Contributions to Zoology, 70 (4) 191-212 (2002)

SPB Academic Publishing bv, The Hague

Systematic revision of the enigmatic Malagasy broad-headed frogs

(Laurentomantis Dubois, 1980), and their phylogenetic position within the

endemic mantellid radiation of Madagascar

¹ ³ 4 4 Vences Frank Glaw Franco Andreone Riccardo Jesu & Giovanni Schimmenti Miguel ²,

national d’Histoire 'Museum naturelle, Lahoratoire des Reptiles et Amphibiens, 25 rue Cuvier, 75005 Paris,

France; address for correspondence: Zoologisches Forschungsinstitut und Museum Alexander Koenig,

2 Adenauerallee 160, 53113 Bonn, Germany; email: [email protected]; Zoologische Staatssammlung,

3 Miinchhausenstr. 21, 81247 Miinchen, Germany; e-mail: [email protected]; Museo Regionale

di Scienze Naturali, Sezione di Zoologia, Via G. Giolitti, 36, 10123, Torino, Italy;

4 email: [email protected]; Acquario di Genova, Area Porto Antico, Ponte Spinola, 16128 Genova, Italy.

Keywords: Amphibia: Mantellidae, Mantidactylus, subgenus Laurentomantis, Aglyptodactylus, Boophis,

Laliostoma, Mantidactylus, Mantella systematics, phylogeny, radiation, new species, tibial

glands, Madagascar.

Abstract Relationships ahd origin of mantellids 206

Acknowledgements 209

References 209 A revision ofspecies included in the subgenus Laurentomantis Appendix: Characters used for analysis 210 information about (genusMantidactylus) yieldednew phylogeny,

taxonomy, and biogeography of the endemic mantellid frog

radiation in Madagascar. Four Laurentomantis species, disting-

uished by morphology and advertisement calls, are recognized: Introduction

Mantidactylus (Laurentomantis ) horridus (Northern and North-

Western biogeographic regions), M. (L.) ventrimaculatus(South- Recent phylogenetic studies based on mitochon- East and East); M. (L.) malagasius (East); and the new species drial DNA sequences suggested that the endemic M. (L.) striatus (North-East). M. striatus and M. malagasius of the based bioacoustic and mor- Malagasy genera are probably sister species on frogs Aglyptodactylus,

phological affinities.A tibial gland, so far unknown in anurans, Boophis, Laliostoma (previously Tomopterna),

is described in M. and M. horridus. A phylogenetic malagasius Mantidactylus and Mantella form a monophyletic analysis of 54 mainly osteological and morphological characters lineage (Richards and Moore, 1998; Bossuyt and in 33 endemic Malagasy anuransresulted in a position of Lau- Milinkovitch, 2000; Richards et al., 2000; Vences rentomantis close to species ofthe subgeneraSpinomantis and et al., 2000), although they had been previously Gephyromantis (genus Mantidactylus), in accordance with its

also the well-established to three different subfamilies in the fam- subgeneric status. However, genus assigned

Based Mantellaresulted to be nested within Mantidactylus, supporting ily Ranidae(Blommers-Schldsser, 1993). on

need of of the latter. the generic partitioning the genetic evidence, Vences and Glaw (2001)

proposed including representatives of the five genera

in a separate family Mantellidae, with three sub- Contents families (Mantellinae, Boophinae, Laliostominae).

Molecular studies on Mantidactylus included Introduction 191 single representatives of eight subgenera (Richards Materials and methods 192 et al. 2000), but morphological phylogenies of this Systematic accounts 193

204 genus based on an adequate number of characters Key to species ofLaureniomantis

Phylogenetic analysis 205 and terminal taxa have so far not been published

206 Discussion (see Glaw et al., 1998). While Mantella, Aglypto- Phylogenetic relationships of Laureniomantis 206 dactylus and Laliostoma are well defined lineages Biogeography 206

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which with a limited number of species, Boophis and mantis, was considered to merit genus rank

and Blanc and Mantidactylus are speciose, with about 40 and 75 by Blommers-Schldsser (1991) Du-

nominal species, respectively. Especially Manti- bois (1992). Glaw and Vences (1994), however,

dactylus, currently partitioned into 12 subgenera concluded that no phylogenetic data exist to consider

Laurentomantis in addition to (Glaw and Vences, 1994), contains very diverse as a separate genus

the frogs both in size and morphology as well as in speciose Mantidactylus, and consequently con-

sidered Laurentomantisas a of Mantidac- habits and reproductive modes. Basic data on ecol- subgenus

and tylus. ogy reproductive biology are incomplete or

for of During the last numerous additional speci- totally lacking many species Mantidactylus years, of Laurentomantis collected and Boophis. To understand how the mantel!id mens were during sur- in several of In the radiation could give rise to its present extraordi- veys regions Madagascar. present

we review the Laurentomantismaterial avail- nary diversity in Madagascar, it is crucial to gather paper

able to us (more than 45 specimens), provide de- information on its less known lineages. tailed morphological and bioacoustic data, and Despite of the important recent progress in knowl- describe one new species. We furthermore under- edge on the batrachofauna of Madagascar (Glaw

take a of 33 and phylogenetic analysis species, repre- Vences, 2000), a number of groups remain all senting mantellid to assess the position largely unknown. Such is the case for the frog genera,

of Laurentomantis relative to them, and to draw species classified in Laurentomantis, which at pre- hypotheses on the origin and evolution of this ra- sent (Glaw and Vences, 1994) is considered as diation. subgenus of Mantidactylus : Mantidactylus (Lau- rentomantis) horridus (Boettgcr, 1880), M. (L.)

malagasius (Methuen and Hewitt, 1913) and M. Materials and methods (L.) ventrimaculatus (Angel, 1935). The monograph of Blommers-Schldsser and Blanc (1991) contained Vocalizations were recorded using portable tape recorders with no information on habitat, biology or life colora- external microphones and were analyzed either with the MEDA V tion of Laurentomantis Blommers- any species. sound analyzing system Spektro 3.2 (M. malagasius, M. striatus,

Schldsser and Blanc (1993) showed, for the first M. ventrimaculatus) or on a PC using the software Cool Edit (Syntrillium Software Corp.) (M. horridus). time, a photograph of a living M. malagasius. Glaw

The following morphological measurements were taken with and Vences (1994) provided photographs, call des- a calliper to the nearest 0.1 millimeter: SVL (snout-vent length), criptions and natural history notes for two differ- HW HL ED dia- (head width), (head length), (horizontal eye ent morphs referred to this from the Central species meter), END (eye-nostril distance), NSD (nostril-snout tip East and in the and Marojejy North-East, reported distance), NND (nostril-nostril distance), TD (horizontal tym-

diameter), HAL (hand length), FORE the discovery of an adult M. horridus at Montagne panum (forelimb length), 111L (hindlimb length), FOL (foot length), FOTL (foot length d’Ambre in northern Madagascar. including tarsus), IMTL, IMTH (length and height of inner The nomenclatural and taxonomic history of metatarsal tubercle), TLI (length of first toe). Laurentomantis is extensive and still confusing. Institutional abbreviations are as follows: BMNH (The Na-

While Boettger(I880) described the taxon horridus tural History Museum, London); FAZC (Franco Andreone Zoo-

as Hemimantis horrida, Boulenger (1882) treated logical collection; preliminary numeration ofspecimens which

will be deposited in MRSN); FMNH (Field Museum,Chicago); this species as Arthroleptis horridus. Methuenand MNHN (Museum National d’Histoire Naturelle, Paris); MRSN Hewitt erected the (1913) genus Microphryne to (Museo Regionale di Scienze Naturali, Torino); MSNG (Museo accomodate their taxon malagasius (as M. mala- di Storia Naturale di Genova); NMBE (Naturhistorisches Mu- and assumed that horrida to this gasia), belonged seum Bern); PBZT (Parc Botanique et Zoologique de Tsim-

as well. As the TM genus name Microphryne was pre- bazaza, Antananarivo); (Transvaal Museum, Pretoria);

UADBA de Biolo- occupied, Methuen (1920) created the replacement (Universite d’Antananarivo, Departement gie Animale); ZFMK (Zoologisches Forschungsinstitut und name Trachymantis. However, as demonstrated by Museum Alexander Koenig, Bonn); ZMA (Zoologisch Museum, Dubois also (1980), Trachymantis was preoccu- Amsterdam); ZSM (Zoologische Staatssammlung Miinchen). pied (by Trachymantis Giglio-Tos, 1917). Dubois Statistical analyses were carried out using SPSS for Windows, thus created the Laurento- (1980) replacement name version 9. We performed Mann-Whitney U-tests to assess sig-

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nificance of intersexual and interspecific differences in size inner single and two outer metacarpal tubercles; a and morphometric ratios (relative tympanum length and head distinct inner and a small outer metatarsal tubercle; width, ratios TD/SVL and I1W/SVL; relative size of inner single subarticular tubercles. Intercalary element metatarsal tubercle, IMTL/SVL and 1MTH/SVL).Temporal and between ultimate and metric measurements with standard present penultimate are given as range, mean ± phalanges ofall deviation in parentheses. Terminology for the description of fingers and toes. Ultimate phalanges Y-shaped.

femoral and tibial glands follows Glaw et al. (2000). Number Omosternum ossified and unforked. Vertebral col-

ofcrossbands on hand and foot are given to the tip ofthe longest umn diplasiocoelous. Maxillary teeth present, vome- finger or toe, respectively. The term “tibial gland” is here coined rine teeth present (vomerine odontophore for present), the prominent gland structures on the dorsal surface ofthe but generally concealed undermucous skin ofbuccal tibiae ofMantidactylus horridus and M.malagasius (see descrip- roof. A tions in the corresponding sections). The terms femur, tibia, complete circummarginal groove ventrally

humerus and in the radius, descriptive species accounts, are on finger and toe pads. Males with a slightly dis- used refer to the to corresponding external limb segments, and tensible single subgular vocal sac and distinct femo- not to the bones. of ral glands, type 2 according to Glaw et al. (2000) Coordinates and altitude of collecting localities are listed as assessed detailed in by macroscopic examinations, Vences and Glaw (submitted). Full names and geographic of to nine references ofthe campsites visited by F. Andreone and quoted composed up single large granules; some

in the text are as follows: = with (I) Masoala Campsite 2 Foret de species tibial glands; no nuptial pads.

Beanjada, 15°16.8’S, 49°59.8’E, 620 m, Masoala Peninsula,

Antalaha Fivondronana, Antsiranana (Diego Suarez) Faritany Included species. - Mantidactylus horridus, M. (Province); (2) Masoala Campsite 3 =Foret d‘Andasin‘iGover- M. malagasius, striatus sp. n., M. ventrimaculatus. nera, 15° 18.3‘S,50°01.2‘E, 700 m, Masoala Peninsula, Antalaha

Fivondronana, Antsiranana(Diego Suarez) Faritany (Province);

(3) llampy = Masoala Peninsula, Campsite 4 (Antsarahan’Amba-

rarato), 15°23.52’S, 510 Antalaha 50°02.82’E, m, Fivondronana, Mantidactylus (Laurentomantis) horridus (Boettger, Antsiranana Faritany (Diego Suarez Province); Tsararano (4) 1880) Campsite 1 = Tsararano Forest, Campsite 1 (Antsarahan’ny Fig. la-b Tsararano),AndapaFivondronana, Antsiranana Faritany (Diego

Suarez Province), 14°54.4’S, 49°41.2’E, 700 m. Material. - SMF Osteological data refer to the cleared and stained specimens 7177 (holotype, Nosy Be, collected by C. MSNG 49125A-C and listed in Table III.Terminology and character definitionslargely Ebenau); UADBA 10001 -10002 (Manari- koba Forest, Tsaratanana followthe accounts of Clarke(1981), Drewes (1984), Blommers- Massif, 14°02’24”S, 48°47‘02'’E;

R, G. and J. C. Schlosser (1993), Glaw and Vences (1994), and Glaw et al. Jesu, Schimmenti Piso, 15-22 February 1997);

ZFMK. 57433 d’Ambre; (1998). Characters were analyzed with PAUP*, version 4 beta (Montagne F, Glaw, N. Rabibisoa and Ramilison, (Swofford 1998). We performed Maximum parsimony (MP) O. 14-17 March 1994),

analyses (heuristic searches with TBR branch swapping), coding

all characters as unordered. Multistate characters were coded Original name. - Hemimantis horrida Boettger, as polymorphisms. Species ofHeterixalus (Hyperoliidae) were 1880 used as outgroup (see Blommers-Schlosser, 1993; Glaw et al.,

1998),

Identity. - The holotype is a subadult specimen of 16.4 it mm SVL; agrees with the other specimens here Systematic accounts assigned to the taxon in general morphology

and coloration (coarsely granular without distinct

elements Laurentomantis Dubois, 1980 ridge on the dorsum, with two indistinct

broad dark transversal bands), by having short

Status. - hindlimbs A subgenus ofMantidactylus Boulenger, (tibiotarsal articulation reaching ante-

1895 Glaw and Vences rior following (1994). eye corner), lacking an unpigmented area on the posterodorsal part of the femur (on which the

Diagnosis. - Small to medium sized 20-35 (SVL original pigmentation is still rather well preserved).

scansorial anurans with a to coarse- The mm) moderately area above the cloaca has a immediately very

ly granular dorsum and connected lat- short completely light marking which may be homologous to eral metatarsalia. absent from hand and Webbing the vertebral on stripe the posterior dorsum typi- feet. Third toe distinctly longer than fifth toe. A cal for the new species M. striatus (see below).

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- from otherknown Lau- with faint Diagnosis. Distinguished a very and poorly delimited dark pat- rentomantis by larger body size (male SVL 26-28 tern which forms 2-3 indistinct broad transversal

female mm vs. 20-25 mm; SVL 35 mm vs. 23-29 bands. Limbs with dark crossbands of variable

duration 2-4 mm), long note (1271-2521 ms vs. 407- width: on femur, 2-3 on tibia, 7-8 on tarsus

1468 and low in ms) pulse repetition rate advertise- and foot, 1-2 on humerus, 4-5 on radius and hand.

calls ment (13/s vs. 18-40/s). Further distinguished Ventral side uniformly diffuse greyish-brown with from M. ventrimaculatus absence small and by of prominent light grey (throat belly) or cream (limbs) dorsal ridges, from M. ventrimaculatus and M. markings. In life similar. The light ventral mark- striatus and The by a granular belly, from M. malagasius ings were rather indistinct. iris was yellowish

absence of red color on limbs. with a more intense circle by brown, narrow, orange

around the pupil.

- For and Morphology. measurements see Tables 1

II. The skin of the dorsum is coarsely granular; Distribution. - Known from (1) the type locality granules are only sometimes indistinctly arranged Nosy Be, (2) Montagne d’Ambre, and (3) Tsara-

the as continuous ridges on anterior back (e.g. tanana (altitude 1300 m) (Fig. 5). At Montagne

UADBA The vocal far the 10001). sac, as as recog- d’Ambre, species has also been recorded at an nizable in preserved specimens, is single subgular. altitude of 1200 m by Raxworthy and Nussbaum

the Throat and limbs are ventrally smooth, belly is (1994). The highest elevation of the island of Nosy

A all male Be is tibial is in 430 m in the Lokobe Altitudinal granular. gland present speci- reserve. range

Tsaratanana mens from (although less prominent is therefore 430 m (probably also lower altitudes

M. in than in malagasius), but absent the single at Nosy Be) to 1300 m. Blommers-Schlosser and female from Montagne d’Ambre. Femoral glands Blanc (1991) listed three further localities: Marojejy, in 49125C 3.6 1.9 in MSNG (size x mm; Fig. 2) Fenerive and Tampolo. While the latter two lo- internal view consist of five granules on one limb calities both refer to the specimen MNHN 1953.130

other limb is and six granules on the (granule diam- which here assigned to M. malagasius (see sec- eter 0.8-1.3 The tibial in this tion distribution of mm). gland speci- on that species), we did not

in internal has a similar find voucher for men, view, structure to any the locality Marojejy in the that found in femoral MNHN and glands of type 2 sensu Glaw ZMA collections on which the distri- et al. (2000). It consists of a densely packed field butional data of Blommers-Schlosser and Blanc of diameter 60 0.2-0.4 - ca. granules (granule mm). (1991) were largely based; the Marojejy locality about 60 distinct visible Externally, pores are on which possibly refers to M. striatus - is therefore the surface from the gland (absent surrounding skin), considered in need of confirmation. The specimen

indicating that each granule may have one sepa- shown as M. horridus in Hofrichter (1998) is ac-

M. rate secretion pore (Fig. 3). Size of the tibial gland tually a ventrimaculatus (NMBE 268/96, see is 8.0 x 2.2 mm. below).

No significant sexual dimorphism in relative

tympanum size and relative size of inner metatar- Natural history. - On the Tsaratanana Massif, M. sal tubercle detected female avail- was in the single horridus was collected within the primary rainforest able. Mean of male size was 76% of female size. Manarikoba, which was described by Perrier

de la Bathie (1927) and characterised by trees hea-

Coloration. - In covered preservative, dorsally greyish brown vily with mosses and herbaceous under-

Dorsolateral Fig. I. and ventral views of species in the subgenus Laurentomantis in life, a-b, Mantidactylus (Laurentomantis ) horridus, female ZFMK 57433 from c-d, Laurentomantis Montagne d’Ambre; Mantidactylus ( ) malagasius, male from Andasibe; e- f, Laurentomantis Mantidactylus ( ) striatus. male from Marojejy; g-h, Mantidactylus (Laurentomantis) ventrimaculatus, male from Vohiparara.

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HT, are

IMTH 0.9 0.7 0.9 0.8 0.7 0.9 0.9 0.7 0.8 0.8 0.4 0.5 0.8 0.4 0.7 0.7 0.7 0.8 subadult; hindlimbs - _ IMTL 1.9 1.4 1.5 1.6 1.3 1.8 1.4 1.4 1.3 1.2 1.0 1.1 1.0 1.0 1.0 0.9 1.4 1.1 - SA, the

TL1 2.5 2.7 2.6 2.6 2.7 2.5 2.3 2.2 2.4 2.4 2.6 2.7 2.4 2.1 2.2 1.7 2.2 1.8 when -

female; 1 1 RHLRHL 1 1 1 1 2 2 0 3 2 3 5 4 5 3 3 2 3 2 1 2 2 4 2

F,

1 7.3 articulation FOLFOL 12.5 11.7 11.3 12.3 12.2 15.5 10.7 13.8 11.3 11.4 12.0 10.5 11.11.1 10.8 11.7 11.4 11.1 11.1 10.4 10.6 12.2 12.4 male, tip. M, snout FOTL tibiotarsal FOIL 11.5 19.8 18.1 17.7 19.7 18.9 24.1 16.8 22.5 18.6 17.6 18.2 16.3 17.4 17.517.5 18.2 17.6 16.5 16.6 16.9 16.3 19.0 19.4

characters. the beyond HIL 26.3 42.6 41.4 40.8 43.0 41.8 52.6 38.7 47.0 40.840.8 40.3 40.6 37.6 38.1 38.338.3 40.1 38.5 36.3 38.338.3 39.7 35.9 42.0 42.5

by of (5) HAL 4.8 10.0 9.1 8.9 9.3 9.4 10.8 7.5 9.7 8.18.1 8.4 8.6 6.5 8.0 7.77.7 8.2 8.0 7.9 7.77.7 8,2 7.6 8,48.4 9.2 reached tip, snout abbreviations FORLFORL 11.6 20.5 19.5 18.9 20.4 20.1 23.7 17.2 22.022.0 18.2 18.2 18.4 13.913.9 16.6 16.916.9 17.5 16.6 16.2 16.716.7 16.2 16.3 17.6 19.0 position (4) for NNDNND 1.9 2.8 3.0 2.7 2.9 3,43.4 2.12.1 2.4 2.5 1.8 1.61.6 2.3 2.3 2.1 2.22.2 2.3 2.2 2.22.2 2.1 - the nostril,

(3) section gives NSDNSD 1.5 2.2 2.2 2.1 2.2 1.8 2.2 1.7 2.5 1.61.6 1.6 1.7 1.4 1.61.6 1.41.4 2.1 1.61.6 1.7 2.02.0 1.6 2.2 1.71.7 1.8

nostril, ENDEND 2.0 3.0 3.2 2.9 3.0 2.8 3.7 2.5 3.2 2.62.6 2.8 3.0 2.5 2.62.6 2.32.3 3.0 3.13.1 2.4 2.72.7 2.6 2.4 2.42.4 2.7 Methods length) and EDED 2.5 3.9 3.6 3.7 4.0 3.8 3.9 3.2 3.6 3.6 3.2 3.3 2.8 2.9 2.52.5 3.0 2.7 3.0 3.03.0 3.7 3.2 3.33.3 3,4 and hindlimb eye TDID 1.2 1.5 1.6 1.3 1.5 1.5 1.9 1.4 1.6 1.71.7 1.7 1.7 1.3 1.8 1.21.2 1.2 1.2 1.5 1.61.6 1.4 1.6 1.41.4 1.6 between Materials 1 7.4 9.3 1.8 9.7 9,49.4 8.6 8.78.7 8.68.6 9.9 9.89.8 9.0 9.49.4 9.6 9.2 9.09,0 9.8 HLHL 11.0 10.5 10.6 10.7 11.11.1 13.8 11.8 10.110.1 (relative 1 See (2) RHL comer, HWHW 6.8 10.6 10.7 10.3 10.4 10.6 13.9 9.0 12.5 10.310.3 9.2 9.6 8.3 8.78.7 8.4 9.9 9.29.2 9.0 9.4 9.4 8,68.6 9.4 9.3

specimens. eye SVL 16.4 27.1 27.3 26.0 28.1 26.5 35.4 22.0 29.1 25.025.0 22.7 23.0 20.2 22.222.2 21.6 25.6 24.224.2 22.5 24.9 25.8 22.2 24.0 25.7 paralectotype; anterior 2 2

(1) d’Ambred’Ambre Andranobe AndranobeAndranobe Laurentomantis PLT, Camp CampCamp centre, Be of saratanana saratanana Isaka-Ivondro Ambohitantely Locality Nosy Tsaratanana Tsaratanana Tsaratanana Tsaratanana Tsaratanana Montagne Isaka-Ivondro Andasibe Vohiparara Vohiparara Folohy AmbatovakyAmbatovaky AmbatovakyAmbatovaky Masoala, Masoala,Masoala, Masoala, Masoala, Ambohitantely Andasibe Ankeniheny lectotype; eye T T - mm) (0) (F?)(F?)-- - 17.2(F?)(F?) - 25.8 (in LT, - -- 18.9 -- Sex SA SA SA M M M M M F SA F M M M M M F F M M M M F body: F F F F F F paratype; the

A Measurements along (HT)(HI) (HT) FT, 125 49I25A 49125B 49125C 10001 10002 1935.172 1935.173 268/96 62273 1988.590 1976.250 233/96 5743457434 57435 49 57433 62281 7236 72547254 7700 77307730 57435 I. horridushorridus 71777177 ventrimaculatus malagasius 10076 83668366 (PLT)(PLT) (LT) Table holotype; adpressed ZFMK SMFSMF MSNGMSNG MSNG MSNG UADBA UADBA ZFMK MNHN MNHN NMBE ZFMK ZFMK BMNH ZMA MNHN NMBE ZFMK ZFMK M. M. ZFMK ZFMK M.M. TM ZMA FAZC FAZCFAZC FAZC FAZCFAZC

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IMTH IMTH 0.6 0.7 2.1 0.7 0.4 0.5 0.4 0.6 0.4 0.4 0.4 0.5 0.4 0.5 0.4 0.5 0.5 0.6 0.5 0.6 0.5 0.5 - - 0.5 0.5

IMTLIMTL 1.1 1.2 0.4 1.1 0.9 1.1 1.1 1.3 0.9 1.0 1.1 1.2 1.0 1.2 1.1 0.9 1.0 0.8 0.8 0.7 - - 0.6 0.9 0.9 0.9 1.2 1.0 1.2 1.1

TL1 2.1 1.8 1.2 2.6 2.9 2.4 2.3 2.2 2.7 2.5 2.4 2.3 2.6 2.4 2.8 2.9 2.9 3.0 2.4 2,42.4 2.5 2.1 - - 2.5

5 3 3 I RHL 3 3 2 4 1 2 1 2 2 2 2 3 3 2 4 2 5 2 5 4 5 5 5

FOL 11.8 11.4 11.2 11.5 11.8 11.4 11.0 10.8 10.7 11.0 11.2 10.9 11.8 11.3 10.6 12.0 12.3 11.3 12.3 12.3 11.7 12.2 11.9 11.4

FOIL 18.3 17.2 18.2 18.0 19.5 17.5 17.1 16.4 16.216.2 17.0 16.3 16.4 19.4 18.5 16.8 19.3 19.8 18.3 19.3 19.1 18.3 18.5 18.7 18.6

HIL 40,5 39.7 41.0 39.7 42.742.7 40.2 39.0 36.4 36.636.6 41,441.4 35.6 38.3 43.5 43.5 37.837,8 44.6 42.9 42.0 43.0 44.4 39.6 42.7 42.2 44.0

HAL 8.2 8.2 8.1 8.7 8.2 7.5 8.0 7.5 7,0 7.8 7.5 7.3 8.1 8.3 7.3 8.9 8.2 8.6 9.0 8.6 8.1 8.8 8.5 8.0

FORL 16.7 17.0 17.6 17.6 19.5 17.417.4 16.2 15.6 17.1 16.8 15.7 16.3 19.0 18.8 16.1 18.4 17.5 18.7 19.8 19.1 17.0 18.6 19.4 18.4

. NND 2.0. 2.1 2.1 1.8 2.2 2.0 2.0 1.81,8 2.0 1.91.9 2.2 1.9 1.9 2.0 2.1 2.0 2.1 2.2 2.1 2:12-1 1,8 2.0 2.1 2.2 2.1

NSD 1.51,5 1.5 1.7 1.0 1.8 1.8 1.7 1.3 1.51.5 1.8 1.6 1.6 1.8 1.61.6 1.2 2.1 1.3 1.4 2.0 1.7 1.2 1.2 1.5 1.4

END 2.3 2.7 2.4 2.5 2.5 2.9 2.5 2.5 2.4 2.5 2.6 2.6 2.5 2.8 2.6 2.3 2.8 2.8 3.1 2.8 2.3 2.4 2.6 2.7

ED 3.0 2.7 2.8 3.3 2.3 3.0 2.8 2.9 2.9 2.8 2.8 2.5 2.5 3.0 2.7 2.8 2.6 3.0 3.2 2.8 3.2 3.3 3.2 3.4

TD 1.2 1.3 1.7 1.4 1.5 1.2 1.3 1.4 1.2 1.3 1.3 1.3 1.3 1.4 1.4 1.3 2.0 1.3 1.5 1.3 1.3 1.2 1.5 1.3

8.2 9.0 9.0 9.2 9.1 9.4 9.1 8.7 8.8 9.6 8.6 8.3 9,6 8.9 8.8 9.29,2 8.8 8.9 9.0 9.4 HL 9.1 9.1 9.1 10.0 9.1 8.8 8.9 9.0 9.4

HW 8.2 8.7 8.6 8.3 8.0 9.1 8.6 8.7 8.3 8.3 8.3 8.4 7.2 8.0 7.5 8.0 8.2 7.9 8.6 7.8 7.6 8.0 8.3 8.7

SVL 22.0 23.0 23.2 23.8 25.3 25.6 23.9 22.7 22.7 24.3 22.2 22.2 25.1 25.4 22.5 24.9 25.0 23.4 26.9 23.8 22.8 23.4 23.6 24.8

2 3 3 3 3 3 3 3 Camp Camp Camp Camp Camp Camp Camp Camp Masoala Masoala Masoala Masoala Masoala Locality Andasibe Andasibe Vohidrazana Marojejy Masoala, Masoala, Masoala, Masoala, Masoala, Masoala, Masoala, Masoala, Ilampy, Ilampy, Ilampy, Ilampy, Ilampy,Ilampy, Tsararanosararano Tsararano Tsararanosararano Marojejy Marojejy Marojejy Marojejy - - T T Sex M M F M F F F M M F M M M M M M F F F F F M F F M F M M M M F

(HI) (PT) (PT) (PT) (PT) (PT) (PT) (PT) (PT) (PT) (PT) (PT) (PT) (PT)(PT) Continued. (PT) (PT) (PT) (PT) (PT) (PT) (PT) 8 59876 60039 A1937.1 A1937.2 59876 324/2000 938/2000 7645 7711 7796 7840 7858 78597859,790s’7905 7910 10299 10313 10378 10424 1044110441 A1937.1 A193 57437 57438 59930 59931 I. striatus ZFMK ZFMK ZFMK MRSNA1937.2 MRSNA1938 Table ZSM M. ZSM FAZC FAZC FAZC FAZC FAZC FAZC FAZC FAZC FAZC FAZC FAZC FAZC FAZCFAZC MRSN MRSN MRSN ZFMK ZFMKZFMK ZFMK ZFMK

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II. characters of Table Differential Laurentomantis species. Morphometric ratios were calculated from data in Table I.

TT is used as abbreviation for libiotarsal articulation.

M. horridushorridus M. ventrimaculatus M. malagasius M. striatus

Male SVL 26.0-28.1 mm 23.0-25.0 mm 20.2-24.0 mm 22.2-23.8 mm

Female SVL 35,4 mm 29.1mm29.1 mm 23.2-25.7 mm 23.9-26.9 mm

Head width

(ratio HW/HL) 0.97±0.03 (0.92-1.02) 1,00±0.051.00±0.05 (0.95-1.06)(0.95-1.06) 0.98±0.03 (0.94-1.04) 0.90±0.06 (0.79-1.01)

Tympanum size 0.057±0.008 0.067±0.008 0.0610.061±0.0I0±0.010 0.057±0.007

(ratio TD/SVL) (0,050-0.073)(0,050-0,073) (0.050-0.075) (0.047-0.080) (0.047-0.080)

Hindlimb short length (TT(TT reaching at most long (TT reaching up to mostly long (IT(TT reaching mostly long (TT

nostril) beyond snout tip) sometimes beyond snoutsnout reaching sometimes

tip) beyond snout tip)

Tibial in absent absent in males andand in males andand absent inin males gland present males, present and

in the singlesingle known female females femalesfemales of some popula- females

tions, absent in males

and females ofof other

populations

Granules in femoral

glandsglands (per(per femur) 5-6 9 1-4 3-6

Dorsal skin texture strongly granular, but strongly granular, mostly moderately toto strongly slightly to moderately

without ridge elements on with ridge elements on granular, without or with granular, without or with

anterioranterior dorsum anterior dorsum only weak ridge elementselements only weak ridge ele-

on anterior dorsum ments on anterior dorsum

Ventral skin texture granular smooth slightly granular largely smooth

Red color on hindlimbs absent absent present absent

VentralVentral colorcolor in life brown/blue dark dark/grey marbling marbling brown/light marbling uniformly grey-

(with small white spots) brown with fewfew lighter

markings

VertebralVertebral absentabsent or indistinct absent absent and distinct stripe or very present Note length in

advertisement calls 1271-2521 ms 407-455 ms 768-1468 ms 440-1266 ms

PulsePulse repetition raterate inin

advertisement calls 13/s 21-24/s 18-36/s 29-40/s

forest is growth. The physionomy of this unspoilt of intervals between pulses was 52-64 ms (58± also marked the of of 4 Pulse by presence high numbers ms, n=15). repetition rate was 12.6-13.3

ferns and Pandanus. second. Pulse increased tree ( Cyathea) During the study (12.9±0.3; n=7) per intensity

period (15-22 February 1997), which occurred in at the beginning of one note, and decreased again

2300-4300 the peak of the rainy season, only calling males towards its end. Frequency was Hz,

were observed; they were spotted at night on bushes dominant frequency 2300-3300 Hz.

at 50-150 cm from the ground in the vicinity of

a mountain stream. The female from Montagne

in Laurentomantis d’Ambre was found hidden under deadwood Mantidactylus ( ) ventrimaculatus

primary forest (Glaw and Vences, 1994). (Angel, 1935)

Fig. lg-h

- Calls. Recorded on the Tsaratanana Massif on

Material. - MNHN 1935.173 and MNHN 1935,172 17 February 1997, at 17.5°C air temperature. Calls (lectotype) (paralectotype), both from Isaka-Ivondro, collected by R. Catala; were series of long notes with widely spaced pulses ZFMK 62273 and 62281 (Vohiparara;F. Glaw, D. Rakotomalala (Fig. 4a). Note duration was 1271 -2521 ms (2116± and F. Ranaivojaona, 27 February 1996); NMBE 268/96 (Anda- 440 ms, n=7), duration of intervals between notes sibe-Analamazaotra; D. Vallan, 25 January 1996).

was 1271-3113 ms (1846±589 ms, n=7). Each note

consisted of 16-33 (27.3±5.9; n=7) pulses. Pulse

duration was 11-17 ms (I4±2 ms, n=15), duration

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Original name. - Trachymantis malagasia var. ven-

trimaculatusi Angel, 1935

Identity. - The morphological and chromatic char-

acters of the types (light marbling on venter; rela-

tively long hindlimbs; dorsal ridges) leave little

attribution doubts on the correct of the Vohiparara

and Andasibe specimens to M. ventrimaculatus.

Diagnosis. - Distinguished from other known Lau- Fig. 2. Femoral gland of.Mantidactylus (Laurentomantis) ventri- rentomantis by short note duration in advertise- maculatus (NMBE 268/96) in ventral (external) view. Arrows ment calls ms 440-2521 Further (407-455 vs. ms). indicate the external central which apparent depressions may from M. smaller distinguished horridus by body be used by the surrounding gland granules for secretion.

size (male SVL 23-25 mm vs. 26-28 mm), smooth

belly, and higher pulse repetition rate in advertise-

ment calls (21-24/s vs. 13/s); and from M. striatus

and M. malagasius by dark belly with distinct light

marblings (which are bluish in life).

- For Morphology. measurements see Tables I and

II. The skin of the dorsum is strongly granular.

Tubercles on the posterior head and anterior dor-

sum fuse to form a symmetrical pattern of ridges.

Theseridges are prominent in most specimens, rigid

and of tibial is absent a sharp appearance. A gland

in NMBE and ZFMK material, both in the males

and in the subadult (female) specimens. Venterand

throat are smooth, very slightly granular areas are

the present on belly close to the inguinal region.

Femoral glands are very distinct and prominent;

in ZFMK 62281, the ovoid gland consists of 9 large Fig. 3. Tibial gland of.Mantidactylus (Laurentomantis) horridus granules which in internal view are regularly packed (MSNG 49125C) in external view (above) and internal view but in external view to enclose two median Note the number of small secretion in appear (below). large pores

external view. Scale bars = 1 depressions. This external configuration is even mm.

better visible in NMBE 268/96 (Fig. 5), in which

the 5.6 3.0 7 1-2 gland measures x mm. Diameter ofsingle on tarsus and foot, on humerus, 6 on radius

granules is 0.9 mm. and hand. Ventral side dark brown, distinctly

No in marbled with In significant sexual dimorphism relative grey. life, the dorsum was reddish

size and relative size of inner metatar- brown and the ventral tympanum light marbling was light blue

sal was Mean male was 82% on a tubercle detected. size deep black venter and throat. The iris was

of mean female size (only adult specimens con- greyish brown, with a narrow vertical black streak

in its lower The crossbands sidered). part. on the flanks were deep black.

Coloration. - In preservative, dorsally greyish brown

with two indistinct, faintly recognizable dark cross- Distribution. - Known from (1) the type locality bands which are largely discontinuous dorsally but Isaka-Ivondro (at an altitude of 700 m according rather distinct the flanks. Limbs with on distinct to original description), (2) Vohiparara, and (3) and crossbands: 2-3 on 3 on Andasibe Altitudinal regular femur, tibia, (Fig. 3). range 700-1000 m.

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- Laurentomantis Natural history. A single calling male was heard Mantidactylus ( ) malagasius (Me-

in forest thuen and from near the ground at night at Vohiparara, Hewitt, 1913) at a distance of more than 10 m from a brook. Fig. Ic-d

- - Material, BMNH 1988.590 400 C. J. Rax- Calls. Recorded at Vohiparara on 27 February (Ambatovaky, m, worthy 13 March 1990); FAZC 7236 and 7254 (Andranobe, 1996. Calls were series of unharmonious notes (Fig. Masoala; J, E, Randrianirina, 28 October- 6 November 1998); ± 4d). Note duration was 407-455 ms (442 19 ms, FAZC 7700 and 7730 (Masoala, Campsite 2; F. Andreone and n = 7), duration of intervals between notes was J. E. Randrianirina, 30 November 1998); MNHN 1976.250

±1281 = 2673-6274 ms (3452 ms, n 7). Each note (locality unknown); MNHN 1953.130 (Tampolo Est); NMBE

consisted of 9-11 (9.9±0.7, n = 7) pulses. Pulse 233/96 (Ambohitantely; D. Vallan); TM 10076 (holotype; Fo-

collected M. Herschell-Chauvin, in duration 5-19 ± 4 = dura- lohy; by apparently 1911); was ms (13 ms, n 75), ZFMK 57434 and 59876 (Andasibe; F, Glaw and M. Vences, tion of intervals between pulses was 28-48 ms (36 1-4 January 1994); ZFMK 57435 (Ankeniheny; F. Glaw, N.

= ± 4 ms, n 66). Pulse repetition rate was 21-24 Rabibisoa and O. Ramilison, 19 February 1994); ZFMK 60039

± n = second. (22 1, 7) pulses per Frequency ranged (Andasibe; F. Glaw, 1 February 1995); ZMA 8366 (Ambatovaky; between 2000-5750 Hz, the poorly defined domi- C. J. Raxworthy, March 1990); ZSM 324/2000 (Vohidrazana,

F. Glaw, 10 April 2000). nant frequency between 2300-4250 Hz.

Original name. - Microphryne malagasia Methuen

and Hewitt, 1913

Fig. 4. Sonagrams and oscillograms of calls of species in the subgenus Laurentomantis. a, Mantidactylus horridus, recorded at

at Tsaratanana; b, Mantidactylus malagasius, recorded at Andasibe; c, Mantidactylus striatus, recorded Marojejy; d, Mantidactylus

ventrimaculatus, recorded at Vohiparara.

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of (composed 1-2 granules) indicate that the name

malagasius is correctly applied to the red-legged

Laurentomantis species from eastern Madagascar.

Diagnosis. - Distinguished from other known Lau-

rentomantis by red color on posterior and ventral

surface of limbs in (appearing pigmentless preser-

vative vs. pigmented brown-greyish in the other

species). Further distinguished from M. horridus

shorter by note duration (768-1468 ms vs. 1271-

2521 ms) and higher pulse repetition rate in ad-

vertisement calls (18-36/s vs. 13/s), and smaller

size body (male SVL 20-24 vs. 26-28 mm; female

SVL 23-26 mm vs. 35 mm); from M. ventrima-

culatus by absence ofregular light marbling (bluish

in life) on a dark venter, and longer note duration

in advertisement calls (768-1468 ms vs. 407-455

For ms). a distinction from M. striatus, see diag-

nosis of that species below.

Remark. - The specimen MNHN 1976.250differs

in and femoral general appearance gland morp-

hology from the remaining specimens; it is only

tentatively assigned to M. malagasius, and not con-

sidered in the morphometric calculations.

5. Distribution of in the Laurento- Fig. map species subgenus

- For measurements see Tables I and mantis. of localities in is in Morphology. Positioning the maps part only II. The dorsum is the approximate. Numbers of localities correspond to those given moderately granular, gran- in the sections of each “Distribution” species. ules only occasionally form ridge-like structures

which, however, are never very prominent. The

- The male is characterized Identity. holotype by throat is smooth, the belly slightly to moderately a dorsal with coarsely granular skin, two ridge- granular. Very distinct and prominent tibial glands

in like granules the shoulder region and two larger are present in the males from Andasibe and Ambo- rounded the anterior throat well granules on dorsum; hitantely as as in the female from Ankeniheny and chest are smooth, venter and ventral surface (ZFMK 57435). They are also present in the dubi- of femur are slightly granular. Femoral glands are ous specimen MNF1N 1976.250, but absent in the

The the left femur consists of male prominent. gland on holotype and in specimens from Ambatovaky

1.8 and one single diameter ca. Andranobe This enlarged granule, mm; (Masoala). may indicate a the on the femur consists of gland right two gran- constant difference between mid-altitude and low ules (see also Methuen and Hewitt, 1920). There altitude but localities, more material is necessary are no tibial Color about 90 in to assess whether it bear glands. (after years may taxonomic relevance. preservative) has faded into an almost uni- In ZFMK largely 57434, the tibial gland measures 7.1 x form brown. light According to the original de- 1.4 mm and consists (in internal view) of about 75

“hidden surface of the and tibiae scription, thighs granules (0.2-0.4 mm in diameter). The external with white blotches” which almost large certainly pores of the gland are clearly visible.

to the areas in correspond unpigmented (red life) Femoral glands are only present in males; on typical for this the species. Furthermore, general each femur, the gland generally consists of two morphology, size, and structure of the femoral gland large granules (a single granule in the holotype);

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in BMNH 1988.590 from Ambatovaky, the gland (1991). Despite its short hindlimbs(tibiotarsal ar-

each femur consists of of ticulation anterior consider on two groups two gran- reaching eye corner) we ules each (total number of granules on both fe- it as M. malagasius because on the posterodorsal

Femoral size in NMBE 233/96 of the murs: eight). gland part femur, unpigmented areas are recog-

ofeach nizable the red life is 3.1 x 1.4 mm in external view (diameter (which probably correspond to

in internal 1.2 granule view ca. mm). color typical for this species). The dorsum of the

No significant sexual dimorphism in relative specimen is coarsely granular without ridge ele- tympanum size and relative size of inner metatar- ments. According to the MNHN catalogue, two sal tubercle was detected. Meanmale size was 92% specimens were originally granted the number of size. MNHN the information mean female 1953.130; locality reads

“Fenerive, Tampolo Est”. Blommers-Schldsserand

Coloration. - Blanc this In preservative, dorsally greyish (1991) apparently interpreted as sepa- brown, with indistinct dark pattern. Except short rate informations for the two specimens, and cor- brown crossbands, the femur is pigmentless. There respondingly plotted two localities on their

4-5 for are 3-5 crossbands on femur, on tibia, 8-10 on distribution map, one Fenerive, a second one tarsus and foot, 6-7 on radius and hand; no distinct for Tampolo. There are at least two localities named

of bands are seen on humerus. Ventrally grey-brown Tampolo along the eastern coast Madagascar, on throat, chest and anterior belly, pigmentless- one close to Mananara and north of Fenoarivo ( =

A small cream on posterior belly. pattern of white Fenerive), a second one south of Toamasina(=Ta- spots, partly forming aggregations, is present on matave). As specimens from different localities the On of these dark ventral areas. the belly, each generally were not catalogued under the same

spots coincides with a single granule. Along the number in the MNHN, we interpret that “Fenerive”

form that Est” refers lower lip, the light spots sometimes crossbands was added to indicate “Tampolo to (e.g. ZFMK 59876). Ventral side of forelimbs brown the locality north of Fenoarivo.

with light spots. Femur ventrally pigmentless cream, tibia cream with brown markings. In life, the pos- Natural history. - Calling males were observed at

in in terior and ventral surfaces of the femur are deep night forest the vegetation, 5-50 cm above

is ofthe the concentrated red, as part inguinal region and of a small ground, not around waterbodies. area around the forelimb insertion. The femurs and The vocal sac was single subgular and only slightly posterior belly are more or less intensively shaded distensible.

in red. The iris is orange brown its upper part,

in its lower Calls. - 23.5°C greyish part. The dorsal surface can Recorded at Ankeniheny at air

have an olive-greenish shade. temperature and at Andasibe. Series of unharmo-

nious notes (Fig. 4b). Note duration was 936-1468

= Distribution. - The species is known from (1) the ms (II69 ± 205 ms, n 6) at Andasibe and 768-

= type locality Folohy, (2) Andranobe(Masoala), (B) 995 ms (864 ± 72 ms, n 12) at Ankeniheny.

Masoala (other localities), (4) Ambatovaky, (5) Duration of intervals between notes was 769-1049

± = Fenoarivo, (6) Andasibe, (7) Ankeniheny, (8) ms (876 108 ms, n 5) at Andasibe, and 1126-

The 1698 ms ± 200 = Vohidrazana, (9) Ambohitantely (Fig. 3). (1433 ms, n 11) at Ankeniheny. known altitudinal range is 300-900 m. Each note consisted of 22-43 (35 ± 9, n = 6) re-

The specimens BMNH 1928.5.9.13(Brickaville; spectively 15-20 (17 ± 2, n = 12) pulses. Duration purch. Rosenberg) and BMNH 1925.7.2.92(Antsi- of intervals between pulses at the two localities hanaka; purch Rosenberg) were not available at was 30-80 ms. Pulse repetition rate was 23-36 (30

the time of and be confirmed = our study cannot as ± 5, n 6) respectively 18-22 (20 ±11, n=13)

M. until examined. The from second. malagasius specimen pulses per Frequency was 2500-4500 re- is subadult Fenerive (MNHN 1953.130) (SVL 15.2 spectively 2650-4400 Hz. of femoral and in mm; traces glands recognizable) mediocre state of preservation. It was considered

and as M. horridus by Blommers-Schlosser Blanc

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Laurentomantis striatus Mantidactylus ( ) n. sp. species is found in head width (U-test ofratio HW/

Fig. le-f SVL, PO.OOl; P<0.005 considering males only),

in this although some range overlap character ex-

Material. -1 teletype: ZSM 938/2000 ZFMK (formerly 57436), ists. On the Masoala peninsula, the two species adult male, collected by F. Glaw, N. Rabibisoa and O. Ramilison and come into close contact even syntopy: while on 27-31 March 1994 at the Marojej'y Massif, Campsite 1 (ca, only M. striatus was collected at Ilampy and only 300 m altitude). Paratypes: FAZC(MRSN) 10299,10313,10378, M. malagasius at Andranobe, at Campsite 2 both 10424, 10441,collected by F. Andreone and J, E. Randrianirina

at Ilampy (Masoala), in November and December 1999. MRSN species were captured.

A1937.I-2 and A1938, collected by F. Andreone at Tsararano

Forest (Campsite 1) on 3-4 December 1996. FAZC 7581 and Description of holotype. - Adult male, SVL 23.8 7645, collected by F. Andreone and J. E. Randrianirina at mm. For measurements, see Table I. Body slen- Masoala (Campsite 2) on 28 November 1998;FAZC 7711,7796, der; headlonger than wide, slightly wider thanbody; 7840, 7858, 7859, 7905, 7910, collected by F. Andreone and J. rounded in dorsal E. Randrianirina snout and lateral views; nostrils at Masoala (Campsite 3) on 2-9 December

1998. ZFMK57437-57438,same collectors and collecting date directed laterally, slightly protuberant, much nearer

ZFMK as holotype; 59897 (cleared and stained specimen), to of tip snout than to eye; canthus rostralis indis- collected by F. Glaw and O. Ramilison at the Marojejy Massif, loreal tinct, concave; region concave; tympanum Campsite 1, on 21-23 February 1995. ZFMK 59929 (cleared small, distinct on left side of head, indistinct on and stained specimen), collected by F. Glaw and O. Ramilison

right side, rounded, 42% of eye diameter; supra- at the Marojejy Massif (Campsite 3) on 25-26 February 1995; fold indistinct ZFMK 59930-59931, collected by F. Glaw and O. Ramilison tympanic very and rather irregular,

at the Massif 1) on 22 1995. Marojejy (Campsite February not clearly curved; tongue ovoid, distinctly bifid

posteriorly; vomerine teeth not visible on the buc-

Diagnosis. - A member of the Mantidactylus but genus cal roof, present under the mucous skin, choa-

based on the absence of and nuptial pads presence nae small, rounded. Arms slender, subarticular

of femoral in males. to the sub- glands Assigned tubercles single; two outer, and one inner metac-

genus Laurentomantis based on the moderately, arpal tubercles present; fingers without webbing; dorsum, the irregularly granular single subgular relative length of fingers 1<2<4<3, finger 2 dis- vocal absence of foot and sac, webbing, completely tinctly shorter than finger 4, only slightly shorter connected lateral metatarsalia. from Distinguished than finger 1; finger disks distinctly enlarged; nuptial all other Laurentomantis of by presence a light pads absent. Hindlimbs slender; tibiotarsal articu-

brown vertebral the to orange stripe on posterior lation reaches between and lateral eye nostril; meta- dorsum. Further distinguished from M. horridus tarsalia connected; inner metatarsal tubercle distinct,

by smaller size SVL 22-24 mm vs. 26- body (male outer metatarsal tubercle present; only traces of

28 female SVL 24-27 mm vs. 35 lower mm; mm), webbing between toes; relative toe length 1<2<5<

note duration (440-1266 ms vs. 1271-2521 and 5. ms) 3<4, toe 3 distinctly longer than toe Skin on the

higher repetition rate (29-40/s vs. 13/s) in pulse upper surface coarsely granular; granules are ar- advertisement calls; from M. ventrimaculatus by ranged asymmetrically except for the anteriorback

absence of regular light (blueish in and head where marbling life) regions, some symmetrical larger

on the venter, and by a longer note duration in tubercles and short ridges are present; a number advertisement calls 55 (440-1266 ms vs. 407- ms). of also above the granules eyes; no distinct en- Most similar (and probably closely related) to larged tubercles in the cloacal region; ventral skin M. from which it differs absence of malagasius by smooth. Femoral distinct glands very and promi-

reddish color on hindlimbs (versus red color on nent. On the right femur, in internal view (after

posterior and ventral surface of femur in life), largely reflection of skin), four large granules are visible, dark and smooth ventral surface (ver- uniformly three of which are arranged symmetrically, appar- sus light surface with dark markings, and granular ently encircling a weakly marked external central texture with white-stippled granules), and presence depression. Tibial glands absent. The folded vocal

(vs. absence) of a light vertebral stripe on the pos- sac marks two distinct longitudinal lateral folds terior dorsum. A further difference between both on the posterior throat.

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in colora- After six years preservative, dorsal

- from the tion is uniformly greyish brown, with some darker Distribution. Known (1) Marojejy Massif, coloration the dorsal tubercles and Masoala bordering gran- (2) Tsararano, (3) (Campsite 2), (4) Ilampy

Known altitudinal 300-700 ules. The limbs are light brown with distinct dark (Fig. 3). range m. brown crossbands (3 on femur, 3 on tibia, 6-7 on tarsus and foot, 1-2 on humerus, 6-7 on radius and Natural history. - At Marojejy, calling males were hand). A short (3.5 mm) beige vertebral stripe starts observed at night in the vegetation 5-50 cm around

the All this on the posterior back and ends at the cloaca. Lower ground. specimens at locality were found

of lip with indistinct dark and light crossbands. Ven- along small brooks, at a maximum distance 5 m

brown, with from the water. FAZC 10313 contained two trally uniformly greyish a few small very

4 and light spots on chest. large unpigmented oocytes (diameter mm)

two smaller unpigmented oocytes (diameter 3 mm).

Variation. - The vertebral stripe on the posterior dorsum short in the Calls. - Recorded at Series of unhar- can be very as holotype, or Marojejy.

monious notes Note duration was 440- run along the whole posterior halfof the dorsum (Fig. 4c).

1266 Durationof intervals (e.g. ZFMK 59931, stripe length 12.5 mm). It is ms (823±324 ms, n=14).

in between notes was 162-432 ms (238±72 absent only a single specimen (MRSN A1937.2) ms, n=13).

Numberof notewas 15-39 examined. Limb crossbands are 2-4 on femur, 3-4 pulses per (28±9, n=14).

Intervals between lasted 21-23 ms on tibia, 6-8 on tarsus and foot, 1-2 on humerus, pulses (n=5).

Pulse repetition rate was 29-40 n= pulses 7-8 radius and hand. Indistinct brown patches are (35±4, 14)

note. was 3000-4500 Hz. generally present behind the forelimb insertion and per Frequency Excep- 2000 in the inguinal region. ZFMK 59930 is exception- tionally, a very long note type (duration ca.

colored but in The ms, consisting of 75 pulses) was heard. ally light agrees pattern. ven- tral ground color is always a diffuse and uniform greyish brown. In the female ZFMK 59931, the Key to species of Laurentomantis (see also table throat is of a darker brown shade than the venter. II) A number ofwhite markings can be present on the chest and on the throat along the lower lip. The male SVL 26-28 female SVL to (la) Larger species; mm, up hindlimbs diffuse are ventrally uniformly grey- 35 mm. Hindlimb mostly short; when adpressed along body, brown. Dorsum the coarsely granular, granules tibiotarsal articulation reaches at most nostril. Pulse repetition generally not forming ridges (the holotype show- rate in notes of advertisement calls low (13/s), note duration

long (1.2-2.5 s) horridus ing an exceptional state). The belly skin is smooth Mantidactylus (Laurentomantis)

to (lb) Smaller species; male SVL 25 mm, remale SVL in all specimens. The femoral glands measure be- up up to 29 mm. Hindlimb mostly long; when adpressed along body, tween 3.8 x 1.8 mm (FAZC 10378) and 4.8 x 2.1 tibio tarsal articulation reaches at least nostril, often snout tip mm (MRSN A1938). They consist of 3 (FAZC or beyond. Pulse repetition rate in notes of advertisement calls

to 5-6 7905) granules per gland (MRSN A1938). higher (18-40/s), note duration mostly shorter (0.4-1.5 s)

Granule diameter is 0.9-1.2 mm. 2

Color in life reddish brown dark with blue No significant sexual dimorphism in relative (2a) dorsally, marbling ventrally. Notes of advertisement calls short (0.4-0.5 s) but the inner tympanum size was detected, meta- Mantidactylus (Laurenlomanlis) ventrimaculatus tarsal tubercle was significantly higher in males (2b) Color in life without reddish brown tone dorsally, and (U-test, and hadalso a P<0.005), possibly tendency without blue marbling ventrally. Notes of advertisement calls

in Mean of being larger males (U-test, P=0.09). generally long (0.4-2.5 s) 3

color. male size was 92% of mean female size. (3a) Ventral surface of hindlimbs in life with bright red

No vertebral stripe. Ventrally with irregular dark/light marbling. In life, the color was rather similar to that in Ventral skin slightly granular, granules marked with small while The iris reddish in its preservative. was upperpart, Head about wide spots. generally as as long brown in its lower part. The vertebral stripe was Mantidactylus ( Laurenlomantis ) malagasius The ventral side was dark to generally orange. light (3b) Ventral surface of hindlimbs without red color. Posterior without of red reddish color. grey, any trace or back with beige-orange vertebral stripe. Ventrally uniformly

Downloaded from Brill.com10/09/2021 09:33:27PM via free access Contributions to Zoology, 70 (4) - 2002 205

Fig. 6. Maximum parsimony cladogram of a phylogenetic analysis of 34 Malagasy anuran species based on the 54 characters listed

in Appendix 1, The tree is a strict consensus of 12 equally most parsimonious trees; 324 steps, consistency index 0.364. Species of

values in below Heterixalus were used as outgroup. Numbers are bootstrap percent (2000 replications; values 50% not shown).

dark sometimes with few Ventral skin grey, light markings. The maximum parsimony analysis (Fig. 6) failed

smooth. Head generally slightly longer than wide to place Boophis as monophylum: all Boophis striatus Mantidactylus (Laurentomantis) sp, n. species were paraphyletically arranged along the

lineage leading to Mantidactylus and Mantella.

Mantidactylus and Mantella together formed a Phylogenetic analysis monophyletic group, the two Laurentomantis in-

cluded were sister taxa nested within Mantidactylus. We analysed a total of 54 osteological, morpho- A cluster of three species of the subgenus Gephyro- logical, etho-ecological and karyological charac- mantis were the sister of Laurentomantis. in group ters as listed Appendix 1. Osteological data are Mantella was also nested within Mantidactylus, with based on cleared and stained specimens listed in a species of the subgenus Guibemantis (M. liber) and TableIII, on informations published previously sister as group. Laliostoma and Aglyptodactylus (Claw et ah, 1998; Vences et ah, 1998). Non-os- sister were taxa. Bootstrap support for most group- teological data were taken from the information ings was low, indicating their relatively low reli- summarized in Claw et ah (1998) and Glaw and ability. Vences (1994). Three species ofHeterixalus (family

Hyperoliidae) were used as outgroup.

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Discussion tion on the biogeographic regions within Mada-

defined Glaw and gascar by Angel (1942), Vences

Phylogenetic relationships of Laurentomantis (1994) and Raxworthy and Nussbaum (1995). The

presence of M. ventrimaculatus at Isaka-Ivondro

Phylogenetic analysis placed the two Laurento- in the South-East and at Andasibe in the Central

East both 675 mantis species as sister species deeply nested within (distance between localities ca. km)

the The a further of a genus Mantidactylus. phylogenetic posi- provides example species occupying

tioning of Mantella and Laurentomantis, nested a large and probably more or less continuous dis-

tribution the rainforests. within Mantidactylus, agrees with the (much bet- area along eastern On the

ter supported) results of the molecular study of other hand, the sharp distribution border between

M. and M. striatus Richards et al. (2000). Based on these data, we malagasius is a further example

continue to use Laurentomantis as a subgenus of of the faunal turnover between the Eastern and

On North-Eastern which at least takes Mantidactylus rather than as a valid genus. the regions partly

other be in the hand, Mantidactylus appears to paraphyle- place Masoala-Tsararano-Anjanaharibe cor-

because the well established Mantella ridors al. without tic, genus (Vences et 1999), conspicuous

resulted to be phylogenetically related to the sub- recent distribution barriers. Finally, the presence

of horridus in Sambirano the genera Blommersia and Guibemantis. The mono- M. (Nosy Be), Cen-

of defined here tral North and the North phyly Laurentomantis as appears (Tsaratanana) (Montagne

to be ascertained considering the similar advertise- d’Ambre) emphasizes the similarities between the

ment calls (long notes of widely spaced short pulses) herpetofaunas occurring in these regions.

and similar of the four included morphology spe-

cies. Within Laurentomantis, M. malagasius and

M. striatus almost certainly are sister species based Relationships and origin of mantellids

on their similarities in morphology and advertise-

calls. results corroborate that ment Our the two genera Boophis

Due to the derived mating behaviour (absence and Mantidactylus (classified in two separate sub- of strong mating amplexus), sexual dimorphism families, Boophinae and Mantellinae; Vences &

in affects different character Mantidactylus com- Glaw, 2001) belong to a monophyletic lineage en-

plexes in comparison to most other frogs. As in demic to Madagascar and adjacent islands (Richards other of Mantidactylus, males the subgenus Lau- and Moore, 1998; Bossuyt and Milinkovitch, 2000; rentomantis lack dimor- nuptial pads. Sexual size Vences et al., 2000) which has been defined as

to be rather in at least phism appears pronounced family Mantellidae (Vences and Glaw, 2001). A

M. horridus and two species, M. ventrimaculatus, major difference between the two genera is the

in which male size is 76-82% of female size. In loss of the apomorphic anterolateral process of the

contrast, it is faint in M. malagasius and M. strickus hyoid plate in most Boophis (Glaw et al., 1998),

size >90% of female least in (male size). At M. but this process is present in B. tephraeomystax

striatus, males have a (slightly) larger inner meta- according to the new datapresented herein (Tables

tarsal tubercle, a state shared with species of the 111 and IV). We also ascertained that the number

and a of used both subgenera Phylacomantis Gephyromantis; tarsals, previously to distinguish genera

survey ofthis character in more subgenera ofManti- (Blommers-Schlosser, 1993; Glaw et al., 1998) is

is its dactylus necessary to assess phylogenetic variable within Boophis (a small but distinct third

value. free tarsal in B. tephraeomystax and B. guibei; only

two tarsals in the remaining species examined) and

in possibly also Mantidactylus (if our observation

Biogeography ofonly two tarsals in M. horridus is not an artifact

caused alcian blue of the often by poor staining

The distribution patterns of the four Laurentomantis cartilaginous third tarsal). This also stresses that

of species as revised herein provide some informa- the value osteology and morphology to assess

Downloaded from Brill.com10/09/2021 09:33:27PM via free access Contributions to Zoology, 70 (4) - 2002 207

faintly 21 1 1 0 2* 2 7 i1 2 2 7 ?7 ?7 7 1! 0 0 2 2 2 0I1 0 0 0 1 1 0?7 7 0 0 ?7 7 2 2 2 numbers 0 2 2 2 2 2 2 2 2 7 2 2 2 2 2 2 2 7 2 10 20 1/0 22 02 02 02 02 02 2 2 2 2* 2 1 two 2 2 2 2 7 2 2 2 7 7 7 7 2 2 2 2 2 2 2 2 2 2 2 2 2 7 2 2 2 2 7 2 2 2 2 2 2 2 '? 22 2 2 2 2 7 2 0 0 0 0 0 0 7 0 represent - 27? 27? 220 27? 27? 27? 19 0 2 2 2 0 00 0 0 0 2 1 2 2 2 2 2 2 2 7 3 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 between 2 2 2I1 I12 2 2 2 2 2 2 2

(0)(0)

18 2 1 11 1 2 00 0 7 0 0 7 0 0 0 7 7 0 0 0 parentheses 0 0 00 7 00 0 7 00 1 2 2 ii 0 0 0 02 2 7 7 7 7 7 7 hyphen 10 in 17 01 01 01 02 00 00 7? 10 00 7? 10 20 7? 7? 10 10 10 7? 10 10 7? 20 20 10 10 10 22 22 22 22 20 0 0 0 0 0 0? 21 7? 1? 21 01 2? 7? 7? 0? 0 0 0 0 07 7 i 7 ii i1 i i i i 0 0 2 7 7 1 7 i 1 i i 7 2 2 2 I 1 12 A 2 2 7 2 1 2 0 2 2 7 7 0

16 0 0 0 00 3 ?7 3 1 1 7 I1 1 1 2 1 1 1 1 1 Numbers 1 1 2 2 2 (2) 7 7 7 2 2 3 2 1 2 ?7 7 1 i i1i1 2 2 7 2 7 1 0 3 2 3 3 3 3 3 3 species. 1 3 3 7 3 11 13 1 1 I1 1 0 1 11 1-21. 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 11 I1 1 1 1 15 0 1 1 1 1 1 1 1I 1 1 1 1 I 1 1 I1 1 1 1 1 same 1 1 1 1 1

the 14 01 0 0 0 0 7 0 1 1 7 7 0 0 0 0 i 0 1 0 1 ii i 00 0 I 0 1 1 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 00 7 0 characters of 13 21 71 7 7 7 2 i1 i 1 2 7 0 0 0 0 0 0 0 7 7 i 7 2 1 I 2 1 7 iIi i 2 0 00 0 1 I I 1 2 2 2 0 1 1 2 0 1 1 2 7 of 100 12 2 2 2 1 0-10-1 0-10-1 7 2 3 3 3 3 3 3 3 3 1 1 2 3 3 2 3 3 0 0 2 2 2 2 2 2 2 1 2 1 1 2 2 2 1 0 0 specimens 0 0 0 0 7 0 states 0301 1200 1200 0020 1 121 11 1 1 1 1 1 i1 i 1 7 00 1 7 0 7 0 0 1 7 i i i 1 1 i 1ii1 i i i i i 0? 0 i i 1 i i 0? (left/right). 00 0 0 0 0 0 0 0 0 7 0 012?0 7021? . further 70020 00010 0002 10 (1) 0 0 1 7 0 (0) (0) (0) observed 0 1 0 0 7 7 7 0 0 0 0 7 (0) 0 0 0 00 77 0 7 7 7 ?’7 0 7 7 0 i1 i 0 7 7 0? 7 7 0 0 0 0 in states the (1) 9 (1)(I) but 9 0 0 0 0 1 1? (1) (1) 7 0 0 0 0 1 0 0 00 0 0 0 0 1 1 (1)7(1)? (0) (0)(0) (1) 7 7 7 1 1 1 1 1 two 0 0 0 7 0 7 0 1 0 0 by of 20001270 20001270 07 8 0 0 0 0 0 0 1 0 1 0 0 0 70171200 0 00 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 10 0 00 0 0 0 0 0 0 0 specimen 20 001110210 000

7 2 7 2 2 1 1 I 11 1 0 0 0 I 2 2 0 0 7 0 0 0 0 0 070? 1 expressed occurrence 2 202000. 0 7 01001?0 0 0 0 0 0 00 0 0 0 0 0 0 0 0 (I)(1) (1)(1) (1) as 6 (1) 0 1 6 0 7 10 00 1? (0) (1)(i) (1)(1) 0 0 0 0 0 0 0 7 7 0 0 7 0 7 0 7 i 7 i i i1 0 0 7 00 0 0 0 0 0 0 010000 0 (0)(0) (0) (1)

5 0 1 0 0 0 1 0(1) I1 1 I 1 0 0 0 1 1 0 1 1 1 (0) 1 7 1 1 1 1I I 1 1 1 1 1 II 0 0 0 0 0 0 1 1 corresponding 101010107010 0 0 1 1 1 0 specimens

2 2 0 4 0 0 7 40 0 0 7 2 0 0 7 7 2 00 00 0 0 0 0 0 0 0 0 7 2 2 2 2 0 3 2 2 7 0 1 1 simultaneous 0 2 2 0 1 11 the 210I70070I 101 in 2I0210100712I0I 3 0 0 00 71770770777777 0 2 22200011000301 12200020000300 1 2277101000137? 1 7 2 2 7 2 7 7 100270011073111 11101770700371 1 1 1 I 1 0 22771010071301 22201010077301 22101010071300 22771010001301 0 0 120200000003710 0 021001000001010 0 00 100011001711100 0 021021100101210 0 0 0 0 7 10000000101211 7 11021000770000 0 0 70021000070110 0177100070012071001000070110 0 0 021021000170010 0 7701100000????0 stained 721710000000020 marks 2220002100 110007000 1770 1021 i 1i 11001700 1-2 2 2 7 i 0 2 2 1 1 1 and 2 0 22 2 2 2 22 0 1 01011000 02001000 0 2 1 1 1 10 0 1 1 0 1 10 0 1 1 1 0 0 0 0 0 I 1 i verified ?2000 27020 71020 10000? 0002 01010 2 I2101 7 1-21-2 0/2 11 2/0 1 2 i1 1 0/2 2/0 2/0 7 7 7 2 2 2 2 2 2 2 1 1 1 1 10 0 2 1 1 1 1 1 1 1 not 1 1 7 7 i 0 2 0 0 cleared numbers 2 2 0 0 0 2 7 2 of were two Nr. (73)(73) (679)(679) 054 60018 52647 1973.5351973.535 49125C 1972.1875 1972.1904 1973.8831973.883 1973.886 1972.393 analysis which 60018 52647 53606 52682 69967 68806 68807 62794 62791 62790 62793 62792 A437.1 17655 8877 46021 FN A2012 59897 52726 52712 68804 68805 A380.2A380.2 25373 A2013 A2014 68810 A2015 50651 53660 48166 53631 50648 between 6769 69726972 6810 6766 6786 6853 states Catalogue ZFMK ZFMK ZFMK ZFMK ZFMKZFMK ZFMK MNHNMNHN ZFMK ZFMK ZFMK ZFMK ZFMK MRSN MRSN MRSN MSNG slash ZMA .ZFMK ZFMK ZFMK ZFMK ZFMK ZFMK ZMAZMA ZFMK ZFMK ZFMK ZFMK MSNG ZFMK MNHN MNHNMNHN ZFMK ZMA ZFMK ZFMK MRSNMRSN MNHN MNHN ZFMKZFMK MRSN MRSN ZMA ZFMK ZMA ZMA MRSN ZFMK MNHN ZFMKZFMK ZFMK ZFMK ZFMK

osteologica! A marks ensis

the star states. of opiparisopiparis A depressiceps madagascariensis blommersae blommersae blommersae blommersae blommersae liber depressicepspulcherpulcher bicalcaratus aglaveiredimitusredimitus rivicola striatus horridus webbi boulengeriboulengeri boulengeri luteus biporus ulcerosus ulcerosus majori lugubris lugubrislugubris guttulatus grandidieri punctatus madagascari variabilis labrosum labrosum cf. madagascariensis Results states, madagascariensis crocea laevigata aurantiaca tephraeomystaxtephraeomystax tephraeomystax microtympanummicrotympanum intermediate us guibei boehmei laurentilaurenti miniatus miniatus viridis guibeicf. III. Table expressed marks Species HelerixalusHeterixalus HeterixalHeterixalus Heterixalus Aglypt.Laliostoma Laliostoma Mantella MantellaMantella MantellaMantella Mantidactylus MantidactylusMantidactylus Mantidactylus Mantidactylus Mantidactylus Mantidactylus MantidactylusMantidactylus Mantidactylus Mantidactylus MantidactylusMantidactylus Mantidactylus Mantidactylus Mantidactylus Mantidactylus MantidactylusMantidactylus Mantidactylus MantidactylusMantidactylus Mantidactylus MantidactylusMantidactylus Mantidactylus Mantidactylus MantidactylusMantidactylus Mantidactylus Mantidactylus Mantidactylus BoophisBoophis BoophisBoophis Boophis BoophisBoophis Boophis Boophis BoophisBoophis Boophis Boophis BoophisBoophis Boophis

Downloaded from Brill.com10/09/2021 09:33:27PM via free access 208 M. Vences et al. - Revision o/Laurentomantis (Amphibia, Mantellidae) ofMadagascar

0 •’S’ in 9 0 9 0 0 0 0 0 0 0 ’t 0 0 0 0 9 ? 9 9 9 9 0 9 0 6 i i1 i 0!?6? 9 ?6 ? ? =

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sC ■n 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 0 0 t'A'Cr'Xff'0'-Nr, 0 10020000000 10020210700 10020200000 1002001077? 0 0 1 0 0 1102000IOOI0 0 100170727000 0 0 0 0 0 1000007077?0 0 0 r, 0 0 0 0 0 0 0 0 0 0 • 0 1 I I T^^^^'t^i/iiflinifii/) 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 00 0 0 0

T■*r 1 1 1 0 0 0 1 0 0 0 0 10 10 1 1 ZFMK 3 3 2 2 3 1 1 13 1I1 2 13 2 I 0 0 1030120002777 101012000277? 1031020000000 13

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f o •'t 1 I 0 0 2 3 3 3 2 1 1 2 12 3 2 1 1 verified , 2 3 3 3 2 10 0 2 0 0 0 I 0 1 1 1 I 0

. i 121 121 1 1 1 O' m 00 0 I 1 1I 1 I 1 I 1 1 1 1 1 1 1 1 1 1221310100020000000 1221310100020000700 1121201020021202777 I 1 1 1 1 1 10 0 0 1221210000020200000 1001000020020000100 1000000730010000000 0 001000100010020010??? 1000000030020010??? 1 ,

ac f*3 2 2 2 1 0 2 2 2 12 0 2 9 2 2 2 0 0 0 0 ■ I 2 0 1 2 0 0 0 00 00 0 0 0 0 0 encountered either i 12212101 12012101 h f*5 1 12010101 0 2 2 2 0 2 00 0 2 2 2 2 2 2 2 0 0 2 1 1 122131 1221311 1 12 0 1 0 2 10 11213 100121 0 0 0 0 I 1101 1201 1700100101 1201 121 II i sc 0 2 2 1 1 1 1 1 1 1 0 1 i i ii i 00 0 9 9 0 i i 0 i i Xi i i i 9i ii

were 1

m i i i 0 i1i 9 i1i i i i i i i i1 i1 i code 0 i i i i 1i i i i 1 1i1Ii Ii1 ii i ii i

’’t 2 I 1 2 2 0 0 2 1 12 2 2 2 2 2 2 1 I 2 1 1I 1 1 1 1 1 to 1 2 I I 2 2 2 2 2 2 2 2 i i^nnnnnni

rn 2 2 2 0 0 0 2 0 1 2 P9 1 2 2 2 I I 0 0 0 1 1 I 1 1 1 101 101 0 0 0101 0 0 2 2 2 2 2 2 2 2 , used rs f*i 2 2 2 2 0 0 2 2 1 2 0 1 characters 9 9 2 2 9 2 2 0 0 2 1 ? 2 1 2 2 i ? 79 79 9 2 2 2 2 2

m 0 0 0 0 2 2 1 0 0 0 0 000 3 9 3 3 9 3 3 1 1 I 1 1 1 0 0 are 1 1I 1 I 1 1 1 9

o m 6 9 96 0 0 0 0 i 0 9 ? 4 4 9 4 0 2 000 i 4 0 79 3 ?7 796?6 0 0 . 1 9 9 9

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«N fN 0 0 1 2 2 2 2 0 0 0 1 I 1 1 2 22 2 11022120001771777121717137023002007177?2 2 2 3 3 3 3 3 4 4 0 0 0 0 0 00 0 III. | 1

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Table O' 10012210001132000221 0 0 0 0 0 2 1 2 2 2 2 2 2 2 I 11720200000721707221 3 2 2 2 2 2 2 2 2 2 2 11022020001132430 2 12 2 2 2 2 2 2 2 i 117722020001777127221 17772102000I137I27221 127722720001132770I21 the X i A 1 i 2 0 0 0 0 0 0 9 ?6000 0 9 ? 0 i 0 0 2 2 2 i 1210220200017324321 i 100221200011 1103A02000 60 9 0? 9 ? from r- 12122231 12212241 in 0 0 0 0 0 0 0 1 2 i i ii i i i 0 0 1 1 1202219 9 9 ? C2022232 2 11022232 2 2 I 2 0 2 2 , i'i 96 ir-'C^x5'0-Nf 10121022120001732732121

1201301022000000000020002001 1 — 1 1 SC 0 0 0 3 3 2 2 9 2 1 2 3 2 22003020770000070000200022 1200221 1 1 i 2 1 9 c i i ii 0 D 1 3 3 3 3 33 are

i

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1 Tf « 0 0 0 0 1 1 A 1 I1 1 1 0 1 1 1 included 0 0 00 1 0 0 0 0 0 0 0 0 1-21) 1I 0 0 0 0 0 0 0 0

r*'. 2 2 — 9 9 9 0 0 9 i1 i ? 1 12 1 i i 0 0 2 0 0 1 9 200 0 10 1 c 0 2 1 2 1 22 N 2 2 2 1 1 A A 2 3 3 3 3 3 3 2 3 3 10 0 2 2 12 2 2 10210102212241 taxa 2 1 2 I 0 0 0 0 0 0 0

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)

sc 1 1 1 1 0 0 0 i i i 000810000001C01D222220000000000222I 01010I00000020132I202000000220I2221 appendix) 0 000 0 9 96 0 9 6 9 9 1 i i 21710000000020130720200000777772221 6 100007007012001210227200011324321 A 0 96 0 0 0 0 0 1 0 0

n I 0 0 0 10 0 0 1 0 i i 0 1 i 79 i 1 10 A 01 A A I I 1 1 1010000013201 0 0 0 0 1 1 1 101770700371 I0A020AG121 0 T 0 2 2 0 4 0 0 00 2 0 0 0 0 2 0 0 0 0 1000700001A710 2 2 2 0 3 2 2 1 10 1 (see B 10210007700001 00 2 02A100 1 Osteological , 122000200003001 r*i 10010001 0 22C0101000130A1 1 1 0210101010701011 1 1G02AI00I712001 c B2101700701A21 0 0 0 0 2 2 0 22200021000301 2 0 0 0 0 0 0 1 0 0 0 0 0 0 1100170017021 1 0 0 0 0 210010000010 1000000010121 0 0 0 0 ? i 00000100101 r-i 2 2 2 001307001 i i 1 1 1 2 9 A 2 2 0 1I 0 2 2 1 1 1 I1 I 0 0 00020001 1 1 I 0 0 0 3). 10001 00 I 010112 21011 i — 9 2 9 1 2 1 2 2 1 1 1 1 1 0 0 B 2 1 Cc B 1 2 1 00 1 characters — 1 1 1 1 B 0 10 2 0 0 2 2 and (1994). 54 2 = iceps all D of 2; Vences blommersaeblommersae depressdepressicepspulcherpulcherbicalcaratus aglaveiredimitusredimitus rivicola horridus boulengeri biporus ulcerosus guttulatusguttulalusgrandidieri madagascariensis a liber striatus webbi luteus biporus majorilugubris opiparis and and punctatuspunctalus variabilisvariabilis labrosum States 1 crocea laevigata aurantiaca tephraeomystax madagascariensis madagascariensis croce guibeiguibei boehmeiboehmei laurenti microtympanum miniatus viridis IV. C Glaw ella 2; Helerixalus Heterixalus HelerixalusHelerixalus Laliostoma Mantella Mantella Mantella MantidactylusMantidactylusMantidactylus MantidactylusMantidactylusMantidactylusMantidactylus MantidactylusMantidactylusMantidactylusMantidactylus MantidactylusMantidactylusMantidactylusMantidactylus MantidactylusMantidactylus Mantidactylus MantidactylusMantidactylusMantidactylus MantidactylusMantidactylus Mantidactylus Mantidactylus Mantidactylus Table and from Aglypt. Mant Mantella BoophisBoophisBoophisBoophisBoophisBoophisBoophis BoophisBoophisBoophisBoophis

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phylogenetic relationships among frogs is limited Acknowledgements by the high degree of homoplasy affecting many characters. We areindebted to N.Rabibisoa, D. Rakotomalala, O. Ramilison,

F. Ranaivojaona, and J. E. Randrianirina for their help in the Vences et al. (2000) noted that basal relict groups field. W. Bdhme (Bonn). B. Clarke and C. McCarthy (London), in amongMalagasy anurans mainly occur dry wes- B. van Tuijl (Amsterdam), D. Vallan and K. Grossenbacher and that the tern Madagascar, hypothesized ances- (Bern), G. Kohler (Frankfurt), R. Poggi and G. Doria (Genova), tors of extant rainforest in were frogs Madagascar and A. Dubois and A. Ohler (Paris) allowed examination and

to arid conditions. In adapted contrast, specializa- partly dissection of specimens held in their care. W. Bohme tion to constant humid conditions is probably found additionally allowed the exchange ofthe holotype ofM. striatus

the ZSM. M. Matasconi carried ofthe osteo- in the Mantidactylus species without free-swim- to (Bonn) out part

logical examinations. We are gratefulto the Malagasy authorities ming larvae: direct development is known in the for research and export permits. The work of FG and MV was subgenus Gephyromantis, and is probable at least made possible by a cooperation accord between the Departe- in some Laurentomantisbased on the data presented ment de Biologie Animale, Universite d’Antananarivo and the in the in- present paper (large eggs, calling largely ZoologischesForschungsinstitut und Museum A. Koenig, Bonn,

dependent from water bodies). As no terrestrial and financially supported by the Deutscher AkademischerAus-

tauschdienst (DAAD). The work of FA at Masoala was Tertiary fossils are known from Madagascar, the survey

carried out within a project in collaboration with the Parc classification of the Latest Cretaceous remains as de Tsimbazaza Botanique et Zoologique (Antananarivo), sup- described by Asher and Krause (1998) is of im- ported by the Wildlife Conservation Society (WCS) (Anta- to reconstruct the of these portance timing adap- nanarivo), and the Museo Regionale di ScienzeNaturali (Torino)

The aimed at tive radiations into rainforest environments. monitoring some rainforest areas in northern Ma-

- dagascar. For their to the WCS main fragments - an atlas and a sacrum belong assistance, he is very grateful staff at Antananarivo, especially M. Flatchwell, and to all the to large frogs characterized by a procoelous verte- people ofWCS at Maroantsetra. bral column and narrowly separated atlantal cotyles.

As mantellids are generally characterized as dipla-

siocoelous and having widely separated cotyles, References Asher and Krause (1998) concluded that the fos-

sils were related to the more basal Pelobatidae Angel F. 1935. Batraciens nouveaux de Madagascar, recoltes (which has no extant representatives in Madagas- M. Catala. Zool. 60: 202-207. par R. Bull. Soc. France car and Subsaharan Africa). This conclusion is Angel F. 1942. Les lezards de Madagascar. Mem. Academic

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it of the ceous) record of Anura from Madagascar. J. Vertebrate Pa- However, must be noted that one largest leontology 18: 696-699. extant endemic Malagasy frogs, Mantidactylus gut- Blonimers-Schlosser RMA. 1993. Systematic relationships of tulatus, is characterized by procoely (Guibe, 1978). the MantellinaeLaurent 1946 (AnuraRanoidea). Ethol. Ecol. to data this and According presented here, species Evol. 5: 199-218.

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todactylus andLaliostoma, also have narrowly sep- (premiere partie). Faune de Madagascar 75 (1): 1-379.

Blonimers-Schlosser RMA, Blanc CP. 1993. Amphibiens arated cotyles. In these species, the space between (deuxieme partie). Faune de Madagascar 75 (2): 385-530. cotyles is slightly larger thanthe width of one cotyle, Bocttgcr O. 1880. Diagnoses Reptilium et Batrachiorum agreeing with the state shown by Asher and Krause novorum a Carolo Ebenau in insula Nossi-Be Madagas- (1998) in their fossil specimen FMNH PR-1961. cariensi lectorum. Zooi Anz. 3: 279-283. The that the remains belong to Manti- hypothesis Boulenger GA. 1882. Catalogue of the Batrachia Salientia s.

dactylus, and that mantellids were already present Ecaudata in the collection of the British Museum. Addenda.

London, Trustees of the British Museum. on Madagascar during the Latest Cretaceous, can

Bossuyt F, Milinkovitch MC. 2000. Convergent adaptive ra- thus not be ruled out at the present state. diations in Madagascan and Asian ranid frogs reveal

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Sci. USA 97: 6585-6590.

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lionships in the African Raninae (Anura Ranidae). Monit. Richards CM, Nussbaum RA, Raxworthy CJ. 2000. Phylo-

zool. Hal. (N.S.) suppl. 15: 285-331. genetic relationships within the Madagascan boophids and

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Dubois A. 1992. Notes sur la classification des Ranidae simony (* and other methods), Version 4. Sunderland,

(Amphibiens Anoures). Bull. mens. Soc. linn. Lyon 61: 305- Massachusets: Sinauer Associates.

352. Vallan D. 2000. Influence of forest fragmentation on amphib-

Glavv F, Vences M. 1994. A fieldguide to the amphibians and ian diversity in the nature reserve of Ambohitantely, high-

reptiles ofMadagascar. 2nd edition. Kdln: Vences and Glaw land Madagascar. Biol. Conserv. 96: 31-43.

Verlag. Vences M, Glaw F. 2001. When molecules claim for taxonomic

Glavv F, Vences M. 2000. Current counts of species diversity change: New proposals on the classification of Old World

and endemism of Malagasy amphibians and reptiles. In: treefrogs. Spixiana 24: 85-92.

Lourenco WR, Goodman SM (eds.). Diversite et endemisme Vences M, Glaw F. submitted. Revision of the subgenus

de Madagascar. Memoires de la Societe de Biogeographie, Chonomantis (Anura: Ranidae: Mantidactylus) from

Paris, 243-248. Madagascar, with description of two new species.

Glaw F, Vences M, BohmeW. 1998. Systematic revision ofthe Vences M, Glavv F, Mausfeld P, Bohme W. 1998. Comparative

1919 of of the genus Aglyptodactylus Boulenger, (Amphibia: osteology Malagasy poison frogs genus Mantella

Ranidae), and analysis of its phylogenetic relationships to (Amphibia: Ranidae: Mantellinae). Bonn. zool. Beitr 48:

other ranid Madagascan genera (Tomopterna, Boophis, 205-215.

Manlidactylus, and Mantella). .1 Zool. Syst. Evol. Res. 36: Vences M, Ziegler T, Visscr S, Andreone F. 1999. New data

17-37. on the zoogeography and genital morphology of the lizards

of Glavv F, Vences M, Gossmann V. 2000. A new species Zonosaurus brygooi Lang and Bdhme 1990 and Z. aeneus

Mantidactylus (subgenus Guibemantis) from Madagascar, (Grandidier 1872) from Madagascar (Reptilia Squamata

with of internal femoral a comparative survey gland struc- Gerrhosauridae). Tropical Zool. 12: 145-155.

in ture the genus (Amphibia: Ranidae: Mantellinae). J. Nat. Vences M, Glavv F, Kosuch J, Das I,Veith M. 2000. Polyphyly

Hist. 34: 1135-1154. of based of Tomopterna (Amphibia: Ranidae) on sequences

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II: 1-140. of relict Monogr. biogeography Malagasy amphibian groups. In:

Hofrichter R. 1998 (ed.). Biologic der Amphihien. Augsburg: Lourenco WR, Goodman SM (eds.). Diversite et endemisme

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vertebrates from Madagascar. Implications for biotic

changes in deep time. In: Goodman SM, Patterson BD (eds). Received: 14 May 2001

Natural change and human impact in Madagascar.

Smithsonian Institution Press, Washington and London, 3-

43. Appendix: Characters used for analysis Methuen PA. 1920. Descriptions ofa new snake from Transvaal,

together with a new diagnosis and key of the genus

Character I. Medial contact of nasals: (0) close medial con- Xenocalamus, and of some Batrachia from Madagascar. between nasals Proc. Zool. Soc. London 25: 349-355. tact nasals, (1) narrowly separated medially,

(2) nasals Methuen PA, Hewitt .1. 1913. On a collection of Batrachia widely separated medially.

Character 2. of from made Ann. Transvaal Squamosal: (0) zygomatic process squamosal Madagascar duringthe year 1911. of similar length or longer than Museum (4) 2: 49-64. posterior (otic) process, (1) zygomatic of shorter than Perrier de la Batbic II. 1927. Le Tsaratanana, PAnkaratra et process squamosal slightly posterior

(otic) of much shorter PAndringitra. Mem. Acad, malgache 3: 1-70. process, (2) zygomatic process squamosal than Nussbaum RA. 1994. A rainforest of posterior (otic) process. Rawvorthy CJ, survey Character 3. of Maxilla preorbital fascialis: a amphibians, reptiles and small mammals at Montagne process pars (0)

distinct triangular or elongate at the level of palatines, d’Ambre, Madagascar. Biol. Conserv. 69: 65-73. process

mostly directed posteriorly, (I) a small distinct triangular or Raxworthy CJ, Nussbaum RA. 1995. Systematics, speciation

elongate at the level ofpalatines, (2) absent or and biogeography of the dwarf chameleons ( Brookesia; process process reduced. Reptilia, Squamata, Chamaeleontidae) of northern Mada- very Character 4. Maxilla anterior maxilla J. Zool. Land. 235: 525-558. preorbital process, part: (0) gascar.

anteriorly not especially high, (1) maxilla anteriorly with a Richards CM, Moore WS. 1998. A molecular phylogenetic

distinct separated from preorbital by a deep study of the Old World Treefrogs, family Rhacophoridae. process, process

notch, (2) maxilla anteriorly high, with a similar height to the Herpetol. J. 8: 41-46. which is still preorbital process protruding higher, (3) maxilla

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the level anteriorly high, continuing to of the palatines, wit- known, strong mating amplexus and release calls absent). hout additional maxilla preorbital process, (4) anteriorly very Character 27. Egg deposition: (0) aquatic, (1) terrestrial.

with two anterior high, protruding an oneand a second Character 28. Maximum number processes, of eggs laid by one female:

one at the level of the palatines. (0) >1000, (I) 500-1000, (2) 100-500, (3) <100. Character 5. Anterior end ofmaxilla: (0) (1) convex. Character 29. Maximum of concave, diameter eggs or mature oocytes

Character 6. Palatines: to vomerine in (0) reaching medially gravid females: (0) 1-2 mm, (1) larger than 2 mm.

odontophore, (1) reaching further than Character medially odontophore. 30. Tadpole morphology: (0) generalized tadpole, Character 7. Vomerine odontophore and vomerine teeth: (0) (1) specialized phytotelmous tadpole, (2) funnel-mouthtadpole, teeth and odontophorepresent, teeth absent or (1) rudimentary, (3) specialized tadpole with probably filtering mouthparts, (4) but thin, odontophore present very (2) teeth and odontophore (probably) direct development. absent. Character 31. Larval development in: (0) free lentic water, (1) Character 8. Maxillary teeth: (0) present, (1) absent. free lotic water, (2) phytotelmes or tree holes, (3) (probably)

Character 9. Parasphenoid: (0) not reaching palatines anteriorly, outside free water.

(1) reaching palatines anteriorly. Character 32. Calling activity; (0) mainly diurnal, (1) diurnal Character 10. Separation ofatlantal cotyles: (0) wide (distance and nocturnal, (2) mainly nocturnal.

between cotyles than their narrow larger width), (1) (distance Character 33. Habits: (0)terrestrial/semiaquatic, (1) scansorial, between cotyles smaller than their width). (2) arboreal.

Character II. Vertebral column: (0) procoelous, (1) diplasio- Character 34. Circummarginal groove ventrally on finger and coelous. toe pads: (0) absent or incomplete, (1) complete. Character 12. Base of omosternum: unforked, Character 35. (0) (1) slightly Fingertips: (0) not enlarged, (1) slightly enlarged, the between forked, greatest space the arms is less than half (2) distinctly enlarged. the width ofone the arm, (2) moderately forked, greatest space Character 36. Relative finger length: (0) second finger longer between the is to arms once twice the width of one arm, (3) than first finger, (1) first and second finger of similar length,

the is first broadly forked, greatest space between the arms two to (2) finger longer than second finger.

four times the with of one arm. Character 37. Relative toe length: (0) fifth toe longerthan third

Character 13. Relative omosternum length: (0) longer than toe, (1) fifth and third toe of similar length, (2) fifth toe shorter

of shorter sternum, (1) same lengthas sternum, (2) than sternum. than third toe.

Character 14. Base of sternum: (0) unforked, (1) forked. Character 38. Lateral metatarsalia: (0) separated by webbing, Character 15. Intercalary element: (0) absent, (1) present. (1) partly connected, (2) connected.

Character 16. Free distal tarsals: (0) three free the third Character 39. between tarsals, Webbing fingers: (0) present and distinct, tarsal rather free being large, (1) three tarsals, the third tarsal (1) absent or almost absent. being small, three free tarsals, the third tarsal Character (2) beingextremely 40. Inner foot webbingat fourth toe; (0) 0-1 phalanges small, (3) two free tarsals. free ofweb, (1) 1.5-2 phalanges free ofweb, (2) 2.5-3 phalanges 1 7. Number of Character prehallux elements in one free of arranged web, (3) no foot webbing. row: (0) two, (1) three, (2) four. Character 41. Outer metatarsal tubercle: (0) absent, (1) pre- Character 18. Additional prehallux element, leading to an axe- sent. shape: (0) absent, (1) present, small, (2) present, distinct. Character 42. Outer tubercles: very metacarpal (0) single orindistinct, Character 19. Terminal phalanges; (0) not bilobed nor forked (I) double and prominent.

(more or less knob-like), (1) 43. very slightly bilobed, (2) distinctly Character Relative foot length: (0) foot longer than tibia, bilobed or slightly Y-shaped, foot and tibia (3) distinctly Y-shaped. (1) more or less of same length, (2) foot shorter

Character 20. Anterolateral of than process hyoid plate: (0) absent, tibia.

(1) rudimentary, (2) and 44. present distinct. Character Relative hindlimblength: tibiotarsalarticulation

Posterolateral Character 21. process ofhyoid plate: (0) absent, mostly reaches: (0) between and tympanum eye, (I) beyond

(I) rudimentary, (2) present and distinct. not eye, beyond nostril, (2) beyond nostril, not beyond snout Character 22. Femoral in males glands (according to Glaw et tip, (3) beyond snout tip. al. 2000): (0) absent, (1) gland 45. Sexual type 1, (2) gland type 2, (3) Character dimorphism in relative tympanum size: gland type 3, (4) gland type 4. tympanum ofsimilar size (0) in males and females, (1) tympanum Character 23. distal femoral section with external Enlarged gland distinctly larger in males than in females. central in males: median depression (0) absent, (1) present. Character 46. Dermal above spines or prominent granules eyes:

Character 24. Femoral in glands females: (0) absent, (1) pre- (0) absent, (1) present. sent rudiments. as Character 47. Position of than nostrils: (0) nearer to eye to

Character 25. Tibial in glands (0) absent, (1) present males snout between and tip, (1) midway eye snout tip, (2) nearer to absent in (present or females). than snout tip to eye.

Character 26. Nuptial pads and reproductive behaviour; Character (0)* 48. Vocal sac: (0) single or slightly bilobed subgular, nuptial pads present (as far as known, strong mating amplexus (1) paired subgular, and release calls absent far present), (1) nuptial pads (as as Character 49. Vocal sac color; (0) no conspicuous color (not

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clearly differing from general ventral coloration), (I) black, Character 52. Cellular DNA content: (0) around modal value,

(2) white. (1) distinctly lower than modal value.

Character Vocal sac inflation: vocal disten- Character 50. (0) sac slightly 53. Karyotype: (0) 2n=26 chromosomes, (1) 2n=24 sible, (1) vocal sac largely distensible. chromosomes.

Character 51. Dorsal skin: (0) smooth, (1) slightly granular, Character 54. Acrocentric chromosomes: (0) absent, (1) present.

(2) moderately to coarsely granular.

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