European Journal of Taxonomy 9: 1-9

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2012 • Frank Glaw et al.

Research article

A tiny new species of Platypelis from the Marojejy National Park

in northeastern Madagascar (Amphibia: Microhylidae)

Glaw R, Kohler J. & Vences M. 2012. A tiny new species of Platypelis from the Marojejy National Park in

northeastern Madagascar (Amphibia: Microhylidae). European Journal of Taxonomy 9: 1-9. httD://dx.doi.

org/lQ.5852/eit.2Q12.9

Frank GFAW^, Torn KOHFER^ & Miguel VENCES^^

^Zoologische Staatssammlung Mtinchen, Mtinchhausenstr. 21, 81247 Mtinchen, Germany.

^Hessisches Eandesmuseum Darmstadt, Friedensplatz 1, 64283 Darmstadt, Germany.

^Division of Evolutionary Biology, Zoologieal Institute, Teehnieal University of Braunsehweig,

Mendelssohnstr.4, 38106 Braunsehweig, Germany.

"^Email: [email protected]

Abstract. We deseribe a tiny new frog speeies of the genus Platypelis (Anura: Mierohylidae: Cophylinae)

from Marojejy National Park, northeastern Madagasear. Platypelis ravus sp. nov. differs from all other

known Platypelis and Cophyla speeies by its small size (17-19 mm snout-vent length) and a eombination

of other morphologieal and bioaeoustie eharaeters. The new speeies seems to be most elosely related to

P. milloti with whieh it shares the prineipal eolour pattern, but exhibits a yellow rather than red posterior

venter. Uneorreeted pairwise sequenee divergenee in a 16S rRNA gene fragment to all other known

speeies of the genus (exeept P. cowanii for whieh no genetie data is available) is greater than 6%. We

suggest the inelusion of the new speeies in the lUCN threat eategory “Data Defieienf’.

Key words. Mierohylidae, Cophylinae, Platypelis ravus sp. nov., Madagasear, Marojejy National Park.

Introduction

Narrow-mouthed frogs, family Mierohylidae Gunther, 1858, are a speeies-rieh elade of almost

eosmopolitan distribution and only partly elarified phylogenetie relationships (Van Boexlaer et al. 2006;

Van der Meijden et al. 2007; Kurabayashi et al. 2011). Madagasear’s mierohylids are elassified in three

endemie subfamilies, Cophylinae Cope, 1889, Dyseophinae Boulenger, 1882 and Seaphiophryninae

Eaurent, 1946 (Blommers-Sehlosser & Blane 1991) of whieh the Cophylinae is by far the most speeies-

rieh group and sister to the Seaphiophryninae (Van der Meijden et al. 2007). Cophylines are eharaeterized

by the presenee of endotrophie (non-feeding) tadpoles that develop either in water-filled tree holes

or similar eavities, in foam nests, or in subterraneous jelly nests (Blommers-Sehlosser 1975; Glaw &

Venees 2007). With few exeeptions, the advertisement ealls of eophyline speeies are highly stereotyped,

eonsisting of regularly repeated single notes whieh typieally are melodious whistles or elieks (Venees

et al. 2006).

Aeeording to Glaw & Venees (2007) and subsequent speeies deseriptions (summarized in AmphibiaWeb

2012), eophylines eurrently eomprise seven genera with altogether 58 speeies: Anodonthyla Muller,

European Journal of Taxonomy 9 : 1-9 ( 2012 )

1892 (11 species), Cophyla Boettger, 1880 (3 species), Madecassophryne Guibe, 1974 (1 species),

Platypelis^oulQngQX, 1882 (10 species), Plethodontohyla BoulQngQX, 1882 (10 species), Rhombophryne

Boettger, 1880 (10 species), and Stumpffia Boettger, 1881 (13 species). These frogs have radiated into a

variety of niches and contain arboreal, terrestrial and fossorial frogs (Andreone et al 2005; Wollenberg

et al. 2008). The subfamily contains species as large as 100 mm and miniaturized species as small as

10 mm snout-vent length (Glaw & Vences 2007). Field surveys and molecular assessments of diversity

(e.g. Vieites et al. 2009) have provided evidence that the species inventory of the cophylines is far

from being completed, and a large number of undescribed candidate species is known from most of the

cophyline genera.

Arboreal cophylines include the gmeraAnodonthyla, Cophyla, Platypelis and a few species of the genus

Plethodontohyla. Anodonthyla is relatively well characterized by the presence of a distinct prepollex in

males and the absence of vomerine teeth. The distinction of Cophyla and Platypelis is less clear and

mainly based on osteological characters (Blommers-Schlosser & Blanc 1991). Cophyla is the sister

group of Platypelis based on molecular data (Wollenberg et al. 2008).

During fieldwork in 2005, we collected specimens of a Platypelis from the Marojejy National Park in

northeastern Madagascar that distinctly differ from all cophylines known so far. We describe this new

species in the following.

Material and Methods

Specimens were collected at night by opportunistic searching and localizing calling males, using torches

and head lamps. They were euthanized in a chlorobutanol solution, fixed in 95% ethanol and preserved in

70% ethanol. Locality information was recorded with GPS receivers. Type specimens of the new species

are deposited in the Zoologische Staatssammlung Mtinchen (ZSM) and the Universite d’Antananarivo,

Departement de Biologic Animale, Madagascar (UADBA). FGZC refers to F. Glaw field numbers.

Terminology for biogeographic regions of Madagascar follows Glaw & Vences (2007).

Morphological measurements (in millimetres) were all taken by M. Vences with a digital caliper (precision

0.01 mm) to the nearest 0.1 mm. Used abbreviations are: SVL (snout-vent length), HW (greatest head

width), HL (head length), ED (horizontal eye diameter), END (eye-nostril distance), NSD (nostril-snout

tip distance), NND (nostril-nostril distance), TD (horizontal tympanum diameter), TE (tibia length),

HAE (hand length), HIE (hindlimb length), FE (foot length), FOTE (foot length including tarsus), FORE

(forelimb length), and RHE (relative hindlimb length). Terminology and description scheme follow

Vences et al. (2010).

Calls were recorded in the field using a Sony WM-D6C tape recorder and external microphones

(Sennheiser Me-80, Vivanco EM 238). Recordings were sampled at 22.05 kHz and 16-bit resolution and

computer-analysed using the software Adobe Audition. Frequency information was obtained through

Fast Fourier Transformation (FFT; width 1024 points). The spectrogram was obtained at Hanning

window function with 256 bands resolution. Temporal measurements are given as range, with mean ±

standard deviation in parentheses. Terminology in call description follows Vences et al. (2010).

Molecular genetic analyses were conducted by Wollenberg et al. (2008) and Vieites et al. (2009). See

those studies for exact methodology applied.

GLAW F. et al. , New Platypelis from Madagascar

Results

Class Amphibia Gray, 1825

Order Anura Fischer von Waldheim, 1813

Family Microhylidae Gunther, 1858

Subfamily Cophylinae Cope, 1889

GQxms Platypelis 1882

Platypelis ravus sp. nov.

Remark

This species has been referred to as Platypelis sp. (Marojejy) by Vences et al. (2006), as Platypelis

sp. afif. mavomavo 2 by Glaw & Vences (2007), as Platypelis sp. 2 by Wollenberg et al. (2008) and as

Platypelis sp. 4 by Vieites et al. (2009).

Etymology

The specific epithet is an adjective derived from Latin ravus = grayish-yellow, referring to the diagnostic

ventral colour of the species.

Holotype

ZSM 349/2005 (field number FGZC 2813), adult S, collected at Marojejy National Park, ‘Camp

Simpona’, 14°26.199'S 49°44.601'E, 1326 m above sea level, Antsiranana Province, northeastern

Madagascar, on 15 Feb. 2005 by F. Glaw, M. Vences, andR.D. Randrianiaina.

Paratypes

ZSM 350/2005 (field number FGZC 2823) and ZSM 351/2005 (FGZC 2824), two adult SS, with same

data as holotype, except that ZSM 351/2005 was collected on 16 Feb. 2005. Three additional paratypes,

all SS collected while calling, were deposited in the UADBA collection (still uncatalogued) and are

labelled with the field numbers FGZC 2822, 2825 and 2869. Their collection data are the same as in the

holotype except for FGZC 2869 (collected on 16 May 2005).

Diagnosis

The new species is assigned to the genus Platypelis based on enlarged terminal finger discs, absence

of prepollex, and molecular phylogenetic relationships (Wollenberg et al. 2008). It differs from other

arboreal cophyline microhylids of the genus Anodonthyla by the absence of a prepollex in males,

and from Plethodontohyla with enlarged finger and toe discs [P. guentheri Glaw & Vences, 2007, P.

inguinalis Boulenger, 1882, P. mihanika Vences, Raxworthy, Nussbaum & Glaw, 2003, P. notosticta

(Gunther, 1877)] by much smaller size (SVL 17-19 mm vs. 26-100 mm). Within the Platypelis /

Cophyla clade, the species is distinguished from Cophyla berara Vences, Andreone & Glaw, 2005,

C. phyllodactyla Boettger, 1880, Platypelis alticola (Guibe, 1974), P. cowanii Boulenger, 1882, P.

grandis (Boulenger, 1889), P. mavomavo Andreone, Fenolio & Walvoord, 2003, P. milloti Guibe, 1950,

P. pollicaris Boulenger, 1888, P. tsaratananaensis Guibe, 1974, andP. tuberifera (Methuen, 1920) by

distinctly smaller size (SVL of adult males 17-19 mm vs. 23-88 mm), and from P. tetra Andreone,

Fenolio & Walvoord, 2003 by presence of a regular pattern of moderately-sized tubercles on the back

(vs. four symmetrically arranged and distinctly enlarged white tubercles), a third toe of the same length

than the fifth (vs. toe three longer than toe five), by presence of yellow ventral colouration (vs. absence),

and by a much higher note repetition rate of the advertisement calls. It differs from Cophyla occultans

(Glaw & Vences, 1992) by smaller size (SVL 17-19 mm vs. 18-21 mm), and equal length of toe 3 and 5

(vs. toe 3 < 5). It is distinguished from Platypelis barbouri Noble, 1940 (and most likely from its junior

European Journal of Taxonomy 9: 1-9 (2012)

synonym Paracophyla tuberculata Millot & Guibe, 1951, described from the well sampled type locality

Andasibe) by absence of red colour on the ventral side of the hindlimbs and belly, and a largely smooth

dorsal surface (vs. strongly tuberculate).

Summarizing, the new species is most similar to P. barbouri (the only other Platypelis without vomerine

teeth and in addition with a rather similar advertisement call, but with reddish instead of yellow ventral

surface, and more strongly expressed and irregularly arranged tubercles on dorsal surface), and to P.

mavomavo (with yellow ventral surface but distinctly larger SVL). It differs, however, from these and

all other nominal species of Platypelis and Cophyla (except P. cowanii for which no genetic data is

available) by a high genetic divergence (see Wollenberg et al 2008; Vieites et al 2009 and unpublished

data for P. alticola, P tsaratananaensis, P tetra and Cophyla occultans). Phylogenetically, it appears to

be sister to P. milloti (see below) which however is larger, has a much more contrasting dorsal pattern,

red colour ventrally, and a much faster note repetition rate in advertisement calls (106/min vs. 18-19/

min).

Description of the holotype

Specimen in good state of preservation, some muscle tissue removed from right thigh, snout-vent length

19.1 mm. Body slender; head as wide as long, not wider than body; snout bluntly rounded in dorsal and

lateral views; nostrils directed dorsolaterally, not protuberant, nearer to tip of snout than to eye; canthus

rostralis indistinct, very slightly concave; loreal region plain; tympanum moderately distinct, 56% of

eye diameter; supratympanic fold moderately distinct, straight; tongue ovoid, not bifid or notched;

weakly expressed maxillary teeth present; vomerine teeth absent; choanae rounded. Forelimbs slender;

subarticular tubercles single, fiat, and hardly recognizable; outer metacarpal tubercle probably large and

fiat, but very difficult to recognize; inner metacarpal tubercle large, forming distinct protuberance at

base of first finger; hand with traces of webbing only between fingers 3 and 4; fingers distinctly fiattened

and relatively broad along entire length; relative length of fingers 1 < 2 < 4 < 3, fourth finger distinctly

longer than second; finger discs distinctly enlarged, slightly triangular; nuptial pads absent. Hindlimbs

slender; tibiotarsal articulation reaching tympanum when hindlimb adpressed along body; tibia length,

39% of SVL; lateral metatarsalia strongly connected; inner metatarsal tubercle small and fiat, difficult

to recognize; outer metatarsal tubercle absent; webbing between toes weakly developed, with traces of

web between second and third toe, and some webbing between third and fouth, and fourth and fifth toe;

webbing formula difficult to assess because subarticular tubercles on toes are hardly recognizable; toes

fiattened and relatively broad along their entire length; relative length of toes 1<2<3 = 5<4; third toe

of similar length as fifth. Dorsal skin smooth, without dorsolateral folds. Ventral skin slightly granular

on throat, smooth on chest and moderately granular on belly.

After five years in 70% ethanol, dorsum light brown with distinct and well-delimited symmetrical

dark brown markings: a heart-shaped marking starting between eyes and extending on posterior head,

bordered anteriorly by a light brown band between eyes; a W-shaped marking on anterior dorsum; and

a chevron-shaped marking on posterior dorsum. Shank and foot with one distinct dark crossband each,

another dark crossband on anterior hand. Ventrally, the throat is dark brown and this dark colour fades at

the level of the chest into a whitish colour on posterior belly. Limbs light with some dark brown pigment.

In life (Fig. 1), two series of regularly arranged and light-coloured small tubercles recognizable on the

dorsum, with five and three tubercles bordering the anterior edges of the W-shaped and chevron-shaped

dark markings, respectively. In life (Fig. 1 A), the colour was very similar to that in preservative but more

contrasted. The interorbital band was posteriorly beige. The iris was uniformly bronze. Throat and chest

were grayish. The yellow ventral colour extended onto the lateral part of the inguinal region. Also the

finger and toe discs (especially of first finger) were yellowish.

GLAW F. et al ., New Platypelis from Madagascar

Variation

The two ZSM paratypes are very similar in external morphology and colour pattern when compared

to the holotype (Table 1), but have a thin light middorsal line, and (in preservative) ventrally less dark

pigment, especially in ZSM 351/2005 which is mostly light coloured ventrally. The UADBA paratypes

were not available for detailed studies but were in general similar to the ZSM paratypes.

Fig. 1. Platypelis ravus sp. nov. in life. A. Dorsolateral view of S paratype (ZSM 350/2005) from

Marojejy National Park. B. Ventral view of same specimen (ZSM 350/2005). C. Dorsolateral view of S

holotype (ZSM 349/2005).

European Journal of Taxonomy 9: 1-9 (2012)

Table 1. Morphometric measurements (all in mm) ofholotype (HT) and two paratypes (PT) of Platypelis

ravus. For abbreviations of measured variables, see Materials and methods; further abbreviations used:

M (male); F (female). Tympanum diameter (TD) values in brackets indicate that the tympanum was not

distinct. In all specimens, third and fifth toe were of equal length, and the tibiotarsal articulation reaches

tympanum when hindlimbs are adpressed along body.

ZSM 349/2005

ZSM 350/2005

ZSM 351/2005

Field number

FGZC 2813

FGZC 2823

FGZC 2824

Status

Sex

SVL

19.1

17.9

17.2

5.8

5.4

5.6

5.5

1.4

1.1

1.2

2.5

2.4

END

1.5

1.4

1.5

NSD

1.1

1.4

1.2

NND

2.2

HAL

5.5

5.2

4.8

LORE

11.2

10.9

10.7

HIE

25.8

25.7

21.7

EOTL

11.8

11.4

10.4

7.4

7.1

6.3

7.5

7.8

7.1

^ 250 500 750 -iqOO

Time (milliseconds)

Fig. 2. Audiospectrogram and corresponding oscillogram of the advertisement call of the holotype of

Platypelis ravus sp. nov. (ZSM 349/2005), recorded at Camp Simpona, Marojejy National Park, on 15

February 2005 (19:20 h) at 21.1°C air temperature.

GLAW F. et al. , New Platypelis from Madagascar

Natural History

Numerous specimens of the new species were heard in the evening in dense primary forest directly next

to Camp Simpona campsite, calling from leaves in bushes and on trees at perch heights of 1-2 m above

the ground.

Advertisement calls

Calls of the holotype (ZSM 349/2005) were recorded at Camp Simpona, Marojejy National Park, on 15

February 2005 (19:20 h) at 21.1°C air temperature. They have been published by Vences et al (2006;

CDS, Track 64). The call is a single melodious note that is repeated at regular intervals. Note duration

ranges from 384-443 ms (418 ± 16; n = 11), duration of intervals between notes varies between 2504-

3200 ms (2797 ± 243; n = 10). Note repetition rate is approximately 18-19 notes/minute. The dominant

frequency ranges between 3990-4028 Hz (4010 ± 13; n = 11).

Molecular differentiation

DNA sequences of various mitochondrial genes determined from the holotype of Platypelis ravus were

included in a comprehensive molecular analysis of cophyline phylogeny, under the name Platypelis

sp. 2 (Wollenberg et al 2008), and are deposited in GenBank under accession numbers EU341101

(12S and 16S genes) and EU341035 (cytochrome b gene). According to this study, the species is

deeply genetically divergent from other Platypelis and was grouped by some analyses as sister species

of Platypelis milloti. According to Vieites et al (2009) the pairwise uncorrected sequence divergence

(p-distance) in a fragment of the 16S rRNA gene of P. ravus (as P. sp. 4) to all other cophylines was

above 6%.

Discussion

With the description of Platypelis ravus sp. nov. we add a distinctive new species to the genus Platypelis

which represents - together with P. tetra (see Andreone et al 2003) - the smallest species of the

Platypelis / Cophyla clade. Phylogenetic analyses (Wollenberg et al 2008) placed the new species

with some confidence sister to P. milloti Although at first glance these two species are very distinctive

in numerous characters, including size and advertisement calls, it is interesting that they bear some

similarities in the dorsal colour pattern. Platypelis milloti has a dorsal pattern of strongly contrasting

black blotches of almost the same shape as the poorly contrasting dark blotches in P. ravus sp. nov. (see

pictures in Glaw & Vences 2007). Also the light bar between the eyes and the vertebral line typical for

P. ravus sp. nov. are seen more strongly expressed in P. milloti. Furthermore, the general ventral pattern

is similar, except that the posterior venter is coloured red in P. milloti and yellow in P. ravus sp. nov. It

is therefore appealing to hypothesize that this similarity of colour pattern between the two species is due

to common ancestry.

Platypelis ravus sp. nov. is so far only known from its type locality. Own surveys in rainforests of other

mountain massifs in northern Madagascar at similar elevations (e.g. Montagne d’Ambre, Manongarivo,

Tsaratanana, Makira, at elevations of 900-2500 m) have so far failed to yield additional specimens

attributable to P. ravus sp. nov., and we therefore cannot exclude that this species is indeed endemic to

the Marojejy massif, as it seems to be the case for several other frog species occurring at comparable

elevation in this mountain massif (e.g. Gephyromantis tahotra Glaw, Kohler & Vences, 2011, G. tandroka

Glaw & Vences, 2001, G. schilfi Glaw & Vences, 2000). In another survey at Marojejy, Raselimanana

et al (2000) identified five species of the Platypelis / Cophyla clade {P. barbouri, P grandis, P

tsaratananaensis, P tuberifera and C. occultans) and we cannot exclude that their high altitude records

of C. occultans (recorded form 350-1300 m altitude) include P. ravus sp. nov., a similarly tiny species.

However, as we are aware of only six specimens from a single locality, we suggest to consider P. ravus

sp. nov. ‘Data Deficient’ according to the lUCN Red Eist criteria (lUCN 2001).

European Journal of Taxonomy 9: 1-9 (2012)

Acknowledgements

Fieldwork was carried out in the framework of cooperation accords among the Departement de Biologic

Animale, Universite d’Antananarivo, the Association Nationale pour la Gestion des Aires Protegees,

the Zoological Museum, University of Amsterdam, and the Zoologische Staatssammlung Mtinchen.

Permits for collection and export of specimens were kindly issued by the Ministere des Eaux et Forets

of Madagascar. We are grateful to Roger-Daniel Randrianiaina for his help in the field.

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Manuscript received on: 10 November 2011

Manuscript accepted on: 26 January 2012

Published on: 14 March 2012

Topic editor: Rudy Jocque

In compliance with Article 8.6 of the ICZN, printed versions of all papers are deposited in the libraries

of the institutes that are members of the EJT consortium: Museum national d’Histoire naturelle, Paris,

France; National Botanic Garden of Belgium, Meise, Belgium; Royal Museum for Central Africa,

Tervuren, Belgium; Natural History Museum, London, United Kingdom; Royal Belgian Institute of

Natural Sciences, Brussels, Belgium; Natural History Museum of Denmark, Copenhagen, Denmark.