of knowledge, ie. of facts and theories about these facts, it is highly questionable

Source : MNHN, Paris

ne)

ALYTES 21 (1-2)

whether any scientific publication ever becomes “obsolete”: it is part of a complex edifice, and suppressing or ignoring.

it may be like removing a stone at the basis of a wall. Stating that current science can be understood and mastered

thout connection with the past of science is similar to saying that the architecture, painting, music or literature of the

past are now “obsolete” and should be forgotten or destroyed 10 replace them by the works of our time.

In some scientific fields, the progress of research is so quick, and the competition between researchers, research

teams or even countries so strong, that many new results or theories, at the time when they are published, are already

“obsolete”, in the sense that they are already known of several other researchers and teams. In such research fields, it is

striking that the bibliography of a paper often contains many references 10 works quoted as “personal communication”,

“unpublished data”, “work in progress”, “in preparation” or “in press”. Of course, this may cause problems later if

subsequent authors Wish 10 trace these “phantom publications”, which not rarely happen not to have ever been actually

published after having been quoted (see Duois, 1999): hence the recommendation, which is even an editorial rule in

some periodicals, not to mention such “references” in a bibliography. The same problem applies to the growing practice

consisting in quoting internet sites as bibliographic references in scientific papers or books. In fact, such references are of

the same nature as a ‘personal communication”: in the future, they won”t be available to readers of the publication where

they are quoted. An internet it has no permanency, as can be modified, “updated”; "correted” or suppresed at any

moment. No long-term trace is usually kept of what was available on a site at a given date, and even if such a memory is

Kept privately by the owner or editor of a site, it is not directly available to customers.

The function of a list of “references cited” at the end of a scientific paper is double: (1) to provide the sources of

information used by an author to support some of the scientific statements considered in the paper as valid or discussed

in the paper: (2) to allow any reader of the paper to go back personally to these sources and 10 study them exactly as they

were when they were quoted. In order for a bibliography to be useful all references that appear there should be accessible

Lo any reader, and should remain so in the future, whatever the fate of the authors of the works or of the paper where

these references appear, This condition does not apply to works quoted as “personal communication” or “in press"

except if the precise reference of the future publication can be provided with certainty), and similarly does not apply to

internet sites. The problem here does not come from internet sites being on a support other than paper, but on their

having no permanency. Some non-paper publication systems, such as CD-Roms, audio or video cassettes, can be as

permanent and non-modifiable as paper publications, and qualify for “publications” that can be quoted in a bibliogra-

phy: for this reason, such works can be considered as publications for the purposes of the International Code of =ological

Nomenclature (ANONYMOUS, 1999), which is not the case of internet sites.

No one knows what will be the long-term fate of our society, of the activity we call science and of the corpus of

results and theories produced by this activity. However, as soon as a scientific perodical places itself within the frame of

long-term science, “as if science was still to exist for many decades and centuries”, it should care for publishing only

bibliographie information that will be available for readers in the long-term future, For this reason, the periodical A/ytes.

does not accept the mention of internet sites among the references listed in the Lirerature cired section of a paper. IF

absolutely necessary for the understanding of the text, or to provide some information that would not be available

otherwise, exceptional mention of an internet site in the corpus of the text may be acceptable, just like in some cases it

is acceptable to mention a “personal communication” or “unpublished data”. But this reference won”t be repeated in the

bibliography. In most cases, information that may roday be easier to find on an internet site may also be available in

aper-published works: although it may be à little more time-consuming for an author to trace such a published

information, mention of such a published reference will, in the long run, be much more useful to the future readers of the

paper and is worth the effort to find it.

For the same reason, ie. the need of a long-term accessibility of all the information mentioned in the papers it

publishes, the journal A/ytes does not put a priori limitations on the length of papers or of tables of data and results of

ntific works submitted to the journal: *Alyres 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.” (DUBOIS, 1997: 188). This also means that A/yres does not encourage authors

to state that additional detailed data are available at their address and can be obtained by writing directly to them, or are

available online (e.g.. as “supporting online material"): what will be the fate of such pieces of information in 50 or 100

years? Either this information is useful for the understanding of the paper and evaluation of its merits, and then it should

be included in the paper (even as a long table or appendix), or it is not, and then it should not be mentioned at all.

LITERATURE CITED

ANONYMOUS [International Commission on Zoological Nomenclature], 1999. — /nternational code of =oological nomen-

clature. Fourth edition. London, International Trust for zoological Nomenclature: i-xix + 1-306.

Dumois, A.. 1997. — Instructions to authors of papers submitted to Ares. Alytes, 14 (4): 175-200.

1999. - Miscellanea nomenclatorica batrachologica. 19. Notes on the nomenclature of Ranidae and related groups.

Alvtes, A7 (1-2): 81-100.

VAGE, J. M. 1990a. - Meetings of the International Commission on Zoological Nomenclature. Sysr. Zool.. 39 (4):

424-425.

1990, — ICZN meetings. Copeia, 1990 (4): 1205-1208.

1991. — Meetings of the International Commission on Zoolog

16-118.

cal Nomenclature. Amphibia-Reptilia, V2 (1)

Source : MNHN, Paris

Alytes, 2003, 21 (1-2): 3

ntaddress: US Geolog

FL 32653, USA; <steve_johnson@usg:

22.

Orientation and migration distances

of a pond-breeding salamander

(Notophthalmus perstriatus,

Salamandridae)

Steve A. JOHNSON!

Department of Wildlife Ecology and Conservation,

IFAS, University of Florida,

Box 110430, Gainesville, Florida 32611, USA

Habitat loss and modification have played a significant role in the

decline of amphibian populations and species. Loss of wetlands, which are

used as breeding sites for many amphibians, has contributed to the decline.

The protection of small, isolated wetlands and core areas of associated

uplands is one way in which population declines in certain species can be

slowed or prevented. Nevertheless, migration distances of individuals of

most amphibian species from their breeding sites are unknown. Using drift

fences and pitfall traps, 1 studied migration distance and orientation of

striped newts (Notophthalmus perstriatus) at a breeding pond in northern

Florida, USA. Newts entered (immigration) and exited (emigration) the pond

basin in a nonrandom fashion but no obvi.

apparent. Patterns of emigration and immigration differed significantly

between sexes, life-history stages, and migration events. Individuals tended

to exit and enter the pond basin within the same quadrant, sometimes

leaving and returning at the same point. Newts moved hundreds of meters

into the sandhill uplands surrounding the pond. 1 found an inverse rela-

tionship between the proportion of newts migrating and distance from the

pond. Nonetheless, 1 estimated that at least 16 % of individuals breeding at

the pond migrated in excess of 500 m from the pond. Thus, a core of

protected upland with a radius of approximately 800 m from the pond

would be needed to preserve the area used by the vast majority of

individuals that breed at the pond. These data underscore the need to study

upland habitat requirements for amphibians; findings for one taxon (e.g.

ambystomatids) may not be applicable to others (e.g., salamandrids)

Without such data, designating terrestrial core habitat to conserve aquatic-

breeding amphibians will be difficult or impossible. However, without better

protection of small, isolated wetlands, arguments to preserve surrounding

uplands are irrelevant.

INTRODUCTION

al Survey, Florida Integrated Science Ceni

7920 NW 71° Strect, Ga

During the past two decades, amphibian declines have received considerable attention

(BARINAGA, 1990; Wake et al., 1991; WAKkE, 1991; ALFORD & RICHARDS, 1999; HoUL.

AHAN et

ainesville,

Source : MNHN, Paris

4 ALYTES 21 (1-2)

al., 2000). Although pathogens have been implicated in several die-off events (BERGER et al.,

1998; Lips, 1998, 1999), there is a consensus among herpetologists that the global decline is a

result of multiple factors (ALFORD & RICHARDS, 1999). Habitat modification and destruction

have been identified as significant factors contributing to the global decline (DoDD, 1997;

ALFORD & RICHARDS, 1999; DUELLMAN, 1999; SemLirsCH, 2000). Although they do not

attract the media attention that mass mortality or deformed amphibians receive, habitat

modification and loss are insidious processes that must be addressed if amphibians are to

persist. The effects of habitat changes on amphibian populations are of particular concern in

areas that are characterized by a high density of small, isolated wetlands (DeLis et al., 1996:

HECNAR & M’CLosKkEY, 1996; KNUTSON et al., 1999; BABBITT & TANNER, 2000; SEMLITSCH,

2000; SxopGrass et al., 2000; RussELL et al., 2002). In these areas (e.g., the Southeastern

Coastal Plain of North America), amphibian diversity is high (DUELLMAN & SWEET, 1999)

and many species rely solely on small, isolated wetlands as breeding sites (DopD, 1997;

SEMLITSCH & BODIE, 1998; BABBITT & TANNER, 2000).

Despite their size (ï.e., less than a few hectares), small, isolated wetlands are of tremen-

dous biological importance, particularly for amphibians. They play a vital role in amphibian

metapopulation dynamics and therefore are essential in maintaining viable populations of

amphibians at a landscape scale (SEMLITSCH & BODIE, 1998; SEMLITSCH, 2000; SNODGRASS et

al., 2000). In addition to amphibians, numerous other vertebrates and a suite of invertebrate

species depend on small, isolated wetlands (BROWN et al., 1990; MorErR & FRANZ, 1988;

BURKE & GIBBONS, 1995; HART & NEWMAN, 1995; SemLITsCH & BODIE, 1998; RUSSELL et al.,

2002).

Preserving a wetland alone may not result in protection of many of the organisms that

depend upon the wetland. Many amphibians have complex life cycles in which they require

ponds to breed but spend the majority of their lives in surrounding upland habitats (DODD,

1997; Dopp & CADE, 1998; SemLITsCH, 1998; SEMLITSCH & JENSEN, 2001). If sufficient upland

habitat surrounding isolated breeding-ponds is not preserved, amphibians with complex life

cycles are not likely to persist at a local scale. Therefore, at some point the loss of uplands may

lead to extirpation of some amphibian populations because of disruption of metapopulation

dynamics (SEMLITSCH & BODIE, 1998; SEMLITSCH, 2000; MARSH & TRENHAM, 2001), even when

the ponds themselves are preserved.

One strategy to curtail the loss of amphibians associated with habitat alteration around

small, isolated wetlands is to preserve “core habitat” and “buffer zones” consisting of

protected uplands surrounding the wetlands (SEMLITSCH & JENSEN, 2001). These zones

provide habitat for retreats and foraging for those species with complex life cycles, many of

which are now considered common. Without preservation of appropriate upland habitat,

even common species will decline.

Little is known, however, about the extent of upland “core habitat” required by pond-

breeding amphibians. DopD (1996) summarized the literature on upland movements of

amphibians in North America and found that this distances amphibians migrate from

breeding sites are poorly known. From this summary and a review by SEMLITSCH (1998) on

migration distances of ambystomatid salamanders, it is apparent that many amphibians move

considerable distances from breeding ponds. Unfortunately, migration distances are only

available for a few species and usually are based on a single or a few individuals. Clearly there

Source : MNHN, Paris

JOHNSON 5

is need for data on migration distances from breeding sites for most North American

amphibians. These data are essential to justify establishing adequate “core habitat” of

uplands around amphibian breeding ponds.

Tcollected data on orientation and migration distances for striped newts (Notophthalmus

perstriatus) at a breeding pond and in the surrounding uplands in north-central Florida.

Striped newts breed exclusively in small, isolated wetlands that lack fish. They have a complex

life cycle and individuals spend much of their lives in uplands surrounding breeding ponds

(CHRISTMAN & MEANS, 1992; DopD & LACLAIRE, 1995; JoHNsON, 2001, 2002; Dopp et al., in

press). Striped newts are restricted to xeric uplands (1.e., sandhill and scrub communities) and

are endemic to southern Georgia and northern Florida, USA (fig. 1). The species has declined

throughout its range (DopD & LACLAIRE, 1995; FRANZ & SmrrH, 1999) and its biological

Status is under review by the US Fish and Wildlife Service (L. LaClaire, pers. comm.). The

objectives of my study were (1) to determine orientation patterns of striped newts into and

away from a breeding pond, and (2) to determine migration distances of individuals into the

surrounding upland habitat.

MATERIALS AND METHODS

STUDY SITE

The study was conducted on the Katharine Ordway Preserve-Swisher Memorial Sanct-

uary, Putnam Co., Florida, USA (29°41°N, 82°00°W; fig. 1). EISENBERG & FRANZ (1995),

LACLAIRE (1995) and Dobp (1996) provided descriptions of the preserve and its habitats.

Data were collected from 7 October 1996 to 11 September 1998 at One Shot Pond (OSP). OSP

is a small, isolated pond with a variable hydroperiod (hydroperiod refers to the number of

days a pond holds water between periods when it is dry) and is located in xeric sandhill

uplands dominated by longleaf pine (Pinus palustris), turkey oak (Quercus laevis) and

wiregrass (Aristida beyrichiana). Small stands of planted slash pine (Pinus elliottii) are located

north and southwest of the pond basin (fig. 2). Several water bodies are located near OSP

(fig. 2). These water bodies are isolated from one another and only receive water from rainfall

and ground water seepage; their hydroperiods are dictated by fluctuations in the water table.

Fox Pond held water from 26 November 1997 until the end of the study, whereas OSP, Berry

Pond, Lake McCloud and the Anderson Cue Lakes held water throughout the entire study

period. During the study, striped newts were only present in OSP and Fox Pond. However,

only 32 newts (16 adults and 16 juveniles) were captured at Fox Pond (S. A. Johnson,

unpublished data). McCloud and the Anderson Cue lakes support predatory fishes, and

striped newts do not breed there. No striped newts were captured during periodic sampling

throughout the study period in Berry Pond. Because there were no other breeding ponds

within several kilometers of OSP, I assumed that striped newts caught in upland fences

around OSP originated from within OSP.

Source : MNHN, Paris

6 ALYTES 21 (1-2)

South .

Carolina

Atlantic

Ocean

Fig. 1. - Geographic range of striped newts, which are endemic to Georgia and Florida, USA. Note the

hiatus (?) between the western and eastern portions of the range. This area likely represents a true gap

in the species distribution, rather than an artifact of inadequate survey effort. The black dot (+)

shows the location of study area, Katharine Ordway Preserve, Putnam Co., north-central Florida,

USA.

ORIENTATION AT ONE SHOT POND

L'encireled OSP with a 190-m drift fence made of galvanized metal flashing that was

buried ca. 15 em below the ground, with ca. 35 cm extending above the ground. Thirty-eight

pitfall traps (19-1 plastic buckets) were buried flush with the ground. Pitfall traps were placed

Source : MNHN, Paris

JOHNSON 7

à] Sandhill Uplands

One Shot Pond

FR À

Che approximate locations of the 500 m drift fence sections (see fig. 3) are indicated. Dirt

roads appear as thin, white lines.

in pairs, one on each side of the fence, at intervals of about 10 m. I usually checked traps three

to five days per week, depending on weather and movements of animals. I weighed and

measured newts caught in pitfall traps at the pond and in the surrounding uplands. Each newt

was individually marked by toe clipping (DONNELLY et al., 1994) and released on the opposite

side of the fence. Sex of adults was determined by the presence of a conspicuous whitish gland

visible at the posterior edge of the vent in mature males. Recently transformed newts were

recognized by the presence of gill vestiges visible for several days after metamorphosis.

Recently transformed newts with swollen vents were presumed to be mature (JOHNSON, 2001),

and aquatic sampling in the pond showed that such individuals represent paedomorphic

Source : MNHN, Paris

8 ALYTES 21 (1-2)

animals that recently bred. These newts are referred to as paedomorphs. Transformed newts

without swollen vents (i.e., immatures) are referred to as efts.

I obtained a compass orientation for each pair of pitfall traps surrounding OSP. To do

this, I stood in the center of the pond and took a bearing on each pair of traps at the drift

fence. Following the methods of Dopb & CADE (1998), I used Rao’s spacing test (RAO, 1976;

BATSCHELET, 1981) to determine if captures were distributed uniformly around the drift fence

(i.e., random orientation). I analyzed orientation of newts into and away from the pond by sex

and life history stage. I made comparisons between distinct migration events (JOHNSON, 2001)

within the adult and eft life-history stages. For comparisons between sexes, life-history stages,

and migration events, I ran the same multirespsonse permutation procedure (MRPP; MIELKE

& BERRY, 2001) used by DopD & CADE (1998). Orientation analyses were performed with the

statistical software package BLOSSOM, which was developed by the US Geological Survey

(CADE & RICHARDS, 1999). BLOSSOM is available free at www.fort.usgs.gov./products/

software/software.asp.

UPLAND MIGRATION

Migration distances of newts in the sandhill uplands around OSP were determined

through captures in pitfall traps associated with drift fences. Drift fences were oriented to

capture newts during movements to and from the pond (fig. 3). In year one, five fence sections

were established at each of four distances from OSP (20 m, 40 m, 80 m and 160 m). Fence

sections at each distance totaled 20 % of the circumference at that distance from the pond.

Fence sections were distributed evenly at each distance, and they did not overlap with fence

sections at the other distances (fig. 3a). Fence sections at 20 m were 10.0 m long with 4 pitfalls

(2 on each side of the fence); at 40 m fence sections were 15.1 m with 6 pitfalls; at 80 m sections

were 25.1 m with 8 pitfalls; at 160 m sections were 45.2 m with 10 pitfalls. Pitfall traps were

installed on both sides of the upland fences (i.e., pond side and upland side: fig. 3a). This

upland fence array was monitored from 7 October 1996 to 5 December 1997, and fences were

constructed similarly to the fence at the pond.

Results from year one demonstrated that striped newts regularly moved more than

160 m. Therefore, a new upland fence array was installed in year two, with upland drift fences

erected much farther away from OSP. On 5 December 1997, the upland drift fences described

above were replaced with a different array of fence sections (fig. 3b) and the new fences were

in place by 7 December 1997. These fences were constructed of heavy-gauge silt-fence

material buried ca. 15 cm into the ground — ca. 40 cm extended above ground. Two fence

sections were installed at each of five distances (100 m, 200 m, 300 m, 400 m and 500 m) from

the pond. Fence sections at each distance totaled 13.4 % of the cireumference at that distance

from the pond, and fence sections overlapped (fig. 3b). The two fence sections at 100 m were

each 42 m long with 6 pitfalls (3 on each side of the fence), installed evenly throughout each

section; at 200 m sections were 84 m long with 10 pitfalls; at 300 m sections were 126 m long

with 14 pitfalls: at 400 m sections were 168 m long with 18 pitfalls; at 500 m sections were

210 m long with 22 pitfalls. Pitfall traps were oriented in the same manner as year one:

pond-side traps were on the side of the fences toward OSP and upland-side traps were away

Source : MNHN, Paris

JOHNSON 9

upland-side

tal aps

pond-side upland-side

Pitall aps

Pitall traps.

160m

One Shot Pond

Fig. 3. — Upland drift fence arrays around One Shot Pond, Putnam Co., Florida, USA. The upland

array design in year one of the study is depicted in À and the year two design is depicted in B. One

Shot Pond is shown as a solid circle, and the circle around it represents the drift fence at the pond.

from OSP (fig. 3b). The upland fence array in year two was monitored until the study ended

on 11 September 1998.

In total, 280 pitfall traps were installed at upland fence sections and were monitored

during the 2-year study, for a total of 98,140 trap-nights (one trap-night means one pitfall trap

open for 24 hours). Upland traps were checked on the same schedule as those at the pond and

newts were processed as described above.

OSP, I estimated the proportion of the newt population that migrated different dis!

the pond. Data used in the estimates were confined to 7 December 1997 through 31 March

1998. During this period, there was a mass migration of newts toward the pond and very little

movement away from the pond (JOHNSON, 2001). Ninety-one percent of upland fence captures

during year two occurred during this period. These captures, however, only represented newts

that migrated through a subset of the surrounding uplands. Because upland drift fences

sampled only 13.4 % of the uplands at each distance, I multiplied the number of captures in

the outside pitfalls by 7.5. The product of this calculation is an estimate of the number of

captures expected at each distance had the upland fence sections sampled 100 % of the

uplands at each distance. For each upland fence section, I divided the estimate by the number

of total newt captures on the outside of the fence at OSP to approximate the proportion of

individuals that had migrated various distances (ï.e., 100 m to 500 m, at 100 m intervals). I

assumed there was no strong nonrandom orientation of newts moving through the uplands.

Nonetheless, movement of newts into and away from the pond was nonrandom (see below),

but there was no overwhelmingly strong directionality that would violate this assumption.

Source : MNHN, Paris

10 ALYTES 21 (1-2)

However, estimates of the proportion of newts that migrated various distances from the pond

are probably conservative.

Tuse the term “migration” to indicate seasonal, two-way movements of newts away from

and toward a breeding pond. “Immigration” indicates a general pattern of migration toward

the breeding pond, whereas “emigration” indicates migration away from the pond (SEMLITSCH

& RYAN, 1999). “Dispersal” refers to “once-in-a-lifetime” movement away from a pond and

infers that the dispersing individual will not return to its natal pond.

RESULTS

ORIENTATION AT ONE SHOT POND

All patterns of adult immigration and emigration were significantly nonrandom (fig. 4;

Rao’s spacing tests, all P < 0.001). Adult striped newts entered and exited the pond in all

directions. They tended to enter the pond basin primarily from the east and west (fig. 4).

Adults emigrated in all directions but there was a single, distinct angle of emigration, as

indicated by the relatively high number of captures in a pitfall trap located at a south-

southeast direction (fig. 4). Emigration of paedomorphs and efts also was nonrandom (fig. 5;

Rao’s spacing tests, both P < 0.001). There was no obvious pattern to paedomorph emigra-

tion, but emigrating efts exited the pond basin most often in the southwest quadrant (fig. 5).

Overall patterns of immigration differed significantly from emigration for females and

males (tab. 1). Although the directionality of immigrating adults appeared similar between

the sexes (fig. 4), patterns were significantly different (MRPP test, P = 0.002). There were three

distinct immigration events of adults, but orientation patterns were significantly different

between the sexes only during the third, and largest of these events (tab. 2). Differences in

emigration between males and females (fig. 4) were not significant overall or when distinct

emigration events were compared (tab. 1-2).

There were two distinct emigration events of recently transformed striped newts com-

prising the 1996-97 cohort. The first emigration event took place from October through

November 1996, and the second event from April through June 1997 (JOHNSON, 2002).

Immature newts (i.e., efts) comprised the first event.. whereas emigration later consisted

mostly of recently transformed paedomorphs (JOHNSON, 2002). Patterns of emigration were

significantly different between the eft and paedomorph life-history stages of the same cohort

(tab. 1). In addition to the eft emigration of 1996, a second emigration event of efts took place

from June through early September 1998 (JOHNSON, 2002). Patterns of eft captures at OSP

differed significantly between these two emigration events and, considering all efts and all

adults, efts exited the pond basin in a different pattern from adults (tab. 1-2).

Data for 44 individually marked efts initially caught leaving the pond in the winter of

1996 and recaptured when they returned to breed in the winter of 1997 indicated that

individuals tended to enter and exit the pond within the same quadrant. Sixty-four percent of

these newts left and returned to OSP in the same quadrant, and four individuals (9 %) were

caught leaving and returning to the pond at the same pair of pitfall traps. The vast majority of

Source : MNHN, Paris

JOHNSON 11

Immigrating males Emigrating males

N N

Immigrating females Emigrating females

W E W

Fig. 4. - Orientation patterns of immigrating and emigrating striped newt adults captured in pitfall traps

at a drift fence encircling One Shot Pond, Putnam, Co., Florida, USA. Orientation was significantly

different from random for all four patterns. The length of the lines indicates the number of newts

entering and exiting the pond basin at each pitfall trap.

Source : MNHN, Paris

12 ALYTES 21 (1-2)

Emigrating paedomorphs

Fig. 5. Orientation patterns of emigrating striped newt paedomorphs and efts captured in pitfall traps

at a drift fence encircling One Shot Pond, Putnam, Co., Florida, USA. Orientation was significantly

different from random for both patterns. The length of the lines indicates the number of newts

exiting the pond basin at each pitfall trap.

Source : MNHN, Paris

JOHNSON

Table 1.— Overall comparisons of directional orientation patterns for striped newts entering

Gimmigrating) and leaving (emigrating) One Shot Pond, Putnam Co., Florida, USA.

USE L Standardized >

pe test statistic

Immigrating vs. emigrating males 1159, 486 =15317 <0.001

Immigrating vs. emigrating females 1489, 645 -3.798 0.008

Immigrating males vs. females 1159, 1489 -5.524 0.002

Emigrating males vs. females 486, 645 0.437 02

Emigrating efts vs. emigrating adults 5008, 1131 = 67.639 <0.001

Emigrating efts vs. emigrating 9,506

pacdomorphs ofthe same cohort 0 ï F0

Table 2. — Comparisons of directional orientation patterns for striped newts entering (immigrating) and

leaving (emigrating) One Shot Pond, Putnam Co. Florida, USA.

metamorphie Event 3

Comparison n RUE P

Pi test statistic

Immigrating males vs.

immigrating females ; à

— Immigration Event 1 23,13 0.697 07

— Immigration Event 2 22, 66 —0.130 03

— Immigration Event 3 1049, 1290 —4.008 0.006

Emigrating males vs.

emigrating females ? è

— Emigration Event 2 15,68 0.686 0.7

— Emigration Event 3 430, 484 — 0.005 03

Emigrating efs during metamorphic

Event 1 vs. emigrating efts during 745, 4237 — 3.599 0.01

individuals (84 %) entered the pond basin within the same half they had exited from the

previous year.

MIGRATION INTO UPLANDS

1 captured 831 newts in the upland drift fences during year one (fig. 3a, tab. 3). Pond-side

captures accounted for 73 % of total captures, and migration in year one consisted primarily

of recently transformed efts that were moving into the uplands. I captured newts at all of the

upland fence sections (fig. 3a: tab. 3) and in most (91.4 %) of the pond-side pitfall traps.

Source : MNHN, Paris

14 ALYTES 21 (1-2)

Table 3. — Numbers of striped newts captured in pitfall traps at drift fence arrays in the sandhill uplands

surrounding One Shot Pond, Putnam, Co., Florida, USA. Drift fences were located at various

distances from the pond. See fig. 3 for a depiction of the arrays.

Year 1 Year 2

d 20m 40m 80m 160 m | 100m | 200m | 300m | 400m | 500 m

Pond-side 140 126 169 172 Il 6 10 12 7

Upland-side | 79 39 64 42 121 108 86 86 48

Total 219 165 233 214 132 114 96 98 55

During each period of migration most newts were captured on the same sides of upland drift

fences. However, for some movement events, a few newts were captured in pitfalls on the

opposite side of fences from the majority of captures. I believe this is because there was a small

degree of wandering by some newts in the uplands as they moved to or from OSP. Pond-side

captures at upland fences in year one represented three distinct periods of newt migration, two

emigration events and one immigration event (tab. 4). Most newts captured on the pond-side

of upland fences in year one (76% of pond-side captures) were caught during the first

emigration event (i.e., El), which occurred from October 1996 through February 1997 (tab. 4).

Emigration during this period consisted almost exclusively of immature efts that had recently

transformed. I captured far fewer newts (15 % of pond-side captures) during emigration event

two (E2), which occurred from April through July of 1997 (tab. 4). This emigration event was

comprised of recently transformed paedomorphic newts (54 % of the migrating newts), as

well as recently transformed efts and several adults that likely had finished breeding and were

moving back into the uplands. The third period of migration, indicated by pond-side fence

captures in year one, was the result of an immigration event (1.e., 13) that began in October

1997 (tab. 4). There was a major breeding migration of adults to the pond that began in

October 1997 and pond-side captures at this time probably resulted from adults that were

moving toward the pond but happened to be captured on the pond-side of the upland drift

fences (tab. 4).

Upland-side captures of striped newts accounted for 27 % of captures in year one. I

captured newts at each of the five fence sections (fig. 3a), at each distance from OSP (tab. 3)

and in most (81,4%) of the pitfall traps on the upland-side of the fences in year one.

Upland-side captures occurred during three distinct periods of migration, all of which were

immigration events. These migration events (11, 12 and 13; tab. 4) occurred during the same

time periods as described above for pond-side captures (tab. 4). Immigration event 13

accounted for the largest proportion (54 %) of upland-side captures in year one, followed by

event I (29 %) and 12 (17 %). AI of these migration events consisted of adult newts moving

toward OSP to breed (tab. 4).

Icaptured 495 newts in the upland drift fences during year two (fig. 3b, tab. 3). In contrast

to year one, migration consisted primarily of immigrating adults. Pond-side captures accoun-

ted for only 9 % of total captures. I captured newts at each of the two fence sections (fig. 3b)

and at each distance from OSP (tab. 3), but captures were recorded in less than half of the

pitfall traps (42.8 *%) on the pond-side of the upland fences in year two. Pond-side captures at

Source : MNHN, Paris

Table 4. — Captures of striped newts in upland fences around One Shot Pond, Putnam Co., Florida, USA, during distinct periods of movement.

arrays modified in early December 97.

: fence

Predominant Number

Fence side Migration direction of Time period of event ofnewts Description

ofcaptures EERe newt movement captured

Year 1

Pond-side El Away from pond | October 96 through February 97 461 Emigrating efts

Pond-side E2 Away from pond April 97 through July 97 91 Primarily emigrating paedomorphs and efts

Pond-side B Toward pond | October 97 through December 97* 55 Immigrating adults

Upland-side nl Toward pond October 96 through January 97 65 Immigrating adults, some emigrating efts

Upland-side m2 Toward pond April 97 through July 97 36 Immigrating adults

Upland-side B Toward pond | November 97 through December 97* 123 Immigrating adults

Year 2

Pond-side 5 Toward pond December 97* through March 98 16 Immigrating adults

Pond-side E3 Away from pond June 98 through September 98 25 Emigrating efts

Upland-side 5 Toward pond December 97* through March 98 449 Immigrating adults

NOSNHOS

Source : MNHN, Paris

16 ALYTES 21 (1-2)

upland fences in year two represented two distinct periods of newt migration, one immigra-

tion event (i.e., 13) and one emigration event (i.e., E3). I captured few newts during both of

these events; 16 during 13 and 25 newts during E3 (tab. 4). Captures during migration event 13

were adults that were moving to the pond to breed but were captured in pond-side traps as

they wandered toward the pond. Captures during E3 were recently transformed newts that

were leaving OSP.

In year two, I captured far more newts (91 % of total upland captures) on the upland-side

of drift fences than on the pond-side (tab. 3). I captured newts at all sections of drift fence and

in almost all of the upland-side pitfalls (88.6 %). Captures occurred during a single immigra-

tion event (13; tab. 4) and were exclusively of adults that were immigrating to OSP to breed.

The number of captures declined as the distance from the pond increased (tab. 3). Based on

estimated values, at least 360 newts (16 % of the breeding migration) migrated more than 500

m from OSP (fig. 6). I estimated that 645 newts (29 % of the breeding migration) migrated at

least 400 m. The estimate was the same for 300 m (645 newts). Iestimated that 810 (36 % of the

breeding migration) and 908 (41 % of the breeding migration) of newts migrated from the

pond at least 200 and 100 m, respectively (fig. 6). Based on these estimates, it appears that

roughly 60 % of the striped newts emigrated less than 100 m. However, as indicated by

captures at the 500 m fences, a substantial percentage of individuals comprising the 1997-98

breeding migration immigrated to OSP from farther than 500 m. In fact one newt that was

marked leaving OSP as an eft on 18 November 1996 was recaptured on 4 February 1998 as it

colonized Fox Pond, a dispersal distance of approximately 685 m.

DISCUSSION

ORIENTATION

The distribution of habitats surrounding a breeding pond should influence patterns of

immigration revealed by captures of salamanders at the pond. Habitat preferences among

species and/or differential survivorship in various habitat types might be apparent as individ-

uals arrive at the breeding pond. For example, imagine an amphibian breeding-pond in which

one half of the uplands surrounding the pond were pine plantation (i.e., marginal habitat)

whereas the other half remained native uplands (i.e., preferred habitat). The pattern of

captures at the pond would be expected to reflect the distribution of upland habitats. One

might predict significantly fewer captures along the half of the pond adjacent to the pine

plantation as compared to the native upland half. This is because pond-breeding salamanders

have the ability to select appropriate upland habitats and accurately navigate through uplands

during migration, often using specific habitat types (SHOoP, 1968; HURLBERT, 1969;

SemLirsCH, 1981; STENHOUSE, 1985; MaDISON, 1997; MADISON & FARRAND, 1998; DEMaY-

NADIER & HUNTER, 1999; MALMGREN, 2002; ROTHERMEL & SEMLITSCH, 2002).

In this study, although newts entered and exited the pond basin from all directions,

migration was nonrandom. Some directions were preferred over others, but there were no

obvious upland habitat features that could explain the newts’ orientation behavior. However,

I did not measure habitat variables in the uplands and individuals could have used micro-

Source : MNHN, Paris

JOHNSON 17

Estimated numbers of captures

0 100 200 300 400 500 600 700 800 900 1000

Distance from pond (m)

Fig. 6. - Estimated numbers of striped newt captures in pitfall traps at drift fences in the sandhill uplands

around One Shot Pond, Putnam Co., Florida, USA. Drift fences were located at 100 m intervals up

to 500 m from the pond. The zero point represents captures at a drift fence encircling the pond. See

Materials and methods for an explanation of how the estimated numbers were calculated.

topographic features as cues to navigate toward the pond. In a similar study, Dobp & CADE

(1998) concluded that movements of striped newts and narrowmouth toads were a reflection

of the distribution of favorable upland habitats around the pond. Although the uplands at

OSP were primarily sandhill habitat, a small plantation of slash pine (with intact groundco-

ver) was well within the dispersal capabilities of migrating newts (fig. 2). In year one I often

caught newts at a section of drift fence in the pine plantation. Newts could have resided within

the plantation or have traveled through it en route to native sandhill. Nevertheless, this

plantation represented only a small portion of the uplands and had no detectable effect on

striped newt movements.

Although upland-habitat preferences and microenvironmental features 1 did not

measure could have influenced the nonrandom pattern of immigration observed at OSP, if

measured over several seasons, orientation may in fact be random. It is possible that striped

newts are roughly evenly distributed in the uplands around OSP but that only a portion of the

population migrates to the pond during any particular breeding event. If the portion of

individuals moving was not indicative of the whole population, then what truly should be

random orientation would appear as nonrandom because data were collected for a relatively

short time.

Patterns of newt emigration were also nonrandom, and newts exited the pond basin in all

directions. Efts emigrated predominantly in the south quadrant of the pond. The slope of

the pond basin was shallowest in this quadrant, and water depth during metamorphic events

Source : MNHN, Paris

18 ALYTES 21 (1-2)

could have influenced the behavior of recently transformed efts as they left the pond. On the

other hand, adults emigrated most often in the south-southeast portion of the basin.

Differences in aquatic habitat preference (e.g., depth) between adult and immature newts

might explain the varying emigration patterns, although habitat preferences of both life

history stages are unknown.

UPLAND MIGRATION

Using upland drift fence arrays in year two, I was able to estimate the percentage of the

striped newt breeding population that migrated different distances (in increments of 100 m)

from the pond. Captures at drift fences in the sandhill uplands surrounding OSP indicated

that many striped newts (16%) migrated more than 500 m from the pond. This is a

conservative estimate because newts captured in traps closer to the pond may have migrated

further than indicated by the traps. Captures at the drift fence surrounding the pond and at

upland drift fences at the end of year one showed that a breeding migration of newts into OSP

had begun before the installation of fences for year two (JoHNsoN, 2001, 2002). Although the

proportion of individuals caught at the pond before the new upland fence arrays were

established was small (7 % of the total), some newts already had moved toward the pond

before the upland arrays were in place. Moreover, immigrating adults did not arrive at the

pond in a random fashion during this breeding migration. The upland fence arrays in year two

were located north and southeast of OSP and newts were caught at the pond with lowest

frequency toward the north. Therefore, the proportion of the breeding population caught at

each distance from the pond in year two is likely an underestimate of the actual proportion

that migrated to that particular distance.

Many pond-breeding amphibians have complex life-cycles and spend much of their adult

lives in terrestrial habitats away from breeding sites. Distances that individuals disperse or

migrate from breeding ponds have been reported for some species (DODD, 1996; SEMLITSCH,

1998 and references therein). It is clear that individuals disperse and migrate hundreds of

meters from breeding sites into upland habitats, some even thousands of meters. With few

exceptions, however, distance values usually have been presented for less than 10 individuals

per species. The results from my study appear to be the first estimates of migration distances

for a breeding population of North American amphibians based on a substantial sample size.

CONSERVATION IMPLICATIONS

Central to a successful amphibian conservation strategy is the protection of sufficient

breeding and nonbreeding habitat (i.e., the pond and appropriate “core habitat”: SEMLITSCH

& JE 2001). Studies of amphibian migration and dispersal can provide the scientific basis

for determining directional and distance components that can be used to establish protected

areas around breeding ponds. BROWN et al. (1990) used spatial requirements (i.e., distance

moved from a wetland), among other data, to recommend width of “buffer zones” for wildlife

protection at wetlands in Florida. Nevertheless, lack of data for amphibians forced them to

use rough estimates for most of the species considered. Further utility of movement distance

data can be found in regulations to protect the flatwoods salamander (Ambvstoma cingula-

Source : MNHN, Paris

JOHNSON 19

tum) which, as a result of severe population decline (MEANS et al., 1996), was federally listed

as threatened in the USA (ANONYMOUS, 1999). The US Fish and Wildlife Service restricts

specific silvicultural practices within 450 m of flatwoods salamander ponds. Additionally,

only selective timber harvest at specific times is allowed within a primary radius of 164 m

around breeding ponds (ANONYMOUS, 1999). The width of the primary zone (164 m) was

derived from a review of migration distances for pond-breeding salamanders of the genus

Ambystoma (SEMLITSCH, 1998), despite the fact that no data for À. cingulatum were presented.

This example underscores the need to determine migration and dispersal distances for all

pond-breeding amphibians. SEMLITSCH (1998) acknowledged that the extent of protected

upland recommended for Ambystoma species may apply to some species of pond-breeding

amphibians, but certainly not all. My data show that recommendations for protecting

terrestrial habitat for ambystomatid salamanders are inadequate for Notophthalmus perstria-

tus. Therefore, it is not defensible to extrapolate data across taxa. Clearly, a 164 m protected

zone would not protect all of the striped newts breeding at OSP. Based on extrapolation of

migration distances revealed by upland drift fences, a protected area of “core habitat”

extending ca. 1000 m from OSP would likely be needed to encompass almost all of the newts

that breed there.

Although they have great value as wildlife habitat, small, isolated wetlands in the United

States are afforded little protection from development. Overall, more than 50 % of wetlands

have been destroyed by development in the United States (DAHL, 1990), and much of this loss

has been small wetlands. In Florida, a state with an extremely large number and diversity of

wetlands, isolated wetlands less than 0.2 ha receive no protection from development. This size

threshold was adopted by the state’s water management districts “based on a consensus of

scientific and regulatory opinion rather than on biological and hydrological evidence” (HART

& NEWMAN, 1995). Small wetlands are just as vulnerable at the national level as they are in

Florida.

There is strong evidence that protection of core areas of terrestrial habitat surrounding

breeding sites is crucial for persistence of amphibian populations and species. Data from OSP

demonstrate that small, isolated wetlands can support breeding populations of salamanders

that migrate hundreds of meters into the surrounding uplands. Similar studies at other ponds

and in different upland types are necessary because data on upland habitat requirements

(quality and quantity) of most amphibian species are lacking. Without this information,

designating terrestrial “core habitat” to conserve aquatic-breeding amphibians will largely

remain guesswork, with generalizations made from data on relatively few individuals of a few

species. However, unless more protection is afforded to small, isolated wetlands, arguments to

preserve uplands surrounding the wetlands are irrelevant.

ACKNOWLEDGMENTS

This study was funded by Grant No. 1448-0004-96-985 from the US Fish and Wildlife

1 would like to especially thank Linda LaClaire for administering the grant. [thank Brad Austin.

Barichivich, Cheryl Cheshire, Richard Franz, C. Kenneth Dodd, Jr., Dale Johnson, Kenneth D. Krysko,

Rick “Bubba” Owen, Joe Sexton, Matt Seguin, Jennifer Staiger, Chad Truxall and Travis Tuten for help

Source : MNHN, Paris

20 ALYTES 21 (1-2)

installing and/or checking pitfall traps. 1 am grateful to Dale Johnson for drafting several of the figures

and to C. Kenneth Dodd, Jr., Richard Franz, Holly Freifeld, David Leonard, Betsie Rothermel and two

anonymous reviewers for comments on earlier versions of the manuscript. Brian Cade provided help with

the BLOSSOM program and data analysis. I thank John Eisenberg and the Board of the Katharine

Ordway Preserve-Swisher Memorial Sanctuary for allowing me to conduct the research on the preserve.

This project adhered to guidelines of the Institutional Animal Care and Use Committee of the University

of Florida. This paper comprised one portion of my Ph.D. dissertation research conducted through the

Department of Wildlife Ecology and Conservation at the University of Florida. This research was

supported by the Florida Agriculture Experiment Station, and approved for publication as Journal Series

No. R-09692.

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Source : MNHN, Paris

Alvtes, 2003, 21 (1-2): 23-44. 23

Revision of the genus Ophryophryne

Boulenger, 1903 (Megophrvyidae)

with description of two new species

Annemarie OHLER

Vertébrés: Reptiles & Amphibiens,

USM 0602 Taxonomie & Collections,

Département de Systématique & Evolution,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

<ohler@mnhn.fr>

Study of type specimens and neuly collected specimens of frogs of the

oriental genus Ophryophryne Boulenger, 1903 lead to the redefinition of

the recognized taxa and the recognition of two new species, one reported

from Vietnam and Laos, and the second only from Vietnam. Ophryophryne

poilani Bourret, 1937 is put to the synonymy of Ophryophryne micros-

toma Boulenger, 1903. The range of the species O. microstoma, which

includes Vietnam and China, is extended to Thailand. Ophryophryne

pachyproctus Kou, 1985 is confirmed by a new specimen from Vietnam.

Ophryophryne Boulenger, 1903 is a member of the family Megophryidae Bonaparte,

1850 as defined by FORD & CANNATELLA (1993). Sister-group of Xenophrys Günther, 1864 as

defined by Dugois & OuLer (1998), it is either regarded as a separate genus (recent references:

YEet al., 1993; MANTHEY & GROSSMANN, 1997) or as a subgenus of Megophrys Kuhl & Van

Hasselt, 1822 (Dugois, 1980).

The genus Ophrvophryne was described in 1903 by BOULENGER (19034) who mentioned

both its close morphological resemblance to Megophrys, and its lack of maxillary and

vomerine teeth and the presence of a horizontal pupilla, characters defining the family

Bufonidae Gray, 1825. Although BOULENGER (19034) was aware of the problem of using

isolated characters in making taxonomic decisions, he included his new genus into the

Bufonidae (BOULENGER, 1903b). Despite the work of NoBLe (1926), who clearly recognized

this genus as a member of the Pelobatidae (including then the subfamily Megophryinae),

nus Megophrys, most authors continued to consider it to be a member of the

Bufonidae (see review in DuBois, 1980: 470; recent reference: NGUYEN & Ho, 1996). The

description of the tadpole (Lru & HU, 1962) and the study of external morphology indicated

rvophryne to be a megophryid and closely related to Megophrrs (DuBois, 1980). Recent

d on cytology and adult and larval morphology confirmed Ophryophryne to be

roup of Xenophrys, a group formerly included in the genus Megophrys (TIAN & HU,

; Rao & YANG, 1997; DuBois & OHLER, 1998).

Source : MNHN, Paris

24 ALYTES 21 (1-2)

The genus was described to include one species, Ophryophryne microstoma Boulenger,

1903. In 1937, BouRRET described a second species, Ophryophryne poilani, based upon a

single, badly preserved specimen. Not until 1985 was a third species described by Kou,

Ophryophryne pachyproctus from southern China (Yunnan).

In recent years important new material of amphibians was collected in South-East Asia.

INGER et al. (1999) reported on the three species of Ophryophryne from southern Vietnam.

Beside the material collected in Vietnam, I had the opportunity to study the type-specimens of

the three nominal species and other material stored in museum collections. The analysis of

these data led me to describe two new species in this genus and to put Ophryophryne poilani in

the synonymy of Ophryophryne microstoma.

MATERIALS AND METHODS

The material studied is listed under the appropriate species and is deposited in the

museums mentioned below. My study is based on morphology and morphometry of

formalin-fixed and alcohol-preserved specimens. Data on live animals are added when

available. The descriptions, methodology and terminology follow OHLER (1996) and DuBois

& OHLER (1998). The webbing formula used is that of Myers & DUELLMAN (1982). AIl

measurements are in millimetres. The tables present measurements and their per mille ratios

to SVL (in brackets) with the associated mean and standard deviation. Abbreviations for

measurements and institutions are as follows:

SVL, snout-vent length.

Head. - HW, head width; HL, head length (from posterior edge of mandible to tip of

snout); MN, distance from posterior edge of mandible to posterior border of nostril; MFE,

distance from posterior edge of mandible to anterior corner of eye; MBE, distance from

posterior edge of mandible to back of eye: IFE, distance between anterior corner of eyes;

IBE, distance between posterior border of eyes: IN, internarial space; EN, distance from

anterior corner of eye to posterior border of nostril; EL, eye length; SN, distance from

anterior border of nostril to tip of snout; SL, distance from front of eye to tip of snout; TYD,

greatest tympanum diameter; TYE, smallest distance from anterior border of tympanum to

posterior corner of eye; IUE, minimum distance between upper eyelids; UEW, maximum

width of inter upper eyelid.

Forearm. - HAL, hand length (from base of outer palmar tubercle to tip of toe); FLL,

forelimb length (from elbow to base of outer tubercle): TFL, third finger length (from base of

first subarticular tubercle).

Hindlimb. — FL, femur length (from vent to knee): TL, tibia length; FOL, foot length

(from base of inner metatarsal tubercle to tip of toe); FTL, fourth toe length (from base of

first subarticular tubercle); TFOL, distance from base of tarsus to tip of fourth toe.

Webbing. - MTTE, distance from distal edge of metatarsal tubercle to maximum

incurvation of web between third and fourth toe; TFTEF, distance from maximum incurvation

of web between third and fourth toe to tip of fourth toe; MTFF, distance from distal edge of

Source : MNHN, Paris

OHLER 25

metatarsal tubercle to maximum incurvation of web between fourth and fifth toe; FFTF,

distance from maximum incurvation of web between fourth and fifth toe to tip of fourth toe.

nm, not measured.

p.m., per mille.

BMNA, Natural History Museum, London, United Kingdom.

CIB, Chengdu Institute of Biology, Chengdu, Sichuan, China.

FMNB, Field Museum of Natural History, Chicago, Illinois, USA.

IEBR, Institute of Ecology and Biological Research, Hanoï, Vietnam.

MNAHN, Muséum national d'Histoire naturelle, Paris, France.

YU, Yunnan University, Kunming, Yunnan, China.

TAXONOMIC ACCOUNTS

Ophryophryne Boulenger, 1903

Ophryophryne Boulenger, 1903: 186. — Type-species: Ophryophryne microstoma Boulenger, 1903, by

monotypy.

Ophryophryne gerti sp. nov.

Holotype. - BMNH 1921.4.1.324, adult male, Cam Ly (river), south-east of Da Lat (11°56N,

108°25°E), Lang Bian Plateau, sLam Dong Province, Vietnam.

Paratypes. - BMNH 1921.4.1.323, young female, Dran (11°50°N, 108°34’E), Lang Bian

Plateau, Lam Dong Province, Vietnam; BMNH 1972.15.2.4, adult male, Huey Sapan, Pak

Maat (precise location not found), Mekong, Laos.

Other specimens examined. — VIETNAM: Buon Luoi, An He District, Gia-Lai Province:

FMNH 252899, 252901.

Diagnosis. - Small-sized Ophryophryne, with relatively long tibia, small tympanum, small

head. Dorsal coloration uniform, dark. Supraorbital horn distinct, no dermal protuberance

bearing anus.

Description of holotype (fig. D. -(A) Size and general aspect. — (1) Specimen of rather small

size (SVL 34.8 mm); body slender.

(B) Head. - (2) Head small, wider (HW 9.1 mm) than long (HL 8.5 mm; MN 7.5 mm;

MFE 6.5 mm; MBE 3.4 mm), convex. (3) Snout truncate, protruding: its length (SL 3.31 mm)

shorter than horizontal diameter of eye (EL 3.95 mm). (4) Canthus rostralis rounded, loreal

region slightly convex, acute in cross section. (5) Interorbital space concave, narrower (IUE

2.59 mm) than upper eyelid (UEW 2.98 mm) and than internarial distance (IN 3.05 mm);

distance between front of eyes (IFE 5.31 mm) about two third of distance between back of

eyes (IBE 7.9 mm). (6) Nostrils oval, with small flap of skin laterally: closer to eye (EN

1.23 mm) than to tip of snout (NS 1.62 mm). (7) Pupil rounded (in preservative). (8)

Tympanum (TYD 2.14 mm) rounded, about half eye diameter and approximately equal to

Source : MNHN, Paris

26 ALYTES 21 (1-2)

Fig. 1. — Ophryophryne gerti sp. nov. BMNH 1921.4.1.324, holotype, adult male, SVL 34.8 mm, Dorsal

view (op): lateral view of head (bottom).

Source : MNHN, Paris

OHLER 27

tympanum-eye distance (TYE 2.07 mm). (9) Pineal ocellus absent. (10) Vomerine ridge

absent. (11) Tongue rounded, with dorsal hollow, largely attached to mouth floor. (12)

Supratympanic fold prominent, from eye to shoulder.

(HAL 8.0 mm), not enlarged. (14) Fingers I, II and IV short and thin; finger III long and thin

(TEL 4.99 mm). (15) Relative length of fingers, shortest to longest: I < II =IV< III. (16) Tips

of fingers rounded, not enlarged. (17) Fingers without dermal fringe; webbing absent. (18)

Subarticular tubercles absent. (19) Prepollex oval, indistinct; palmar tubercles indistinct and

supernumerary tubercles absent.

(D) Hindlimbs. — (20) Shanks three times longer (TL 15.4 mm) than wide (TW 4.8 mm),

about same length as thigh (FL 15.2 mm) and longer than distance from base of internal

metatarsal tubercle to tip of toe IV (FOL 14.8 mm). (21) Toes long and thin, toe IV (FTL 7.0)

about third of distance from base of tarsus to tip of toe IV (TFOL 21.4 mm). (22) Relative

length of toes, shortest to longest: I < II < V < III < IV. (23) Tips of toes rounded, not

enlarged. (24) Webbing absent: I 2-2 %2 II 2-3 II 2 % -41V 4-2 % V (MTTF 3.95 mm;

MTFF 4.61 mm; FTTF 6.97 mm; FFTF 6.97 mm). (25) Dermal fringe along toe V absent.

(26) Subarticular tubercles absent. (27) Inner metatarsal tubercle flat, its length (IMT

2.07 mm) 1.16 times in length of toe I (ITL 2.40 mm). (28) Tarsal fold absent. (29) Outer

metatarsal tubercle, supernumerary tubercles and tarsal tubercle absent.

(E) Skin. — (30) Dorsal and lateral parts of head and body: snout shagreened, granular;

between eyes smooth and shagreened; side of head granular; orbital horn free, pointed, its

length 0.78 mm; anterior part of back shagreened: posterior part of back shagreened,

granular; upper part of flank shagreened, with glandular warts; lower part of flank granular.

(31) Cephalic ridges absent. (32) Latero-dorsal folds, lateral line system and “Fejervaryan”

line absent. (33) Dorsal parts of limbs: forelimb smooth, with few small glandular warts:

thigh and leg with flat glandular warts; tarsus smooth. (34) Ventral parts of head, body and

limbs: throat, chest, belly and thigh smooth. (35) Small pairs of pectoral and femoral glands

present.

(F) Coloration in alcohol. - (36) Dorsal and lateral parts of head and body: dorsal parts

of head and dorsum and upper part of flank dark brown, homogeneous; lower parts of flank

with darker brown spots; loreal region brown with darker brown indistinct bands including

upper lip; tympanum and tympanic region dark brown, tympanic fold underlined by blackish

brown. (37) Dorsal parts of limbs: dorsal part of forelimbs, of thigh, of shank and of foot

dark brown with indistinct darker brown bands; posterior part of thigh brown with blackish

triangle around vent. (38) Ventral parts of head, body and limbs: throat, margin of throat,

chest and thigh rather dark brown; lower part of belly yellow with brown spots and blackish

spots; dark brown spots also on ventral part of thighs; macroglands white.

Coloration in life. - Not known.

(G) Male secondary sexual characters. — (39) Nuptial spines present on fingers I and IT:

numerous small brown spines forming two oval patches. (40) Vocal sacs absent, non visible

either exteriorly or interiorly. (41) Other male secondary sexual characters: dorsally of vent,

presence of a short fleshy flap.

Female sexual characters. - Not observed.

Source : MNHN, Paris

28 ALYTES 21 (1-2)

Table 1. — Measurements (mm) and per mille of snout-vent length (in parenthesis) of five specimens,

including holotype, of Ophryophryne gerti sp. nov.

Collection number |BMNH 1921.4.1.324] BMHN 1972.15.24 [BMNH 1921.4.1.323] FMNH252899 | FMNH 252901

Locality Cam Ly, Vietnam | Huey Sapan, Laos | Dran, Vietnam | Buon Luoi, Vietnam | Buon Luoi, Vietnam

(Status Holotype Paratype Paratype Additional material | Additional material

Sex Adult male Adult male Juvenile female Adult female Adult female

ÉSnout-vent length 348 320 210 414 458

Head length 8.5 (244) 8.5 (266) 64 (305) 11.0 (266) 11.2(245)

[Tympanum diameter| 2.01 (58) 213 (67) 1.16 (55) 2.92 (71) 324 (71)

[Thigh 152 (437) 15.6 (488) 8.7 (414) nm nm

[Shank 154 (443) 15.5 (484) 9.3 (443) 17.4 (420) 18.0 (393)

[Foot 14.8 (425) 13.4 (419) 7.3 (348) 16.8 (406) 16.9 (369)

Variation. — À second male is smaller in body size (tab. 1), but very similar in all body

measurements. Its nuptial pads are translucent, which might indicate that he is sub-

adult.

Distribution. — Ophryophryne gerti is known from Laos and Vietnam.

Etymology. — This species is dedicated to my sister Gerti for her help during fieldwork. The

invariable specific epithet gerti is a noun used in apposition.

Comments. - Two female specimens (FMNH 252899, 252901) from Buon Luoi (Vietnam) are

here tentatively referred to this species. These specimens are distinguished by smooth skin.

Skin is in general smoother in female than in male specimens in Ophryophryne. As these

females are morphologically distinct from the three other species of the genus, but show

similarities to the type-specimens of ©. gerti, they are included in this species.

Ophryophryne hansi sp. nov.

Ophryophryne poilani (non Bourret, 1937: 8): INGER et al., 1999: 9.

Holotype. - FMNH 252880, adult male, Buon Luoi (700-750 m), 20 km west of the town of

Kannack (14°20°N, 108°36'E), An Khe District, Gia-Lai Province, Vietnam (INGER et al.,

1999).

Paratopotypes. - FMNH 252873, 252875, 252878-79, 252882, 252884, 252892-93, 7 adult

males, 1 adult female.

Diagnosis. — Relatively large-sized Ophryophryne, with relatively long shank, small tympa-

num, large head. Dorsal coloration dark, almost black. Supraorbital horn forming small

projection, no dermal protuberance bearing anus.

Description of holotype (fig. 2). (A) Size and general aspect. — (1) Specimen of medium size

(SVL 38.8 mm), body rather stout.

(B) Head. — (2) Head very small, wider (HW 12.4 mm) than long (HL 10.4 mm; MN

9.3mm; MFE 8.1 mm; MBE 4.0 mm), convex. (3) Snout rounded, very protruding, its length

Source : MNHN, Paris

OBLER 29

Fig. 2. — Ophryophryne hansi sp. nov. FMNH 252880, holotype, adult male, SVL 38.8 mm. Dorsal view

(left); ventral view (right).

(SL 3.76 mm) shorter than horizontal diameter of eye (EL 4.86 mm). (4) Canthus rostralis

rounded, loreal region concave, acute in cross section. (5) Interorbital space convex, narrower

(LUE 2.72 mm) than upper eyelid (UEW 3.76 mm) and internarial distance (IN 3.04 mm);

distance between front of eyes (IFE 6.0 mm) three fifth of distance between back of eyes

(IBE 10.0 mm). (6) Nostrils oval, with small flap of skin laterally, closer to eye (EN 1.43 mm)

than to tip of snout (NS 1.69 mm). (7) Pupil diamond-shaped, vertical. (8) Tympanum (TYD

2.14 mm) distinct, oval, vertical, smaller than half diameter of eye; tympanum-eye distance

(TYE 2.46 mm) more than its diameter. (9) Pineal ocellus absent. (10) Vomerine ridge absent.

(11) Tongue large, rounded, not emarginate. (12) Supratympanic fold distinct, present from

eye to shoulder, posterior part slightly enlarged.

Source : MNHN, Paris

30 ALYTES 21 (1-2)

10.9 mm), not enlarged. (14) Fingers long and thin (TFL 5.7 mm). (15) Relative length of

fingers, shortest to longest: I < II < IV< II. (16) Tips of fingers rounded, not enlarged. (17)

Fingers without dermal fringe; webbing absent. (18) Subarticular tubercles indistinct; on

fingers I and II a single proximal, oval tubercle. (19) Prepollex oval, indistinct; a single, oval

palmar tubercle; supernumerary tubercles absent.

(D) Hindlimbs. — (20) Shanks four times longer (TL 18.9 mm) than wide (TW 4.9 mm),

about length of thigh (FL 18.7 mm), but longer than distance from base of internal metatarsal

tubercle to tip of toe IV (FOL 16.9 mm). (21) Toe IV (FTL 9.1) about one third of distance

from base of tarsus to tip of toe IV (TFOL 26.7 mm). (22) Relative length of toes, shortest to

longest: I < II < V < II < IV. (23) Tips of toes rounded, slightly enlarged; discs absent. (24)

Webbing absent: I 2-2 V2 II 2-3 % III 3 —3 2 IV 3 2-2 2 V (MTTF 5.53 mm; MTFF

6.32 mm; FTTF 10.40 mm; FFTF 10.40 mm). (25) Dermal fringe along toe V absent. (26)

Subarticular tubercles indistinct. (27) Inner metatarsal tubercle long, flat, its length (IMT

3.37 mm) 1.06 times in length of toe I (ITL 3.57 mm). (28) Tarsal fold absent. (29) Outer

metatarsal tubercle, supernumerary tubercles and tarsal tubercle absent.

(E) Skin. — (30) Dorsal and lateral parts of head and body: snout, between eyes, side of

head, back and upper part of flanks with glandular warts and horny spinules; on lower part

of flank, some of these warts of large size. (31) Cephalic ridges absent. (32) Latero-dorsal

folds, lateral line system and “Fejervaryan” line absent. (33) Dorsal parts of limbs: forelimb,

thigh, leg and tarsus with glandular warts and horny spinules. (34) Ventral parts of head, body

and limbs: throat, chest with foldings; belly and thigh smooth. (35) Femoral and pectoral

glands present; medium sized glands on flanks and rear part of thigh.

(F) Coloration in alcohol. — (36) Dorsal and lateral parts of head and body: dorsal parts

of head and dorsum grey-brown with dark brown spots, in particular a triangle between eyes;

flanks grey-brown with dark brown and white spots, white ones corresponding to glands;

loreal region dark brown; tympanum and tympanic region dark brown with a light brown

stripe from eye to upper lip; upper lip dark brown with few clear spots. (37) Dorsal parts of

limbs: dorsal part of forelimbs, of thigh, of shank and of foot grey-brown with broad dark

brown bands; posterior part of thigh brown with dark perianal zone and dark zone around

femoral glands; femoral glands and glands near vent whitish. (38) Ventral parts of head, body

and limbs: throat and margin of throat brown with dark brown spots; chest brown with dark

brown spots and whitish pectoral glands; belly and thigh yellowish with brown marblings:

pectoral glands whitish.

Coloration in life. — “In life males vary from black with sharply delimited yellow spots to

grey-brown with obscure lighter spots; females yellowish grey with obscure, small lighter

spots” (INGER et al., 1999: 9).

(G) Male secondary sexual characters. — (39) Numerous, small, dark brown nuptial

spines forming oval pads on fingers I and IL. (40) Single vocal sac present, with rounded

openings posterior on mouth floor. (41) Other male secondary sexual characters: fleshy flap

on vent not present.

Female sexual characters (FMNH 252882, SVL 53.5 mm). - Large (2.40 mm) creamy-

whitish ovocytes in ovary.

Source : MNHN, Paris

OHLER 31

Table 2. - Measurements (mm) and per mille of snout-vent length (in parenthesis) of type-specimens of

Ophryophryne hansi sp. nov., from Buon Luoi, Vietnam. Means and standard deviations are given

between square brackets. Thigh was not measured in these specimens.

Collection number FMNH 252880 FMNH 252873, 252875, 252878-79, 252884, 252892-93 FMNH 252882

Status Holotype 7 paratypes Paratype

Sex Adult male Adult males Adult female

Snout-vent length 38.8 35.3-43.0 [38.84 2.51] 53.5

Head width 124 (320) 11.3-12.9 (291-347) [12.3 4 0.54 (319 4 20.9)] 15.8 (295)

Head length 10.4 (268) 10.6-11.7 (258-315) [11.1 + 0.35 (286 + 20.1)] 14.5 (271)

Tympanum diameter 2.14 (55) 2.59-2.92 (63-74) [2.72 + 0.14 (70 + 4.3)] 3.6 (68)

Shank 18.9 (487) 18.4-19.0 (440-538) [18.9 + 0.22 (488 + 33.9)] 25.6 (479)

Foot 16.9 (436) 16.1-18.2 (395-478) [16.8 + 0.69 (435 +31.0)] 23.9 (447)

Variation. — Table 2 gives variation of body measurements for males and female.

Distribution. - Species known only from the type-locality in Vietnam.

Etymology. - This species is dedicated to my sister Hansi who very kindly “adopts” my son

during periods of fieldwork, thus generously supporting my research. The invariable specific

epithet hansi is a noun used in apposition.

Ophryophryne microstoma Boulenger, 1903

Ophryophryne microstoma Boulenger, 1903a: 186. - Type-specimen: lectotype, by present designation,

BMNH 1947.2.22.52 [ex 1903.4.29.106], adult male (examined). — Type-locality: Mau Son [“Man-

Son Mountains, Tonkin, altitude 3000-4000 feet”] (22°00°N, 106°45°E), Lang Son Province, Viet-

nam

Megophrvs (Ophryophryne) microstoma: DUvoIs, 1980: 473.

Ophryophryne microstoma: Dunois, 1987: 23.

Ophryophryne poilani Bourret, 1937: 8. — Tÿpe-specimen: holotype, by monotypy, MNHN 1948.0113,

adult female (examined). — Tÿpe-locality: Dong-Tam-Ve (16°40°N, 106%45E), concession of the

collector E. Poilane, near the Ailao Pass, Quang Tri Province, Vietnam. — New synonym.

Megophrys (Ophryophryne) poilani: Durois, 1980: 472.

Ophryophryne poilani: Dusois, 1987: 23.

Other specimens examined. - THAILAND: NE Thailand: BMNH 1974.2334 (hands and limbs

missing); VIETNAM: Man-Son Mountains, Tonkin, 3000-4000 feet: BMNH 1947.2.22.50-51

[ex 1903.4.29.104-105], 1947.2.22.53 [ex 1903.4.29.107]; Ben En, Tanh Hoa province: MNHN

1997.5258. , IEBR D.23l; Tam Dao, Viet Tri FMNH 254250-254251, MNHN

1997.4931-4933.

Diagnosis. — An Ophryophryne of relatively large size, with relatively short shank, moderate-

sized tympanum, small head. Dorsal coloration ochre, rather clear, with distinct pattern.

Supraorbital horn distinct, no dermal protuberance bearing anus.

Description of the lectotype (fig. (A) Size and general aspect. — (1) Specimen of medium

size (SVL 39.1 mm), body elongate.

Source : MNHN, Paris

32 ALYTES 21 (1-2)

.— Ophrvophryne microstoma Boulenger, 1903. BMNH 1947.2.22.52, lectotype, adult male, SVL

39.1 mm. Dorsal view (top}; lateral view of head (bottom).

Source : MNHN, Paris

OHLER 33

(B) Head. — (2) Head very small, as wide (HW 9.5 mm) as long (HL 9.4 mm; MN 8.0 mm;

MFE 7.0 mm; MBE 4.6 mm), convex. (3) Snout truncate, protruding; its length (SL 3.56 mm)

shorter than horizontal diameter of eye (EL 4.54 mm). (4) Canthus rostralis rounded, loreal

region concave, acute in cross section. (5) Interorbital space convex, narrower (IUE 2.53 mm)

than upper eyelid (UEW 3.37 mm) and than internarial distance (IN 3.05 mm); distance

between front of eyes (IFE 5.6 mm) about two thirds of distance between back of eyes (IBE

8.8 mm). (6) Nostrils oval, with small flap of skin laterally, at equal distance to tip of snout

(NS 1.62 mm) and to eye (EN 1.56 mm). (7) Pupil indistinct, shape not visible. (8) Tympanum

(TYD 2.59 mm) rounded, rather distinct, more than half eye diameter; tympanum-eye

distance (TYE 1.94 mm) two thirds of its diameter. (9) Pineal ocellus absent. (10) Vomerine

ridge absent. (11) Tongue moderate, rounded, thick, emarginate, adhering largely to mouth

floor. (12) Supratympanic fold prominent, present from eye to shoulder.

hand (HAL 9.2 mm). (14) Fingers I, II and IV short and thin; finger III long and thin (TFL

5.57 mm). (15) Relative length of fingers, shortest to longest: I < II = IV< IIL. (16) Tips of

fingers rounded, not enlarged, without distinct grooves. (17) Fingers without dermal fringe;

webbing absent. (18) Subarticular tubercles absent. (19) Prepollex oval, distinct; palmar

tubercles indistinct; supernumerary tubercles indistinct.

(D) Hindlimbs. — (20) Shanks four times longer (TL 16.0 mm) than wide (TW 3.9 mm),

longer than thigh (FL 15.8 mm) and than distance from base of internal metatarsal tubercle

to tip of toe IV (FOL 15.4 mm). (21) Toe IV (FTL 7.4) about one third of distance from base

of tarsus to tip of toe IV (TFOL 23.3 mm). (22) Relative length of toes, shortest to longest: I

<IT< V < II < IV. (23) Tips of toes rounded, not enlarged, without distinct grooves. (24)

Webbing absent: 12-24 112-31113-41V 4-3 V(MTTF 4.47 mm; MTFF 5.39 mm; FTTF

7.89 mm; FFTF 8.03 mm). (25) Dermal fringe along toe V absent. (26) Subarticular tubercles

absent. (27) Inner metatarsal tubercle indistinct, its length (IMT non measurable, about

2 mm) 1.4 times in length of toe I (ITL about 2.8 mm). (28) Tarsal fold absent. (29) Outer

metatarsal tubercle, supernumerary tubercles and tarsal tubercle absent.

(E) Skin. — (30) Dorsal and lateral parts of head and body: snout, between eyes, side of

head and back shagreened; upper and lower part of flank shagreened covered with granules.

(1) Cephalic ridges absent. (32) Latero-dorsal folds, lateral line system and “Fejervaryan”

line absent. (33) Dorsal parts of limbs: forelimb, thigh, leg and tarsus shagreened, with

granules. (34) Ventral parts of head, body and limbs: throat, chest, belly and thigh smooth.

(35) Pectoral glands, femoral glands and glandular tympanic fold present.

(F) Coloration in alcohol. - (36) Dorsal and lateral parts of head and body: dorsal parts

of head and dorsum and upper part of flank dark ochre, homogeneous; dorsal folds with

some small darker spots: lower part of flank beige with ochre spots and some dark brown

spots; loreal and tympanic regions dark ochre; tympanum transparent; upper lip dark ochre.

(37) Dorsal parts of limbs: dorsal part of forelimbs, of thigh, of shank and of foot ochre with

darker bands; posterior part of thigh ochre with distinct dark brown spots. (38) Ventral parts

of head, body and limbs: ground light yellow, almost entirely covered by ochre on throat,

chest and upper vent; with large ochre flecks on central part of belly, leaving light yellow

ground out; posterior part of vent light yellow; thigh light yellow with dense ochre spots:

webbing ochre; macroglands whitish.

Source : MNHN, Paris

34 ALYTES 21 (1-2)

Coloration in life (specimen IEBR D231).- Dorsal parts of head and dorsum ochre, with

darker brown grey pattern and sand colored crests. Flanks brown grey and sand colored with

black spots, inferior part with white warts. Loreal region coppery brown, iris copper colored

with network of melanophores, tympanic region anteriorly dark reddish brown and poste-

riorly coppery brown, tympanum coppery brown, upper lip continuous with other parts of

side of head. Forelimbs, dorsal part of thigh, dorsal part of leg and foot brown grey and sand

color with black spots. Posterior part of thigh dark and whitish grey with brownish spots and

white warts: femoral glands ivory. Throat brownish, chest coppery brown with ivory colored

pectoral glands, belly coppery brown with white spots anteriorly, and whitish grey with dark

brown spots in its posterior part. Ventral side of thigh whitish grey with brown spots.

(G) Male secondary sexual characters. — (39) Traces indicate that nuptial spines have

been present on fingers I and II, with spines small, numerous. (40) Vocal sacs: not observed (to

preserve holotype). (41) Other male secondary sexual characters: dorsally of vent, presence of

a small fleshy flap.

Female sexual characters (adult female MNHN 1997.5259). - Large glandular circum-

voluted oviduct; creamy white, small sized, immature ovocytes (this female probably just laid

eggs).

Variation. — Sexual dimorphism of body size is rather pronounced (tab. 3). Females show

smaller heads and shorter shanks than males. The differences of head size of the holotype of

O. poilani (fig. 4) to the other females should be imputed to poor condition of this specimen,

as well as absence of supraorbital horn, a dermal structure which might easily be destroyed. In

life, the pupil of specimen IEBR D.231 showed horizontally enlarged diamond-shape,

thus confirming the horizontal shape of pupil mentioned in the original description of the

genus.

Distribution. — China, Thailand, Vietnam. Species previously not cited from Thailand.

Etymology. —- The Greek term microstoma describes the small mouth, characteristic for the

species of this genus.

Ophryophryne pachyproctus Kou, 1985

holotype, by original designation, YU

hushihe (alt. 1000 m), Mengla Xian

Ophryophryne pachyproctus Kou, 1985: 41. — Type-specimen:

A.8311032, adult male (not examined). — Tipe-locali

@1°29N, 101°33'E), Yunnan, China.

Specimens examined. — CHiNA: Zhushihe (alt. 1000 m), Mengla Xian (21°29°N, 101°33'E),

Yunnan: CIB A.8311038, male, paratype; VIETNAM: Nghe An Province: MNHN 2000.9087,

male.

Diagnosis. - Small-sized Ophryophryne, with relatively short shank, large tympanum, small

head. Dorsal coloration dark. Supraorbital horn small, anus terminal on a distinct dermal

protuberance.

Description of paratype CIB A.8311038 (fig. 5). - (A) Size and general aspect. — (1) Specimen

of rather small size (SVL 30.0 mm), body rather slender.

Source : MNHN, Paris

Table 3. - Measurement (mm) and per mille of snout-vent length (in parenthesis) of specimens of Ophryophryne microstoma Boulenger, 1903, including

lectotype of Ophrvophryne microstoma and holotype of Ophrvophryne poilani Bourret, 1937. Means and standard deviations are given between

square brackets.

Collection number | BMNH 1947.2.22.52 MNHN 1997.5258, 1997.5260; FMNH 254251 MNHN 1948.0113 Rd en ENT E S

Locality Mau Son, Ben En & Tam Dao, Dong-Tam-Ve, Ben En, Mau Son & Tam Dao,

Vietnam Vietnam Vietnam Vietnam

Eu ones Dre pan ont at MN) ?

Sex Adult male 3 adult males Adult female 6 females 3

Snout-vent length 39.1 381-444 [4144 3.17] 471 24.6-56.5 [46.1 4 11.5]

Head width 9.5 (243) 9.7-124 (255-285) (11.3 # 1.44 (273 4 16.0)} 11.5 (244) 8.1-15.1 (242-329) [12.1 # 2.47 (269 4 33.0)]

Head length 94 (240) 9.0-11.9 (236-270) [10.7 + 1.53 (258 4 19.1)] 9.5 (202) 74-14.0 (236-401) [11.5 # 2.59 (257 4 26.8)]

Tympanum diameter 2.44 (62) 2.59-3.37 (68-78) [3.07 à 0.42 (74 4 5.07)] 2.72 (58) 1.524,60 (59-81) [3.09 # 1.13 (66 + 8.71)]

Thigh 15.8 (404) 19.0-19.6 (428-469) [19.3 + 0.42 (448 à 29.0)] 19.8 (420) 11.5-25.2 (382-467) [19.6 4 5.79 (434 + 36.4)]

Shank 16.0 (419) 16.2-19.4 (423-464) [18.1 & 1.70 (438 4 23.0)] 19.4 (412) 11.4-23.9 (372-463) [18.9 4 4.17 (414 31.5)]

Foot 15.4 (394) 14.0-19.2 (367-438) [17.2 & 2.78 (413 à 39.2)] 18.6 (395) 9.6-20.8 (368-390) [17.2 + 4.58 (378 4 9.46)]

Source : MNHN, Paris

36 ALYTES 21 (1-2)

Fig. 4. — Ophryophryne microstoma Boulenger, 1903. MNHN 19480113, holotype of Ophrvophryne

poilani Bourret, 1937, adult female, SVL 47.1 mm, Dorsal view (top); lateral view of head (bottom).

Source : MNHN, Paris

OHLER 37

.— Ophryophryne pachyproctus Kou, 1985. CIB A.8311038, paratype, subadult male, SVL 30.0 mm.

Dorsal view (top): lateral view of head (bottom).

Source : MNHN, Paris

38 ALYTES 21 (1-2)

(B) Head. — (2) Head very small, slightly wider (HW 7.9 mm) than long (HL 7.4 mm;

MN 6.5 mm; MFE 5.3 mm; MBE 2.7 mm), flat. (3) Snout truncate, very protruding, its length

(SL 2.9 mm) shorter than horizontal diameter of eye (EL 3.4 mm). (4) Canthus rostralis

sharp, loreal region concave, acute in cross section. (5) Interorbital space flat, narrower (IUE

1.5 mm) than upper eyelid (UEW 2.2 mm) and internarial distance (IN 2.4 mm); distance

between front of eyes (IFE 4.4 mm) two third of distance between back of eyes (IBE 6.7 mm).

(6) Nostrils rounded, without flap of skin laterally, at equal distance from eye (EN 1.0 mm)

and tip of snout (NS 1.0 mm). (7) Pupil indistinct diamond. (8) Tympanum (TYD 2.2 mm)

very distinct, rounded; smaller than half diameter of eye; tympanum-eye distance (TYE

1.2 mm) half its diameter. (9) Pineal ocellus absent. (10) Vomerine ridge absent. (11) Tongue

rounded, not emarginate. (12) Supratympanic fold distinct, present from eye to above

shoulder, posterior part enlarged.

(HAL 7.9 mm), not enlarged. (14) Fingers long and thin; finger III long, thin (TFL nm). (15)

Relative length of fingers, shortest to longest: I < II < IV< III. (16) Tips of fingers rounded,

slightly enlarged. (17) Fingers without dermal fringe; webbing absent. (18) Subarticular

tubercles indistinct, all absent. (19) Prepollex oval, distinct; a single, oval palmar tubercle:

supernumerary tubercles absent.

(D) Hindlimbs. — (20) Shanks four times longer (TL 13.1 mm) than wide (TW 3.4 mm),

a little shorter than thigh (FL 14.3 mm), but longer than distance from base of internal

metatarsal tubercle to tip of toe IV (FOL 12.6 mm). (21) Toe IV long (FTL »m; TFOL

19.4 mm). (22) Relative length of toes, shortest to longest: I < II < V < III < IV. (23) Tips of

toes rounded, scarcely enlarged; discs absent. (24) Webbing absent (MTTF 6.1 mm; MTFF

6.2 mm; FTTF 7.1 mm; FFTEF 6.50 mm). (25) Dermal fringe along toe V absent. (26)

Subarticular tubercles absent. (27) Inner metatarsal tubercle long, scarcely distinct, its length

(MT 1.3 mm) 1.77 times in length of toe I (ITL 2.3 mm). (28) Tarsal fold absent. (29) Outer

metatarsal tubercle, supernumerary tubercles and tarsal tubercle absent.

(E) Skin. — (30) Dorsal and lateral parts of head and body: snout, between eyes and side

of head shagreened; back shagreened with fine glandular folds and few glandular warts

posteriorly; flanks with few glandular warts. (31) Cephalic ridges absent. (32) Latero-dorsal

folds, lateral line system and “Fejervaryan” line absent. (33) Dorsal parts of limbs: forelimb,

thigh, leg and tarsus shagreened. (34) Ventral parts of head, body and limbs: throat, chest,

belly and thigh smooth. (35) Femoral and pectoral glands small, present; supra-anal protu-

berance distinct, bearing at its distal part a ventral cloacal opening.

(F) Coloration in alcohol. — (36) Dorsal and lateral parts of head and body: dorsal parts

of head and dorsum almost uniformly dark brown; flanks slightly clearer, their lower parts

clear greyish brown with few dark spots, more or less in a line; loreal region dark brown:

tympanum orange brown; tympanic region dark brown; upper lip dark brown with an

indistinct darker band. (37) Dorsal parts of limbs: dorsal part of forelimbs, of thigh, of shank

and of foot brown with fine dark brown transversal bands; posterior part of thigh light

greyish brown with a large dark triangular perianal zone extending to knees. (38) Ventral parts

of head, body and limbs: throat brown; margin of throat dark brown; chest and upper part of

belly brown: lower part yellowish with dark brown flecks: thigh yellowish; macroglands

whitish.

Source : MNHN, Paris

OHLER

Table 4. — Measurements (mm) and per mille of snout-vent length (parenthesis) of two specimens,

including paratype, of Ophryophryne pachyproctus Kou, 1985.

Collection number CIB A.8311038 MNHN 2000.9087

Locality Zhushihe, China Nghe An, Vietnam

Status Paratype Additional material

Sex Subadult male Adult male

Snout-vent length 30.0 28.9

Head width 7.9 (263) 8.3 (287)

Head length 74 (247) 8.3 (287)

Tympanum diameter 22(73) 2.59 (90)

Thigh 143 (478) 11.7 (405)

Shank 13.1 (437) 12.0 (415)

Foot 12.6 (420) 12.1 (419)

Coloration in life (according to the original description). - Dorsal surface grey white or

brown grey; a dark, distinct or indistinct triangle between eyes; side of head, lips, throat, chest

and belly dark colored; sides of head and body bearing 4-6 black spots; posterior part of

thighs and ventral surface of hands black brown, fore and hind legs dark colored, forearm and

thigh generally with a large black spot; dorsal surface of hands and external surface of legs

usually with black spots, palm and upper surface of hands orange red, tips of fingers light red,

tips of toes dark. When put to alcohol, coloration gets darker, reddish parts turn white.

(G) Male secondary sexual characters. - Not observed (male not adult?).

Female sexual characters. - Not observed.

Variation. — The measurements of the two specimens studied are given in tab. 4. Only larger

samples might inform about the significance of the differences observed between the specimen

from Vietnam and from China.

Dis!

ibution. — China (Yunnan) and Vietnam (Nghe An Province).

Etymology. - The term pachyproctus, “having a thick vent”, is derived from the Greek pakhys,

“thick” and proktos, “vent”.

KEY TO THE KNOWN SPECIES OF THE GENUS Ophryophrvne

18: Palmar tuberclé distinct. : 024% + à emhetée states ee OR ent age alete ste pe dB 2

1b. Palmar tubercle indistinct .

Za. Dorsal skin with glandular ‘wart and horny pinules: cloacal appendag

adult males: SVL 35.5-43.0 mm. TL 18.4-19.0 mm; adult female: SVL 53.5 mm, TL

25.6mm ©. hansi

2b. Dorsal skin shagreened; cloa 0.0 mm, TL

12.0-13.0 mm ....................44,44.4.. esse sesssese ©. pachyproctus

Source : MNHN, Paris

40 ALYTES 21 (1-2)

3a. Dorsum dark brown; posterior part of shank with fine brownish marbling; body size

small: adult males SVL 32.0-34.8 mm, TL 15.4-15.5 mm; adult females SVL 41.4-45.8

mm, TL 17.4-18.0 mm; vocal sacs absent .................................. ©. gerti

3b. Dorsum greyish-brown; posterior part of shank with large brown spots; body size large:

adult males 38.1-44.4 mm, TL 16.0-19.4 mm; adult females SVL 46.3-56.5 mm, TL

18:1-23.9 mn; vocal: Sacs present. : ns à 5 4 0 4 sois diere n à 5 a ae e ola ere murs eus dl O. microstoma

DISCUSSION

In Ophryophryne, several trends, which can also be observed in the sister-group

Xenophrys, occur, such as diminutive size, reduction of palpebral horn and variation of

structure in dorsal skin. Characters used to differentiate these two taxa are mostly linked to

reduction of the size of the of skull in Ophryophryne.

The geographic distribution of the genus Ophryophryne includes Laos, Thailand, Viet-

nam (northern and central), as well as China (Yunnan and Guangdong Provinces) (fig. 6). It

is completely included in the distributional range of its sister-group Xenophrys, but shows a

much reduced extension. This range may still be increased by further observations, when field

data are accumulated, as all over its range only few specimens have been collected since its

description a century ago.

The species referred to the genus show a complex pattern of distribution. The range of ©.

pachyproctus seems to be similar to that of ©. microstoma, as are the ranges of ©. hansi and O.

gerti. The distribution areas of both species couples are allopatric with a large area without

data. The new species ©. hansi is known only from its type-locality. However, additional field

data are likely to extend its range, as has occurred for ©. pachyproctus, which was known for

some years from its type-locality only, but now can be considered as a member of the fauna of

northern Vietnam. For ©. microstoma the distributional range can be extended to Thailand.

There is no evidence for syntopic occurrence of two or more species of Ophryophryne. ©. gerti

and ©. hansi have the same collecting locality, but the females of ©. gerti were collected in a

swampy area, whereas ©. hansi males were actively calling along a stream (INGER et al., 1999).

This does not confirm that the two species occur in different habitats, as obviously one species

was reproductively active, but not the other. Mature O. microstoma were collected along a

small stream (personal observation), as was described for ©. hansi. O. microstoma and ©.

pachyproctus share a large distribution range. On the opposite, ©. pachyproctus and ©. gerti do

not share distribution range according to the current data.

The synonymy of ©. poilani is based on morphological data, especially on measurement

of tibia length, which should be relatively precise despite the poor state of preservation of its

type-specimen. ©. hansi is the species showing the largest head size, but the type of ©. poilani

has a head width smaller than all studied specimens of ©. hansi. The type-specimen of ©.

poilani ly is not conspecific with the specimens that INGER et al. (1999) mentioned as ©.

poilani. This type-specimen has definitely shorter tibia than ©. hansi, but its tibia length is

included in the range of variation observed in ©. microstoma.

Source : MNHN, Paris

OHLER 41

OPHRYOPHRINE

[ gert

Le microstoma

D pccpproctus

@ 5:

——

Fig. 6. - Collection localities of species of the genus Ophryophrvne Boulenger, 1903

Source : MNHN, Paris

4 ALYTES 21 (1-2)

Ophryophryne hansi is relatively distinct from the other three species and shows plesio-

morphic characters, such as relatively large head (320 p.m. of SVL) and large body size. The

cloacal appendage of ©. pachyproctus is a unique character, present only in this species. The

tubercles on the vent observed in some of the specimens of other species of Ophryophryne do

not form a cone including the anal opening. They form dermal structures which surround the

anal opening. Size reduction is probably independent in ©. pachyproctus and O. gerti as both

only share this character. The two species are distinct for presence of palmar tubercles, length

of inner toe and size of tympanum.

Ophryophryne species show an important reduction of head size. These species have a

small mouth and a tongue fixed to the mouth floor which does not allow them to capture and

swallow large-sized prey. Such morphological adaptations can be observed in various groups

of frogs and has been studied in Rhinophrynus Duméril and Bibron, 1841 and in the

Microhylidae Noble, 1931 (TRUEB & Gans, 1983: BLUM & MENZIES, 1988). À small head is

usually linked to a special nutritional mode and a fossorial life (DUELLMAN & TRUEB, 1985).

The shape of the hindlimbs, in particular the feeble development of the inner metatarsal

tubercle, does not indicate any particular adaptation to fossorial life in Ophryophryne. The

particular buccal anatomy might be linked to the nutritional mode of these species. Their

digestive tract is well developed, showing a large gastric pouch, but it was found empty in the

specimens examined (personal observations). Nothing is known about foraging in nature. In

laboratory conditions they accept young insects as well as earthworms (R. Boistel, unpub-

lished observations). The structural similarity of buccal anatomy in Ophryophryne and

Xenobatrachus and Xenorhina (Microhylidae) might indicate that these frogs have similar

diets. Xenobatrachus and Xenorhina feed on earthworms (BLUM & MENZIES, 1988) and have

well developed vomerine teeth on the buccal roof, which seem to be crucial for ingestion of

this food. Such structures are absent in Ophryophryne. New data on the biology of these

species will be necessary to understand the ecological role and niche of small mouthed

megophryids in the amphibian communities of southeast Asian primary forests.

RÉSUMÉ

L'étude des spécimens-types et des spécimens récemment collectés de crapauds du genre

oriental Ophryophryne Boulenger, 1903 (Megophryidae) nous mène à la redéfinition des taxa

reconnus et de la définition de deux nouvelles espèces dont une est rapportée du Vietnam et du

Laos et l’autre seulement du Vietnam. Ophryophryne poilani Bourret, 1937 est mis en

synonymie d’'Ophryophryne microstoma Boulenger, 1903. L’aire de distribution de l'espèce O.

microstoma, qui incluait le Vietnam et la Chine, est élargie à la Thaïlande. Ophrrophryne

pachyproctus Kou, 1985 est confirmé du Vietnam.

Source : MNHN, Paris

OHLER 43

ACKNOWLEDGEMENTS

T'express all my thanks to the staff of the Field Museum for sending me in loan the specimens so

promptiy. Barry T. Clarke and E. Nick Arnold welcomed us at the British Museum in London very

warmly, giving us the possibility to study this important type-collection. Fei Liang, Ye Chuanyuan,

Zhong Shengxian and Wang Yuezhao from the Academia Sinica at Chengdu friendly invited me to

examine the Chinese frogs. Stéphane Grosjean, Julio Mario Hoyos, Steven Swan and all the other

volunteers of Frontier Vietnam participated in the field work. IEBR, Hanoi, is thanked as our collabo-

ration partner for collection and exportation permits in Vietnam. Edouard-Raoul Brygoo, Roger Bour,

Renaud Boistel and the staff of the Laboratoire des Reptiles et Amphibiens, Paris, were helpful at various

stages of preparation of this paper. My friendly recognition to Xiaohua Tu, who kindly translated the

Chinese descriptions. Thanks to Alain Dubois and Craig Guyer for comments on the manuscript. This is

publication N° 03-78 of the Programme Pluriformation “Faune et Flore de l'Asie du Sud-Est” of the

Paris Museum (N° 03-77, see DEUVE, 2003).

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from New Guinea with description of nine new species. A/ytes, 7: 125-163.

BouRRET, R., 1937. —- Notes herpétologiques sur l’Indochine française. XIV. Les Batraciens de la

collection du Laboratoire des Sciences Naturelles de l'Université. Descriptions de quinze espèces

ou variétés nouvelles. Annexe Bull. Inst. publ. Hanoi, 1937: 5-56.

BOULENGER, G. A., 1903a. - Description of three new Batrachians from Tonkin. Ann. Mag. nat. Hist., (7),

12:186-188.

—— 1903b. — IV. Reptilia and Batrachia. Zool. Rec., 40: 1-38.

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Carabus et Cychropsis d'Asie (Coleoptera, Carabidae). Coléoprères, 9: 215-240.

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

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

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

Dugois, A. & OHLER, A., 1998. - A new species of Leprobrachium (Vibrissaphora) from northern

Vietnam, with a review of the taxonomy of the genus Leptobrachium (Pelobatidae, Megophryi-

nae). Dumerilia, 4: 1-32.

DUELLMAN, W. E. & TRUEB, L., 1985. - Biology of amphibians. New York, McGraw-Hill: i-xix + 1-670.

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NOBLE, G. K.

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Pyxicephalinae). Aves, 13: 141-166

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Source : MNHN, Paris

Alytes, 2003, 21 (1-2): 45-58. 45

A redescription of the external

and buccopharyngeal morphology

of the tadpole

of Ophryophryne microstoma

Boulenger, 1903 (Megophrvidae)

Stéphane GROSIEAN

Vertébrés (Reptiles & Amphibiens),

JSM 0602 Taxonomie & Collections,

Département de Systématique & Evolution,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

<sgrosjea@mnhn.fr>

The external morphology and the buccopharyngeal features of the

tadpole of Ophryophryne microstoma Boulenger, 1903 are redescribed.

Morphometric data are provided. Few morphological deviations as compa-

red to the previously described tadpoles from China are noted. The

taxonomic status of the genus Ophryophryne within the Megophryidae is

discussed in the light of the tadpole morphology.

INTRODUCTION

Ophryophryne Boulenger, 1903 is a small group of species from southeast Asia with a

rather limited distribution, still little studied, and including now four species after a recent

taxonomic revision of the genus (OH: 2003). The status of Ophryophryne has always been

confused. Though its descriptor noticed its overall resemblance with members of the Peloba-

tidae (BOULENGER, 19034), he included this genus within the Bufonidae (BOULENGER, 1903b)

and it is still yet considered as such by some authors (NGUYEN & Ho, 1996). It was then put

within the Pelobatidae by NoBce (1926). The status of the taxon Ophryophryne among the

Megophryidae has already been discussed in previous works (NOBLE, 1926; DuBois, 1980,

1987; Rao & YANG, 1997; DuBois & OHLER, 1998). Rao & YANG (1997) rose Ophryophryne,

Tian & Hu, 1983, Brachytarsophrys Tian & Hu, 1983, Megophrys Kuhl & Van

Hasselt, 182 1864 to the generic level on the basis of cytological,

morphological and ecological characters. DUBoIs (1980) put first Ophryophryne within the

genus Megophrys then on the basis of new subdivisions within the genus Megophrys S. Str.

re-evaluated his proposition and gave Ophryophryne a generic status (Dugois, 1987). Though

its position seems now to be clearly within the Megophryidae and the genus Xenophrys has

Atympanophr.

and Xenophrys Günth

Source : MNHN, Paris

46 ALYTES 21 (1-2)

been identified as its sister-group by several authors (TIAN et al., 1985; RAo & YANG, 1997), its

generic or subgeneric status (within the genus Megophrys) is still discussed (MATSUI in FROST,

1985; YE et al., 1993; MANTHEY & GROSSMANN:; 1997).

Ophryophryne adults bear autapomorphies, which distinguish them from other members

of Megophrys (sensu lato including four subgenera, i.e., Atympanophrys, Brachytarsophrys,

Megophrys, Xenophrys): absence of maxillar and vomerine teeth and presence of a horizontal

pupil. However, are these characters sufficient to consider Ophryophryne as a valid genus?

Anuran tadpoles are, as a general rule, well intergenerically differentiated on the basis of the

buccopharyngeal morphology (VIERTEL, 1982; Grosjean, unpublished data). The aim of this

work is, besides a description of the morphology and buccopharyngeal features of the only

known tadpole of the genus, to compare these data with those of known tadpoles of the four

subgenera of the genus Megophrys, in order to discuss the status of Ophryophryne in the light

of larval data.

MATERIALS AND METHODS

A total of 52 tadpoles, all Ophryophryne microstoma Boulenger, 1903, were collected in

Ben En National Park, Thanh Hoa Province, Vietnam (19°30'-40°N, 105°21°-35’E) during

August 1997. This national park is situated in a region of low hills surrounding the Song Muc

Lake. Altitude ranges from 20 to 497 m above sea level, with most areas being below 200 m.

The vegetation is that of a tropical semi-evergreen forest partly degraded by human activity.

The climate is subtropical, with the heaviest rainfall between July and October. The average

temperatures between 1961 and 1990, in the months of July and August, were 28.9 and 27.8°C

respectively (TORDOFF et al., 1997).

Some tadpoles were preserved soon after capture whereas others were reared in bowls of

24 cm of average diameter and 10 cm depth, and were fed with baby fish food (TetraMin).

Tadpoles in developmental stages 25-43 (GOsNER, 1960) were preserved in a mixture of equal

parts of 4% formaldehyde and 70 % ethanol (GriLLrrscH, 1984). Some tadpoles reached

metamorphosis assuring the identity of the species and were preserved in the same solution.

This material is deposited in the collections of the Muséum national d'Histoire naturelle of

Paris (MNHN 1999.0521-0572).

Morphological terminology follows ALTIG & MCDiaRMID (1999), developmental stages

were determined according to GOsnER (1960) and terminology of buccopharyngeal features

follows WassERSUG (1976). Measurements were taken with a graduated ocular attached to a

stereomicroscope except for the total length and the distance from opening of vent to tip of

tail which were measured with a hand caliper. The distance from tip of snout to nares was not

taken into account because the tip of snout was hidden beneath the oral funnel. For exact

location of measurement landmarks, see GROSIEAN (2001: fig. 2), except for tip of snout which

is taken from the point where the funnel originates with the head anteroventrally. Drawings

were made with the aid of a camera lucida.

Preparation for SEM examination (JEOL JSM 840) comprised dehydratation (ethanol),

critical-point-drying (liquid carbondioxide) and gold sputter surf

Source : MNHN, Paris

GROSJEAN 47

RESULTS

ECOLOGICAL CONSIDERATIONS

The tadpoles studied were collected along a little stream running in the forest. The stream

was 1-2 m wide and its depth ranged from a few centimetres in fast-flowing parts to up to 70

em in the deepest pools. The bottom was mainly made up of rocks and stones, and of sand

covered by dead leaves in the quiet pools. These tadpoles belong to the lotic-neustonic

ecomorphological guild of ALTIG & JOHNSTON (1989). They were found in the lower part of

the stream where the cover of vegetation was more open, in shallow water, hidden between the

stones with their funnel open at the interface air-water. Tadpoles were also found near the

banks of the stream, where the water ran between the stones, the tadpoles not being visible (as

already noticed for tadpoles of the genus Megophrys by HoRA, 1928) and in a little natural

dam made up by branches and dead leaves. AII developmental stages (from stage 25 to imago)

were found all together at the same time and in the same part of the stream. This suggests that

the mating and/or spawning period might be prolonged (parsimonious assumption) or that

larval development is very slow and extends all over the year rather than being limited to a

circumscribed period (in accordance with other observations). Indeed the tadpoles reared for

one month did not show evolution (except for the latest stages). Slow development is not rare

among Megophryidae and was observed in other genera like Leptobrachium Tschudi, 1838,

Leptolalax Dubois, 1980, Scutiger Theobald, 1868 and Oreolalax Myers & Leviton, 1962

(SMirH, 1917; Liu & HU, 1960; CHEN et al., 1984; ZHao et al., 1994; pers. obs.) and in

rheophilous species in general. Few tadpoles of Leptolalax sp. were found among them.

Numerous tadpoles of Leptobrachium chapaense (Bourret, 1937) lived in the pools of the

same stream but not in the same niche (no tadpole of Ophryophryne microstoma was found in

pools). Clutches of Rana (Sylvirana) nigrovittata (Blyth, 1855) were found in crab holes, in

the bank.

During the two months spent in the field, the reproduction of the species which produced

the tadpoles was not observed: advertisement calls were not heard and neither mates in

amplexus nor egg clutches were found. Non-calling adults of O. microstoma were occasionally

found among the vegetation of the bank and on emerged rocks in the stream bed.

DESCRIPTION OF TADPOLE

External morphology

Gross morphometric parameters (SVL and TL) of all tadpoles are presented in table 1.

The following description is based on four tadpoles at stages 35-38 (MNHN 1999.0532-0535),

except where specified. Detailed morphometric data are presented in table 2. In dorsal view

(fig. la), body elliptical. Eyes of moderate size (diameter about 0.1 time body length), bulging,

separated by a distance which equals about 1.3 times the internarial distance, directed and

positioned laterally, visible in ventral view. Nares tubular, of moderate size, directed laterally

Source : MNHN, Paris

Table 1.- Variation of snout-vent length (SVL) and total length (TL) with stage (GOSNER,

1960) in tadpoles of Ophryophryne microstoma. Number of tadpoles (7) examined,

ALYTES 21 (1-2)

mean value + standard deviation in mm (range in parentheses).

Stage n SVL TL

25 24 5.87 + 0.74 (5.06-8.55) 17.49 4 2.19 (15.00-24.30)

26 7 7.83 + 0.50 (7.06-8.55) 22.75 + 1.34 (20.65-24.75)

27 2 8.62 + 0.28 (8.42-8.82) 26.35 + 0.07 (26.30-26.40)

28 2 9.14 1.21 (8.29-10.00) 27.45 + 2.05 (26.00-28.90)

31 3 9.56 + 0.20 (9.34-9.74) 27.28 + 0.88 (26.35-28.10)

34 1 10.92 31.05

35 1 947 28.10

37 2 10.66 + 0.00 (10.66-10.66) 30.70 + 3.54 (28.20-33.20)

38 1 10.66 32.40

40 1 11.18 31.60

a 5 9.87 0.57 (9.21-10.53) 28.42 + 1.54 (25.85-29.65)

4 1 10.26 29.75

43 1 11.18 25.55

Table 2.— Morphometric data for tadpoles of Ophryophryne microstoma in avanced

developmental stages (35-38, GOSNER, 1960). BH, maximum body height; BW,

maximum body width; ED, maximum eye diameter; HT, maximum tail height; LF,

maximum height of lower tail fin, MNHN, collection number, Muséum national

d'Histoire naturelle, Paris; #m, no measurement; NN, internarial distance; NP, naro-

pupillar distance; PP, interpupillar distance; SS, distance from tip of snout to opening of

spiracle; SU, distance from tip of snout to insertion of upper tail fin; SVL, snout-vent

length; TL, total length; UF maximum height of upper tail fin; VT, distance from vent

opening to tip of tail.

Stage MNEN SVL TL ss SU VT HT UF

35 1999.0535 9.47 28.10 4.21 11.71 1930 4.99 128

37 1999.0534 10.66 28.20 4.67 10.79 18.60 4.86 1.24

37 1999.0533 10.66 33.20 444 14.34 21.50 4.86 131

38 1999.0532 10.66 32.40 4.93 8.03 21.70 4.99 1.34

Stage MNEHN LF BH BW ED PP NN NP

35 1999.0535 1.40 nm nm 101 3.24 2.61 0.75

37 1999.0534 1.18 4.15 4.73 1.09 3.50 2.72 0.93

37 1999.0533 1.18 4.15 4.73 1.09 3.64 2.71 0.93

38 1999.0532 1.24 4.08 4.99 1.10 3.48 2.49 0.96

Source : MNHN, Paris

GROSJEAN 49

and horizontally, and positioned rather dorsally, closer to pupils than to tip of snout. In

profile (fig. 1b), body depressed. Spiracle sinistral, conical, very short, attached to the body

wall except its tip which is free, positioned just beneath the longitudinal axis, oriented

posteriorly. Spiracular opening situated slightly closer to pupil than to the end of the body,

and at a level situated just beneath the apex of the caudal myotomes (fig. 1b) to between the

apex of caudal myotomes and the hindlimbs; its opening from rounded to oval. Tail muscu-

lature strong, gradually tapering, almost reaching tail tip (very near to the end). Tail fins

shallow, moderately developed, not extending onto body, dorsal fin slightly higher than

ventral fin in the distal half of the tail; free margin of dorsal fin horizontal and very shallow

in the proximal half of the tail then following tail musculature; free margin of ventral fin

parallel to tail musculature. Point of maximum height of tail located just before halfway of the

tail, in the proximal part, tail tip bluntly pointed. Vent tube of moderate size, medial, tubular

(often slightly bulging in the middle), directed posteriorly, not linked to ventral tail fin,

opening medial. Neither skin glands nor neuromasts visible.

Oral disk subterminal (fig. 2), lips expanded vertically forming a dorsally oriented

funnel; lateral corners pronounced; upper lip smaller than lower; lips lacking keratodonts, but

furnished with a few short, low ridges (variable number among specimens), more densely

arranged on the upper labium than on the lower one: arrayed in 20 longitudinal rows (in

mean) and 2 (on the upper labium) to 4 (on the lower labium) transversal rows (without clear

limits); no marginal papillae. Width of expanded funnel about 50 % of body length (35 %

when folded, which is often the case in the preserved specimens). At stage 40 the funnel began

to be resorbed, at stage 42 the posterior part of the funnel disappeared while the anterior part

still persisted. At stage 43 the funnel entirely disappeared. Lower jaw sheath (fig. 3a) straight

anteriorly, radically curved backward laterally, entirely white, its free margin bearing fine,

pointed, hair-like serrations; upper jaw sheath (fig. 3b) nearly straight, notched medially,

bearing fine and elongate serrations only in the medial third of its free margin, brown grey

with a white margin; both beaks soft.

Colour in preservative

Dorsal side of body and upper part of flanks brownish-khaki. Lower part of flanks

weakly mottled with the same colour; ventral side of head (from snout to posterior part of

eyes) intensely coloured. Belly white, intestine not visible through body wall. Caudal muscle

weakly coloured (more on upper than on lower portion, with emphasis on the myotome

apexes); tail fins translucent with few spots (more on upper than on lower fin); half to a

quarter of distal part of tail often intensively coloured in dark brown. Oral funnel greyish

with brownish-khaki papillae. There is considerable intrapopulational variation in colour:

certain individuals are almost unpigmented whereas others are strongly coloured, others have

à tail strongly coloured except in the distal quarter.

Tadpole at stage 45 (just metamorphosed, MNHN 1999.0524, SVL 11.97 mm) in

preservative. Head and dorsum brown; flanks and dorsal part of thigh and tibia with large

white tubercles. Dorsal folds present. Supratympanic fold underlined with white above and

black below, the black line extending as far as the armpit. Limbs white below, faintly tinted

with brown above. The above features began to appear from stage 42 on.

Source : MNHN, Paris

0ç

(T-D 17 SALATV

Fig. 1. Ophryophrynemicrostoma(stage38, TL 32,1 mm; MNHN 1999.0532):a, dorsal view:b,lateralview.

Source : MNHN, Paris

GROSJEAN 51

Fig. 2. Funnel-like mouth of Ophryophryne microstoma (stage 38; MNHN 1999.0532). Scale line: 1 mm.

Internal buccal features

The description is based on a single tadpole at stage 37 (MNHN 1999.0534).

Buccal floor (fig. 4a): a soft lower jaw sheath present at entry of buccal cavity, straight

anteriorly, radically curved backward laterally; a nodule on the inner wall of the beak at the

level of curvatures. Prelingual arena deep, with two successive pairs of palps directed

medially, followed by a fifth palp in median position. These structures are thick, fleshy and

smooth lobes, concave anteriorly, the first pair reaching almost the lower jaw sheath being

visible without dissection. Tongue anlage distinct, not very prominent, drop-shaped, position-

ed on an elevated mass; lingual papillae absent. Buccal floor arena forming an elongate oval,

in a depression, without buccal floor arena papillae but its anterior half bounded laterally by

a thick ridge on each side, these two ridges merging with the elevated part bearing the tongue

anlage anteriorly. The ridges ending posteriorly in front of the buccal pockets with elevated

knobs each wrapped by an anteriorly concave flap; posterior half of arena bounded by an

undulating ridge on each side. Anterior part of arena with a median groove of about a third

of the length of the buccal floor arena, 2-5 pustules on each side between the ridges and the

median groove; posterior part of arena lacking ornamentations. Buccal pockets short,

transversely oriented, arched anteriorly, with fine openings: area anterior to buccal pockets

with 1-3 pustules on each side: area posterior to buccal pockets with less than 10 small

Source : MNHN, Paris

52 ALYTES 21 (1-2)

Fig. 3. Close-up view of the jaw sheaths of Ophryophryne microstoma (stage 37; MNHN 1999.0534): a,

lower part; b, upper part. Scale line: 0.1 mm.

papillae. Ventral velum slightly undulated, its medial part extending backward; two minor

projections laterally, medial notch absent; spicular supports present laterally; secretory pits of

ventral velum absent; glottis slightly exposed. Branchial baskets large; a single common filter

cavity on each side; filter rows wide with tertiary or higher-order ramifications, mesh size

about 100 ym.

Source : MNHN, Paris

GROSJEAN 53

Fig. 4. — Ophrvophryne microstoma (stage 37: MNHN 1999.0534): a, buccal floor arena; b, buccal roof

arena.

Source : MNHN, Paris

54 ALYTES 21 (1-2)

Buccal roof (fig. 4b): upper jaw sheath nearly straight, bearing fine and elongate

serrations only in the medial third. Prenarial arena wide, trapezoidal, in medial position a

large U-shaped ridge with a posteromedial knob, its anterior arms almost reaching the beak:;

a little knob just posterior to the U-shaped ridge (hidden by the median ridge); on the wall of

the prenarial arena two pairs of well pronounced dorsoventrally compressed ridges aligned

longitudinally. Choanae transverse, short; anterior narial wall smooth, with a short, stocky

papilla on the internal end and another even stockier on the external end; narial valve smooth

with a small stocky triangular projection located rather medially. Postnarial arena bounded

laterally by a pair of pronounced ridges (which could be the homologue of postnarial

papillae), fused anteriorly with the medial posterior wall of choanae, ending behind the

medial ridge, their posterior ends curved medially. Median ridge a tall straight projection

directed anteriorly plus three little prominent lobes, reaching as far as the posteromedial knob

of the prenarial U-shaped ridge. Postnarial arena filled by the body of the median ridge.

Lateral ridge papillae a large dorsoventrally compressed flap; above their anterior part a

similar but smaller flap. Buccal roof arena with about fifty pustules arranged more densely

posteriorly; no buccal roof arena papillae but two large nodules fused on each side bounding

anterolaterally the buccal roof arena and abutting the ridges lateral to the medial ridge. About

ten pustules on each lateral wall of the buccal roof, at the level of the anterior half of the

buccal roof arena. Posterolateral ridges present but not very prominent laterally and medially,

and rather distinct lateromedially. Glandular zone rather indistinct, absent medially; secre-

tory pits absent. Dorsal velum curved ventrally, interrupted medially.

DISCUSSION

Among the four species in the genus Ophryophryne known at present, only the tadpole of

©. microstoma has been described previously. I redescribe here the tadpole of the species based

on specimens belonging to a much more southern population.

Liu & HU (1962) reported on the external morphology of the tadpole from Kwangsi

Province, China. These authors did not specify the developmental stage of the tadpole upon

which their description was based but just noticed that it bore buds of hind limbs and had a

total length of 33 mm. The only tadpole in our sample which reached this size was a tadpole

of stage 37. Hence, tadpoles in equivalent developmental stages were larger in the Chinese

sample than in the Vietnamese sample. The other characters differing between the two

samples were: oral disk bigger, eyes larger and tip of tail rounded (not bluntly pointed as in

this sample) in the Chinese population. Differences were also observed in the number of

papillae of the funnel: 15 longitudinal rows and three transversal rows in the Chinese

population, 20 longitudinal rows (in mean) and 2-4 transversal rows (without clear limits) in

our sample. Finally, the coloration varied in a few points: in the tadpole of the Vietnamese

population the upper fin was rimmed with dark margins in its anterior part and the anterior

part of the lower fin was coloured with large dark marks whereas it was white in the Chinese

one. The imago collected by Liu & Hu (1962) was comparatively large (14 mm body length

and 3.5 mm tail, whereas an imago entirely metamorphosed collected in the Vietnamese

population was about 12 mm in length). However in both cases the adult characters appeared

Source : MNHN, Paris

GROSJEAN 55

early since the skin folds and the tubercles were present before resorption of tail. The eyelid

processes seemed to appear after total resorption of tail.

The buccopharyngeal features of a specimen from Longjin, Guangxi Province, China

were reported by HUANG et al. (1991). Their illustration showed some differences with our

data. In the buccal floor of the HUANG et al. (1991) specimen the posterior pair of palps as

well as the single medial palp are shorter than in our specimen and a medial notch is present.

Within the buccal roof, our specimen had a bigger prenarial ridge, two papillae on anterior

narial wall and another on the narial valve whereas only one was present on the anterior narial

wall in the Longjin specimen; moreover, pustulations in the buccal roof arena were present in

the Vietnamese tadpole.

The tadpoles of the genus Megophrys sensu lato (including the subgenera Arympa-

nophrys, Brachytarsophrys, Megophrys and Xenophrys) have been little studied and their

detailed descriptions are rare: (1) for the external morphology, M. (X.) aceras (Boulenger,

1903) (LaiDLAW, 1900; ANNANDALE, 1912, 1917; SmirH, 1926 [all under the name M.

montana)), M. (X.) major (Boulenger, 1908) (ANNANDALE, 1912), M. (X.) parva (Boulenger,

1893) (ANNANDALE, 1912), M. (X.) boettgeri (Boulenger, 1899) (ANNANDALE, 1917; POPE,

1931; Liu, 1940), M. (X.) longipes (Boulenger, 1885) (LEONG & CHoU, 1998), M. (X.) minor

Stejneger, 1926 (Liu, 1950), M. { M.) montana Kuhl & Van Hasselt, 1822 (BOULENGER, 1908),

M. (M.) nasuta (Schlegel, 1858) (INGER, 1985); (2) for the buccopharyngeal features, M. (B.)

carinensis (Boulenger, 1899) (HuANG et al., 1991), M. (À. ) minor (WASSERSUG, 1980; HUANG

et al., 1991), M. (M.) nasuta (INGER, 1985), M. (X.) omeimontis Liu, 1950 (HUANG et al.,

1991) and M. (A.) shapingensis Liu, 1950 (HUANG et al., 1991). The tadpoles of the genus

Megophrys sensu lato are very similar to those of Ophryophryne and nothing allows to

distinguish them externally. The most peculiar structure of these animals, the funnel-like oral

disc surrounding the mouth, is very conservative and its typical morphology isencountered in

all species. The small differences reported in terms of variation in number of rows or of ridges

are not greater between Ophryophryne and Megophrys sensu lato than between the members

of the genus Megophrys sensu lato themselves (Grosjean, unpublished data). Of all the species

on which the width of the funnel was measured, the tadpole of Ophryophryne microstoma is

the one which possesses the smallest [50 % of SVL vs. 74.4 % in M. minor (Liv, 1950) and M.

boettgeri (Liu, 1940), and 82.3 % in M. aceras (LAIDLAW, 1900)]. LEONG & CHou (1998) did

not give a measurement comparable but noticed that the funnel of M. longipes is the largest of

all the species known. The tadpoles of these two genera bear another peculiar structure,

unique among the Megophryidae: soft jaw sheaths with long serrations at the opening of the

mouth (INGER, 1985; HUANG et al., 1991; this paper). This structure, although externally

visible, is not always conspicuous. It is now largely accepted that buccopharyngeal features are

very conservative within genus (e. g. VIERTEL, 1982) as, e.g., within the Megophryidae (INGER,

1983, for a taxonomic review of three genera with larval characters; HUA tal., 1991). Each

megophryid genus has a typical buccopharyngeal morphology, which distinguishes it clearly

from all other megophryid genera. However, in contradiction to HUANG et al. (1991)'s

conclusion (not detailed in their work), I failed to find any significant differences in their

buccopharyngeal features. The morphology of these tadpoles does not contribute to clarify

e status of Ophrvophryne, but the lack of clear differences between Ophrrophrine and the

s of the four taxa included in Megophrys (sensu lato) does not justify a different

taxonomical level for the former. A recent study based on cytology, morphology and ecology

Source : MNHN, Paris

56 ALYTES 21 (1-2)

(Rao & YANG, 1997) suggested two clades within this group, one including large species

(Atympanophrys, Brachytarsophrys and Megophrys), the other including small and moderate-

sized species (Ophryophryne and Xenophrys). Furthermore, RAo & YANG (1997) proposed

that these five taxa should be risen to generic level. The results of the present study do not

conflict with the opinion of these authors.

RÉSUMÉ

La morphologie externe, incluant des données morphométriques et l'anatomie bucco-

pharyngée du têtard d’Ophryophryne microstoma Boulenger, 1903 sont présentées. Les indi-

vidus de cet échantillon présentent quelques différences morphologiques avec des têtards

rapportés à cette espèce mais appartenant à d’autres populations. Le statut taxinomique du

genre Ophryophryne au sein des Megophryidae est discuté à la lumière des caractéristiques

morphologiques et buccopharyngées du têtard d’Ophryophryne microstoma et de celles du

genre Megophrys au sens large publiés dans la littérature.

ACKNOWLEDGEMENTS

1 am grateful to the Frontier stafT (London) for their technical assistance in the field and to the

Institute of Ecology and Biological Resources (Hanoi) for the export permit for the specimens studied.

Many thanks to Dr. Wen-hao Chou (Taiwan) for his translation of Chinese text. Thanks to Dr. Heinz

Grillitsch (Wien) and two anonymous reviewers for critical evaluation and language improvement on

earlier versions of the manuscript. This is publication N° 03-81 of PPF “Faune et flore du sud-est

asiatique” (N° 03-80, see DEUVE & MOURZINE, 2003).

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Gex, H.-J., Li, F-L. & X1A0, H., 1984. — Preliminary observations on ecology of Vibrissaphora

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Dusois, A. & OuLer, A., 1998. — À new species of Leprobrachium (Vibrissaphora) from northern

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GRosyEAN, S., 2001. — The tadpole of Lepobrachium (Vibrissaphora) echinatum (Amphibia, Anura,

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their classification and phylogenetic relationships. Asiatic herp. Res., 7: 93-102.

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Coran 1926. - The function of the “funnel” mouth of the tadpoles of Megalophrys, with a note on M. aceras

Boulenger. Proc. ool. Soc. London, 1926: 983-988.

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with descriptions of a new subfamily and subdivision of Bombina. Acta herp. Sin, 4: 219-224. [In

Chinese]

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Vaerte, B., 1982. - The oral cavities of Central European anuran larvae (Amphibia). Morphology,

ontogenesis and generic diagnosis. Amphibia-Reptilia, 4: 327-360.

WasserSUG, R. J., 1976. - Oral morphology of anuran larvae: terminology and general description. Occ.

Pap. Mus. nat. Hist. Univ. Kansas, 48: 1-23.

—- 1980. — Internal oral features of larvae from eight anuran families: functional, systematic, evolutio-

nary and ecological considerations. Misc. Publ. Mus. nat. Hist. Univ. Kansas, 68: 1-146.

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Zmao, E.-M., INGER, R. F., Wu, G.-F. & SHAFFER, H. B., 1994. - Morphological variation and ecological

distribution of co-occuring larval forms of Oreolalax (Anura: Pelobatidae). Amphibia-Reptilia, 15:

109-121.

Corresponding editor: Heinz GRILLITSCH.

Source : MNHN, Paris

Alytes, 2003, 21 (1-2): 59-65. 59

Larval stages, habitat and distribution

of the hyperoliid frog

Heterixalus rutenbergi (Boettger, 1881)

Liliane RAHARIVOLOLONIAINA”, David R. VtaTes"”, Frank GLaAwW°”

& Miguel VENCES""""

* Université d’Antananarivo, Département de Biologie Animale, Antananarivo, Madagascar

** Departamento de Bioloxia Animal, Laboratorio de Anatomia Animal, Buzôn 137, Facultade de Ciencias

Biolôxicas, Universidade de Vigo, Apdo. 874, 36200 Vigo, Spain

*#* Zoologische Staatssammlung, Münchhausenstr. 21, 81247 München, Germany

###* Institute for Biodiversity and m Dynamics, Zoological Museum, University of Amsterdam,

PO Box 94766, 1090 GT Amsterdam, The Netherlands

<vences@science.uva.nl>

We describe the hitherto unknown external larval morphology of Hete-

rixalus rutenbergi, a reed frog from highlands in central Madagascar.

Tadpoles were collected in a sun-exposed pond in a swampy savannah at the

Itremo Massif. Their morphology is similar to that of other Heterixalus,

with a labial tooth row formula of 1/1+1:2. They differ by a distinct marbled

or spotted color on the proximal third of the caudal musculature. Metamor-

phosing juveniles have the distinctive pattern of adult frogs with five white

stripes on a green dorsum, unlike other Heterixalus who show a juvenile

coloration with two dorsolateral stripes. Contrary to other Heterixalus

species, H. rutenbergi seems to be restricted to highland savannahs and

has 50 far not been recorded in high densities; its status should therefore be

more carefully monitored.

INTRODUCTION

The genus Heterixalus Laurent, 1944 contains the endemic Malagasy representatives of

the family Hyperoliidae. Heterixalus is the sister group of the Seychellean Tuchycnemis

Fitzinger, 1843 in this otherwise exclusively African family (RicHARDS & MooRE, 1996:

VENCES et al., 2003). Currently 10-11 species of Heterixalus are known (VENCES et al., 2000),

two of which are endemic to the highlands of central and central-eastern Madagascar: /1.

betsileo (Grandidier, 1872) and H. rutenbergi (Boettger, 1881).

Heterixalus species are typical inhabitants of open areas, and often oceur in secondary

habitats such as rice fields. Their larvae are of a rather generalized pond type, with a single

uninterrupted row of labial teeth on the upper lip, and one interrupted and two uninterrupted

rows on the lower lip. This morphology has been ascertained by BLOMMERS-SCHLÔSSER (1982)

and GLAW & VENC 1993, 1994) for H. betsileo, H. madagascariensis (Duméril & Bibron,

1841), H. boertgeri (Mocquard, 1902) and A. luteostriatus (Andersson, 1910).

Source : MNHN, Paris

60 ALYTES 21 (1-2)

Heterixalus are also very uniform in adult morphology, and a reliable species distinction

is only possible by combining advertisement calls and coloration in life (GLAW & VENCES,

1993). However, one species, H. rutenbergi, has a number of highly divergent traits: its call

bears no resemblance to that of any other species, the gular gland on the vocal sac of males

has blackish color, and the dorsal pattern (green with five longitudinal white bands) is unique.

We recently started with intensive herpetological surveys in the montane areas of central

Madagascar (see VENCES et al., 2002). During the fieldwork, we discovered tadpoles that could

be unambiguously assigned to H. rutenbergi by the pattern of metamorphosing juveniles. In

the present paper, we describe the morphology of these tadpoles and review the published

information on distribution and habitat of H. rutenbergi.

MATERIALS AND METHODS

Tadpoles were collected on 11 March 2001 at Ambatomenaloha, Itremo Massif, central

Madagascar (19°58'S, 46°55°E; 1820 m above sea level). They were found in a shallow

sun-exposed pond on a large unforested plain that partially was flooded by a river. Most

ponds on this plain were fed by the river and had relatively cold water, whereas the ponds

populated by Heterixalus tadpoles were much warmer, but no measurements of temperature

could be effectuated. The pond had a depth of ca. 60 cm, and was bordered by grass only. The

specimens were preserved in 5% formalin after capture, but were damaged during the

transport. À batch of 14 tadpoles were deposited in the herpetological collection of the

Zoologische Staatssammlung München under the number ZSM 789.2001. One additional

specimen (field number LR 271) used for the detailed morphological description and

drawings will be incorporated in the ZSM later on. Developmental stages are described after

Gosner (1960). Morphological measurements were taken by L. R. using a digital caliper to

the nearest 0.1 mm, following landmarks, terminology and definitions of McDiarmid &

ALTIG (1999). The formula of labial tooth rows follows Dupois (1995).

We use the following abbreviations: BL, head and body length (in tadpoles: from the tip

of the snout to the junction of the posterior body wall with the axis of the tail myotomes:

McDiarMiD & ALTIG 1999); TAL, tail length; BW, maximum body width; ODW, maximum

width of oral disc; DGMP, dorsal gap of marginal papillae; IOD, interorbital distance

between centers of pupils; ED, eye diameter; TH, tail height at beginning of tail, MTH,

maximum tail height including the caudal fin; TMH, height of caudal musculature at

mid-tail, TMW, caudal muscle width; SVL, snout-vent length (in adult and juvenile frogs):

UTR, upper tooth row; LTR, lower tooth row.

RESULTS

The series of tadpoles assigned to Hererixalus rutenbergi had a conspicuous color

pattern. They were brownish with green-olive, and had a very distinct silvery white marbling

on the proximal third of the caudal musculature (fig. la-b). In late developmental stages

Source : MNHN, Paris

RAHARIVOLOLONIAINA et al. 61

(42-45) the typical adult coloration (green dorsum with five white longitudinal stripes, each

bordered by two black lines) became visible (fig. 1e).

The following morphological description is based on one tadpole in stage 37 (field

number LR 271; fig. 2a-c). Tail only partly preserved, part of the skin detached. A rather

compressed tadpole of ORTON’s (1953) type 4; eyes directed laterally; spiracle sinistral and

positioned closer to the anus than to the tip of snout; caudal fin, as far as recognizable,

dorsally and ventrally with straight edges, starting directly behind body (fig. 2b): intestine not

visible through the ventral skin. Further proportions and detailed characters of body and tail

not reliably assessable because of poor state of preservation.

Oral disc apparatus in excellent state of preservation (fig. 2c), generalized, small, almost

terminal, oriented ventrally; labial tooth row formula 1/1+1:2; tooth rows distinct but

relatively small; LTR1 with a small gap (< 0.1 mm); UTRI with approximately 80 labial teeth

(ca. 34 per mm). Oral disc without a recognizable lateral notch; beak distinct and black, both

jaw sheaths with serrations at their cutting edges. Oral papillae present around the oral disc

except for its upper part; 1-2 rows of submarginal papillae, restricted to lateral parts of oral

disc; marginal papillae in one row: altogether about 37 marginal and 8-10 submarginal

papillae, all shorter than | mm.

Morphometric measurements: BL 14.4 mm; TAL (incomplete) 14.9 mm; BW 9.4 mm;

ODW 1.7 mm; DGMP 1.7 mm; IOD 5.6 mm; ED 2.1 mm; TH 5.4 mm; MTH 8.8 mm;

TMH 4.8 mm; TMW 3.5 mm; UTRI 1.9 mm; each part of LTRI 0.8 mm; LTR2 1.8 mm;

LTR3 0.7 mm.

In preservative, anterior lateral surface of body dark brown with yellowish shade,

posterior part dark brown with some larger silvery shades; whole dorsum dark brown with

many black spots of 0.4-1.6 mm diameter (fig. 2a-b). Similar spots also on dorsal and ventral

caudal fins, and on caudal musculature (0.6-2.3 mm in diameter). Belly white with many

smaller black spots.

In a just metamorphosed juvenile (fig. Ic), the color pattern typical for adult H.

rutenbergi was already fully expressed. SVL of one specimen in stage 41-42 belonging to the

series ZSM 789.2001 is 15.5 mm.

No adult H. rutenbergi were found during our survey at Itremo. Other frog species

collected or observed by us were Boophis ankaratra Andreone, 1993, B. goudoti Tschudi, 1883,

B. luteus (Boulenger, 1882) (call record), B. microtympanum (Bocttger, 1881), Mantidactylus

aff. brevipalmatus Ahl, 1929, M. domerguei (Guibé, 1974) (call record), M. femoralis (Boulen-

ger, 1882), M. lugubris (Duméril, 1853), M. sp. A. aff. curtus (Boulenger, 1882), M. sp. B aff.

curtus and Ptychadena mascareniensis (Duméril & Bibron, 1841). Furthermore, a coll

made by D. Rakotomalala included a subadult specimen of Scaphiophryne madagascariensis

(Boulenger, 1882).

Source : MNHN, Paris

62 ALYTES 21 (1-2)

Fig. 1. - Drawing and photographs of larval stages of Heterixalus rutenbergi from Ambatomenaloha,

Itremo, central Madagascar. (a), drawing of a tadpole in life, based on a color photograph: (b),

photograph of another tadpole specimen in life (c), photograph of a metamorphosing juvenile (tail

not yet fully reduced), already showing the typical adult coloration. Both photographs were made on

12 March 2001.

DISCUSSION

Assigning tadpoles to adult frogs is a difficult endeavour, and the decisions often remain

tentative, except for cases in which (1) tadpoles are identified by means ol

tadpol d from clutches deposited by well-identified adult specimens, (3) metamor-

phosed juveniles are raised to the adult stage, (4) metamorphosed juveniles already show

characters that are fully diagnostic for the particular species. The case of the tadpoles

described herein belongs into the fourth category, and their assignment to Heterixalus

rutenbergi is based on the following rationale. (1) They have the typical Heterixalus tooth

formula (see below) which is not found in any other group of Malagasy frogs (GLAW &

VENCES 1994). (2) No other Heterixalus species is known from Itremo (GLAW & VENCES

1994), and during our survey we did not hear any call asignable to a Heterixalus species. (3)

The larval color pattern on the flanks is different from that of the other known Heterixalus

tadpoles, among which the only other species known to occur in the central highlands and

adjacent western savannahs, A. bersileo and H. luteostriatus. (4) One specimen of the batch

ZSM 789.200! in stages 41-42 (forelimbs fully emerged, but larval mouthparts still present)

still has the characteristic larval color pattern on the flanks but also shows a central, two

Source : MNHN, Paris

RAHARIVOLOLONIAINA et al.

Fig. 2.

Itremo, central Madagascar, specimen LR 271 (developmental stage 37). (a) lateral view: (b) dorsal

view; (c) mouthparts. Not to scale. Sharply delimited white patches in lateral and dorsal views

Drawings of preserved tadpole specimen of Heterixalus rutenbergi from Ambatomenaloha,

symbolize detached skin

Source : MNHN, Paris

64 ALYTES 21 (1-2)

dorsolateral and two lateral light stripes. (5) This five-striped pattern, which is fully developed

in specimens in stages 45-46, is absent in all other Heterixalus and indeed also in all other

Malagasy frog species (GLAW & VENCES, 1994), including all taxa reported from Itremo.

Hence, this character reliably characterizes the tadpole specimens as Heterixalus rutenbergi.

According to the data provided herein, general larval morphology of A. rutenbergi is

similar to that of other Heterixalus. However, tadpoles of this species have a conspicuous

color and distinct differences regarding the transition to the adult pattern. The adult colora-

tion in Heterixalus is very diverse, and important differences can be observed within and

among conspecific populations (GLAW & VENCES, 1993, 1994). Some species are character-

ized by a pair of light dorsolateral stripes. This pattern seems to be always present in adult H.

betsileo, H. carbonei and H. luteostriatus (with the exception of H. betsileo from Ankaratra, in

which the stripes are almost unrecognisable). Other species do not display this pattern as

adults. However, two species of uniform adult coloration (H. boettgeri and H. madagasca-

riensis) have dorsolateral stripes as juveniles, as do H. andrakata, H. betsileo and H. “varia-

bilis” (GLAW & VENCES, 1993). A. rutenbergi differs from this trend because its five stripes

appear simultaneously already at metamorphosis. Also the final color (dark green) was

present from stage 45 onwards. This means that, in contrast to other Heterixalus species, a

typical juvenile coloration is lacking in H. rutenbergi.

A second aspect that merits attention is the conspicuous silvery white marbling on the

proximal portion of the tail of Æ. rutenbergitadpoles. This pattern is not known from any other

Heterixalus tadpoles (BLOMMERS-SCHLÔSSER, 1982; GLAW & VENCES, 1994), but it reminds the

tadpoles of the African hyperoliid genus Kassinathatarealso pond-dwellers(with veryhighfins,

however) and display brightly striped or mottled patterns (e.g., CHANNING, 2001).

Heterixalus rutenbergi is known from six precise localities, all on the central high plateau

of Madagascar: Ambohitantely, Mantasoa, Ambatolampy, Tsinjoarivo, Itremo and Amba-

tofitoharanana (BLOMMERS-SCHLÔSSER & BLANC, 1991; GLAwW & VENCES, 1994; VALLAN,

2000). Because it is not a forest species, H. rutenbergi has not been recorded in most

herpetological highland surveys, which did not focus on unforested areas. It seems clear,

however, that the habitat choice of this species is more specialized than in its congeners that

populate in huge densities all types of secondary habitats and even occur in flooded areas

within towns. At Ambatolampy, we found H. rutenbergi in low densities in a moorland area,

whereas A. betsileo was very common in the rice fields around the town (VENCES et al., 2002).

AtMantasoa we were not able to confirm the presence of the species despite its characteristic

calls that can be recognized over long distances (pers. obs.). Our findings in Itremo also refer

to a relatively special highland savannah habitat. Certainly, the species is widespread over

central Malagasy highlands, but its populations may have low densities and be vulnerable to

transformation of moorland into rice fields. Additional fieldwork is needed to ascertain its

habitat requirements and conservation status.

ACKNOWLEDGEMENTS

We are grateful to D. Rakotomalala for assistance in the field. The Malagasy authorities issued

collection and export permits. This work was carried out in the framework of a cooperation accord

Source : MNHN, Paris

RAHARIVOLOLONIAINA et al. 65

between the Département de Biologie Animale, Université d’Antananarivo, and the Zoologische Staats-

sammlung München, and received financial assistance by the Deutscher Akademischer Austauschdienst

(DAAD) and by the Volkswagen Foundation.

LITERATURE CITED

ALriG, R. & McDiariD, R. W., 1999, — Body plan. Development and morphology. Jr: R. W.

McDiarMiD & R. ALTIG (ed.), Tadpoles: the biology of anuran larvae, Chicago University Press:

24-51.

BLOMMERS-SCHLÔSSER, R. M. A. 1982. - Observations on the Malagasy frog genus Hererixalus Laurent,

1944 (Hyperoliidae). Beaufortia, 32 (1): 1-11.

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

Madagascar, 75 (1): 1-379.

CHaNNING, À. 2001. — Amphibians of Central and Southern Africa. Cornell University Press.

Durots, A., 1995. — Keratodont formulae in anuran tadpoles: proposals for a standardization. J. zoo!

Syst. Evol. Res., 33 (1): I-XV.

GLaw, F. & VeNcES, M. 1993. - Zur Bioakustik, Biologie und Systematik der Gattung Hererixalus aus

Madagaskar (Anura: Hyperoliidae). Salamandra, 29 (3-4): 212-230.

_— 1994. — À fieldguide ro the amphibians and reptiles of Madagascar. 2nd edition, Kôln, Vences & Glaw

Verlag: 1-480, 48 pl.

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

identification. Herpetologica, 16: 183-190.

Ortow, G. L., 1953, - The systematics of vertebrate larvae. Syst. Zool., 2: 63-15.

RicHARDs, C. M. & MOoRE, W. S., 1996. - A phylogeny of the African treefrog family Hyperoliidae based

on mitochondrial DNA. Mol. Phylogenet. Evo. 5 (3): 522-532.

VALLAN, D. 2000. - Influence of forest fragmentation on amphibian diversity in the nature reserve of

Ambohitantely, highland Madagascar. Biol. Conserr., 9%: 31-43.

Vexcrs, M., ANDREONE, F., GLAW, F., RAMINOSOA, N., RANDRIANIRINA, J. E. & VIEITES, D. R., 2002. —

Amphibians and reptiles of the Ankaratra Massif: reproductive diversity, biogeography and

conservation of a montane fauna in Madagascar. Jtal. J. Zool., 69: 263-284.

Vexces, M., GLAW, F., JESU, R. & SCHIMMENTI, G., 2000. — À new species of Heterixalus (Amphibia:

Hyperoliidae) from western Madagascar. African Zool. 35 (2): 269-276.

Vexces, M. KosUCH, J., GLAW, F., BôHME, W. & Vert, M. 2003. — Molecular phylogeny of hyperoliid

treefrogs: biogeographic origin of Malagasy and Seychellean taxa and re-analysis of familial

paraphyly. Z zool. Syst. Evol. Res, 41: 205-215.

Corresponding editor: Alain DUROIS.

Source : MNHN, Paris

Alytes, 2003, 21 (1-2): 66-76.

Morfologia del tubo digestivo en larvas

de Dermatonotus muelleri

y Elachistocleis bicolor

(Anura, Microhylidae)

del Noroeste argentino

Zandra E. ULLOA KREISEL

Cätedra de Embriologia y Anatomia Comparadas de la Fac. de Cs. Naturales e Inst. M. Lillo.,

Univ. Nac. de Tucumän, M. Lillo 251, 3° Piso, San Miguel de Tucumän (4000), Argentina

<zulloa@esnat.unt.edu.ar>

Morphology of the digestive tube in larvae of two species of Microhyli-

dae from the Argentine Northwest, Dermatonotus muelleri and Elachis-

tocleis bicolor, is described. The digestive tube length is standard, with two

coiled nucleï in the midgut. Histologically the foregut presents a dorsal fold

in the esophagus with many ciliated cells in the mucosae. The gastric region

is characterized by a “manicotto glandulare” with a thick glandular layer.

The epithelium of the midgut and posterior intestine is simple, showing

columnar cells with brush border in the former and cylindric cell

caliciform cells in the latter. The anatomical characteristics of the c

the midgut have not been described for other larvae of neotropical Micro-

hylidae, but the dorsal fold in the esophagus is common to other American

microhylid genera.

INTRODUCCION

Las larvas de Microhylidae de vida libre se caracterizan por tener labios flexibles; el

inferior puede ser protusible a modo de cuchara o en forma de embudo, en otros casos. La

microfagia es el häbito alimenticio mäs comün y las estructuras bucofaringeas colaboran en la

captura de las particulas alimenticias. ORTON (1953) clasifica a las larvas de Microhylidae

como larvas de tipo IT, las cuales presentan, entre otros caracteres, ausencia de piezas bucales

queratinizadas, una câmara opercular simple y un espiräculo medio ventral. ALTIG &

JOHNSTON (1989) clasifican a algunas larvas de Microhylidae como suspensivoras de tipo I,

con pliegues labiales semiesféricos sobre la boca, cuerpo muy deprimido y cireular en vista

dorsal.

Entre las descripciones de las estructuras orales internas en larvas de Microhylidae estân

adas por SAVAGE (1952) quien describié la anatomia de las estructuras bucofaringeas

en larvas de cuatro especies de Microhylidae asiäticos (Calluella guttulata, Glyphoglossus

molossus, Chaperina fusca y Kaloula pulchra), WassersuG (1980) en tres especies del género

las

Source : MNHN, Paris

ULLOA KREISEL 67

Microhyla (M. berdmorei, M. heymonsi y M. ornata), WaASSERSUG & PYBURN (1987) en

Otophryne robusta que es un Microhylidae neotropical, entre otros autores. GRADWELL (1974)

observé las estructuras branquiales en Phrynomantis annectens.

La informaciôn referida al tubo digestivo en los Microhylidae neotropicales es inexis-

tente. NELSON & CUELLAR (1968) realizaron estudios comparativos de la cavidad oral y del

tubo digestivo en 4 especies de dos géneros de anuros americanos neärticos, Hypopachus

barberi, H. variolosus, Gastrophryne olivacea ÿ G. usta.

Existen dos especies de Microhylidae en el Noroeste argentino que son Dermatonotus

muelleri y Elachistocleis bicolor. La morfologia externa de la larva de D. muelleri fue descripta

por Ce1 (1980) y ALTIG & JONHSTON (1986). LAvILLA (1992) re-describi6 la morfologia externa

de la larva y analizé el condrocräneo y el esqueleto visceral. De acuerdo con estas descripcio-

nes la larva presenta un cuerpo en vista dorsal cuadrangular y en vista lateral triangular, ojos

pequeños dirigidos dorsolateralmente, una boca ventral, sin papilas marginales ni pico

crneo, un espiräculo medial posterior cercano al tubo proctodeal, una cola triangular,

gruesa y corta, con aletas dorsal y ventral delgadas.

Elachistocleis bicolor fue citada para otras zonas neotropicales por Cet (1980) y GUDYy-

NAS (1983). WizLiams & GupyNas (1987) describieron la larva: presenta un cuerpo globoso,

ojos pequeños que se ven dorsalmente, una boca pequeña, de ubicaciôn anteroventral, con

pliegues a modo de faldas y sin estructuras queratinizadas ni papilas marginales, un espirä-

culo terminal, inmediatamente por delante y a la izquierda de la abertura anal, una cola

musculosa con aletas bajas que alcanzan el cuerpo.

Las estructuras orales internas en larvas de ambas especies fueron caracterizadas por

ECHEVERRIA & LAVILLA (2000).

LaviLLa & LANGONE (1991, 1995) describieron cambios ontogenéticos en la orientaciôn

del espiräculo y tubo proctodeal, asi como las estructuras del condrocräneo en las larvas de E.

bicolor.

En base a los antecedentes sobre el grupo de Microhylidae y la necesidad de contar con

descripciones que posibiliten comparaciones intra e interfamiliares, se ha planteado el

objetivo de este trabajo en la descripcién del aparato digestivo post-faringeo en las larvas

suspensivoras micréfagas de Microhylidae que se encuentran en el Noroeste argentino.

MATERIAL Y MÉTODOS

Se utilizaron larvas de Dermatonotus muelleri ÿ Elachistocleis bicolor entre los estadios

equivalentes a los 30-37 de la tabla de desarrollo normal de GosNER (1960), provenientes de

colectas realizadas en la localidad de Las Lajitas, Departamento Anta, Salta, Argentina, en el

mes de enero de 1998 y de la coleccién del Museo de Ciencias Naturales de Salta MCN 123,

respectivamente.

El material se fijé en una solucién de formaldehido al 10 % y se postfijé en liquido de

Bouin. Las observaciones macroscopicas se hicieron por medio de disecciones bajo lupa,

efectuando un corte en la linea media ventral para exponer el tubo digestivo.

Source : MNHN, Paris

ALYTES 21 (1-2)

Tab. 1. - Morfometria del tubo digestivo en larvas de Microhylidae suspensivoras micréfagas

(WDermatonotus muelleri y Elachistocleis bicolor). %: sobre la longitud total del tubo

digestivo.

Dermatonotus muelleri

Elachistocleis bicolor

Tubo digestivo

@=7)

Longitud del tubo digestivo

Eséfago

Manicotto glandulare

Intestino medio

Intestino posterior

840.21 cm

= 10 veces la longitud

del cuerpo del renacuajo

0.3 +0.01 cm = 3.750 %

0.25 + 0.01 cm = 3.12%

5.95 + 0.15 cm = 74.37 %

1.5 + 0.04 em = 18.75 %

4.45 + 0.15 cm

& 6 veces la longitud

del cuerpo del renacuajo

0.1 + 0.01 cm = 2.25 %

0.15 + 0.01 cm = 3.37 %

3.5 + 0.11cm = 78.65 %

0.7 + 0.02 cm = 15.73 %

Los estudios morfométricos se realizaron en larvas equivalentes al estadio 35 de GOSNER

(1960), se tomaron la longitud del cuerpo de cada renacuajo correspondiente a la medida

entre el hocico y el vientre para comparar con el largo total del tubo digestivo y se midieron

con un calibre Vernier cada porciôn del tubo digestivo. Las mediciones del intestino medio se

realizaron sobre un papel milimetrado.

Para los estudios histolégicos se extrajeron las distintas regiones del tubo digestivo, el

material se deshidraté y se utilizé xilol como intermediario para su inclusién en paraplast. Se

efectuaron cortes seriados de 6 1m de espesor con micrétomo de deslizamiento. La técnica de

coloracién utilizada fue Hematoxilina-Eosina (H&E). Las observaciones se efectuaron en

microscopio estereoscopico Leica MPS30 y documentadas en cämara fotogräfica fotoauto-

mat MPS30.

RESULTADOS

DERMATONOTUS MUELLERI

Descripciôn anatémica

La longitud del tubo digestivo es aproximadamente diez veces la longitud del cuerpo

(hocico-vientre) (tab. 1).

En D. muelleri el eséfago se ubica en la linea media del cuerpo en la parte anterior de la

cavidad abdominal, es relativamente corto y comprende un 3.75 % de la longitud total del

tubo digestivo.

El manicotto glandulare estä ubicado en la regién dorsal de la cavidad, con paredes mäs

gruesas y blanquecinas, y representa un 3.12 % de la longitud total del tubo digestivo (fig. IA,

©).

Source : MNHN, Paris

ULLoA KREISEL 69

Fig. 1. Tubo digestivo en Dermatonotus muelleri. (A) Vista lateral di

Vista lateral izquierda de la cavidad abdominal. (C) Vista dorsal del intestino anterior. (D) Vista

dorsal del intestino posterior. C, corazén; C.H, condueto hepätic séfago; H, higado: LM,

intestino medio; L.P, intestino posterior; N, nücleo de enrollamiento; Mn, manicotto glandulare; P,

päncreas: V.B, vesicula biliar. Escala: ! mm.

cha de la cavidad abdominal. (B)

Source : MNHN, Paris

70 ALYTES 21 (1-2)

La primera porcién del intestino medio (duodeno) se curva alrededor del pâncreas en el

lado derecho. El päncreas es mediano, de forma circular y color blanquecino, y el higado esta

ubicaciôn algo anterior al pâncreas. El intestino medio es largo (74.37 % de la longitud total

del tubo digestivo) y ocupa gran parte de la cavidad abdominal. Se caracteriza por presentar

dos nücleos de enrollamiento. El primero es mäs corto, teniendo 1.5 vueltas y estä ubicado

lateralmente en el lado izquierdo. El segundo es mäs largo, con 4.5 vueltas, ocupa la parte

central de la cavidad y su eje estâ orientado en sentido dorso-ventral; ambos forman una

espiral doble al enrollarse (fig. 1A-B).

El intestino posterior es mäs corto, presenta mayor diâmetro y se enrolla internamente

con posicién mäs dorsal que el intestino medio. Representa un 18.75 % de la longitud total del

tubo digestivo (fig. ID).

Observaciones histolégicas

Eséfago

Presenta un septo longitudinal en la pared dorsal que abarca el primer tramo del eséfago,

tiene una submucosa con abundante tejido conectivo laxo y un epitelio con células cilindricas

ciliadas. À ambos lados del septo, en la base del mismo, se encuentran células cilindricas bajas

secretoras de mucus.

El resto de la mucosa del eséfago presenta una capa simple de células cilindricas ciliadas

yescasas células caliciformes en la pared ventral. La submucosa es una capa delgada de tejido

conectivo laxo y la capa muscular es muy delgada con fibras circulares (fig. 2A-B).

En la porcién posterior se forman varios plegamientos longitudinales con concentracio-

nes de células cilindricas secretoras de mucus en forma de penachos. La capa muscular es mâs

gruesa con fibras circulares y externamente fibras longitudinales junto a la capa serosa (fig.

2C).

“Manicotto”

La mucosa tiene células cilindricas secretoras de mucus con nücleo grande medio basal

basôfilo y citoplasma apical eosinéfilo homogéneo. Se alterna con escasas zonas de células

cilindricas ciliadas. La mucosa tiene un espesor de 15 um.

El epitelio con células cilindricas bajas de nücleo redondo basôfilo puede estar interrum-

pido por la formacién de criptas en algunas zonas. Las gländulas se disponen longitudinal

mente con células cübicas de nücleo redondo basôfilo y citoplasma homogéneo cosinôfilo.

Esta capa tiene un espesor de 55 m. Rodeando esta estructura se encuentran fibras muscu-

lares circulares y la serosa con un espesor de 2.5 um (fig. 2D-E).

Intestino medio

Tiene un epitelio monoestratificado con abundantes células cilindricas de borde estriado

con nücleo redondo medial baséfilo y citoplasma homogéneo eosinéfilo y escasas células

caliciformes, pequeñas y apicales. La capa mucosa tiene un espesor de 15 m y la muscular de

2.5 um (fig. 2F).

Source : MNHN, Paris

Microfotografias del tubo digestivo en Dermatonotus muelleri. (A) 0 con septo longitu-

a: 100 um. (B) Septo longitudinal. Escala: 5 m. (C) Eséfago, zona de tran:

100 ym. (D) Manicotto glandulare. E a: 100 um. (E) Manicotto glandulare. Escala: N

Intestino medio. Escal ino posterior. Escala: 5 Ca, células caliciformes:

re pa glandul ulas de borde iliadas:

lulas secretoras , septo longitudinal; T.C,

Source : MNHN, Paris

72 ALYTES 21 (1-2)

Intestino posterior

Elepitelio es simple con células cilindricas de nücleo redondo basôfilo y citoplasma poco

teñido, de 11 ym de espesor. La submucosa y la capa muscular son muy delgadas, y en total

tienen 4 ym de espesor (fig. 2G).

ELACHISTOCLEIS BICOLOR

Descripciôn anatémica

El tubo digestivo presenta caracteristicas semejantes a las descriptas en D. muelleri.

La longitud total es algo mayor de seis veces la longitud del cuerpo. El eséfago y el mani-

cotto glandulare son porciones cortas del tubo digestivo y representan un 2.25 % y 3.37 %

de la longitud total del tubo digestivo, respectivamente. La porciôn mäs larga es el intes-

tino medio con un 78.65 % del largo total, al intestino posterior le corresponde un 15.73 %

(tab. 1).

Observaciones histolôgicas

Eséfago

Presenta un septo longitudinal en la pared dorsal con un epitelio monoestratificado de

células cilindricas ciliadas y las cilindricas bajas secretoras de mucus en la base a ambos

lados. En la pared ventral las células cilindricas ciliadas se intercalan con células caliciformes.

La submucosa forma una capa delgada de tejido conectivo laxo rodeada por una capa

muscular muy delgada con fibras circulares (fig. 3A).

La zona posterior tiene pliegues con grupos de células cilindricas secretoras de mucus en

forma de penachos. La capa muscular a este nivel es mäs gruesa con fibras circulares y

externamente fibras longitudinales junto a la capa serosa.

“Manicotto”

La mucosa tiene un epitelio simple con células cilindricas bajas y escasas células ciliadas:

ambos tipos de células estän poco diferenciadas, con un éspesor de 10 4m. Las glandulas se

disponen longitudinalmente con células cübicas de nücleo redondo basôfilo y citoplasma

homogéneo con poca afinidad a la eosina, formando una capa gruesa de 113 um. Las

musculares cireulares tienen un espesor de 2.5 um (fig. 3B-C).

Intestino medio

Tiene un epiteilo monoestratificado de 16 1m de alto, con abundantes células cilindricas

de borde estriado con nücleo redondo medial basôfilo y citoplasma granular levemente

cosinéfilo y escasas células caliciformes. La capa muscular es mäs gruesa que en otras

porciones del tubo digestivo y mide aproximadamente 8 ym de espesor (fig. 3D).

Source : MNHN, Paris

ULLOA KREISEL 73

- Microfotografias del tubo digestivo en Elachistocleis bicolor: (A) Eséfago con septo longitudinal.

cala: 25 um. (B) Manicotto glandulare. Escala: 100 um. (C) Manicotto glandulare. Escala: 25

(D) Intestino medio. Escala: 5 um. (E) Intestino posterior, Escala: 5 um. C.G, capa glndular

ulas de borde estriado; C.C.C, células cilindricas cilia

L, septo longitudunal

Source : MNHN, Paris

74 ALYTES 21 (1-2)

Intestino posterior

Elepitelio es simple con células cilindricas con nücleo redondo basôfilo y citoplasma con

gränulos eosinôfilos, de 17 ym de espesor. Las capas muscular y serosa son muy delgadas, con

5 um de espesor (fig. 3E).

DISCUSION Y CONCLUSIONES

Las dos especies de Microhylidae del Noroeste argentino estudiadas presentan a nivel de

tubo digestivo caracteres morfolégicos muy semejantes entre si, coincidiendo con lo obser-

vado previamente por ECHEVERRIA & LAVILLA (2000) con respecto a las estructuras internas

de la cavidad oral.

La longitud del tubo digestivo en las larvas de Dermatonotus muelleri es de aproximada-

mente 10 veces la longitud del cuerpo del renacuajo y en Elachistocleis bicolor es mäs corto,

representando 6 veces la longitud del cuerpo.

ALTIG & JOHNSTON (1989) clasifican a las larvas de varias especies de Microhylidae como

formas que se alimentan de materia en suspensiôn, häbito que corresponde también para las

especies aqui estudiadas. La presencia de todas estas caracteristicas tanto de la cavidad oral

como del tubo digestivo, corroboran el hâbito alimenticio suspensivoro en estas especies,

siendo de tipo micréfago.

La es informacién disponible en relaciôn con la estructura del tubo digestivo en

larvas de Microhylidae neotropicales permite solamente comparar las observaciones realiz:

das en los géneros Æypopachus y Gastrophryne (NELSON & CUELLAR, 1968) y especificamente

en Hypopachus aquae (SAVAGE, 1955), los cuales presentan a nivel del eséfago el septo dorsal

y en la regiôn gästrica el “manicotto glandulare” (LAMBERTINI, 1929) con caracteristicas muy

semejantes a las observadas en las larvas de D. muelleri y E. bicolor.

La longitud del eséfago en relaciôn con el resto del tubo digestivo es muy corta, hecho

que estä relacionado con las especies suspensivoras (BARRINGTON, 1946; GRIFFITHS, 1961).

También presenta una Zona de transiciôn entre el eséfago y el manicotto glandulare, carac-

teristica que comparte con Ranidae y Rhacophoridae (VIERTEL & RICHTER, 1999).

El manicotto glandulare en las larvas de ambas especies presenta estructuras un poco

mäs complejas, en comparacién con otras especies suspensivoras que lo presentan, como

Rana ridibunda (GriFriTHs, 1961) y Pleurodema borellii (ULLOA & TERAN, 1998) entre otras.

Las diferencias particularmente radican en la disposiciôn de las gländulas tubulares y la

presencia de numerosas criptas que interrumpen la continuidad de la capa epitelial. Estas

caracteristicas también son compartidas con otras especies de Microhylidae (GRIFFITHS,

1961).

El intestino medio es la porcién del tubo digestivo mâs larga y la caracteristica mâs

sobresaliente es la presencia de dos nücleos de enrollamiento. El primero es mäs pequeño y

lateral, el segundo es mâs grande y ventral. Esta ca stica no ha sido observada o

descripta en formas de Microhylidae americanas (SAVAGE, 1955; NELSON & CUELLAR, 1968),

nien larvas de otras especies de anuros neotropicales (ULLOA KREISEL, 2001; ULLOA & TERAN,

Source : MNHN, Paris

ULLOA KREISEL 75

1998; GIMÉNEZ et al., 1991; RaDA & BELLO, 1988; JORQUERA et al., 1982), ni en observaciones

propias realizadas en larvas de distintas especies de anuros del Noroeste argentino.

El intestino posterior es mäs grueso y corto que el intestino medio, formado por un

epitelio simple de células cilindricas, una capa muscular muy delgada y serosa.

RESUMEN

Se describe la morfologia del tubo digestivo en las larvas de dos especies de Microhylidae

del Noroeste argentino, Dermatonotus muelleri y Elachistocleis bicolor. La longitud del tubo

digestivo es eständar, con dos nücleos de enrollamiento en el intestino medio. Histologica-

mente el intestino anterior presenta un pliegue dorsal en el eséfago, con numerosas células

cilindricas ciliadas en la mucosa. La regiôn gästrica tiene un “manicotto glandulare” con una

espesa capa glandular. El epitelio del intestino medio y posterior es simple con células

cilindricas estriadas en el primero y células cilindricas y caliciformes en el ültimo. Las

caracteristicas anatémicas del enrollamiento del intestino medio no han sido descriptas para

otras larvas de Microhylidae neotropicales, pero el pliegue dorsal en el eséfago es comün a

otros generos de microhylidos americanos.

AGRADECIMIENTOS

Deseo agradecer a la Dra. Ernestina S. Teisaire por sus aportes y lectura critica del trabajo, también

por colaborar en la elaboraciôn de las fotogafas del material, y a là Dra. Marissa Fabrezi por f

parte del material de estudio.

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Source : MNHN, Paris

Alytes, 2003, 21 (1-2): 77-99. 77

The ‘‘pseudofirmisternal””

pectoral girdle of anurans

Moises KAPLAN

Division of reptiles and amphibians, Museum of Zoology, University of Michigan,

Ann Arbor, MI 48109, USA

<moiseska@umich.edu>

The pectoral girdles of the following species were studied histologically:

(1) Brachycephalidae: Brachycephalus ephippium; (2) Bufonidae: Atelo-

pus subornatus, Frostius pernambucensis, Osornophryne bufoniformis;

(3) Leptodactylidae: Geobatrachus walkeri, Insuetophrynus acarpicus;

(4) Pipidae: Hymenochirus boettgeri; (5) Ranidae: Rana sylvatica; (6)

Rhinodermatidae: Rhinoderma darwinii, Of the non-ranoid frogs, only A.

subornatus, F permanbucensis, H. boettgeri and O. bufoniformis have

completely fused, non-overlapping, epicoracoids (i.e., pseudofirmisterny).

The girdle morphologies of G. walkeri and R. darwinii are unique in

anurans. The girdles of B. ephippium and I. acarpicus are arciferofirmi

ternal. Morphological differences between the pseudofirmisternal and fir-

misternal girdles suggest they are not homologous.

TINTRODUCTION

Pseudofirmisterny is the term describing the pectoral girdles of frogs, other than Ranoi-

dea (i.e., Dendrobatidae, Hyperoliidae, Microhylidae, Ranidae and Rhacophoridae sensu

ForD & CANNATELLA, 1993, or Hyperoliidae, Microhylidae, Ranidae, Mantellidae and

Rhacophoridae sensu VENCES & GLAW, 2001), having the epicoracoid cartilages completely

fused to (i.e., from anterior to posterior) and not overlapping one another.

The following genera of non-ranoid frogs are considered to have pseudofirmisternal

girdles: Brachycephalus (including Psyllophryne; KAPLAN, 2002), Atelopus, Frostius, Osor-

nophryne, Atopophrynus, Geobatrachus, Insuetophrynus, Hymenochirus, Pseudhymenochirus

and Rhinoderma (BARRIO, 1970; TRUEB, 1973; RUIZ-CARRANZA & HERNANDEZ-CAMACHO,

1976; LyNcH, 1978; ARDILLA-ROBAYO, 1979; LYNCH & RUIZ-CARRANZA, 1982; CANNATELLA,

1985, 1986; DUELLMAN & TRUEB, 1985; MYERS & FORD, 1986; CANNATELLA & TRUEB, 1988:

GRAYBEAL, 1997; however, see MCLACHLAN, 1943; GrirrirHs, 1957, 1963; MCDIARMID,

1969). The non-ranoid, pseudofirmisternal family Dendrobatidae (LYNCH, 1973; Hay et al.,

1995; FELLER & HEDGES, 1998; VENCES & GLAW, 2000, 2001) was not examined. The above

distribution of pseudofirmisterny in anurans is questionable, because only the girdles of

Brachycephalus ephippium and Rhinoderma darwinii have been examined histologically and

Source : MNHN, Paris

78 ALYTES 21 (1-2)

observations of the pectoral girdle in cleared-and-stained specimens frequently are mislead-

ing (KAPLAN, 1993).

Generally, it is accepted that the character “epicoracoids completely fused and not

overlapping one another” evolved independently in both non-ranoid and ranoid anurans

(NOBLE, 1926; GRIFFITHS, 1963; DUELLMAN & TRUEB, 1985; FORD & CANNATELLA, 1993).

Moreover, it is thought that this character evolved several times in non-ranoid frogs — viz., in

Insuetophrynus, Rhinoderma and Brachycephalus (GrirriTHs, 1963; LYNCH, 1978; DUELLMAN

& TRUEB, 1985; FORD & CANNATELLA, 1993), in the ancestors of Frostius, Atelopus and

Osornophryne (CANNATELLA, 1986: GRAYBEAL, 1997), in Hymenochirus and Pseudhymenochi-

rus (CANNATELLA & TRUEB, 1988), and in Atopophrynus and Geobatrachus (MYERs & FORD,

1986). However, it is still unclear if these hypotheses are parsimonious, because there is no

available cladistic analysis of the taxa with this girdle morphology. The character pseudofir-

misterny supported the following monophyletic groups: Brachycephalus and Atelopus (GRIF-

FITHS, 1963), Brachycephalus and Psyllophryne (FORD & CANNATELLA, 1993), Arelopus,

Frostius and Osornophryne (CANNATELLA, 1986; GRAYBEAL, 1997), and Geobatrachus and

Atopophrynus (MYERS & FORD, 1986).

Herein, I describe the ventromedial parts of the pectoral girdles of most frogs that have

been described as pseudofirmisternal, along with one having a firmisternal girdle — i.e., the

pectoral girdles of ranoid frogs having the epicoracoids completely fused (FORD & CANNA-

TELLA, 1993). The descriptions are based on examination of serial sections which were

prepared to determine, first, whether the examined frogs have the epicoracoids completely

fused and not overlapping, and second, whether the hypothesis that pseudofirmisterny

evolved several times from firmisterny is consistent with the morphology (1.e., anatomical

differences between non-ranoid and ranoid frogs). The systematic implications of these

observations will be discussed.

MATERIALS AND METHODS

The midventral parts of the breast-shoulder apparatus of sexually mature individuals of

the following families and species of frogs were sectioned: (1) Brachycephalidae: Brai

phalus ephippium (UMMZ 103568); (2) Bufonidae: Atelopus subornatus (ICN 15820), Fr

pernambucensis (UMMZ 225143), Osornophryne bufoniformis (ICN 11505); (3) Leptodacty-

lidae: Geobatrachus walkeri (ICN 35186), Insuetophrynus acarpicus (UMMZ 225142); (4)

Pipidae: Hymenochirus boettgeri (UMMZ 229751); (5) Ranidae: Rana sylvatica (UMMZ

229752); (6) Rhinodermatidae: Rhinoderma darwinii (UMMZ 143361). The medial part of

the breast-shoulder apparatus was excised by cutting through the procoracoid cartilages,

clavicles and coracoid bones; the epicoracoids and the attached prezonal and postzonal

elements were removed, decalcified (Cal-Ex II, Fisher Scientific), embedded in paraffin

CWESSNER, 1960), sectioned transversely from the anterior tip of the omosternum to the

posterior tip of the sternum, and stained with hematoxylin and eosin.

The names of the muscles of Arelopus subornatus, Brachycephalus ephippium, Hymeno-

chirus boettgeri, Rana sylvatica and Rhinoderma darwinii follow those in TyYsoN'’s (1987) and

Source : MNHN, Paris

KAPLAN 79

BEDDARD's (1895, 1908) studies. The muscles of Osornophryne bufoniformis, Frostius pernam-

bucensis, Insuetophrynus acarpicus and Geobatrachus walkeri are designated by numbers, as

myological studies of these taxa are not available. Histological terminology follows that of

FAWCETT (1986). Drawings of the girdles of Atelopus farci (KAPLAN, 1994) and Pseudhyme-

nochirus curtipes (DE VILLIERS, 1929) are used instead of those of 4. subornatus and H.

boettgeri because the latter are not available.

Herein, I consider the epicoracoid cartilages to be the ventromedial elements of the

girdle extending from the level of the clavicles to the coracoids, including the parts lying

medial to the procoracoid shafts (GRiFFITHS, 1963; but see TYsON, 1987 and DUELLMAN &

TRUEB, 1985) and the coracoid bones (i.e., the ossified portion of the embryonic coracoid-

epicoracoid cartilage: TYSON, 1987; KAPLAN, 1993), and anterior to the clavicles (fig. 1); note

that the medial position of the epicoracoids with respect to the procoracoid is assumed

because they are indistinguishably fused in sexually mature individuals. I consider the

epicoracoid horns to be the part of the epicoracoid cartilages that lie posterior to the

posteromedial part of the coracoids. “Medial ligament” refers to the band of dense connec-

tive tissue ventromedial to the epicoracoids. The term “completely fused” refers to the fusion

of the epicoracoids from their anterior to posterior tips. The term “fused” describes epicora-

coids whose medial ends, or part of them, are united synchondrotically or by connective tissue

that changes gradually from cartilage, near the epicoracoids, to dense regular connective

tissue, at the midline. “Indistinguishably fused” is used to describe absence of a suture

between the epicoracoids, where “suture” is defined as a thin, transverse band of cartilage

with low cell, and high fiber, densities, different coloration, and/or refrective properties. The

epicoracoids are considered “overlapping” when every part of their medial ends are aligned

with one another on the vertical axis. The descriptors “left” and “right” refer to the

organism's left and right sides from the dorsal perspective.

Developmental stages are given according to GosneR (1960). The following abbrevia-

tions are used to designate the collections where the specimens studied are kept: ICN,

Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotä, Colombia;

UMMZ, University of Michigan Museum of Zoology, Ann Arbor, Michigan, USA.

RESULTS

BRACHYCEPHALUS EPHIPPIUM

Anteriorly (fig. 2, 3A), the epicoracoids (e) are fused indistinguishably to one another on

the midline at the level of the clavicle (cl). Slightly posterior (fig. 3B), there is a suture (su)

between the epicoracoids. Posteriorly (fig. 3C), the epicoracoids are marked by a shallow

dorsomedial crevice (er) and a broad ventromedial keel (k): loose regular connective tissue fills

the crevice. The m. supracoracoideus (msc) inserts on the ventral keel, medial ligament (ml)

and ventral surfaces of the procoracoids (p). Posterior to the procoracoids (fig. 3D), two flat,

expanded coracoids (co) flank the epicoracoids; the medial end of each coracoid has a

cartilaginous core, that is indistinguishable from the epicoracoids, surrounded by a bony

layer. Posteriorly (fig. 3E). the epicoracoids are free: the left side of the ventral keel is replaced

Source : MNHN, Paris

80 ALYTES 21 (1-2)

anterior process

of keel omosternum

processus uncinatus

procoracoid

coracoid

fenestra

epicoracoid

clavicle —_=——

coracoid

— sternum

Rana sylvatica

Fig. 1. — The ventral

Dashed mid line: medial

tissue; white: bone.

F the pectoral girdle of

of epicoracoids; 1

vlratica (adapted from TYsoN, 1987).

artilage; dark gray: loose connective

Source : MNHN, Paris

KAPLAN 81

IQ n mOUNw >

ZT OTNTMmONnwxz>

Brachycephalus ephippium Rhinoderma darwinii

IO T7 MmONw >

Insuetophrynus acarpicus Geobatrachus walkeri

Fig. 2. — Ventral view of the pectoral girdle of Brachycephalus ephippium (adapted from Tyson, 1987),

Rhinoderma darwinit (adapted from Cri, 1980), nsuetophrynus acarpicus (adapted from BARRIO,

1970) and Geobatrachus walkeri (adapted from ARDILLA-ROBAYO, 1979) lavicle:

epicoracoid horn: p, procoracoid: s, sternum; su, suture. White: bone; light gray

icoids and epicoracoids are desorganized. A-H: transverse

3-6. A and B in insert: transverse sections corresponding to

epicoracoi

cartilage: dark g

sections corresponding 10 those of

those of fig. 4D-E.

: area where the co

Source : MNHN, Paris

ALYTES 21 (1-2)

Fig. 3. — Histological cross sections, in an antero-posterior direction, of the ventromedial part of the

pectoral girdle of an adult Brachycephalus ephippium (UMMZ 103568). The levels of the sections are

indicated in fig. 2. el, clavicle: co, coracoid: cr, crevice: dl, dorsal ligament: e, epicoracoids; h,

epicoracoid horn; k, ventral keel: le, left epicoracoid: msc, mn. supracoracoideus: re, right epicoracoid:

su, suture; vl, ventral ligament.

Source : MNHN, Paris

KAPLAN 83

by dense, regular connective tissue termed the ventral ligament (vl). The m. supracoracoideus

inserts on the ventral ligament. Posteriorly (fig. 3F), the right epicoracoid (re) slightly overlaps

the left epicoracoid (le). The left epicoracoid abuts against the dorsomedial surface of the right

epicoracoid. At the midlevel of the coracoids (fig. 3G), the left epicoracoid bears a small dorsal

process (dp) that overlaps the right epicoracoid; a dorsal ligament (dl) joins this dorsal process

to the dorsomedial surface of the right coracoid. Each epicoracoid terminates in a minute

horn (h) (fig. 3H). The mn. sternoepicoracoideus inserts on the posterior terminus of the horn.

RHINODERMA DARWINI

Anteriorly (fig. 2, 4A), the epicoracoids (e) are indistinguishably fused at the midline. The

m. supracoracoideus (msc) inserts on the ventromedial surface of the clavicles, epicoracoids

and medial ligament (ml). À small ventral keel (k) is present posteriorly (fig. 4B). The

coracoids laterally flank the epicoracoids (fig. 4C). An oblique suture (su) separates the

epicoracoids (fig. 4D). Posteriorly, a triangular notch (n) in the right side of the ventral keel is

filled with loose connective tissue (fig. 4E); the m. supracoracoideus inserts on this connective

tissue, the ventral surfaces of the right epicoracoid and the medial ligament. At the level of the

coracoids (fig. 4F), the loose connective tissue in the notch is replaced by cartilage (x).

Posteriorly (fig. 4G), each epicoracoid terminates in a horn (rh) that curves posterolaterally.

The horns and the sternum (s) are separated by sutures. Posteriorly (fig. 4H), the mn. sternoe-

picoracoideus (mse) inserts on the posterior ends of the epicoracoid horns.

INSUETOPHRYNUS ACARPICUS

The anterior epicoracoids are indistinguishably fused to one another at the midline (fig.

2, 5A), but slightly posteriorly (fig. 5B) the epicoracoids can be distinguished from one

another; each bears a dorsomedial protuberance (dp). There is a distinct dorsomedial crevice

and a rounded, ventromedial keel (k). Muscle 1 (ml) inserts on the lateral surface of each

epicoracoid and Muscle 2 (m2) on the ventrolateral surface and medial ligament (ml). The

dorsal crevice is extended ventrally as a sigmoid shape; the dorsal part of the left epicoracoid

slightly overlaps the right epicoracoid (fig. 5C). Muscle 3 (m3) inserts on the ventral part of

the medial ligament. In the anterior region of the coracoid fenestra (fig. SD), the epicoracoids

are free; the medial end of the left epicoracoid (le) is wide, and overlaps the right epicoracoid

(re). The left side of the ventral keel is replaced by dense, regular connective tissue, the ventral

ligament (vl), on which muscle 2 inserts. Posteriorly (fig. SE), the left epicoracoid is triangular

in section and the right epicoracoid elliptical. The right epicoracoid abuts against the ventral

ligament and dorsal ligament (dl) extends between the medial end of the left epicoracoid and

the right epicoracoid. Muscle 2 inserts on the ventral ligament and the lateral surface of each

epicoracoid. Muscle 3 inserts mostly on the medial ligament. At the level of the coracoids (fig.

5F), a gap separates the medial ligament (ml) and the right epicoracoid (re), which overlaps

the left extensively. The epicoracoids terminate in a pair of horns (fig. 5G), each of which lies

in a lateral sternal groove (sg). Parts of the horns are fused to the sternum. A laterally directed

ligament (Il) inserts on the posterior tips of the epicoracoid horns (fig. SH); a m. sternoepico-

racoideus is not evident.

Source : MNHN, Paris

84 ALYTES 21 (1-2)

A Gighd msc mi (left

ZE Trop, ELA

TT era ELA

Fig. 4. - Histological cross sections, in an antero-posterior direction, of the ventromedial part of the

pectoral girdie of Rhinoderma darwinit (UMMZ 143361). The levels of the sections are indicated in

fig. 2. co, coracoid: e, epicoracoids; h, epicoracoid horn; K, keel: ml, medial ligament; mse, mr.

supracoracoïdeus, mse, m. sternocpicoracoideus; n, notch at the right side of the ventral kecl: s,

Sternum; su, suture; x, part of the ventral keel replaced by cartilage

Source : MNHN, Paris

KAPLAN 85

Fig. 5. - Histological cross section, in an antero-posterior direction, of the ventromedial part of the

pectoral girdle of Insuetophrynus acarpicus (UMMZ 225142). The levels of the sections are indicated

in fig. 2. dl, dorsal ligament: dp, dorsal protuberances; e, epicoracoids: h, epicoracoid horns: k keel:

le, left epicoracoid: Il, lateral ligament; m1, muscle 1; muscle 2; m3, muscle 3: ml, medial ligament:

Source : MNHN, Paris

86 ALYTES 21 (1-2)

GEOBATRACHUS WALKERI

Anteriomedially (fig. 2, 6A), the epicoracoids (e) are indistinguishably fused to one

another. Muscle 1 (ml) inserts on the lateral and ventral surfaces of the epicoracoids and on

the medial ligament (ml). Posterior to the procoracoids (fig. 6B), the fused epicoracoids bear

a shallow dorsomedial depression. Slightly posteriorly (fig. 6C), the coracoids (co) lie lateral

to the epicoracoids, which are separated by a suture (su) marked by a shallow dorsomedial

depression. Muscle 1 is divided into a wide slip, muscle La (mla) that inserts on the coracoids,

and a thin slip, muscle 1b (m1b) that inserts on the medial ligament. At the level of the

coracoid shafts (fig. 6D), an ovoid ventral keel (k) is evident. The epicoracoids bear a shallow

dorsomedial crevice; only muscle 1b is evident. Slightly posteriorly (fig. 6E), only the ventral

keel remains. The medial parts of the coracoids are replaced by a membrane (me). Muscle 1b

inserts on the lateral and dorsolateral surfaces of the keel and on the medial ligament.

Posteriorly (fig. 6F), the epicoracoids and medial part of the coracoids are evident and the

ventral keel is small. Posteriorly (fig. 6G), the epicoracoids diverge slightly from one another,

and muscle 3 (m3) inserts on their dorsal surfaces. At the posterior terminus (fig. 6H), the

epicoracoids and sternum are indistinguishably fused; neither epicoracoid horns nor the m.

sternoepicoracoideus is evident.

ATELOPUS SUBORNATUS

Anteromedially (fig. 7A-B), the epicoracoids (e) are indistinguishably fused to one

another. Posteriorly (fig. 7C), a small, triangular ventral keel (k) is evident. At the anterior

level of the coracoids (fig. 7D), the fused epicoracoids are oval in cross section and the m.

supracoracoideus (msc) inserts on the medial ligament (ml) and ventrolateral surfaces of the

epicoracoids, the m. coracoradialis (mer) on their dorsolateral and dorsal surfaces. Posteriorly

(fig. 7Ë), the coracoids laterally flank the epicoracoids, which are represented by two ovoid

elements that are fused medially. The m. rectus abdominis (mra) inserts on the dorsomedial

surfaces of the epicoracoids. At the posterior level of the coracoids (fig. 7F), the epicoracoid

horns (h) diverge from one another. The horns are fused partially to the sternum (s) and

sternal grooves are evident. The m. sternoepicoracoideus is absent.

OSORNOPHRYNE BUFONIFORMIS

Anteromedially (fig. 8, 9A-B), the epicoracoids (e) are indistinguishably fused to one

another. Slightly posteriorly (fig. 9C), a suture (su) separates the epicoracoids and a small,

triangular ventral keel (k) is evident. Muscle 1 (ml) inserts on the ventral surface of each

clavicle (cl) and the medial ligament (ml). Posteriorly (fig. 9D), the epicoracoids bear a

shallow, dorsomedial crevice and a rounded ventromedial keel. Muscle 1 inserts on the

ventromedial surfaces of the procoracoids, lateral surfaces of epicoracoids and medial

ligament. At the level of the coracoiïd fenestra (fig. 9E), the epicoracoids are expanded (i.e..

bladelike), flat, and slightly eurved. Muscle 1 inserts on the ventral surfaces of the epicora-

coids and medial ligament. An oblique suture (su) separates the two epicoracoids such that

the left epicoracoid bears the ventral keel. Between the coracoids (fig. 9F), the epicoracoids

Source : MNHN, Paris

KAPLAN 87

A (right) (left) B

mia <o cr 4

= =

Fig. 6. — Histological cross section, in an antero-posterior direction, of the ventromedial part of the

pectoral girdle of Geobatrachus walkeri (ICN 35186). The levels of the sections are indicated in fig,

2. co, coracoid: cr, crevice:e, epicoracoids: k, keel; ml, muscle 1; mla, muscle Ja: mb, muscle 1b; me,

membrane: ml, medial ligament; $, sternum; su, suture

Source : MNHN, Paris

88 ALYTES 21 (1-2)

Fig. 7. — ent view of te pectoral fps el HIER farci {adapted from, Karz AN. 1994) and

girdle of pis subornatus (ICN 5820) c

horns; k, keel; mra, m1. rectus abdominis; mer, m. “aracoudials.

supracoracoideus; p, procoracoids; s, sternur

corresponding transverse sections of drawir

Source : MNHN, Paris

KAPLAN 89

Osornophryne bufoniformis Rana sylvatica

Atopophrynus syntomopus

Fig. 8. Ventral view of the pectoral girdle of Osornophryne bufoniformis (adapted from RUZ-CARRANZA

& HERNANDEZ-CAMACHO, 1976), Rana sylvatica (adapted from TYsoN, 1987) and Aropophrynus

syntomopus (adapted from Myers & ForD, 1986). cl, clavicle; co, coracoid: e, epicoracoid: p,

procoracoid; pu, processus uncinatus: S, Sternum. Light gray: cartilage; dark gray: loose connective

tissue; white: bone. A-H: transverse sections corresponding to those of fig. 9-10.

Source : MNHN, Paris

90 ALYTES 21 (1-2)

Fig. 9. — Histological cross sections, in an antero-posterior direction, of the ventromedial part of the

pectoral girdle of Osornophryne bufoniformis (ICN 11505). The levels of the sections are indicated in

fig. 8. co, coracoid: el, clavicle: e, epicoracoid: er, crevice: K, keel; ml, muscle 1: ml, medial ligament:

s, Sternum; su, suture.

Source : MNHN, Paris

KAPLAN 91

are indistinguishably fused and lack both, a ventral keel and dorsal crevice. Posteriorly (fig.

9G), a suture separates the epicoracoids. At the level of the sternum (fig. 9H), the epicoracoids

are indistinguishably fused to this structure. Neither epicoracoid horns nor the mn. sternoepi-

coracoideus are present.

RANA SYLVATICA

Anteriorly (fig. 8, 10 A), the ventral keel of the epicoracoids bears a process (kp) that lies

in the biforated base of the omosternum (0). Slightly posteriorly (fig. 10B), the anterior

protuberance (pu) of each epicoracoid (1.e., processus uncinatus: FUSCH, 1926) laterally flanks

the omosternum. In posterior sections (fig. 10C), the keel process is joined with the processus

uncinatus. Near the anterior levels of the coracoid fenestra (fig. 10D), the epicoracoids bear a

dorsomedial crevice (cr). The m. supracoracoideus (msc) inserts on the ventral keel and the mn.

coracoradialis (mer) on the ventral surfaces of the clavicles. Posterior to the procoracoids (fig.

10E), the coracoids (co) flank laterally the epicoracoids and the epicoracoids are fused

ventrally. The mn. coracoradialis inserts on the lateral surface of each coracoid. At the midlevel

of the coracoids (fig. 10F), the epicoracoids are surrounded by bone and, except medially and

ventrally, they are eroded; the ventral keel (k) remains cartilaginous. Posteriorly (fig. 10G),

a suture (su) separates the ventral keel and the epicoracoids. Posteriorly (fig. 10H), the

ventral keel is replaced by the sternum (s) which is united synchondrotically with the keel. The

posterior termini of the epicoracoids diverge as horns (h), which are surrounded by bone

and fused to the dorsolateral parts of the sternum and to one another by a dorsal osseous

bridge.

HYMENOCHIRUS BOETTGERI

Anteromedially (fig. 11A-B), the epicoracoids (e) are indistinguishably fused to one

another. At the anterior level of the coracoid fenestra (fig. 11C), the epicoracoids are wide, flat

and expanded in cross section. The m. coracoradialis (mer) inserts on the lateral and ventro-

lateral surfaces of the epicoracoids. At the midlevel of the coracoid fenestra (fig. 11D), the

epicoracoids are narrower. Posterior to the coracoids (fig. 11E), the epicoracoids diverge

laterally as two epicoracoid horns. These horns have densely packed chondrocytes and are

fused to the sternum (s). The m. sternoepicoracoideus is absent.

FROSTIUS PERNAMBUCENSIS

Anteromedially (fig. 12A), the epicoracoids (e) are indistinguishably fused to one an-

other. Muscle 1 (ml) inserts on the ventromedial surface of each clavicle (cl). Posteriorly (fig.

12B), à suture separates the epicoracoids. Posterior to the procoracoids (fig. 12C), the

cpicoracoids are small and ovoid in sections and bear a dorsomedial depression. Muscle 1

inserts on the lateral, dorsolateral and ventrolateral surfaces of the epicoracoids and medial

ligament (ml). At the level of the coracoid (fig. 12D), the »r. rectus abdominis (mra) inserts on

the dorsomedial surface of the epicoracoids. Posteriorly (fig. 12E), the epicoracoids horns

Source : MNHN, Paris

ALYTES 21 (1-2)

Fig. 10. - Histological cross section, in an antero-posterior direction, of the ventromedial part of the

ee girdle of Rana sylvatica (UMMZ). The levels of the sections are LE me in fs 8.b, bone:

oracoid: cr, crevice; e, epicorac

ms, m. supracoracoideus:

coracoradialis:

suture.

Source : MNHN, Paris

KAPLAN 93

Hymenochirus curtipes

Fig. 11.- Ventral view of the pectoral girdle of Hymenochirus curtipes (adapted from DE ViLLiERs, 1929)

and histological cross section, in an antero-posterior direction, of the ventromedial part of the

pectoral girdle of Hymenochirus boettgeri (ÜMMZ). cl, clavicle; co, coracoid; e, epicoracoid: h,

epicoracoid horn; mer, m. coracoradiali , sternum. Gra tilage: white: bone.

A-D: corresponding cross sections of drawing and photos.

Source : MNHN, Paris

94 ALYTES 21 (1-2)

Fig. 12. - Histological cross section, in an antero-posterior direction, of the ventromedial part of the

pectoral girdle of Frostius permanbucensis (UMMZ 225143). cl, clavicle: e, epicoracoid: h, epicora-

coid horn: ml, muscle 1: mra, m. rectus abdominis: ml, medial ligament: s, sternum; su, suture: x, gap.

Source : MNHN, Paris

KAPLAN 95

diverge from one another. The sternum and horns, which have different coloration and cell

densities, are fused to one another. Posteriorly (fig. 12F), gaps separate the horns and sternum.

The mn. sternoepicoracoideus is absent.

DISCUSSION

Of the pectoral girdles studied, only those of Atelopus subornatus, Frostius pernambu-

censis, Hymenochirus boettgeri, Osornophryne bufoniformis and Rana sylvatica have com-

pletely fused, non-overlapping epicoracoids. However, the epicoracoids of each of these taxa,

are not fused throughout their entire lengths. The condition in Atopophrynus and Pseudhy-

menochirus is unknown. The epicoracoids of Geobatrachus walkeri are completely fused and

not overlapping except for a small area at the level of the coracoid shafts. At this level, the

structure of the epicoracoids and medial part of the coracoids is disorganized; this morphol-

Ogy is unique in anurans.

In Rhinoderma darwiniï, the epicoracoids are indistinguishably fused and not overlap-

ping anterior to the midlevel of the coracoid fenestra, but fused (i.e., through their overlap-

ping surfaces, rather than the medial ends) and overlapping posterior to this level. GRIFFITHS

(1957, 1963) considered the overlap of the epicoracoids to start posterior to the coracoids,

whereas KAPLAN (1993) thought that it started at the posterior level of the procoracoids. The

overlap of the epicoracoids is evident in froglets (stage 46) (KAPLAN, 1993); however, in

sexually mature individuals, it is evidenced only by the presence of an oblique medial suture

and a free (i.e., unfused) medial epicoracoid margin (fig. 2A-B). The girdle morphology in

which the epicoracoids are fused and overlapping in the same cross section, and where the

fused parts are their overlapping surfaces, rather than their medial ends, is unique in anurans.

In Brachycephalus ephippium, the epicoracoids are fused without overlap from their

anterior tips to the posterior level of the coracoid fenestra; posterior to this level, they are free

(i.e., attached by dense connective tissue) and overlapping. The girdle of B. ephippium is

arciferofirmisternal — viz., epicoracoids fused to, and not overlapping, one another from their

anterior tips to a level posterior to the clavicles and free, and overlapping, posteriorly to this

point (DUELLMAN & TRUEB, 1985). However, the girdle of B. ephippium differs from other

arciferofirmisternal girdles (e.g., Dendrophryniscus, Melanophryniscus) in having a very re-

duced area where the epicoracoids are free and overlapping.

TRUEB (1973) and ForD & CANNATELLA (1993) argued that Brachycephalus has com-

pletely ossified epicoracoids; this study shows that the epicoracoids are cartilaginous (i.e., by

definition, the epicoracoids are the cartilaginous remnant of the embryonic coracoid-

epicoracoid cartilage: FUSCH, 1926: TYsoN, 1987; KAPLAN, 1993). TRUEB (1973) considered

the epicoracoids to be juxtaposed, rather than fused, in 8. ephippium. My results indicate that

the epicoracoids are fused to one another up to the anterior level of the coracoids, and firmly

attached to one another posterior to this level.

The girdle of Ansuerophrynus acarpicus is arciferofirmisternal, because the non-

overlapping epicoracoids are fused from their anterior tips to the posterior level of the

procoracoids. They are unfused and overlapping posterior to this level.

Source : MNHN, Paris

96 ALYTES 21 (1-2)

The girdles of Atelopus subornatus, Frostius pernambucensis, Hymenochirus boettgeri

and Osornophryne bufoniformis differ from those of Rana sylvatica and Hoplobatrachus

chinensis (often referred to as Rana rugulosa, but see Kosucx et al., 2001) (KAPLAN, 2000) by

having the most anterior parts of the epicoracoids in contact with one another and indistin-

guishably fused. In R. sylvatica and H. chinensis, the anterior epicoracoids (i.e., the processus

uncinatus, because the anterior process of the ventral keel is of sternal origin; KAPLAN, 1993)

are free (i.e., attached by dense connective tissue) and diverging from one another. This

observation is consistent with the hypothesis that pseudofirmisterny and firmisterny evolved

independently; however, it is still unknown whether the girdle morphologies of R. sylvatica

and H. chinensis are widespread among Ranoidea. The characteristic girdle morphology

known long ago in Hoplobatrachus and related groups (BoLKAY, 1915) and referred to as

arcizony"” (DECKERT, 1938) or “arciferal-like condition” (TRUEB, 1973: 95) is a variant of

firmisterny and has nothing to do with pseudofirmisterny.

The girdles of Atelopus subornatus and Hymenochirus boettgeri differ from those of

Frostius pernambucensis and Osornophryne bufoniformis by having the epicoracoids indistin-

guishably fused. DE ViLLiErs (1929) incorrectly stated that in Hymenochirus the epicoracoids

indistinguishably fused to one another from their anterior tips to the posterior level of the

procoracoids and separated by a suture posterior to this level. These differences are minimal,

and it is still unknown whether the presence or absence of suture between the epicoracoids is

the result of ontogenetic and/or intraspecific variation. Therefore, the structural homology of

the girdles of À. subornatus, F. pernambucensis, H. boettgeri and O. bufoniformis is equivocal.

I found several morphological differences among the girdles of the non-ranoid frogs

studied. The epicoracoids are expanded and flat in A{ymenochirus boettgeri and Osornophryne

bufoniformis, but not expanded and ovoid in Frostius pernambucensis and Atelopus suborna-

but they do on the lateral and dorsal surfaces in 4. subornatus and F pernambucensis and in

the lateral surfaces in A. boettgeri. In O. bufoniformis, a suture separates the epicoracoids

asymmetrically whereas this separation is symmetrical in Æ pernambucensis. The systematic

value of these characters is unknown.

The hypothesis that pseudofirmisterny is a synapomorphy uniting Brachycephalus and

Atelopus (GRIFFITHS, 1963) and Brachycephalus and Psyllophryne (FORD & CANNATELLA,

1993) is false because B. ephippium lacks this character. Similarly, pseudofirmisterny does not

support the monophyly of Geobatrachus and Atopophrynus because Geobatrachus lacks this

character. Moreover, the girdles of these two taxa differ externally, as is evident by compari-

son of figures 2 and 8.

The character “epicoracoids completely fused to, and not overlapping, one another”

unites only Atelopus, Frostius and Osornophryne. However, there are morphological difer-

ences among their girdles (e.g., epicoracoids indistinguishably fused or partially separated by

a suture, symmetrically or asymmetrically separated by a suture, expanded or not expanded)

that may be phylogenctically informative.

Source : MNHN, Paris

KAPLAN 97

RESUMEN

Se estudiaron histologicamente las cinturas pectorales de las siguientes especies: Arelo-

pus subornatus, Brachycephalus ephippium, Frostius pernambucensis, Geobatrachus walkeri,

Hymenochirus boettgeri, Insuetophrynus acarpicus, Osornophryne bufoniformis, Rana sylva-

tica, Rhinoderma darwinü. Dentro de los anuros no ranoideos estudiados, solamente 4.

subornatus, F. permanbucensis, H. boettgeri y O. bufoniformis presentan los cartilagos epico-

racoidales completamente fusionados y sin sobrelaparse (i.e., pseudofirmisternia). Las cintu-

ras pectorales de G walkeri y R. darwinii son unicas dentro de los anuros. Las cinturas

pectorales de B. ephippium y I. acarpicus son arciferofirmisternales. Las diferencias morfolo-

gicas entre las cinturas pectorales firmisternales y pseudofirmisternales sugieren que estas no

son homologas.

ACKNOWLEDGMENTS

I acknowledge Ramon Formas, Arnold Kluge, the late Pedro Ruiz, and John Lynch for loan of

specimens, Arnold Kluge, Linda Trueb, and two anonymous reviewers for revision of the manuscript,

Greg Schneider for assistance with the material, and John Megahan for assistance with the figures.

LITERATURE CITED

ARDiLLA-RoBayo, M. C., 1979, - Status sistematico del genero Geobatrachus Ruthven 1915 (Amphibia:

Anura). Caldasia, 12: 383-495.

BaRiO, A., 1970. — Insuetophrynus acarpicus, un nuevo leptodactylido firmisterno sudamericano

(Amphibia: Anura). Physis, 30: 331-341.

BepparD, FE. 1895. - On the diaphragm and on the muscular anatomy of Xenopus, with remarks on its

affinities. Proc. zool. Soc. London, 1895: 841-850.

2e 1908. - A contribution to the knowledge of the batrachian Rhinoderma darwinit. Proc. zool. Soc.

London, 1908: 678-694.

BoLkAY, S. J., 1915. - Beiträge zur Osteologie einiger exotischer Raniden. Anar. Anz., 48: 172-183.

CANNATELLA, D. C., 1985. — À phylogeny of the so-called primitive frogs (Archaeobatrachians). PhD

Thesis, Lawrence, Univ. Kansas: 1-350.

= 1986. — A new genus of bufonid (Anura) from South America, and phylogenetic relationships of

neotropical genera. Herpetologica, 42: 197-2

CANNATELLA, D. C. & TRUEB, L., 1988. - Evolution of pipoid frogs: intergeneric relationships of the

aquatic frog family Pipidae (Anura). Zool. 4 linn. Soc., 94: 1-38.

Cut, J. M. 1980. - Amphibians of Argentina. Monit. £oal. ital.,

De Vies, C. G. $.. 1929. - The comparative anatomy of the breast-shoulder apparatus of the three

Aglossa Anura gencra: Xenopus, Pipa, and Hymenochirus. Ann. Transvaal mus., 13: 37-69.

Drckert, K. 1938. — Beiträge zur Osteologie und Systematik ranider Froschlurche. Sher. Ges. Narurf

Freunde Berlin, 1938: 127-184.

DuELLMAN, W. E. & Trus8, L., 1985. - Biology of amphibians. New York, McGraw-Hill, 1986": 1-xix +

1-670.

Source : MNHN, Paris

98 ALYTES 21 (1-2)

Fawcerr, D. W, 1986. — Bloom and Fawcett. A textbook of histology. Philadephia, W. B. Saunders,

Philadelphia: i-xi + 1-1017.

FELLER, À. E. & HEDGES, S. B., 1998. - Molecular evidence for the early history of living amphibians. Mol.

Phyl. Evol., 9: 509-516.

Forp, L. $. & CANNATELLA, D. C., 1993. - The major clades of frogs. Herp. Mon., 7: 94-117.

FuscH, H., 1926. - Beitrage zur Entwicklungsgeschichte und vergleichenden Anatomie des Brustschul-

terapparates der Wirbeltiere, Funfte mitteilung. Ueber den Schultergurtel der Amphibia Anura.

JL. Von der naturlichen Unterbrechung der Cartilago procoracoidea und von dem Fenster am

Schultergurtel der Rana fusca. Anat. Anz., 61: 1-34.

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

identification. Herpetologica, 16: 183-190.

GRAYBEAL, A., 1997. - Phylogenetic relationships of bufonid frogs and test of alternative macroevolu-

tionary hypotheses characterizing their radiation. Zoo!. J linn. Soc., 119: 297-338.

Grirrrrus, L, 1957. - The structure and development of the breast-shoulder apparatus in Amphibia,

Salientia. Part 1. PhD Thesis, London, Univ. London: 1-339.

1963. - The phylogeny of Salientia. Biol. Rev., 38: 241-293.

Hay, I M., RUvVINSKY, L., HEDGES, S. B. & MaxsON, L. R., 1995. - Phylogenetic relationships of

amphibian families inferred from DNA sequences of mitochondrial 12S and 16$ ribosomal RNA

genes. Mol. Biol. Evol., 12: 928-937.

KAPLAN, M., 1993. — An evaluation of the current use of the breast-shoulder apparatus characters in

anurans. PhD Thesis, Lincoln, University of Nebraska: 1-464.

1994. — Analysis of some long-standing controverses concerning the pectoral girdle of Arelopus

(Bufonidae) using ontogenetic studies. J. Herp., 28: 128-131.

ie 2000. - The pectoral girdles of Rana rugulosa (Ranidae) and Nesomantis thomasseti (Sooglossidae).

Herpetologica, 56: 188-195.

ans 2002. - Histology of the anteroventral part of the breast-shoulder apparatus of Brachycephalus

ephippium (Brachycephalidae) with comments on the validity of the genus Psy/lophryne (Brachy-

cephalidae). Amphibia- Reptilia, 23: 225-227.

KosucH, J., VENCES, M., Dugois, A., OHLER, A. & BÔHME, W., 2001. — Out of Asia: mitochondrial DNA

evidence for an Oriental origin of tiger frogs, genus Hoplobatrachus. Mol. Phyl. Evol., 21 (3):

398-407.

LyNcH, J. D., 1973. - The transition from archaic to advanced frogs. Jn: JL. ViaL (ed.), Evolutionary

biology of the anurans: contemporary research on major problems, Columbia, Univ. Missouri Press:

133-182.

En 1978. — À re-assessment of the telmatobiine leptodactylid frogs of Patagonia. Occ. Pap. Mus. nat

Hist. Univ. Kansas, 72: 1-57.

LyNCH, J. D. & RUIZ-CARRANZA, P. M., 1982. - A new genus and species of poison-dart frog (Amphibian,

Dendrobatidae) from the Andes of northern Colombia. Proc. biol. Soc. Washington, 95: 557-562.

MCDiarMiD, R. W, 1969. — Comparative morphology and evolution of the neotropical frogs genera

Atelopus, Dendrophryniseus, Melanophryniseus, Oreophrynella, and Brachycephalus. PhD The-

sis, Los Angeles, Univ. Southern Californi vii + 1-282.

MCLACHLAN, P, 1943. - The cranial and visceral osteology of the neotropical anuran Brachycephalus

ephippium Spix. S. af. J. Sci., 40: 164-195.

Myers, W. C. & ForD, L. S., 1986. - On Aropophrynus, a recently described frog wrongly assigned to

Dendrobatidae. Am. Mus. Nov., 2843: 1-15.

Nour, G. K., 1922. - The phylogeny of the Salientia. L. The osteology and the thigh musculature: their

bearing on classification and phylogeny. Bull. am. Mus. nat. Hist., 46: 1-87.

1926. - The pectoral girdle of brachycephalid frogs. Am. Mus. Nov., 230: 1-14.

CARRANZA, P. M. & HERNANDEZ-CAMACHO, JL. 1976. - Osornophryne, genero nuevo de anfibios

bufonidos de Colombia y Ecuador. Caldasia, 1: 93-148.

Truss, L. 1973. - Bones, frogs and evolution. /n: J. L. ViaL (ed.), Evolutionary biology of the anurans

‘ch on major problems, Columbia, Univ. Missouri Press: 65-132

contemporary res

Source : MNHN, Paris

KAPLAN 99

TysoN, H., 1987. - The structure and development of the anuran breast-shoulder apparatus, forelimb, and

associated musculature. PhD Thesis, Edmonton, Univ. Alberta: 1-1275.

Vexces, M., Kosucn, J., LorTERs, $., WIDMER, À. JUNGrER, K. H., KÔHLER, J., & Verri, M., 2000. —

Phylogeny and classification of poison frogs (Amphibia: Dendrobatidae), based on mitochondrial

16$ and 12 ribosomal RNA gene sequences. Mol. Phyl. Evol., 15: 34-40.

Vexces, M. & GLAw, F, 2001. - When molecules claim for taxonomic changes: new proposals on the

classification of Old World tree frogs. Spixiana, 24: 85-92.

WEssner, FE. M., 1960. - General zoological microtechniques. Baltimore, Williams & Wilkins: 1-230.

Corresponding editor: Alain DUBOIS.

Source : MNHN, Paris

Alytes, 2003, 21 (1-2): 100-102. Book review

Comments on a new book on the

Amphibia of Thailand,

with a tentative allocation

of the figured species

Annemarie OHLER

Vertébrés (Reptiles & Amphibiens)

USM 0602 Taxonomie & Collectioi

Département de Systématique & Evolution,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

<ohler@mnhn.fr>

NuTPHUND, Wirot. — Amphibians of Thailand. Thailand, Amarin Printing and Publishing Public Co.,

Thailand, 2001: 1-192. ISBN 974-7751-70-4.

Amphibians of Thailand by Wirot Nutphund is a nice book, with numerous colour drawings and photos for

many of the Thaï species. For each of them the author gives scientific, English and Thai name, followed by a

short description, notes on ecology and distribution data. He also presents drawings of eggs and tadpoles of

various species. However, beside its very nice aesthetic aspect, this book contains many errors. As it is sold in

general book stores and on internet, and will be purchased by many people, I decided to publish corrected

legends to photos, For many photographs it is not possible to allocate species, so only generic names are given.

These tentative allocations, presented in table 1, are exclusively done for the species figured on a photo

(NurHuND, 2001: 60-177). Drawings of this book, that show a more artistic than scientific expression, have not

been studied. The valid names of all species with their author and date of publication are given for allocated

species (for ranids, see Dupois, 1992).

ADDITIONAL COMMENTS

The photo p. 103 (NUrPHUN», 2001) shows a particular colour morph of Rana erpthraea with à fine,

indistinct mid-dorsal line (Bourer, 1942: 331). The overal colour of the frog is darker than the usual bright

grass green.

The photo p. 149 (NUrPHUND, 2001) represents an interesting colour mutation, resulting in a beautiful sky

blue frog with orange webbing on hand and feet and small white dots on the back. This blue colour was found

in other species of frogs like in Rana synkl. esculenta Linnaeus, 1758 and /yla arborea (Linnaeus, 1758). Itis due

to absence of yellow xanthophores in the skin. The frog can with no doubt be allocated to Rhacophorus

bipunctatus Al, 1927.

BOURRET (1942) desci

adult male, B.36 adult fema

covered by large-sized ir

s Kaloula pulchra macrocephala (syntypes. Hanoi University B.35,

Indochina) showing indistinet dorsolateral bands and mid-dorsum

The holotype by monotypy of Kaloula aureata Nutphand, 1989

(MNHN 1997.4923, adult male, SVL 59.9 mm after preservation, donated by Nutphund to the Paris Museum:

iype-locality: Thung Song District, Nakhon Si Thammarat, Thailand) is figured p. 163 (NUTPHUND, 2001) and

shows a similar dorsal pattern as the holotype of Kaloula pulchra macrocephala. The name Kaloula aureata

phand, 1989 is here tentatively considered a subjective junior synonym of Kaloula macrocephala Bourret.

1942

Source : MNHN, Paris

OHLER 101

Table 1. Tentative allocation of specific names to photographs of Amphibia in NUTPHUND (2001).

Page Name in NUTHPUND (2001) Corrected name

60 | Tylototriton verrucosus Tylototriton verrucossus Anderson, 1871

61 Ichthyophis kohtaoensis Ichthyophis kohtaoensis Taylor, 1959

64-65 | Bufo melanostictus Bufo melanostictus Schneider, 1799

71 Bufo asper Bufo asper (Gravenhorst, 1829)

72 | Bufo macrotis Bufo macrotis Boulenger, 1887

73 | Bufo parvas [sic] Bufo parvus Boulenger, 1887

74 | Ansonia malayana Bufo asper (Gravenhorst, 1829), juvenile

75 | Pedostibes hosit Pedostibes hosit (Boulenger, 1892)

76 | Leptobrachium hasseltit Micryletta inornata (Boulenger, 1890)

77 | Leptobrachium hendricksoni Leptobrachium hendricksoni Taylor, 1962

79 | Leptobrachium minimum Kalophrynus interlineatus (Blyth, 1855)

80 | Megophrys monticola Xenophrys sp.

sl Megophrys monticola nasuta Megophrys nasuta (Schlegel, 1837)

82 | Megophrys carienensis [sic] Brachytarsophrys carinensis (Boulenger, 1889)

83 | Megophrys feae Brachytarsophrys feae (Boulenger, 1887)

84 | Megophrys longipes Xenophrys Sp.

85 | Megophrys parva Xenophrys Sp.

86 | Megophrys aceras Xenophrys aceras (Boulenger, 1899)

87 | Megophrys major Xenophrys major (Boulenger, 1908)

89 | Hyla annectens [sic] Rana (Odorrana) andersoni Boulenger, 1882

90 | Ooeidozyga lima Occidozyga lima (Gravenhorst, 1829)

91 Phrynoglossus magnapustulosus Limnonectes sp.

92 | Phrynoglossus martensi Phrynoglossus Sp

97 | Phrynoglossus laevis Phrynoglossus Sp.

98 | Staurois larutensis Amolops (4mo) larutensis (Boulenger, 1899)

99 | Staurois afghanus Rhacophorus feae Boulenger, 1893

100 | Elachyglossa gyldenstolpei Limnonectes laticeps (Boulenger, 1882)

101 | Rana leptoglossa Rana (Sylvirana) sp.

102 | Rana tasanae Limnonectes sp.

103 | Rana tenasserimensis Rana (Hylarana) erythraea (Schlegel, 1837)

104 | Rana andersoni Rana (Pelophylax) lateralis Boulenger, 1920

105 | Rana blythit Limnonectes blythi (Boulenger, 1920)

106 | Rana macrodon Limnonectes kuhlii (Tschudi, 1838)

107 | Rana cyanophlyctis Limnonectes sp.

108 | Rana tigerina Hoplobatrachus chinensis (Osbeck, 1765)

111 | Rana rugulosa Hoplobatrachus chinensis (Osbeck, 1765)

112 | Rana limnocharis Fejervarya limnocharis (Gravenhorst, 1829)

113 | Rana erythraea Rana (Hylarana) erythraea (Schlegel, 1837)

114 | Rana macrodactyla Rana (Hylarana) macrodactyla (Günther, 1859)

115 | Rana cubitalis Limnonectes sp.

116 | Rana miopus Rana (Sylvirana) cubitalis (Smith, 1917)

117 | Rana glandulosa Rana (Pulchrana) glandulosa Boulenger, 1882

118 | Rana chalconota Rana (Chalcorana) chalconota (Schlegel, 1837)

119 | Rana nigrovittata Rana (Sylvirana) nigrovittata (Blyth, 1855)

120 | Rana lateralis Limnonectes sp.

123 | Rana host Same individual as p. 118 as Rana chalconota, ï.e

Rana (Chalcorana) chalconota (Schlegel, 1837)

124 | Rana livida Rana (Eburana) livida (Blyth, 1855)

125 Rana scutigera Rana (Sylvirana) sp

126 | Rana doriae Limnonectes blythi (Boulenger, 1920)

127 | Rana kochangae [sie] Rana (Pulchrana) glandulosa Boulenger, 1882

129 | Rana pileata | Limnonectes gyldenstolpei (Anderson, 1916)

Source : MNHN, Paris

102

ALYTES 21 (1-2)

Table 1. (continued).

Page Name in NUTHPUND (2001) Corrected name

130 | Rana hacheana [sic] Taylorana hascheana (Stoliczka, 1870)

130 | Rana laticeps Philautus sp.

135 | Rana cancrivora Fejervarya cancrivora (Gravenhorst, 1829)

136-137 | Rana fasciculospina [sie] Paa (Quasipaa) fasciculispina (Inger, 1970)

139 | Rhacophorus leucomystax leucomystax | Polypedates leucomystax (Gravenhorst, 1829)

140 | Rhacophorus leucomystax sexvirgatus | Polypedates leucomystax (Gravenhorst, 1829)

141 Rhacophorus robinsoni Rhacophorus robinsoni Boulenger, 1903

143 | Rhacophorus colleti Polypedates sp.

147 | Rhacophorus nigropalmatus Rhacophorus nigropalmatus Boulenger, 1895

149 | Rhacophorus (new sp.) Rhacophorus bipunctatus Ab, 1927

153 | Philautus vittatus Polypedates sp.

154 Philautus hansenae Hyla annectans (Jerdon, 1870)

155 Philautus doriae Philautus sp.

156 | Philautus nongkhorensis Philautus sp.

157 | Philautus bimaculatus Polypedates sp.

158 | Philautus parvulus Polypedates sp.

159 Kaloula pulchra Kaloula pulchra Gray, 1831

161 Kaloula mediolineata Kaloula mediolineata Smith, 1917

162 Kaloula baleata Kaloula baleata (Müller, 1836)

163 Kaloula aureata Kaloula macrocephala Bourret, 1942

164-65 | Calluella guttulata Calluella guttulata (Blyth, 1855)

166 | Glyphoglossus molossus Glyphoglossus molossus Günther, 1868

167 | Kalophrynus pleurostigma Kalophrynus pleurostigma Tschudi, 1838

169 | Microhyla pulchra Microhyla pulchra (Haïlowell, 1860)

170 Microhyla annamensis Microhyla berdmorei (Blyth, 1856)

171 | Microhyla ornata Microhyla heymonsi Vogt, 1911

174 | Microhyla inornata inornata Microhyla ornata (Duméril & Bibron, 1841)

175 | Microhyla inornata lineata Micryletta inornata (Boulenger, 1890)

176 | Microhyla berdmorei Microhyla butleri Boulenger, 1900

177 | Microhyla heymonsi Microhyla heymonsi Vogt, 1911

The error rate of taxonomic allocation is very high in this book: out of 81 species photographed, only 36are

allocated to the proper species. The taxonomy corresponds to the level of TAYLOR (1962) and no recent result

even the most conservative, have been introduced. In conclusion this book should not been considered as

reference for any scientific Work.

LITERATURE CITED

BOURRET, R.. 1942.

1-547,4 pl.

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

Lyon, 61 (10): 305-352

TavyLor, E. H., 1962. - The amphibian fauna of Thailand. Univ. Kansas Sci. Bull., 63

Les Batraciens de l'Indochine. Hanoï, Institut océanographique de l'Indochine: i-x +

65-599,

Source : MNHN, Paris

Alytes, 2003, 21 (1-2): 103-104. Book review

Recent books on the amphibians of Europe

Annemarie OHLER

Vertébrés (Reptiles & Amphibiens),

USM 0602 Taxonomie & Collections,

Département de Systématique & Evolution,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

<ohler@mnhn.fr>

Birgit GOLLMANN & Günter GOLLMANN. — Die Gelbbauchunke: von der Suhle zur Radspur. Bielefeld,

Laurenti Verlag, Beiheft der Zeitschrift für Feldherpetologie, 4, 2002: 1-135, 35 figures and 4 tables.

Softbound. ISBN 3-933066-10-7.

Andrea AMBROGIO & Luca GiLL1. — A tritone alpestre. Tassonimia. Distribuzione. Sviluppo. Ecologia.

Etologia. Protezione. Cavriago, Planorbis, 1998: 1-64. Sofibound.

In the series of monographies on European amphibians, a book on the yellow-bellied toad, Bomibina

variegata (Linnaeus, 1758) was missing. This species seems 10 be particularly endangered and many

populations have disappeared in the last decades. À summary of our knowledge on these toads should be

an important aid for conservation biologists and environmentalists.

The book by Birgit & Günter GOLLMANN is written in German language. It is original in adding to

the known data from the literature many new and unpublished observations. The review is clearly

documented by references and accompanied by many figures. Specimens and habitats are presented in

colour photos.

The phylogeny, taxonomy and systematics are reviewed shortly. Data on size variation and colour

variation try to describe the large differentiation between populations and subspecies, whose phylogenetic

relationships have been investigated by various methods. The phenomenon of hybridization with the

sister taxon Bombina bombina (Linnaeus, 1761) is shortly presented.

Habitat of the species includes breeding places, hibernation and aestivation areas. B. variegata Shows

à large scale of breeding environments which include secondary habitats as road ditches and 1

many other kinds of small water bodies. Not much is known on aestivation and hibernation places.

Yellow-bellied toads are then presented as predators and victims, giving lists of prey and predators

but also treating the behaviour of the toads. A nice description tries to elucidate form and function of the

“Unkenreflex”. B. variegata shows annual and diurnal activity cycles, in particular in the presence of

n the habitat, and in relation to reproduction. Like in most anurans the bulk of data

available concerns reproduction: place and date, regulation, mating call — or advertisement call as it

should better be named -, amplexus, egg laying. Itis particularly difficult to obtain simple data like clutch

size in these toads, as they lay eggs in several patches, Observations of multiple breeding were made both

in males and females. Embryonic and larval development are described, providing data on morphology,

nutrition, influence of temperature and density, as well as results from competition experiments. Finally

various aspects of metamorphosis are discussed to conclude the description of the life cycle

Aninteresting part of the book presents the spatial distribution of the toads. Based on observations

of individual toads — recognised thanks to the particular ventral pattern which is unique and remains

rather constant throughout the life of a toad -, observations of migration, choice of mating places,

immigration in new habitats, periods spent on breeding ponds and fidelity to a certain place are given.

This raises the question of spatial orientation on which no recent investigations have been made.

Source : MNHN, Paris

104 ALYTES 21 (1-2)

Another question concerns longevity, and we learn that individuals of more than 15 years have been

observed in the wild, and that these toads may reach 29 years in captivity. Sexual and age composition of

toad populations are presented and variations between the populations are discussed. Population

dynamics of different life history stages show major differences, the early stages being particularly

affected by loss.

A chapter on conservation discusses legal situations of protection of the toads, and causes of threats.

The use of land by man leading to the loss of wetlands is mentioned as a major factor. Isolation of

suitable habitats, introduction of fish and also pollution might be factors for regression of this species.

The authors claim actions for habitat protection and management in order to protect the yellow-bellied

toad. Introduction of alien specimens in populations should be avoided because it leads to genetic

pollution.

In a final chapter, the methodology of study is presented in order to encourage the reader to study

these interesting animals. This is a basic book, interesting for the herpetologist and the conservationist,

who look for precise data on Bombina variegata, and in particular for the student, as it stimulates

observation and study of these nice and secretive animals.

Andrea AMBROGIO and Luca GiLL1 published a nice booklet on the Alpine newt, Triturus alpestris

(Laurenti, 1768). It is a well done, pleasantly presented book, reflecting the famous Italian sense of taste.

Color photos and figures are of high quality. The pencil sketches recall the notebook of a naturalist, AI

details of the book are thoroughly drawn, like the small figures on right top of the right pages that

illustrates ontogenetical development of the new.

The authors present the species by an introduction to its taxonomy and systematics, giving descrip-

tion of external morphology of the Italian subspecies. Then the geographical and altitudinal distributions

of these subspecies are shown and discussed. The chapter on the habitat is documented by numerous

photos representing as well the ponds and landscapes as the underwater biotopes where the news live.

In a large chapter on eco-ethology, the feeding behavior of adults and larvae is presented in its

different aspects, such as annual variation, nutrition in water and on land, nutrition of adults and larvae,

composition of diet, but also predation of newts. Another aspect shown is the life history and population

structure of Alpine newts. This chapter is completed by the presentation of the amphibian species

syntopic with this newt and their daily activity patterns.

The reproduction of Triturus alpestris is introduced in a well-documented part. Many color photos

illustrate the descriptions of the breeding behavior showing the male-female interactions. The whole

sequence is then summarized and presented in color drawings. À figure explaining the particular terms of

the behavior shows small sketches of the movements and their names. The chapter finishes with the

description of the egg-laying behavior of the female.

Then the book follows the life history of the newt in presenting a chapter on the development, growth

and metamorphosis. Beside normal development and its parameters, the particular case of neoteny or

pedomorphosis is described. À clear scheme depicts the live cycle of Triturus alpes

The book finishes on an important aspect, the conservation of the Alpine newt giving the protection

status of the Italian populations. The content of the book is thoroughly documented by the relevant

literature, so that the reader can find original publications concerning all subjects.

In summary the book is an interesting presentation of this species, in particular for the Italian

peninsula. Itis written in Italian, but the rich illustrations make it attractive for all herpetologists. It seems

to be the first booklet of a series from this editor and a second on the fire-bellied toads of the genus

Bombina is announced.

Source : MNHN, Paris

ADVTES

International Journal of Batrachology

published by ISSCA

EDITORIAL BOARD

Chief Editor: Alain Dupois (Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France; <dubois@mnhn.fr>).

Deputy Editor: Thierry LODÉ (Laboratoire d'Ecologie animale, Université d'Angers, 2 boulevard Lavoisier,

49045 Angers Cedex, France; <thierry.lode@univ-angers.fr>).

Conservation Editor: Stephen J. RicHARDs (Vertebrates Department, South Australian Museum, North Terrace,

Adelaide, S.A. 5000, Australia; <Richards.Steve@saugov.sa.gov.au>).

Editorial Board: Franco ANDREONE (Torino, Italy); Lauren E. BRowN (Normal, USA); Janalee P. CALDWELL

(Norman, USA): Ulisses CARAMASCHI (Rio de Janeiro, Brazil): Günter GOLLMANN (Wien, Austria); Heinz

GriLLrrsCH (Wien, Austria); Tim HALLIDAY (Milton Keynes, United Kingdom); W. Ronald HEvER

(Washington, USA); Esteban O. LAVILLA (Tucumän, Argentina); Masafumi MaTsu1 (Kyoto, Japan);

Donald Ë MCALPINE (Saint John, Canada); Alain PAGANO (Angers, France); John C. POYNTON (London,

England); Miguel VENCES (Konstanz, Germany).

Technical Editorial Team (Paris, France): Alain DUBOIS (texts); Roger BoUR (tables); Annemarie OHLer (figures).

Book Review Editor: Annemarie OHLER (Paris, France).

SHORT GUIDE FOR AUTHORS

(for more detailed Instructions 10 Authors, see Alytes, 1997, 14: 175-200)

Alytes publishes original papers in English, French or Spanish, in any discipline dealing with amphibians.

Beside articles and notes reporting results of original research, consideration is given for publication to synthetic

review articles, book reviews, comments and replies, and to papers based upon original high quality illustrations

(such as colour or black and white photographs), showing beautiful or rare species, interesting behaviours, 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 em. 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: i-x + 1-547,

I. 1-4.

Graf, J.-D. & PoLs PeLaz, M., 1989. - Evolutionary genetics of the Rana esculenta complex. In: R. M. DAWLEY

& J. P. BOGART (ed.), Evolution and ecology of unisexual vertebrates, Albany, The New York State Museum:

289-302.

INGER, R. E, Voris, H. K. & Voris, H. H,, 1974. - Genetic variation añd population ecology of some Southeast

Asian frogs of the genera Bufo and Rana. Biochem. Genet., 2: 121-145.

Manuscripts should be submitted in triplicate either to Alain DuBois (address above) if dealing with

amphibian morphology, anatomy, systematics, biogeography, evolution, genetics, anomalies or developmental

biology, or to Thierry LODÉ (address above) if dealing with amphibian population genetics, ecology, ethology or

life history, or to Stephen J. RICHARDS (address above) if dealing with conservation biology, including declining

amphibian populations or pathology. Acceptance for publication will be decided by the editors following review

by at least two referees. à

If possible, after acceptance, a copy of the final manuscrit on a floppy disk (3 V4 or 5 %) 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 4 high density 1.44 Mo floppy disk only).

Page charges are requested only from authors having institutional support for this purpose, The publication

of colour photographs is charged. For each published paper, 25 free reprints are offered by ISSCA to the

author(s). Additional reprints may be purchased.

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.

Source : MNHN, Paris:

Alytes, 2003, 21 (1-2): 1-104.

Contents

Alain Dugois

Editorial.

Should internet sites be mentioned in the bibliographies of scientific

UDC EDS TS de er nn er A ne 12

Steve A. JOHNSON

Orientation and migration distances of a pond-breeding salamander

(Salamandridae, Norophthalmus perstriatus) .…........................... 3-2

Annemarie OHLER

Revision of the genus Ophryophryne Boulenger, 1903 (Megophryidae)

with description Of two new Species ...................................... 23-44

Stéphane GROSIEAN

A redescription of the external and buccopharyngeal morphology

of the tadpole of Ophryophryne microstoma Boulenger, 1903

MCE ODÉ D TAR) ee CET ct ea. PONS MONS Pa LS APS Lee 45-58

Liliane RAHARIVOLOLONIAINA, David R. VirtTes, Frank GLaw & Miguel VENCES

Larval stages, habitat and distribution

of the hyperoliid frog Heterixalus rutenbergi (Boettger, 1881) ................. 59-65

Zandra E. ULLOA KREISEL

Morfologia del tubo digestivo en larvas de Dermatonotus muelleri y

Elachistocleis bicolor (Anura, Microhylidae) del Noroeste argentino ........... 66-76

Moises KAPLAN

The “pseudofirmisternal” pectoral girdle of anurans . 77-99

BOOK REVIEWS

Annemarie OHLER

Comments on a new book on the Amphibia of Thailand,

with a tentative allocation of the figured species …......................... 100-102

Annemarie OHLER

Recent books on the amphibians of Europe ............................... 103-104

Alytes is printed on acid-free paper.

Alÿtes is indexed in Biosis, Cambridge Scientific Abstracts, Current Awareness in Biological Scien-

ces, Pascal, Referativy Zhurnal and The Zoological Record.

Imprimerie F. Paillart, Abbeville, France.

Dépôt légal: 4 trimestre 2003.

©ISSCA 2003

Source : MNHN, Paris