Describing the smaller majority: integrative taxonomy reveals twenty-six new species of tiny microhylid frogs (genus Stumpffia) from Madagascar

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1 Senckenberg Gesellschft für Nturforschung, (3): Describing the smller mjority: integrtive txonomy revels twenty-six new species of tiny microhylid frogs (genus Stumpffi) from Mdgscr Andollo Rkotorison 1, 2, Mrk D. Scherz 1, 3, Frnk Glw 3, Jörn Köhler 4, Frnco Andreone 5, Michel Frnzen 3, Julin Glos 6, Oliver Hwlitschek 3, 7, Teppei Jono 8, 9, Akir Mori 8, Serge H. Ndrintso 2, Noromll Rsommpionon Rminoso 2, Jn C. Riemnn 6, Mrk-Oliver Rödel 10, Gonçlo M. Ros 11, 12, 13, Dvid R. Vieites 14, Angelic Crottini 15, Miguel Vences 1, * 1 Division of Evolutionry Biology, Zoologicl Institute, Brunschweig University of Technology, Mendelssohnstr. 4, Brunschweig, Germny 2 Mention Zoologie et Biodiversité Animle, Fculté des Sciences, Université d Antnnrivo, Antnnrivo 101, Mdgscr 3 Zoologische Sttssmmlung München (ZSM-SNSB), Münchhusenstr. 21, München, Germny 4 Hessisches Lndesmuseum Drmstdt, Friedenspltz 1, Drmstdt, Germny 5 Museo Regionle di Scienze Nturli, Vi G. Giolitti, 36, I Torino, Itly 6 Zoologicl Institute, University of Hmburg, Mrtin-Luther-King Pltz 3, Hmburg, Germny 7 Institut de Biologi Evolutiv (CSIC- UPF), Psseig Mrítim de l Brcelonet 37, Brcelon, Spin 8 Deprtment of Zoology, Grdute School of Science, Kyoto University, Skyo, Kyoto , Jpn 9 Tropicl Biosphere Reserch Center, University of the Ryukyus, 1 Senbru, Nishihr, Okinw , Jpn 10 Museum für Nturkunde, Leibniz Institute for Evolution nd Biodiversity Science, Invlidenstr. 43, Berlin, Germny 11 Deprtment of Biology, University of Nevd, Reno, NV 89557, USA 12 Institute of Zoology, Zoologicl Society of London, Regent s Prk, London NW1 4RY, UK 13 Centre for Ecology, Evolution nd Environmentl Chnges (CE3C), Fculdde de Ciêncis Universidde de Lisbo, Lisbo, Portugl 14 Ntionl Nturl History Museum, Spnish Scientific Reserch Council (CSIC), C/José Gutierrez Abscl 2, , Mdrid, Spin 15 CIBIO, Reserch Centre in Biodiversity nd Genetic Resources, InBIO, Universidde do Porto, Cmpus Agrário de Virão, Ru Pdre Armndo Quints, Nº 7, Virão, Portugl * Corresponding uthor; m.vences@tu-bs.de Accepted 4.x Published online t on 13.xi Abstrct The genus Stumpffi Boettger, 1881 currently contins 15 nmed, smll to miniturized frog species, clssified in the endemic Mlgsy subfmily Cophyline of the fmily Microhylide. Stumpffi re terrestril frogs with lrgely unknown biology, probbly due to their smll size nd secretive hbits. Previous studies hve suggested lrge proportion of undescribed diversity in the genus. We revise the genus on the bsis of combintion of moleculr, biocoustic, nd morphologicl dt nd describe 26 new species tht re ll geneticlly divergent, lmost ll of them with high pirwise genetic divergences > 4% p-distnce in segment of the mitochondril 16S rrna gene nd concordnt differentition in segment of the nucler Rg-1 gene. The mjority of the new species cn lso be distinguished by the structure of their dvertisement clls (where biocoustic dt re vilble), nd in most comprisons the species cn lso be distinguished morphologiclly. Furthermore, moleculr phylogeny reconstructed from DNA sequences of one nucler nd four mitochondril gene segments reveled tht in mny cses, morphologiclly similr species re not ech other s closest reltives, thus confirming their identity s independent evolutionry lineges nd reveling repeted phenotypic divergence nd convergence mong nd within cldes. The phylogeny distinguishes four min cldes in the genus: Clde A contining 17 species (Stumpffi be, S. hr, S. megsoni, S. stffordi, S. psologloss, S. nlmin, S. gimmeli, S. mdgscriensis, S. pygme, S. ngeluci sp. nov., S. huwei sp. nov., S. ihrn sp. nov., S. lrinki sp. nov., S. mledict sp. nov., S. mmitik sp. nov., S. sort sp. nov., nd S. ynniki sp. nov.) mostly from northern nd northwestern Mdgscr, generlly chrcterized by limited digitl reduction nd divided in subcldes of comprtively lrge, smll, nd miniturized body size, respectively; Clde B with four species (S. miery, S. meikee sp. nov., S. obscoen sp. nov., nd S. dvidttenboroughi sp. nov.) morphologiclly rnging from miniturized with strong digitl reduction to comprtively lrge-sized; Clde C with 18 species (S. grndis, S. kibomen, S. roseifemorlis, S. tetrdctyl, S. nigrorubr sp. nov., S. chillei sp. nov., S. diutissim sp. nov., S. prdus sp. nov., S. edmondsi sp. nov., S. fusc sp. nov., S. jennoeli sp. nov., S. spndei sp. nov., S. grrffoi sp. nov., S. nlnjirofo sp. nov., S. miovov sp. nov., S. mkir sp. nov., S. betmpon sp. nov., nd S. dolchi sp. nov.) mostly distributed in estern nd northestern Mdgscr, contining species of compr- ISSN

2 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs tively lrge size s well s smll-sized species, mny of which re chrcterized by moderte degree of digitl reduction; nd Clde D with two miniturized species (S. tridctyl nd S. contumeli sp. nov.) with strong digitl reduction, which form the sister group of ll other Stumpffi. Two of the newly described species (S. ngeluci nd S. mledict) re not seprted by the 4% threshold in the 16S gene but occur in symptry nd do not shre Rg-1 hplotypes. To chieve comprehensive review of this species-rich genus, we provide simplified differentil dignoses nd descriptions nd bbrevited descriptions of morphologicl vrition. Despite the lrge number of Stumpffi species newly described herein, we identify severl dditionl cndidte species with currently insufficient dt to wrrnt forml description, nd highlight tht some species such s S. nlnjirofo, S. gimmeli, S. kibomen, S. mdgscriensis, S. roseifemorlis nd S. obscoen re composed of two or more deep mitochondril lineges tht might lso turn out to be distinct tx fter in-depth study. We confirm Stumpffi s genus of highly microendemic frogs with mny species pprently restricted to very smll rnges, nd provide evidence tht two of the new species (S. chillei nd S. dvidttenboroughi) do not construct fom nests but ly their eggs in wet plces in the lef litter, or in cvities such s empty snil shells. We propose conservtion sttus for ll the described species ccording to IUCN Red List Criteri, but lso discuss severl problems pplying these criteri to such microendemic nd poorly known frogs. Key words Amphibi; Anur; Microhylide; diversity; systemtics; biocoustics; moleculr genetics; morphology; microendemism. Tble of content Abstrct Introduction Mterils nd Methods Results Moleculr species delimittion Moleculr phylogeny Morphologicl nd biocoustic comprisons Txonomic ccounts Smll-sized or miniturized species of clde A Stumpffi psologloss Boettger, 1881 (type species of the genus Stumpffi) Stumpffi nlmin Klges, Glw, Köhler, Müller, Hipsley & Vences, Stumpffi gimmeli Glw & Vences, Stumpffi mdgscriensis Mocqurd, Stumpffi pygme Vences & Glw, Stumpffi ngeluci sp. nov Stumpffi huwei sp. nov Stumpffi ihrn sp. nov Stumpff lrinki sp. nov Stumpffi mledict sp. nov Stumpffi mmitik sp. nov Stumpffi sort sp. nov Stumpffi ynniki sp. nov Lrge-sized species of clde A Stumpffi be Köhler, Vences, D Cruze & Glw, Stumpffi hr Köhler, Vences, D Cruze & Glw, Stumpffi megsoni Köhler, Vences, D Cruze & Glw, Stumpffi stffordi Köhler, Vences, D Cruze & Glw, Smll-sized or miniturized species of cldes B nd D Stumpffi miery Ndrintso, Riemnn, Vences, Klges, Rminoso, Rödel & Glos, Stumpffi tridctyl Guibé, Stumpffi contumeli sp. nov Stumpffi obscoen sp. nov Stumpffi dvidttenboroughi sp. nov

3 VERTEBRATE ZOOLOGY 67 (3) Lrge-sized species of clde B Stumpffi meikee sp. nov Moderte- to lrge-sized species of clde C Stumpffi grndis Guibé, Stumpffi kibomen Glw, Vlln, Andreone, Edmonds, Dolch & Vences, Stumpffi roseifemorlis Guibé, Stumpffi chillei sp. nov Stumpffi nlnjirofo sp. nov Stumpffi diutissim sp. nov Stumpffi edmondsi sp. nov Stumpffi fusc sp. nov Stumpffi jennoeli sp. nov Stumpffi nigrorubr sp. nov Stumpffi prdus sp. nov Smll-sized species of clde C Stumpffi tetrdctyl Vences & Glw, Stumpffi betmpon sp. nov Stumpffi dolchi sp. nov Stumpffi grrffoi sp. nov Stumpffi mkir sp. nov Stumpffi miovov sp. nov Stumpffi spndei sp. nov Cndidte species not described herein Stumpffi sp. C Stumpffi sp. C Stumpffi sp. C Stumpffi sp. C Stumpffi sp. C Discussion Fst trck txonomy Deep genetic divergences within nd between species Reproductive modes Biogeogrphic ptterns Ptterns of digitl reduction Body size ptterns Color ptterns Cll ptterns Conservtion sttus nd conservtion significnce of newly described species Finl considertions Acknowledgments References Appendix Zoobnk registrtion numbers of new species described in this study

4 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Introduction An ccurte nd complete species inventory is the bsis for ecologicl nd evolutionry studies, s well s for conservtion mngement, nd hs lso been proposed s the morl counterweight tht humnkind cn offer to life s runwy exploittion (Dijkstr, 2016). As with mny other groups of orgnisms, the pst three decdes hve seen steep increse in the discovery nd description of new mphibin species (Köhler et l., 2005), mostly due to intensified explortion of tropicl regions nd ppliction of n integrtive txonomic pproch combining moleculr, biocoustics, nd morphologicl chrcters (Pdil et l., 2010). Prdoxiclly, while more mphibin species re being discovered thn ny time before in txonomic history, mphibins re lso undergoing serious popultion declines nd extinctions worldwide (Hnken, 1999; Sturt et l., 2004; 2008). Among the numerous threts leding to these declines (Mendelson et l., 2006; Wke & Vredenburg, 2008) re the degrdtion nd destruction of nturl hbitts nd the introduction of invsive species (Cushmn, 2006), but lso the rpid spred of emerging infectious diseses such s chytridiomycosis nd rnvirosis (Dszk et l., 2003; Skerrtt et l., 2007; Duffus & Cunninghm, 2010; Olson et l., 2013). Mdgscr is globl hotspot for biodiversity conservtion in which mphibins hve long been minly thretened by deforesttion, which progresses t lrming rtes (Hrper et l., 2007). Disese-relted threts ppered minor (Weldon et l., 2008; Wollenberg et l., 2010; Vredenburg et l., 2012; Crottini et l., 2011; 2014) but fter the recent discovery of the pthogen Btrchochytrium dendrobtidis (Kolby, 2014; Bletz et l., 2015), chytridiomycosis-relted declines my occur t ny time nd t ny rte, s hs been seen elsewhere (e.g., Hirschfeld et l., 2016). Additionl threts re posed by two invsive mphibin species, Hoplobtrchus tigrinus (Guibé, 1953; Glw & Vences, 2007), nd especilly Duttphrynus melnostictus (Moore et l., 2015; Goodmn et l., 2017; Vences et l., 2017). While the conservtion sttus of Mdgscr s mphibins hs continued to deteriorte, the numbers of newly discovered nd described species per yer hve experienced n lmost constnt increse since the 1980s (e.g., Köhler et l., 2005), nd mny dditionl species hve lredy been identified but wit description (e.g., Vieites et l., 2009). As of 2014, in ddition to the 290 described Mlgsy frog species, over 200 undescribed cndidte species of frogs were known (Crottini et l., 2012; Perl et l., 2014). As in mny niml groups, the detection of the mssive proportion of undescribed diversity in Mlgsy frogs ws boosted by the routine ppliction of genetic screening pproches. Such DNA brcoding studies hve often detected high levels of cryptic diversity (e.g., Smith et l., 2005; Monghn et l., 2009; Rmsindrzn et l., 2011), nd in Mlgsy mphibins were bsed both on the mitochondril 16S rrna gene (e.g., Vieites et l., 2009) but lso on the stndrd brcoding mrker cytochrome oxidse subunit I (COI or Cox-1) (Vences et l., 2005; Ngy et l., 2012; Perl et l., 2014). These studies confirmed tht mny species thought to be widespred re geneticlly diverse nd contin dditionl, undescribed lineges (e.g., Boumns et l., 2007; Lehtinen et l., 2007; Gehring et l., 2011; 2012). Mny of the newly discovered cndidte species hd remrkbly smll distribution rnges, fitting n overll pttern of microendemism in Mdgscr (Wilmé et l., 2006; Vences et l., 2009; Brown et l., 2016) nd thereby posing dditionl chllenges for conservtion (Andreone et l., 2008). A lrge proportion of the cryptic nd microendemic mphibin diversity of Mdgscr is mde up by the cophyline frogs. Cophyline Cope, 1889 is Mdgscr-endemic subfmily of the pntropicl fmily Microhylide Günther, 1858, representing morphologiclly diverse group chrcterised by severl osteologicl chrcters, which comprise rborel, terrestril, fossoril, nd rupicolous frogs (Blommers-Schlösser & Blnc, 1991; Glw & Vences, 2007). The typicl mode of reproduction is nidicolous one, with non-feeding tdpoles developing in tree holes nd other phytotelmt, or in terrestril fom or jelly nests (Blommers-Schlösser, 1975; Blommers-Schlösser & Blnc, 1991; Glw & Vences, 1994), nd cophyline dvertisement clls mostly consist of long series of single tonl notes (Blommers-Schlösser, 1975; Vences et l., 2006; Glw & Vences, 2007). Most cophyline gener hve been shown to be monophyletic (Andreone et l., 2005b; Wollenberg et l., 2008; Scherz et l., 2016), but the intergeneric reltionships re lrgely unresolved (Peloso et l., 2015; 2017; Scherz et l., 2016; 2017). Although it ws recently suggested (Peloso et l., 2015; 2017) tht Stumpffi should be synonymized with Rhombophryne, Scherz et l. (2016; 2017) provided genetic, morphometric, nd osteologicl grounds for the rejection of this proposl. We therefore follow the genus-level clssifiction of Scherz et l. (2016; 2017) herein. Of the eight currently recognized cophyline gener, Stumpffi Boettger, 1881, is the most diverse nd most urgently in need of txonomic revision; together, Glw & Vences (2007), Vieites et l. (2009), Köhler et l. (2010), Klges et l. (2013), Perl et l. (2014) nd Scherz et l. (2016) hve recognized over 60 cndidte species of Stumpffi, in ddition to the 15 currently recognized species (AmphibiWeb, 2017). This genus is notble for contining most of the smllest cophyline frogs. These truly miniturized frogs number mong the smllest vertebrtes in the world (Glw & Vences, 2007; Klges et l., 2013; but see lso Scherz et l., 2016 for mention of Stumpffilike gener tht re lso extremely smll but remin to be txonomiclly ddressed) nd re n importnt but underpprecited ecologicl guild (Rittmeyer et l., 2012). Mny Stumpffi show some degree of reduction of digits on fingers nd toes (Glw & Vences, 1994) which hve been flgged s possible txonomic chrcters (Klges et l., 2013). The mode of reproduction is known only for few members of the genus; these ly eggs in fom nests in 274

5 VERTEBRATE ZOOLOGY 67 (3) 2017 the lef litter, where non-feeding tdpoles develop (Glw & Vences, 1994; Klges et l., 2013). The high proportion of cryptic diversity nd the incresing threts fcing Mlgsy frogs demnd the prioritiztion of txonomic revisions (Ndrintso et l., 2015). These mphibins re prime exmple of vertebrtes in which trditionl txonomic prctice my not be ble to ctch up with the rte of cndidte species detection. While uncommon in vertebrtes, such sitution is the rule in invertebrtes, especilly tropicl insects, most groups of which re extremely species-rich. To tckle this diversity, vrious prctices of fst-trck txonomy hve been proposed. At the core of these is the routine use of moleculr screening techniques (e.g., Hebert et l., 2003; Rtnsinghm & Hebert 2007; 2013) but lso focus on moleculr dignoses nd concise nd short morphologicl descriptions (Riedel et l., 2013; Renner et l., 2016), nd on online journls specificlly dpted to fst species descriptions (Smith et l., 2013). Such fst-trck txonomy hs the potentil drwbck of undetected errors: depending on the methods dopted, species my be overlooked, or synonyms produced (Mirlles & Vences, 2013), nd n integrtive pproch using vriety of pproches nd dt sets is most suitble for fst elbortion of species hypotheses without compromising their qulity (Dyrt, 2005; Pdil et l., 2009; 2010; Mirlles et l., 2011). In the present pper, using dt from our own previous studies s well s newly collected mteril, we present complete revision of the genus Stumpffi nd describe 26 new species within this genus. We follow simplified pproch, wherein we present much less morphologicl detil on ech species thn in previous studies (e.g., Klges et l., 2013), nd insted bse our species delimittion on integrting evidence from externl morphology, nturl history, DNA brcoding, congruence between mitochondril nd nucler DNA differentition, nd biocoustic nlyses. We first identify clusters bsed on genetic distnces, then discuss the genetic, morphologicl nd biocoustic evidence for their distinct txonomic sttus, nd proceed with the forml description of the 26 new species. Mterils nd Methods Specimens were collected t night or dy by serching in the lef litter guided by the clling of mles nd through opportunistic serches. The serch ws conducted using torches nd hedlmps t night. Specimens were euthnized in MS-222 solution, fixed in 90% ethnol nd preserved in 70% ethnol. Vouchers were deposited in the Zoologicl Collection of the Kyoto University Museum, Jpn (KUZA), Uni versité d Antnnrivo, Mention Zoologie et Biodiversité Animle, Antnnrivo, Mdgscr (UADBA), Zoologisches Forschungsmuseum Alexnder Koenig, Bonn, Germny (ZFMK), Museum für Nturkunde, Berlin, Germny (ZMB), the Zoologische Sttssmmlung München, Munich, Germny (ZSM), nd the Museo Regionle di Scienze Nturli, Torino, Turin, Itly (MRSN). Additionl voucher specimens were exmined from the N turhistorisches Museum Bern, Bern, Switzerlnd (NMBE), Muséum Ntionl d Histoire Nturelle, Pris, Frnce (MNHN), Senckenberg Museum Frnkfurt, Frnk furt, Germny (SMF), nd the Zoölogisch Museum Am ster dm (ZMA), Netherlnds (collections currently integrted in the Nturlis Biodiversity Center of Leiden). MNCN is used to refer to the Museo Ncionl de Ciencis Nturles in Mdrid, Spin. ACZCV, FAZC nd FN, FG/MV, FGZC, DRV, NSH, JCR, RDR, nd ZCMV refer to A. Crottini, F. Andreone, F. Glw, D.R. Vieites, S.H. Ndrin tso, J.C. Riemnn, R.D. Rn dri niin, nd M. Ven ces field numbers, respectively. In some cses, specimens deposited in UADBA hve not received finl ctlogue numbers yet; these re cited s UADBA followed by the respective field number in prentheses. Tissue smples were tken by cutting pieces of leg muscle, tongue, or whole limbs (depending on specimen size) from the euthnized nimls, nd preserved seprtely in 99% ethnol. Mles nd femles were distinguished bsed on field observtions (clling behvior) or presence of vocl sc in mles, or eggs in femles. Geogrphic regions re nmed ccording to Boumns et l. (2007) nd Brown et l. (2016); we follow Brown et l. (2016) in defining northern Mdgscr s n re roughly delimited by digonl spnning from 15.5 S on the est cost to c S on the west cost. Morphologicl mesurements nd descriptions The following morphologicl mesurements on preserved specimens were tken by the first uthor with digitl cllipers to the nerest 0.1 mm (Fig. 1): snout-vent length (SVL), mximum hed width (HW), hed length (HL), horizontl tympnum dimeter (TD), horizontl eye dimeter (ED), eye-nostril distnce (END), nostrilsnout tip distnce (NSD), nostril-nostril distnce (NND), forelimb length (FORL), hnd length (HAL), hindlimb length (HIL), foot length including trsus (FOTL), foot length (FOL), nd tibi length (TIBL). Terminology nd description scheme follow Vences et l. (2010) nd Glw et l. (2012). Body prts used for the description of color nd pttern re s in Fig. 2. Description of color in life is generlly bsed on single specimen, preferbly the holotype when photos of it were vilble, with some reference to vrition if known. In some cses where color or pttern is especilly vrible, totl-vrition description is given insted, s the pttern of individul specimens is less informtive. Biocoustics Advertisement clls were recorded in the field using different digitl or nlogue devices such s: Sony WM- 275

6 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Fig. 1. Reduced morphologicl scheme indicting the mesurements used herein. Abbrevitions re given in the text. FORL nd HIL re not shown. Fig. 2. Grphic scheme indicting regions of the body of Stumpffi specimens, s referred to in the descriptions of colortion throughout the mnuscript. Lbels nd colors refer to regions of the body nd not necessrily to ntomicl fetures, nd re used to simplify the description scheme of color ptterns. 276

7 VERTEBRATE ZOOLOGY 67 (3) 2017 D6C nd Tensi RCR-3222 tpe recorders with externl microphones (Sennheiser Me-80, Vivnco EM 238), nd Tscm DR07, DR05, Mrntz PMD 660, PMD 661 MkII, or Rolnd EDIROL R-09 digitl recorders, with built-in microphones (Tscm) or ccessorized with semi-directionl (Mrntz nd Rolnd) or supercrdioid (Sennheiser) microphone. Digitl recordings were obtined t smpling rte of 44.1 khz nd 24-bit resolution nd sved s uncompressed files. A number of recordings were done with dmged Tscm DR07 recorder nd contin n rtificil initil high mplitude pek nd possibly other rtifcts; where used, these recordings re clerly indicted s such in the text. Recordings were smpled or resmpled t khz nd 16-bit resolution nd computer-nlyzed using the softwre CoolEdit Pro version Frequency informtion ws obtined through Fst Fourier Trnsformtion (FFT, width 1024 points); the udiospectrogrms were obtined t Hnning window function with 256 bnds resolution using the seewve pckge (Sueur et l., 2008) in R (R Development Core Tem, 2015). Amplitude in spectrogrms is color-coded with reltive db vlues referring to the loudest sound in the recording. Temporl mesurements re given in milliseconds (ms) or seconds (s), s rnge, with men ± stndrd devition in prentheses. Terminology in cll descriptions follows the cll-centered terminology of Köhler et l. (2017). All cll recordings used for nlysis were deposited in the Animl Sound Archive (Tierstimmenrchiv) t the Museum für Nturkunde in Berlin (ccession numbers Ven1_1 Ven1_64). We here lso report dt from survey of clling ctivity in S. nlmin, conducted t dry deciduous forest of Ampijoro, Ankrfntsik Ntionl Prk (S16.3, E46.816). The nnul precipittion of the region is between 1000 nd 1500 mm. The rinfll occurs from November through April (riny seson), without wter bodies t the study site. Men ir temperture nd precipittion were obtined from the Durrell Wildlife Chelonin Breeding Center in Ampijoro sttion. Cll ctivity ws recorded in the forest every night from 2 December 2010 to 7 Februry 2011 nd 13 Jnury to 12 Februry The recordings were obtined t 07:00, 12:00, 17:00, nd 21:00 ech dy, nd were identified mong one of the following six level ctegories: level 0: no vocliztions were herd; level 1: single or few clls were herd, but did not form chorus; level 2: clls or choruses were herd intermittently, but in most of the time no vocliztions occurred; level 3: clls or choruses were frequently herd, but silent intervls were still obvious; level 4: intensive choruses were herd during most of the census, but severl occsionl cesstion of the chorus ws recognized; level 5: choruses were intensive nd never cesed. Moleculr dt sets We ssembled three different but prtly overlpping dt sets for the purpose of ssessing the phylogenetic reltionships mong species of Stumpffi, nd delimiting species. Our rtionle (reiterted in more detil below) is to first delimit species by congruence pproch of integrtive txonomy, i.e., finding evolutionry entities identifible by t lest two, if possible more, independent lines of evidence. For this purpose we seprtely nlyze dt sets of one mitochondril nd one nucler gene nd integrte these with further evidence from biocoustics nd morphology. We then infer the phylogeny mong the delimited species-level entities by conctenting multiple mitochondril nd one nucler gene from single representtives of ech unit. Moleculr dtset A. As mitochondril mrker we used 5 segment of the 16S rrna gene (16S) tht we hve previously used to ssess mitochondril differentition in microhylids, including Stumpffi (e.g., Vences et l., 2010; Köhler et l., 2010; Klges et l., 2013; Rkotorison et l., 2015). This segment differs from the one trditionlly used for DNA brcoding cross ll Mlgsy frogs (e.g., Vieites et l., 2009) but ws chosen to combine newly generted dt with the lrge sequence set of this gene from the work of Klges et l. (2013). We present these dt s phylogenetic tree from simplified nlysis (i.e., mximum likelihood without exclusion of poorly ligned dt) but emphsize tht the gol of this tree is to visulize clustering of geneticlly similr individuls nd their differentition from other individuls, nd not n ccurte reconstruction of the Stumpffi phylogeny. Altogether, 281 sequences of Stumpffi were vilble for this segment (length of ligned sequences: 657 bp), nd Anilny helene (Vlln, 2000) ws used s the outgroup (Scherz et l., 2016). Moleculr dtset B. As nucler mrker, we selected segment of the single-copy protein-coding nucler recombintion-ctivting gene 1 (Rg-1), which is known to often show distinct hplotypes for closely relted species, including Stumpffi (e.g., Klges et l., 2013; Ndrintso et l., 2013). We used nested primer pproch (Rkotorison et l., 2015) to mplify rther long stretch of this gene but trimmed it to 338 bp to llow nlysis together with the extensive set of sequences vilble from the work of Klges et l. (2013). We represented vrition mong the sequences of totl of 163 specimens of Stumpffi s hplotype network (fter seprting lleles by phsing), which in this cse is pproprite due to the smll number of muttions mong species nd the reltively high llele vrition within species. Moleculr dtset C. To resolve the phylogenetic reltionships mong species-level units, we conctented four mitochondril nd one nucler gene segments: two segments of 16S, nd one segment ech of the 12S rrna (12S), cytochrome oxidse subunit 1 (Cox-1) nd Rg-1. A prtitioned Byesin Inference nlysis ws crried out fter removing hypervrible stretches of 12S nd 16S, nd using Scphiophryne s outgroup. This dtset included 49 terminls (of which 48 Stumpffi) for totl of 3325 bp. For series of new specimens collected in November 2016 in the Mrojejy mssif, we sequenced 250 bp 277

8 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs highly vrible segment of 16S by n Illumin mplicon pproch (Vences et l., 2016) to identify them to species level; these short sequences were from different segment of the 16S gene nd re therefore not included in dtset A. DNA sequence nlysis We extrcted genomic DNA from muscle tissue smples preserved in 99% ethnol using stndrd slt extrction protocol (Bruford et l., 1992). All PCR protocols followed Rkotorison et l. (2015). Stndrd PCRs were crried out to mplify the mitochondril DNA segments wheres the Rg-1 segment ws mplified using nested PCR pproch. Primer sequences, primer references, nd cycling protocols re found in Rkotorison et l. (2015). PCR products were clened with enzymtic purifiction: 0.15 units of Shrimp Alkline Phosphtse (SAP) nd 1 unit of Exonuclese I (New Englnd Biolbs, Frnkfurt m Min, Germny) incubted for 15 min t 37 C followed by 15 min t 80 C. Purified PCR products were sequenced on n utomted DNA sequencer (Applied Biosystems ABI 3130XL). Sequencing rection (10 µl) contined 0.2 or 0.3 µl of PCR product, 0.5 µl of BigDye 3.1 (Applied Biosystems, Drmstdt, Germny) nd 0.3 µmol of primer. The mitochondril segment ws sequenced using the forwrd primer while Rg-1 ws sequenced in both directions to llow for more relible identifiction of heterozygote sites. Sequences were checked nd edited, nd heterozygous positions in Rg-1 inferred, in the softwre CodonCode Aligner (Codon Code Corportion, Dedhm, MA, USA). All 167 newly determined sequences were submitted to GenBnk (ccession numbers MF MF768252). Sequences were ligned with those from previous studies in MEGA 6 (Tmur et l., 2013). For dtset A we inferred phylogenetic tree under the Mximum Likelihood optimlity criterion in MEGA 6, under substitution model suggested by the Byesin Informtion Criterion in jmodeltest (Drrib et l., 2012). For dtset B we first trimmed ll sequences to equl length nd removed sequences contining mbiguities tht could not be interpreted s heterozygotes. We then seprted sequences into hplotypes using the Phse lgorithm (Stephens et l., 2001) s implemented in DNASp 5 (Librdo & Rozs, 2009). Phsed sequences were subsequently used to construct hplotype network following the pproch of Slzburger et l. (2011) with the progrm Hplotype Viewer. ( t/~greg/hploviewer) bsed on Neighbour-joining tree computed with MEGA 6 from uncorrected p-distnces. For dtset C we first conctented DNA sequences of four mitochondril nd one nucler gene segments: two non-overlppping segments of 16S, nd one segment ech of 12S, Cox-1, nd Rg-1. We then determined the bestfitting prtition scheme nd substitution models with PrtitionFinder (Lnfer et l., 2012), defining ech codon position of the two protein-coding genes s seprte chrcter sets, nd the whole 12S/16S segments s one further chrcter set. We removed ll gpped nd hypervrible regions of the 12S nd 16S segments fter visul inspection becuse usge of softwre for this tsk ws complicted by the different sequence lengths nd missing dt for some of the species nd segments. We then computed phylogenetic tree by Byesin inference with MrByes 3.2 (Ronquist et l., 2012). Results of two independent runs of 20 million genertions, ech comprising four Mrkov Chins (three heted nd one cold), were smpled every 10,000 genertions. Chin mixing nd sttionrity ws ssessed by exmining the stndrd devition of split frequencies nd by plotting the -lnl per genertion using Trcer 1.5 softwre (Rmbut & Drummond, 2007). Results were combined to obtin 50%-mjority rule consensus tree nd the respective posterior probbilities of nodes, fter discrding 25% of the genertions s burn-in (ll comptible nodes with probbilities < 0.5 were lso kept). Rtionle for species delimittion We here follow n pproch of integrtion by congruence s defined by Pdil et l. (2010), i.e., we define species s independent evolutionry lineges in the frmework of the generl linege or evolutionry species concept (Myden, 1997; De Queiroz, 1997; 2007) if two or more independent lines of evidence support their distinctness. We strt the species delimittion procedure by seeking mitochondril cldes divergent from other mitochondril cldes by sequence divergences > 3% in the 16S gene 5 frgment, given tht this degree of divergence hs been found to often correspond to species-level units in nurns (Fouquet et l., 2007; Mlone & Fonteno, 2008; Vieites et l., 2009). As cophyline microhylids, in generl re chrcterized by quite high intr- nd interspecific divergences (e.g., Vences et l., 2010; Klges et l., 2013), nd s this segment is slightly more vrible thn the 3 segment used for defining cndidte species in Mlgsy frogs (Vieites et l., 2009), we elevted this threshold to 4% uncorrected pirwise sequence divergence (p-distnce), with some exceptions explined below. To identify lineges t this threshold we used the SpeciesIdentifier Cluster lgorithm in Txon DNA 1.7 (Meier et l., 2006) to utomticlly cluster tx ccording to pirwise distnces for every sequence within ech cluster, wherein the mximum pirwise distnce within ech cluster (possible species-level linege) should not exceed given threshold. We subsequently tested if the individuls belonging to these mitochondril lineges re lso seprted in nucler gene (Rg-1) given the importnt differences in inheritnce between the nucler nd mitochondril genomes, nd the bsence of ny recombintion between mitochondril nd nucler genes. Genelogicl concordnce between such lrgely independent genetic mrkers hs been long recognized s n importnt species criterion (Avise & Bll, 1990), nd cn be highly inform- 278

9 VERTEBRATE ZOOLOGY 67 (3) 2017 tive even in the bsence of monophyly (Weisrock et l., 2010), i.e., by strongly different llele frequencies or unique hplotypes not forming clde. Such concordnce between nudna nd mtdna, especilly, cn help to conclusively refute the possibility of mitochondril introgression influencing the species delimittion process. Furthermore, we exmined externl morphology of voucher specimens, using especilly body size, digitl reduction nd specific distinctive color pttern s dignostic chrcters. To compre the highly distinctive dvertisement clls of Stumpffi, we considered in decresing order of importce (1) generl cll structure, i.e., pulsed vs. unpulsed, (2) cll durtion, nd (3) inter-cll intervl. For the lst two chrcters, we considered biocoustic difference s conclusive if, with comprble recording temperture, the respective vlues of two lineges do not overlp. Hence in our species delimittion pproch, we consider groups of individuls s species if they (1) form monophyletic group bsed on mtdna nd (2) differ from other such groups by > 4% sequence divergence (men p-distnce), nd fulfil t lest one of the following dditionl criteri: (3) exclusively privte (unique) hplotypes in the nlysed Rg-1 segment, (3b) biocousticlly distinct from other Stumpffi, (3c) morphologiclly nd/or chromticlly distinct from other Stumpffi. For some species with incomplete dt or smll smple sizes, we restrict the comprisons to their respective sister species, or relted species belonging to n inclusive subclde, nd consider the criteri of unique Rg-1 hplotypes nd morphologicl nd biocoustic distinctness to be stisfied when they pply to this set of reltives rther thn to ll other Stumpffi species. In few cses we furthermore tke number of d hoc txonomic decisions deviting from this species delimittion rtionle. This concerns few species tht we describe despite low smple size or missing dt, if they re chrcterized by extremely high mitochondril divergences nd unclrified reltionships to other Stumpffi. In severl other cses, we llow for substntil mitochondril divergence within our species units to void describing poorly defined species from well-defined mitochondril cldes, thereby leving these probble species complexes for further study. In this we follow the rgument of Mirlles & Vences (2013) who suggested tht flse negtive (filure to detect nd describe species) cn more esily be corrected by future reserchers thn flse positive (wrongly describing n intrspecific linege s species). Results Moleculr species delimittion Mximum Likelihood (ML) nlysis of DNA sequences (657 bp ligned) of the 5 -segment of the mitochondril 16S rrna gene for 281 individuls of Stumpffi yielded tree with cler pttern of clusters of identicl or neridenticl sequences, seprted by long brnches (nd thus high genetic distnces) from other such clusters (Fig. 3). The nlysis with SpeciesIdentifier t the 4% threshold level for uncorrected p-distnces yielded 53 clusters, not counting the outgroup nd short sequences with indequte sequence overlp. All nominl species of the genus were included in the nlysis, ll (except S. miery nd S. tridctyl) with sequences of multiple individuls. No cse of two or more nominl species of Stumpffi lumped in the sme cluster ws detected. Anticipting the results of our species delimittion procedure (specified in detil below), only in one cse were units described s new species lthough they re lumped into one cluster (the symptric S. ngeluci sp. nov. nd S. mledict sp. nov.). On the contrry, SpeciesIdentifier suggested ssignment of individuls to distinct clusters in numerous cses, which we herein consider, in conservtive pproch, deep conspecific lineges (DCLs). Species including dditionl DCLs side from the min species linege were S. nlnjirofo sp. nov. (1 extr DCL), S. gimmeli (2), S. kibomen (1), S. mdgscriensis (1), S. obscoen sp. nov. (1), S. roseifemorlis (1), S. sp. C30 (1). In ll these cses, the mitochondril dt suggest the possibility of dditionl undescribed species hidden under these tx, but we refrin from describing them due to insufficient informtion s discussed in the respective species ccounts below. The Rg-1 hplotype network reconstructed from DNA sequence segment of 338 bp from 163 specimens presented high mount of vrition with totl of 146 hplotypes, including lrge number of singletons (Fig. 4). This vrition ws cused by n overll high number of mbiguities in the chromtogrms interpreted by the phsing lgorithm s heterozygotes. While in mny cses these positions refer to true heterozyotes, others might hve different underlying cuses but were in mny cses confirmed s mbiguous in mplifiction nd sequencing repets. Distinct nd unmbiguous muttions seprted numerous hplotype groups in the network, which in most cses corresponded to lineges identified lso by the mitochondril nlysis. In direct comprison to the mitochondril clustering, nd considering tht for five mitochondril clusters no Rg-1 sequence ws vilble, 26 of the mitochondril clusters were in unique hplogroups (originting from single brnch tht does not lso led to nother hplogroup), three hplogroups contined the representtives of 2 or 3 mitochondril lineges, nd the representtives of few other mitochondril lineges were distributed over vrious hplogroups, or grouped in non-monophyletic hplogroups. Mny of the hplogroups were seprted from ech other by substntil number of five or more muttions. Hplotype shring mong individuls of different mitochondril lineges ws observed in only three instnces: between the linege pirs S. prdus sp. nov./s. diutissim sp. nov., S. gimmeli/s. mmitik sp. nov., nd S. tetrdctyl/s. nigrorubr sp. nov. 279

10 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Tble 1. Between-group men distnce in percent (number of bse differences per site from verging over ll sequence pirs between groups, multiplied by 100) for species nd cndidte species of Stumpffi, bsed on 574 bp of the mitochondril 16S rrna gene (3 end of the gene). All mbiguous positions were removed for ech sequence pir. chillei nlmin nlnjirofo ngeluci be contumeli diutissim edmondsi fusc grrffoi gimmeli grndis hr ihrn jennoeli kibomen kibomen (2) chillei nlmin 7.8 nlnjirofo ngeluci be contumeli diutissim edmondsi fusc grrffoi gimmeli grndis hr ihrn jennoeli kibomen kibomen (2) lrinki mdgsc riensis mdgsc riensis (2) mledict megsoni miery miovov nigrorubr obscoen prdus psologloss pygme roseifemorlis sort sp._c sp._c sp._c sp._c lrinki mdgscriensis mdgscriensis (2) mledict megsoni miery 280

11 VERTEBRATE ZOOLOGY 67 (3) 2017 Tble 1 continued. chillei nlmin nlnjirofo ngeluci be contumeli diutissim edmondsi fusc grrffoi gimmeli grndis hr ihrn jennoeli kibomen kibomen (2) lrinki mdgscriensis mdgscriensis (2) miery mledict megsoni dolchi huwei dvidttenboroughi sp._c spndei stffordi mmitik tetrdctyl tridctyl ynniki miovov nigrorubr obscoen prdus psologloss pygme roseifemorlis sort sp._ C11 sp._ C29 sp._ C30 sp._ C42 tridctyl miovov nigrorubr 9.9 obscoen prdus psologloss pygme roseifemorlis sort sp._c sp._c sp._c sp._c dolchi huwei dvidttenboroughi sp._c spndei stffordi mmitik tetrdctyl tridctyl ynniki sp._ C7 dolchi huwei dvidttenboroughi spndei stffordi mmitik tetrdctyl 281

12 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Fig. 3. Mximum Likelihood tree of individuls of Stumpffi bsed on segment of the 5 -segment of the mitochondril 16S rrna gene (totl lignment length 657 bp). Numbers t nodes re ML bootstrp vlues in percent (only shown if > 50%). Species nd cndidte species re shown in different colors. Inset photos re not to scle. 282

13 VERTEBRATE ZOOLOGY 67 (3) 2017 Fig. 3. Continued. 283

14 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Fig. 3. Continued. 284

15 VERTEBRATE ZOOLOGY 67 (3) 2017 Fig. 4. Hplotype network constructed from sequences of the nucler Rg-1 gene (338 bp, hplotypes inferred using the Phse lgorithm). Colors correspond to the grouping s in the mitochondril nlysis (Fig. 3). Smll dots represent unsmpled or extinct hplotypes; wheres brs represent muttionl steps. Stumpffi tridctyl, S. betmpon sp. nov., S. jennoeli sp. nov., S. dvidttenboroughi sp. nov. nd S. sort sp. nov. could not be sequenced for Rg-1 nd re therefore bsent from this network. Moleculr phylogeny The conctented lignment of 12S, the two 16S frgments, Cox-1, nd Rg-1, fter exclusion of hypervrible regions of 12S nd 16S, consisted of 3325 chrcters, of which 2315 were constnt, 1010 were vrible, nd 714 of these were prsimony-informtive. Prtitioned Byesin Inference nlysis resulted in well-resolved tree in which mny nodes received mximum posterior probbility (PP) support (Fig. 5). Four min cldes were distinguished, of which three received mximum support (PP = 1.0) nd one ws wekly supported (PP = 0.94). These cldes re here nmed A D. Clde A corresponded to previously recovered group tht hs been nmed the northern/northwestern clde (Köhler et l., 2010; Klges et l., 2013; Ndrintso et l., 2013) becuse most of the included species re exclusively distributed in these regions of Mdgscr. All nodes within this clde were resolved with PP > 0.95, nd the mjority of nodes received mximum support. The three subcldes splitting off the bslmost nodes within this clde re here denoted s subcldes A1 A3. Of these, subclde A1 contins S. gimmeli s well s the type species of the genus, S. psologloss, plus eight further deep lineges of which seven re herein described s new species. The species in this subclde re smll but not truly miniturized (see below for definition of size rnges in Stumpffi used herein). Subclde A2 contins the smllest species within clde A, with three described species (S. nlmin, S. mdgscriensis, nd S. pygme), one species newly described herein, nd three further cndidte species. Subclde A3 contins the four lrge-sized species described by Köhler et l. (2010), which probbly re specilized to krst environments nd t lest prtly re cve-dwellers. This subclde contins the Stumpffi species with the lrgest overll body sizes. Clde B is wekly supported ssemblge of morphologiclly nd biogeogrphiclly disprte species. It contins on the one hnd rther lrge-sized new species from high elevtions in the Tsrtnn Mountins (S. meikee sp. nov.), nd on the other hnd, three species from the Southern Centrl Est nd Northern Centrl Est tht re smll to miniturized nd show distinct to extreme digitl reduction. Clde C contins the bulk of remining species of Stumpffi, most of which re distributed in either estern or northestern Mdgscr (including the regions North-Est, Northern Centrl Est, nd Southern Centrl Est). This clde contins 4 previously described species, 14 new species described herein, nd 3 dditionl cndidte species. Reltionships within clde C in generl re poorly resolved, but to fcilitte further discussion we here nme two min subcldes C1 nd C2, both of which received mximum support. Clde C contins rther lrge-sized species (e.g., S. grndis) s well s numerous smll-sized species, some of which show moderte degree of digitl reduction (e.g., S. tetrdctyl). 285

16 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Fig. 5. Mjority-rule consensus tree from Byesin Inference nlysis of the conctented dtset of one nucler (Rg-1) nd four mitochondril (12S nd 16S rrna, Cox-1) gene frgments (3325 nucleotides). Mjor cldes denoted A D nd subcldes (A1 A3, C1 C2) re s discussed in the text. Clde A contins mostly species distributed in northern nd north-western Mdgscr wheres clde C mostly contins species from estern nd northestern Mdgscr. Numbers t nodes re posterior probbilities; not shown for vlues < 0.9. For ech species or cndidte species, up to three representtive collection loclities re mentioned. Inset mps summrize distribution records of ll species in ech of the four cldes. Clde D splits off the bslmost node within Stumpffi nd contins two miniturized species with strong digitl reduction tht re geneticlly highly divergent from other Stumpffi. The genus-level clssifiction of this clde will be ressessed in forthcoming publiction in the light of much-needed comprehensive screening of osteologicl vrition in Stumpffi. Morphologicl nd biocoustic comprisons Given the difficulties in exmining externl morphologicl fetures in these miniturized frogs, nd our gol to rech comprehensive review of the genus in resonble time frme, we focused on limited number of trits tht hd previously proven to be informtive in Stumpffi, nd prtly in other smll to miniturized frogs nd tht re esily reproducible: (1) body size, (2) externlly visible digitl reduction, (3) dorsl nd ventrl color, nd (4) dvertisement cll. In ddition we lso scored (5) reltive length of hnds nd feet, which in the pst hve been found to differ mong species of Stumpffi (Klges et l., 2013), nd (6) the texture of the dorsl skin. Altogether, our result suggest tht the mjority of moleculr lineges of Stumpffi cn be relibly distinguished from ech other by combintion of these six chrcters, lthough some lineges (especilly those with missing dt on biocoustics) were indistinguishble from few others. However, often the morphologiclly most similr lineges were not direct phylogenetic sister lineges, thus not jeoprdizing our txonomic conclusion of ssigning species sttus to them. Furthermore, such lineges tht 286

17 VERTEBRATE ZOOLOGY 67 (3) 2017 Fig. 6. Comprtive view of ventrl surfces of hnds of preserved specimens of smll-sized Stumpffi species of Clde B, C nd D. Not to scle. Fig. 7. Comprtive view of ventrl surfces of hnds of preserved specimens of miniturized/smll-sized species of Clde A. Not to scle. 287

18 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Fig. 8. Comprtive view of ventrl surfces of hnds of preserved specimens of lrge-sized species of Clde B nd C. Not to scle. Fig. 9. Comprtive view of ventrl surfces of feet of preserved specimens of smll-sized species of Clde B, C nd D. Not to scle. 288

19 VERTEBRATE ZOOLOGY 67 (3) 2017 Fig. 10. Comprtive view of ventrl surfces of feet of miniturized/smll-sized species of Clde A. Not to scle. Fig. 11. Comprtive view of ventrl surfces of feet of lrge-sized species of Clde B nd C. Not to scle. 289

20 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Fig. 12. Dorsl nd ventrl views of preserved nme-bering type specimens of newly decribed Stumpffi species. Scle brs equl 5 mm. 290

21 VERTEBRATE ZOOLOGY 67 (3) 2017 Fig. 12. Continued. cnnot be morphologiclly distinguished typiclly do not occur in close geogrphicl proximity to one-nother, mening tht field identifiction is unlikely to be hmpered by this morphologicl similrity. We found body size (Fig. 12) useful mens to define mjor clusters of Stumpffi species. These clusters often corresponded to cldes in the phylogeny, but in some cses species of very different body sizes were grouped in the sme clde. Body sizes rnge from 8 9 mm SVL in dult mles of S. contumeli sp. nov. to mximum of 28 mm in S. stffordi. For the purpose of this study nd restricting our perspective to this genus lone, we define modertely to lrge-sized Stumpffi s those reching regulrly more thn 17 mm snout-vent length. Species tht typiclly rech mm SVL re referred to s smll-sized, nd species below 12 mm re considered s miniturized. This ctegoriztion is however not pplied very strictly becuse there re species rnging inbetween the ctegories nd exceptionlly sized individuls of some species tht we hve disregrded for the ctegories. Our ctegoriztion is used primrily to fcilitte comprisons mong Stumpffi, nd differs from previous work tht defined, for frogs overll, miniture frogs s those 20 mm (Clrke, 1996), or considering species < 14 mm s extremely miniturized (Trueb & Alberch, 1985). In clde A, there is cler distinction of subcldes in body size, with A1 contining mostly smll species, A2 consisting of smll to miniturized species, nd A3 continining moderte-sized to lrge species. Clde D contins only miniturized species, clde C contins smll to moderte/lrge-sized species nd clde B shows lrge vrition despite few included species, from miniturized nd smll species to one lrge-sized species. In Stumpffi, miniturized species < 12 mm SVL re lso those tht most commonly show extreme degrees of digitl reduction (Fig. 6 11). Stumpffi tridctyl nd two new species described herein, hve the most strongly reduced condition, with only one finger nd three toes recognizble nd represent the smllest body sizes in Stumpffi (S. contumeli sp. nov., probble clling mles 8 9 mm). However, digitl reduction is not strictly linked to body size, nd is distinctly expressed in the four mjor cldes. For instnce, species of clde A show only very limited degrees of digitl reduction even if miniturized (e.g., S. nlmin, S. pygme nd S. ynniki sp. nov.) wheres in clde C, the mjority of species, even if lrge-sized, show distinct length reduction t lest of the first toe nd strong or complete reduction of this toe if smll-sized. This chrcter thus llows quick distinction of ll smll-sized species of clde A from those in clde C, while the rther strong reduction pttern t the hnd in the smll-sized species of clde B (S. miery nd S. dvidttenboroughi sp. nov.) llows distinguishing these from both clde A nd clde C species. Aside from these morphologicl chrcters, the dvertisement cll is the most obvious chrcter to dignose species of Stumpffi. Unlike body size nd digitl reduction, the dvertisement cll often differs strongly between closely relted lineges nd direct sister line- 291

22 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Tble 2. Loclity records of Stumpffi species with geogrphicl coordintes. Loclity Coordintes Altitude (m.s.l.) Species 27 km from Ambnj (northest of Ambnj towrds Ambilobe) Stumpffi gimmeli 27 km from Antsohihy S , E Stumpffi nlmin Ambnj S13.670, E Stumpffi gimmeli Ambtolhidimy S , E Stumpffi nigrorubr sp. nov. Ambtorom (cmpsite in the Mnompn / Befnjn forest re) S16.66, E49.59 Stumpffi nlnjirofo sp. nov. Ambodivongy (close to the Mkir Reserve; sometimes spelled Ambodivohngy) S , E c Stumpffi contumeli sp. nov. Stumpffi prdus sp. nov. Stumpffi fusc sp. nov. Stumpffi sp. C57 Ambohitsr S , E Stumpffi miery S , E Stumpffi nigrorubr sp. nov. Ambolo forest frgment S , E Stumpffi miery S , E Stumpffi miery Ambolokoptrik (sometimes spelled Ambolokoptriky) c. S14.52, E Stumpffi sp. C11 Ampombofofo S , E Stumpffi megsoni An Al forest S , E c. 850 Stumpffi kibomen Annitehn S , E Stumpffi nigrorubr sp. nov. Andlngin S , E Stumpffi miery S , E Stumpffi nigrorubr sp. nov. Andp c. S14.66, E49.66 Stumpffi mmitik sp. nov. Andsibe (loclities typiclly in Anlmzotr/Mntdi Ntionl Prk or in Anlmzotr Forest Reserve) S , E Angorony forest frgment S , E Stumpffi obscoen sp. nov. Stumpffi kibomen Stumpffi edmondsi sp. nov. Stumpffi psologloss Stumpffi gimmeli Stumpffi sp. C30 Ankrfntsik Ntionl Prk c. S16.30, E46.81 Stumpffi nlmin Stumpffi lrinki sp. nov. Ankrn Ntionl Prk S , E Stumpffi mmitik sp. nov. S12.95, E Stumpffi be Antsirsir S , E Stumpffi gimmeli Antrmnnvn S , E Stumpffi nigrorubr sp. nov. Benvony S13.7, E Stumpffi psologloss 140 Stumpffi gimmeli Berr (forest in Shmlz Peninsul) S , E Stumpffi gimmeli Betmpon: Betkonn S , E Stumpffi prdus sp. nov. Betmpon: Mintimbto S , E Stumpffi jennoeli sp. nov. Betmpon: Rendrirendry: Piste Fotsimvo S , E Stumpffi jennoeli sp. nov. S , E Stumpffi grrffoi sp. nov. Betmpon: Shbefoz S , E Stumpffi jennoeli sp. nov. Betmpon: Shindrn S , E Stumpffi jennoeli sp. nov. S , E Stumpffi betmpon sp. nov. S , E Stumpffi dvidttenboroughi sp. nov. Betmpon: Shmbendrn S , E Stumpffi jennoeli sp. nov. S , E Stumpffi nlnjirofo sp. nov. S , E Stumpffi betmpon sp. nov. S , E Stumpffi betmpon sp. nov. Betmpon: Vohitsivln S , E Stumpffi jennoeli sp. nov. S , E Stumpffi nlnjirofo sp. nov. Bezvon c. S13.52, E49.83 c. 100 Stumpffi ihrn sp. nov. Fnmbn forest S , E Stumpffi ihrn sp. nov. Fierenn S , E Stumpffi sp. C42 Forest frgment between Belnn nd Antsohihy S , E Stumpffi ynniki sp. nov. Forêt d Ambre Specil Reserve, c. 5 km SW Skrmy S , E Stumpffi ngeluci sp. nov. Gllery forest SE Andrfinkon S , E Stumpffi ihrn sp. nov. Ifndin S , E Stumpffi miery Joffreville S , E Stumpffi ngeluci sp. nov. Mhso Forest S , E Stumpffi spndei sp. nov. Stumpffi grrffoi sp. nov. Mkir plteu, Fotsilnn source (within Mkir Nturl Prk) S , E Stumpffi mkir sp. nov. Mnombo Specil Reserve S , E Stumpffi nigrorubr sp. nov. Mnongrivo Specil Reserve S , E Stumpffi gimmeli 292

23 VERTEBRATE ZOOLOGY 67 (3) 2017 Tble 2 continued. Mrojejy Ntionl Prk: Cmp 0 S , E Mrojejy Ntionl Prk: Cmp Mntell S , E Mrojejy Ntionl Prk: Cmp Mrojeji S , E Mrojejy Ntionl Prk: Cmp Simpon S , E Mrojejy Ntionl Prk, high elevtion site S , E Mrojejy Ntionl Prk, high elevtion Pndnus site S , E Stumpffi diutissim sp. nov. Stumpffi sp. C7 Stumpffi roseifemorlis Stumpffi chillei sp. nov. Stumpffi diutissim sp. nov. Stumpffi chillei sp. nov. Stumpffi diutissim sp. nov. Stumpffi roseifemorlis Stumpffi grndis Stumpffi tridctyl Stumpffi sp. C7 Stumpffi tridctyl Stumpffi tridctyl Mromindr S , E Stumpffi gimmeli Melivinny (cmpsite in the Mnompn / Befnjn forest re) S16.66, E49.59 Stumpffi prdus sp. nov. Montgne d Ambre (loclities refer to Montgne d Ambre Ntionl Prk or directly djcent res) Montgne des Frnçis S , E Stumpffi mdgscriensis S , E c. 975 Stumpffi ngeluci sp. nov. S , E Stumpffi huwei sp. nov. S , E Stumpffi mledict sp. nov. S , E Stumpffi cf. mdgscriensis C25 S12.333, E Stumpffi ngeluci sp. nov. S , E Stumpffi stffordi Ner Ambodimndresy c. S , E c. 778 Stumpffi ihrn sp. nov. Nosy Be S , E Stumpffi psologloss S , E Stumpffi pygme Nosy Borh S , E Stumpffi tetrdctyl Nosy Hr S , E Stumpffi hr Nosy Mngbe Specil Reserve S15.5, E c Stumpffi dolchi sp. nov. Stumpffi nlnjirofo sp. nov. Nosy Skti S , E Stumpffi psologloss Rnomfn villge (villge ner Rnomfn Ntionl Prk; coordintes correspond to villge) S , E Stumpffi nigrorubr sp. nov. Shdikin forest S , E Stumpffi nigrorubr sp. nov. Sort Forest S , E Stumpffi sort sp. nov. Sort Forest S , E Stumpffi miovov sp. nov. Tmpolo (Fenerive Est, Tmtve) c. S , E c. 10 Stumpffi grrffoi sp. nov. Tsrtnn Strict Nture Reserve: Mnrikob Forest, Andmpy Cmpsite S , E Stumpffi gimmeli Tsrtnn Strict Nture Reserve: Antevilm-bzh (Cmp 1) S E Stumpffi sp. C29 Tsrtnn Strict Nture Reserve: Mtsbory Miky (Cmp 2; sometimes spelled Mtsborimiky or Mtsborymik) S , E Stumpffi meikee sp. nov. Stumpffi sp. C29 Vohemr c. S , Stumpffi mmitik sp. nov. Vohidrzn S18.95, E48.5 c Stumpffi edmondsi sp. nov. ges, thereby providing immedite evidence for their sttus s distinct species, if individuls with strongly divergent clls re encountered in symptry (Köhler et l., 2017). At the sme time, the clls of Stumpffi hve very conserved generl structure, consisting of single chirp or whistle repeted t regulr intervls, nd only in one cse (S. psologloss) exhibit distinctly pulsed note structure. Txonomic ccounts In this revision, we review ll of the nominl species of Stumpffi (i.e., the ones with vlid scientific nmes) nd describe totl of 26 new species. Becuse this implies totl of round 40 species, it is imprcticl to provide detiled differentil dignoses. Furthermore, not ll species cn be unmbiguously dignosed by morphology lone. We therefore provide comprtive tbles with detils of importnt biocoustic trits, comprtive pltes with views of hnd nd feet illustrting digitl reductions, nd tble with genetic distnces in the frgment of the 3 -terminus of the mitochondril 16S rrna gene (Tble 1). The im of this study is revision of Stumpffi sensu stricto. We therefore do not consider species of Anilny (A. helene, originlly described s Stumpffi helene) nd do not discuss in detil those cndidte species morphologiclly similr to Stumpffi but flling outside the min Stumpffi clde in moleculr phylogenetics (Scherz et l., 2016; 2017). The species ccounts re orgnized following first the phylogenetic subdivision of the genus into min cldes, 293

24 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Fig. 13. Distribution mp of Stumpffi species of clde A1. The mp shows the remining primry vegettion of Mdgscr ( green colors indicting rinforest, brown/ornge colors deciduous dry nd rid spiny forest. Fig. 15. Distribution mp of Stumpffi species of cldes A3 nd B. The mp shows the remining primry vegettion of Mdgscr ( green colors indicting rinforest, brown/ornge colors deciduous dry nd rid spiny forest. Fig. 14. Distribution mp of Stumpffi species of clde A2. The mp shows the remining primry vegettion of Mdgscr ( green colors indicting rinforest, brown/ornge colors deciduous dry nd rid spiny forest. nd subsequently re subdivided bsed on body size, to fcilitte comprisons. In ech series of ccounts, the nominl species re reviewed first, nd the new species described subsequently. The nominl species in ech cluster re listed lphbeticlly (except for the first cluster in which the type species S. psologloss is listed first). In the dignoses, we briefly provide for ech species short non-differentil dignosis bsed on seven chrcter sets: body size, digitl reduction, terminl phlnges of digits reltive hnd nd foot length, skin texture, color pttern, nd cll. The combintion of these chrcters llows the vst mjority of species to be distinguished from one nother. We then give very brief differentil dignosis. To void exceedingly long differentil dignoses, these were ssembled in the following structure: (1) Ech nominl species is compred only with those nominl species nd newly described species mentioned in preceding ccounts in this pper; i.e., the second species listed is compred only with the first, the third species listed is compred with the first nd the second, nd so on. (2) Ech newly described species is compred with ll nominl species, nd with the newly described species mentioned in preceding ccounts in this pper. Where morphologicl nd/or biocoustic differences re bsent or fint, we extend the respective differentil dignosis (nd thus the species justifiction) to include lso mitochondril nd nucler genetic divergences, nd especilly, the phylogenetic position of the respective species. Figures summrize the distribution of ll species of Stumpffi, wheres living specimens nd spectrogrms/oscillogrms of their clls re represented in Figs A set of representtive preserved voucher specimens of most species is shown s Online Appendix (numbered s Supplementry Figs. S91 S125). 1. Smll-sized or miniturized species of clde A Clde A is unmbiguously supported by different sets of moleculr mrkers nd different nlysis methods. This clde hs previously been nmed the northern/northwestern clde becuse ll of the contined species occur in this re of Mdgscr. Among nominl species of 294

25 VERTEBRATE ZOOLOGY 67 (3) 2017 Fig. 16. Distribution mp of Stumpffi species of clde C1. The mp shows the remining primry vegettion of Mdgscr ( green colors indicting rinforest, brown/ornge colors deciduous dry nd rid spiny forest. Fig. 17. Distribution mp of Stumpffi species of clde C2 nd remining species of clde C. The mp shows the remining primry vegettion of Mdgscr ( green colors indicting rinforest, reddish colors deciduous dry forest, brown/ ornge colors deciduous dry nd rid spiny forest. nd in the next section, the lrge-sized species (clde A3). Stumpffi psologloss Boettger, 1881 (Figures 19 nd S91) Fig. 18. Distribution mp of species of Stumpffi in clde D. The mp shows the remining primry vegettion of Mdgscr ( green colors indicting rinforest, brown/ornge colors deciduous dry nd rid spiny forest. Stumpffi, five smll-sized species nd four lrge-sized species belong to this clde. We here first provide ccounts for the smll-sized species (cldes A1 nd A2) Nme-bering type. One specimen in SMF collected by A. Stumpff on Nossi-Bé (= Nosy Be), ccording to ori ginl description (Boettger, 1881), thus representing the holotype. Boettger (1892) lists six specimens in the SMF collection. Mertens (1967) probbly considered these wrongly s syntypes nd designted SMF 7337 s lec to type; this specimen is here considered to correspond to the originl holotype becuse it fits very well with the holo type drwing in the originl description of Boettger (1881). Identity nd dignosis. Stumpffi psologloss is historiclly the first described species of Stumpffi nd the type species of the genus. Despite some uncertinties regrding the holotype/lectotype of the species, the species identity is cler becuse only two species occur on Nosy Be, nd in only one of them the typicl teddyber -shped 295

26 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Tble 3. Originl morphometric mesurements of representtive specimens of Stumpffi exmined. ND = not determined. Species Collection number-field number Sex SVL HW HL TD ED END NSD NND FORL HAL HIL FOTL FOL TIBL Stumpffi psologloss SMF 7337 ND ND ND 1.8 ND 2.8 ND ND Stumpffi psologloss ZSM 479/2000 (FG/MV ) ND Stumpffi psologloss ZSM 480/2000 (FG/MV ) ND Stumpffi psologloss ZSM 483/2000 (FG/MV ) Juvenile Stumpffi psologloss ZSM 484/2000 (FG/MV ) ND Stumpffi psologloss ZSM 485/2000 (FG/MV ) Juvenile ND ND ND ND Stumpffi psologloss ZFMK ND Stumpffi psologloss ZFMK ND Stumpffi psologloss ZFMK ND Stumpffi psologloss ZFMK ND Stumpffi psologloss ZFMK ND Stumpffi psologloss ZFMK ND Stumpffi psologloss ZFMK ND Stumpffi psologloss ZFMK ND Stumpffi psologloss ZFMK Juvenile Stumpffi psologloss ZFMK ND Stumpffi psologloss ZFMK ND Stumpffi psologloss ZFMK ND Stumpffi psologloss ZFMK ND Stumpffi psologloss ZFMK Juvenile Stumpffi psologloss ZFMK Juvenile ND Stumpffi psologloss ZFMK ND Stumpffi psologloss ZFMK Juvenile Stumpffi nlmin ZSM 542/2009 (ZCMV 11428) Mle ND ND 1.2 ND 2.6 ND ND Stumpffi nlmin KUZA _Ad_059 ND Stumpffi nlmin KUZA _F005 ND Stumpffi nlmin KUZA _F002 ND Stumpffi nlmin KUZA F004 ND ND ND 1.3 ND ND ND ND 2.7 ND Stumpffi nlmin KUZA F003 ND ND ND 1.3 ND ND ND ND 2.8 ND Stumpffi nlmin KUZA Ad-050 ND ND 1.6 ND ND ND Stumpffi nlmin KUZA Ad-049 ND ND 1.8 ND ND ND Stumpffi nlmin KUZA A (= 2013-NoCode) ND ND 1.5 ND ND ND Stumpffi nlmin KUZA Femle ND 1.5 ND ND ND Stumpffi nlmin KUZA Mle? ND 1.5 ND ND ND Stumpffi gimmeli ZSM 597/2001 (MV ) ND Stumpffi gimmeli ZSM 412/2000 (FG/MV ) ND Stumpffi gimmeli ZSM 833/2003 (FG/MV ) Mle Stumpffi mdgscriensis ZSM 3242/2012 (ZCMV 12185) ND ND Stumpffi mdgscriensis ZSM 202/2004 (FGZC 390) ND

27 VERTEBRATE ZOOLOGY 67 (3) 2017 Tble 3 continued. Species Collection number (Field number) Sex SVL HW HL TD ED END NSD NND FORL HAL HIL FOTL FOL TIBL Stumpffi mdgscriensis ZSM 201/2004 (FGZC 389) ND ND Stumpffi cf. mdgscriensis (sp. C25) ZSM 2108/2007 (FGZC 1103) ND Stumpffi cf. mdgscriensis (sp. C25) ZSM 2109/2007 (FGZC 1105) ND Stumpffi pygme ZSM 430/2010 ND ND Stumpffi pygme ZSM 431/2010 ND Stumpffi pygme ZFMK ND ND 1.2 ND ND ND Stumpffi pygme ZFMK ND Stumpffi pygme ZFMK ND Stumpffi pygme ZFMK ND Stumpffi pygme ZFMK ND ND 1.4 ND ND ND Stumpffi pygme ZFMK ND Stumpffi pygme ZFMK Femle ND 1.6 ND ND ND Stumpffi pygme ZFMK ND Stumpffi pygme ZFMK ND Stumpffi pygme ZFMK ND Stumpffi ngeluci sp. nov. ZSM 3241/2012 (ZCMV 12184) Mle Stumpffi ngeluci sp. nov. ZSM 3243/2012 (ZCMV 12197) Mle Stumpffi ngeluci sp. nov. ZSM 300/2004 (FGZC 580) Mle Stumpffi ngeluci sp. nov. ZSM 224/2004 (FGZC 434) Mle Stumpffi ngeluci sp. nov. ZSM 223/2004 (FGZC 433) Mle Stumpffi ngeluci sp. nov. ZSM 301/2004 (FGZC 581) Femle Stumpffi ngeluci sp. nov. ZSM 302/2004 (FGZC 583) ND Stumpffi ngeluci sp. nov. ZSM 303/2004 (FGZC 586) ND Stumpffi ngeluci sp. nov. ZSM 1671/2008 (FGZC 1901) ND Stumpffi huwei sp. nov. ZSM 221/2016 (ZCMV 13618) Mle Stumpffi huwei sp. nov. ZSM 904/2003 (FG/MV ) ND Stumpffi huwei sp. nov. UADBA-A (ZCMV 13619) ND Stumpffi huwei sp. nov. UADBA-A (ZCMV 13515) Mle Stumpffi huwei sp. nov. ZSM 3245/2012 (ZCMV 13514) Mle Stumpffi huwei sp. nov. ZSM 224/2016 (ZCMV 13611) Mle Stumpffi ihrn sp. nov. ZSM 1651/2012 (FGZC 3895) ND Stumpffi ihrn sp. nov. ZSM 1652/2012 (FGZC 3927) Femle Stumpffi ihrn sp. nov. ZSM 1642/2012 (FGZC 3800) ND Stumpffi ihrn sp. nov. ZSM 1650/2012 (FGZC 3893) ND Stumpffi lrinki sp. nov. ZSM 3236/2012 (ZCMV 13525) Mle Stumpffi lrinki sp. nov. ZSM 779/2003- (FG/MV ) ND Stumpffi lrinki sp. nov. ZSM 861/2003- (FG/MV ) ND Stumpffi lrinki sp. nov. ZSM 1669/2008 (FGZC 1620) ND Stumpffi mledict sp. nov. ZSM 2079/2007 (FGZC 1049) ND

28 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Tble 3 continued. Species Collection number (Field number) Sex SVL HW HL TD ED END NSD NND FORL HAL HIL FOTL FOL TIBL Stumpffi mledict sp. nov. ZSM 3244/2012 (ZCMV 13504) Mle Stumpffi mledict sp. nov. ZSM 2169/2007 (FGZC 1244) ND ND Stumpffi mmitik sp. nov. ZSM 3237/2012 (ZCMV 13524) Mle Stumpffi mmitik sp. nov. ZSM 3232/2012 (ZCMV 13528) Mle Stumpffi mmitik sp. nov. UADBA-A (orig. ZSM 3233/2012, ZCMV 13529) ND Stumpffi mmitik sp. nov. UADBA-A (orig. ZSM 3233/2012, ZCMV 13529) ND Stumpffi mmitik sp. nov. UADBA-A (orig. ZSM 3234/2012, ZCMV 13530) ND Stumpffi mmitik sp. nov. ZSM 862/2003 (FG/MV ) ND Stumpffi mmitik sp. nov. ZSM 375/2005 (FGZC 2725) ND Stumpffi sort sp. nov. ZSM 1644/2012 (FGZC 3621) ND Stumpffi sort sp. nov. ZSM 1645/2012 (FGZC 3622) ND Stumpffi sort sp. nov. ZSM 1643/2012 (FGZC 3618) ND Stumpffi ynniki sp. nov. ZSM 629/2014 (DRV 6413) ND Stumpffi ynniki sp. nov. ZSM 1825/2010 (ZCMV 12600) ND Stumpffi tridctyl MNHN ND ND ND 1.4 ND 1.5 ND Stumpffi tridctyl MNHN ND ND ND 1.3 ND 1.4 ND Stumpffi tridctyl ZSM 382/2005 (FGZC 2844) ND ND ND 1.5 ND Stumpffi tridctyl ZSM 527/2016 (ZCMV 15284) ND Stumpffi contumeli sp. nov. ZSM 443/2010 (FGZC 4258) Mle Stumpffi contumeli sp. nov. ZSM 441/2010 (FGZC 4248) ND Stumpffi contumeli sp. nov. ZSM 442/2010 (FGZC 4252) Mle Stumpffi dvidttenboroughi sp. nov. ZSM 204/2016 (ACZCV 106) ND Stumpffi meikee sp. nov. ZSM 1823/2010 (ZCMV 12374) Mle? ND 4.8 ND Stumpffi meikee sp. nov. ZSM 617/2014 (DRV 6163) Mle Stumpffi meikee sp. nov. ZSM 619/2014 (DRV 6171) Mle Stumpffi meikee sp. nov. ZSM 620/2014 (DRV 6175) Mle Stumpffi meikee sp. nov. ZSM 621/2014 (DRV 6177) Mle Stumpffi meikee sp. nov. ZSM 1821/2010 (ZCMV 12372) Mle Stumpffi meikee sp. nov. ZSM 1822/2010 (ZCMV 12373) Femle Stumpffi meikee sp. nov. ZSM 624/2014 (DRV 6185) Femle Stumpffi meikee sp. nov. ZSM 626/2014 (DRV 6189) Femle Stumpffi meikee sp. nov. ZSM 622/2014 (DRV 6179) Femle Stumpffi obscoen sp. nov. ZSM 381/2005 (FGZC 2666) ND Stumpffi obscoen sp. nov. ZFMK ND Stumpffi obscoen sp. nov. ZFMK ND Stumpffi obscoen sp. nov. ZFMK ND Stumpffi obscoen sp. nov. ZSM 380/2005 (FGZC 2664) ND Stumpffi roseifemorlis ZSM 373/2005 (FGZC 2808) ND Stumpffi roseifemorlis ZSM 374/2005 (FGZC 2883) ND

29 VERTEBRATE ZOOLOGY 67 (3) 2017 Tble 3 continued. Species Collection number (Field number) Sex SVL HW HL TD ED END NSD NND FORL HAL HIL FOTL FOL TIBL Stumpffi roseifemorlis ZSM 487/2005 (ZCMV 2047) ND Stumpffi roseifemorlis ZSM 0529/2016 (ZCMV 15172) Mle Stumpffi chillei sp. nov. ZSM 494/2005 (ZCMV 861) Femle Stumpffi chillei sp. nov. ZSM 493/2005 (ZCMV 860) ND Stumpffi chillei sp. nov. ZSM 377/2005 (FGZC 2750) ND Stumpffi chillei sp. nov. ZFMK ND Stumpffi chillei sp. nov. ZFMK ND Stumpffi chillei sp. nov. ZFMK Mle Stumpffi chillei sp. nov. ZSM 0534/2016 (ZCMV 15141) Mle Stumpffi chillei sp. nov. ZSM 0535/2016 (ZCMV 15143) ND Stumpffi chillei sp. nov. ZSM 0536/2016 (ZCMV 15149) Mle Stumpffi nlnjirofo sp. nov. ZSM 489/2005 (ZCMV 873) Femle Stumpffi nlnjirofo sp. nov. ZSM 492/2005 (ZCMV 2178) Femle Stumpffi nlnjirofo sp. nov. ZSM 491/2005 (ZCMV 2104) ND Stumpffi diutissim sp. nov. ZSM 378/2005 (FGZC 2751) Mle Stumpffi diutissim sp. nov. ZSM 376/2005 (FGZC 2742) ND Stumpffi diutissim sp. nov. ZSM 496/2005 (ZCMV 2082) ND Stumpffi diutissim sp. nov. ZSM 495/2005 (ZCMV 2067) ND Stumpffi diutissim sp. nov. ZSM 546/2016 (ZCMV 15064) ND Stumpffi diutissim sp. nov. ZSM 547/2016 (ZCMV 15117) ND Stumpffi diutissim sp. nov. ZSM 549/2016 (ZCMV 15144) Mle Stumpffi edmondsi sp. nov. ZSM 0371/2005 (FGZC 2677) ND Stumpffi edmondsi sp. nov. ZSM 1731/2012 (RDR 1065) Mle Stumpffi fusc sp. nov. ZSM 437/2010 (FGZC 4254) Femle Stumpffi fusc sp. nov. ZSM 436/2010 (FGZC 4253) ND Stumpffi jennoeli sp. nov. ZSM 206/2016 (ACZCV 56) Mle Stumpffi jennoeli sp. nov. MRSN A6386 (FAZC 13903) ND Stumpffi jennoeli sp. nov. ZSM 207/2016 (ACZCV 167) ND Stumpffi jennoeli sp. nov. ZSM 208/2016 (ACZCV 218) ND Stumpffi jennoeli sp. nov. MRSN A6283 (FAZC 13682) Mle Stumpffi nigrorubr sp. nov. ZSM 2448/2007 (ZCMV 5967) ND Stumpffi nigrorubr sp. nov. ZSM 640/2003 (FG/MV ) Mle Stumpffi nigrorubr sp. nov. ZSM 641/2003 (FG/MV ) Mle Stumpffi prdus sp. nov. ZSM 435/2010 (FGZC 4237) Mle Stumpffi prdus sp. nov. ZSM 230/2016 (ACZCV 0147) Mle Stumpffi prdus sp. nov. ZSM 439/2010 (FGZC 4268) Femle Stumpffi prdus sp. nov. MRSN A6276 (FAZC 13498) ND Stumpffi prdus sp. nov. ZSM 432/2010 (FGZC 4210) ND Stumpffi prdus sp. nov. ZSM 440/2010 (FGZC 4278) ND

30 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Tble 3 continued. Species Collection number (Field number) Sex SVL HW HL TD ED END NSD NND FORL HAL HIL FOTL FOL TIBL Stumpffi prdus sp. nov. ZSM 434/2010 (FGZC 4214) ND Stumpffi prdus sp. nov. ZSM 438/2010 (FGZC 4265) ND Stumpffi prdus sp. nov. ZSM 770/2009 (ZCMV 11166) ND Stumpffi prdus sp. nov. ZSM 433/2010 (FGZC 4213) ND Stumpffi tetrdctyl ZSM 594/2006 (ZCMV 3396) ND Stumpffi tetrdctyl ZSM 593/2006 (ZCMV 3395) ND Stumpffi betmpon sp. nov. ZSM 214/2016 (ACZCV 210) ND Stumpffi betmpon sp. nov. ZSM 217/2016 (ACZCV 46) ND Stumpffi betmpon sp. nov. ZSM 219/2016 (ACZCV 103) ND ND Stumpffi betmpon sp. nov. ZSM 218/2016 (ACZCV 47) ND Stumpffi betmpon sp. nov. ZSM 216/2016 (ACZCV 41) ND Stumpffi dolchi sp. nov. ZSM 488/2005 (ZCMV 2143) ND Stumpffi grrffoi sp. nov. ZSM 1752/2008 (ZCMV 8803) Mle Stumpffi grrffoi sp. nov. ZSM 220/2016 (ACZCV 6) ND Stumpffi grrffoi sp. nov. ZSM 632/2009 (ZCMV 8685) ND Stumpffi grrffoi sp. nov. ZSM 631/2009 (ZCMV 8684) ND Stumpffi grrffoi sp. nov. ZSM 633/2009 (ZCMV 8687) ND Stumpffi grrffoi sp. nov. MRSN A6278 (FAZC 13489) Mle Stumpffi mkir sp. nov. ZSM 541/2009 (ZCMV 11257) ND Stumpffi miovov sp. nov. ZSM 1649/2012 (FGZC 3656) Mle Stumpffi miovov sp. nov. ZSM 1640/2012 (FGZC 3650) Femle Stumpffi miovov sp. nov. ZSM 1647/2012 (FGZC 3675) Femle Stumpffi miovov sp. nov. UADBA-A (orig. ZSM 1648/2012, FGZC 3676) Femle Stumpffi miovov sp. nov. ZSM 1646/2012 (FGZC 3751) ND Stumpffi spndei sp. nov. ZSM 1751/2008 (ZCMV 8802) ND Stumpffi sp. C7 ZSM 379/2005 (FGZC 2826) ND Stumpffi sp. C7 ZSM 544/2016 (ZCMV 15181) Mle Stumpffi sp. C7 ZSM 545/2016 (ZCMV 15182) Mle Stumpffi sp. C29 ZSM 627/2014 (DRV 6103) Mle Stumpffi sp. C29 ZSM 1824/2010 (ZCMV 12387) Juvenile ND 1.2 ND ND ND Stumpffi sp. C30 ZSM 634/2014 (DRV 6453) ND Stumpffi sp. C30 ZSM 632/2014 (DRV 6451) ND Stumpffi sp. C30 ZSM 633/2014 (DRV 6452) ND Stumpffi sp. C30 ZSM 635/2014 (DRV 6454) ND

31 VERTEBRATE ZOOLOGY 67 (3) 2017 Tble 4. Body sizes, reltive hnd length, nd reltive length of foot (including trsus) of dults of Stumpffi species, given s rnge with men ± stndrd devition nd smple size in prentheses. See Tble 3 for originl vlues nd differences between sexes. ND = not determined. SVL HAL/SVL FOTL/SVL Stumpffi psologloss mm (12.84 ± 1.52, N = 17) (0.21 ± 0.02, N = 17) (0.70 ± 0.07, N = 16) Stumpffi nlmin mm (12.30 ± 0.79, N = 11) (0.22 ± 0.02, N = 11) (0.69 ± 0.06, N = 10) Stumpffi gimmeli mm (15.97 ± 1.89, N = 3) (0.21 ± 0.02, N = 3) (0.65 ± 0.02, N = 3) Stumpffi mdgscriensis mm (11.50 ± 1.33, N = 5) (0.17 ± 0.01, N = 5) (0.59 ± 0.03, N = 5) Stumpffi pygme mm (10.23 ± 1.17, N = 12) (0.16 ± 0.02, N = 12) (0.62 ± 0.08, N = 12) Stumpffi ngeluci sp. nov mm (13.49 ± 1.73, N = 9) (0.22 ± 0.02, N = 9) (0.69 ± 0.03, N = 9) Stumpffi huwei sp. nov mm (13.43 ± 1.14, N = 6) (0.19 ± 0.02, N = 6) (0.62 ± 0.04, N = 6) Stumpffi ihrn sp. nov mm (14.63 ± 0.65, N = 4) (0.21 ± 0.02, N = 4) (0.66 ± 0.07, N = 4) Stumpffi lrinki sp. nov mm (12.65 ± 0.13, N = 4) (0.23 ± 0.01, N = 4) (0.61 ± 0.09, N = 4) Stumpffi mledict sp. nov mm (14.93 ± 1.63, N = 3) (0.19 ± 0.01, N = 3) (0.65 ± 0.06, N = 3) Stumpffi mmitik sp. nov mm (13.40 ± 1.10, N = 7) (0.21 ± 0.03, N = 7) (0.68 ± 0.05, N = 7) Stumpffi sort sp. nov mm (15.80 ± 0.20, N = 3) (0.2 ± 0.02, N = 3) (0.64 ± 0.01, N = 3) Stumpffi ynniki sp. nov mm (10.20 ± 0.57, N = 2) (0.19 ± 0.00, N = 2) (0.64 ± 0.02, N = 2) Sumpffi be 25.2 mm (N = 1) 0.3 (N= 1) 0.6 (N= 1) Stumpffi hr mm (23.50 ± 1.50 mm, N = 2) (N = 2) (N = 2) Stumpffi megsoni mm (21.30 ± 0.50 mm, N = 2) (N = 2) 0.7 (N = 2) Stumpffi stffordi mm (27.40 ± 0.60 mm, N = 2) 0.3 (N = 2) 0.6 (N = 2) Stumpffi miery mm (14.00 ± 0.50 mm, N = 5) (N=4) (N=4) Stumpffi tridctyl mm (9.85 ± 0.90 mm, N = 4) (0.14 ± 0.05, N = 4) (0.57 ± 0.36, N = 4) Stumpffi contumeli sp. nov mm (8.47 ± 0.45 mm, N = 3) (0.17 ± 0.01, N = 3) (0.74 ± 0.08, N = 3) Stumpffi obscoen sp. nov mm (10.34 ± 0.57 mm, N = 5) (0.15 ± 0.01, N = 5) (0.58 ± 0.04, N = 5) Stumpffi dvidttenboroughi sp. nov mm (N = 1) 0.17 (N = 1) 0.55 (N = 1) Stumpffi meikee sp. nov mm (21.20 ± 1.06 mm, N = 10) (0.23 ± 0.01, N = 10) (0.72 ± 0.04, N = 10) Stumpffi grndis mm (21.60 ± 1.8 mm, N = 4) (N = 4) 0.8 (N = 4) Stumpffi kibomen mm ND ND Stumpffi roseifemorlis mm (17.67 ± 1.00, N = 4) (0.23 ± 0.59, N = 4) (0.68 ± 0.49, N = 4) Stumpffi chillei sp. nov mm (17.10 ± 1.44, N =9) (0.25 ± 0.2, N = (0.74 ± 0.4, N = 9) Stumpffi nlnjirofo sp. nov mm (18.80 ± 2.69, N = 3) (0.24 ± 0.00, N = 3) (0.71 ± 0.06, N = 3) Stumpffi diutissim sp. nov mm (16.48 ± 2.38, N = 8) (0.23 ± 0.21, N = 8) (0.69 ± 0.52, N = 8) Stumpffi edmondsi sp. nov mm (17.35 ± 0.07, N = 2) (0.25 ± 0.00, N = 2) (0.67 ± 0.08, N = 2) Stumpffi fusc sp. nov mm (16.05 ± 2.33, N = 2) (0.25 ± 0.02, N = 2) (0.75 ± 0.04, N = 2) Stumpffi jennoeli sp. nov mm (17.76 ± 1.37, N = 5) (0.21 ± 0.02, N = 5) (0.68 ± 0.07, N = 5) Stumpffi nigrorubr sp. nov mm (18.37 ± 1.42, N = 3) (0.23 ± 0.00, N = 3) (0.67 ± 0.07, N = 3) Stumpffi prdus sp. nov mm (17.63 ± 2.23, N = 10) (0.23 ± 0.03, N = 10) (0.75 ± 0.06, N = 10) Stumpffi tetrdctyl mm (14.00 ± 0.14, N = 2) (0.20 ± 0.01, N = 2) (0.59 ± 0.02, N = 2) Stumpffi betmpon sp. nov mm (12.82 ± 1.03, N = 5) (0.19 ± 0.02, N = 5) (0.69 ± 0.04, N = 5) Stumpffi dolchi sp. nov mm (N = 1) 0.18 (N = 1) 0.75 (N = 1) Stumpffi grrffoi sp. nov mm (13.90 ± 1.38, N = 6) (0.19 ± 0.03, N = 6) (0.68 ± 0.05, N = 6) Stumpffi mkir sp. nov mm (N = 1) 0.16 (N = 1) 0.64 (N = 1) Stumpffi miovov sp. nov mm (16.28 ± 1.38, N = 5) (0.19 ± 0.03, N = 5) (0.71 ± 0.06, N = 5) Stumpffi spndei sp. nov mm (N = 1) 0.21 (N = 1) 0.67 (N = 1) Stumpffi sp. C mm (15.16 ± 1.11, N = 3) (0.11 ± 3.3, N = 3) (0.3 ± 11.09, N = 3) Stumpffi sp. C mm (N = 1) 0.19 (N = 1) 0.65 (N = 1) Stumpffi sp. C mm (9.78 ± 0.22, N = 4) (0.19 ± 0.03, N = 4) (0.63 ± 0.10, N = 4) dorsl mrking occurs tht lso chrcterizes the S. psologloss holotype ccording to the originl description. Incidentlly, this species is lso dignosed from ll other nominl species of Stumpffi (s well s from ll cndidte species, s fr s known) by its unique cll consisting of pulsed trill note. (1) Miniturized to smll-sized species (dult SVL mm); (2) mnus with four fingers (first finger slightly reduced in length) nd pes with five toes (first toe slightly reduced in length); (3) terminl phlnges of fingers nd toes without clerly enlrged discs but very slightly enlrged nd distlly pointed terdrop-shped in some specimens; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum, especilly in life, often not completely smooth but slightly grnulr; (6) dorslly often brown with distinct drk brown ptches on the bck, sometimes with lrge centrl drk teddybershped mrking, nd with more or less complete drk brown bnds long the flnks; without contrsted ventrl colortion, red color elements on ventrl side; (7) unique cll mong Stumpffi, consisting of single regulrly repeted long nd strongly pulsed note. 301

32 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b c Fig. 19. Stumpffi psologloss from Nosy Be in life: () specimen for which no voucher number is known; (b c) ZSM 479/2000 (FG/ MV ). b Fig. 20. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi psologloss from its type loclity Nosy Be: () 1 s durtion section, (b) 6 s durtion section. Specimens exmined. Holotype SMF 7337 collected prior to 1881 on Nosy Be by A. Stumpff. Referred specimens: ZFMK collected in Mrch 1991 on Nosy Be by F. Glw nd M. Vences. ZFMK , , collect ed by F. Glw nd J. Müller in Jnury/Februry 1992 on Nosy Be. ZSM 479/2000 (FG/MV ), collected on 11 Mrch 2000, t estern edge of Lokobe reserve on Nosy Be (S , E , 110 m.s.l.) by F. Glw, K. Schmidt, nd M. Vences; ZSM 480/ /2000 (ll with field number FG/MV ), collected on 9 Mrch 2000 on Nosy Skti (S , E , < 10 m.s.l.) by F. Glw nd K. Schmidt. ZSM 616/2014 (DRV 6486), collected on 30 June 2010 in Mromindr forest frgment ner Ankrmy (S , E , 283 m.s.l.) by F.M. Rtsovin, S. Rsmison, T. Rjofirison, nd F. Rndrinsolo. Distribution. This species is known from (1) Nosy Be (type loclity), (2) Nosy Skti ( smll islnd west of Nosy Be), (3) Benvony, nd (4) Mromindr (two loclities on the Mdgscn minlnd) (Fig. 13). These four loclities re ll confirmed by moleculr genetics, with the minlnd popultions being only slightly differentited from the islnd popultions (Fig. 3). A fifth loclity, Nosy Komb (between Nosy Be nd minlnd Mdgscr) ws recently reported (Blumgrt et l., 2017) nd seems to be supported bsed on photogrph provided in tht pper, but specimens from this loclity hve not yet been studied from moleculr 302

33 VERTEBRATE ZOOLOGY 67 (3) 2017 Tble 5. Bsic temporl nd spectrl vribles of dvertisement clls of Stumpffi. Species Cll durtion Inter-cll intervl durtion Dominnt frequency Stumpffi psologloss ms ms Hz (831 ± 25 ms; N = 11) (2691 ± 860 ms; N = 10) (4955 ± 520 Hz) Stumpffi nlmin (cptive) ms ms Hz (260 ± 23.6 ms; N = 5) (2242 ± 136 ms; N = 4) (6210 ± 80.4 Hz; N = 5) Stumpffi nlmin (wild/ankrfntsik) ms (153 ± 13.5 ms; N = 10) ms (1120 ± 30 ms; N = 9) Hz (5399 ± 91 Hz, N = 10) Stumpffi gimmeli ms ms Hz (126.8 ± 30 ms; N = 14) ( ± ms; N = 14) ( ± Hz; N =146) Stumpffi mdgscriensis ms ms Hz (195 ± 5 ms; N = 5) (4224 ± 153 ms; N = 4) (4231 ± 197 Hz; N = 5) Stumpffi pygme ms ms Hz (196 ± 10 ms; N = 5) (3447 ± 419 ms; N = 4) (6054 ± 49 Hz; N = 5) Stumpffi ngeluci sp. nov ms ms Hz (182 ± 6 ms; N = 6) (3596 ± 254 ms; N = 5) (4651 ± 47 Hz; N = 6) Stumpffi huwei sp. nov ms ms Hz (70 ± 2.4 ms; N = 6) ( ± 267 ms; N = 6) (5020 ± 39 Hz; N = 6) Stumpffi lrinki sp. nov ms ms Hz (138 ± 21 ms; N = 3) (2216 ± 103 ms; N = 2) (2914 ± 124 Hz; N = 3) Stumpffi mledict sp. nov ms ms Hz (102 ± 2.8 ms; N = 4) (5927 ± 735 ms; N = 3) (4833 ± 21.5 Hz; N = 4) Stumpffi mmitik sp. nov ms ms Hz (103 ± 9 ms; N = 20) (955 ± 264 ms; N = 19) (4749 ± 257 Hz; N = 20) Stumpffi be ms ms Hz (174 ± 3 ms; N = 7) (919 ± 84 ms; N = 6) (3912 ± 11 Hz; N = 7) Stumpffi tridctyl ms ms Hz (132 ± 23 ms; N = 18) (1012 ± 39 ms; N = 17) (7244 ± 200Hz; N = 18) Stumpffi miery ms ms Hz (73 ± 12; n= 10) (3102 ± 456; n = 10) (8057 ± 136.9; N = 10) Stumpffi contumeli sp. nov ms ms Hz (42 ± 4 ms; N = 7) (543 ± 27 ms; N = 6) (7493 ± 50 Hz; N = 7) Stumpffi obscoen sp. nov ms ms Hz (144 ± 8 ms; N = 6) (4619 ± 990 ms; N = 5) (8361 ± 69Hz; N = 6) Stumpffi kibomen ms ms (73 ± 2 ms; n=9) (797 ± 15 ms; n = 9) Hz Stumpffi nigrorubr sp. nov ms ms Hz (67 ± 10 ms; N = 19) (1472 ± 104 ms; N = 18) (5283 ± 70.5 Hz; N = 19) Stumpffi chillei sp. nov ms ms Hz (43 ± 3.4 ms; N = 20) (530.4± 19.3 ms; N = 20) ( ± 235 Hz; N = 20) Stumpffi diutissim sp. nov ms ms Hz (54.6 ± 1 ms; N = 10) ( ± ms; N = 10) ( ± 51.8 Hz; N = 10) Stumpffi prdus sp. nov ms ms Hz (38.3 ± 2.3 ms; N = 12) (479.3 ± 7.6 ms; N = 12) ( ± 73.3 Hz; N = 12) Stumpffi jennoeli sp. nov ms ms Hz (44.2 ± 0.8 ms; N = 6) ( ± 5 ms; N = 5) ( ± 76.9 Hz; N = 6) Stumpffi tetrdctyl ms ms Hz (92 ± 9 ms; N = 11) (959 ± 244 ms; N = 10) (5355 ± 138 Hz; N = 11) Stumpffi spndei sp. nov ms ms Hz (46 ± 1.8 ms; N = 11) (801.8 ± 48.6 ms; N = 10) (5755 ± 69.8 Hz; N = 11) Stumpffi grrffoi sp. nov ms ms Hz (103.4 ± 2.5 ms; N = 7) ( ± 150 ms; N = 6) ( ± 84.2Hz; N = 7) Stumpffi roseifemorlis ms ms Hz (278 ± 1.6 ms; N = 6) ( ± 152 ms; N = 6) ( ± 35.1 Hz; N = 6) Stumpffi C ms ms Hz (293.8 ± 2.9 ms; N = 10) ( ± ms; N = 10) ( ± 20.7 Hz; N = 10) perspective nd the loclity is therefore not included in Figure 13. Nturl history. Specimens were observed in the lef litter of forested res nd plnttions. On Nosy Be, often syntopic with S. pygme. Clling ctivity is predominntly in the evening nd night (Vences & Glw, 1991). Cll. Clls were recorded t the type loclity Nosy Be by F. Glw in Februry 1992 (Vences et l., 2006: CD3, trck 35) nd hd lredy been previously described by Vences & Glw (1991) bsed on other recordings. Among clls of Stumpffi species, the cll of S. psologloss is unique in consisting of rther long nd distinctly pulsed note, with pulses/note emitted t repetition rte of pproximtely 17 pulses/second (Fig. 20). Additionl prmeters were s follows: cll durtion (= note durtion) ms (831 ± 25 ms; N = 11), inter-cll intervls ms (2691 ± 860 ms; N = 10), pulse durtion ms (17.2 ± 1.2; N = 23), nd dominnt frequency t Hz 303

34 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b c f g d e Fig. 21. Stumpffi nlmin, specimens in life. () ZSM 3224/2012 (ZCMV 12159) from Ankrfntsik Ntionl Prk; (b) holotype ZSM 542/2009 from ner Antsohihy; (c) uncollected specimen from Ankrfntsik; (d) KUZA 0011 from Ankrfntsik; (e) dditionl specimen from ner Antsohihy (ctlogue number uncertin); (f g) UADBA-A (ZCMV 12160) from Ankrfntsik. (4955 ± 520 Hz, N = 11). Clls from Benvony recorded in 1992 generlly gree in ll temporl nd spectrl chrcters. Stumpffi nlmin Klges, Glw, Köhler, Müller, Hipsley & Vences, 2013 (Figures 21 nd S92) Nme-bering type. Holotype ZSM 542/2009 (ZCMV 11428), collected t bridge locted t km 27 on the ntionl rod from Antsohihy to Mndritsr ccording to the originl description. Identity nd dignosis. Stumpffi nlmin ws described from loclity ner Antsohihy in north-western Mdgscr, nd geneticlly slightly divergent specimens ssignble to this species re lso known from second north-western loclity (Ankrfntsik Ntionl Prk). (1) Smll-sized species (SVL mm); (2) mnus with four fingers (first finger slightly reduced in length) nd pes with five toes (first toe slightly reduced in length); (3) terminl phlnges of fingers nd toes without enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum smooth or slightly tuberculr; (6) gryish or brownish colortion with vrible but typiclly indistinct pttern nd without contrsted ventrl colortion, red color elements on ventrl side, or shrp color border between dorsum nd flnks; (7) regulrly repeted short singlenote chirp cll. Distinguished from S. psologloss by shorter unpulsed cll (Fif. 22), nd in most specimens by different color pttern, different skin texture, nd slightly stronger reduction of first toe. Specimens exmined. Holotype ZSM 542/2009 (ZCMV 11428) collected on 18 June 2009 by M. Vences, D.R. Vieites, F.M. Rtsovin, R.D. Rndriniin, E. Rjeririson, T. Rjofirison, nd J.L. Ptton, t km 27 on the ntionl rod from Antsohihy to Mndritsr (S , E , 140 m.s.l.). Prtypes ZSM 2829/2010 (ZCMV 10034), ZSM 2830/2010 (ZCMV 10036), ZSM 2831/2010 (ZCMV 10037), ll collected on 29 June 2010 t the type loclity by M. Vences, E. Rjeririson, T. Rjofirison, A. Rkotorison, S. Rsmison, F. Rndrinsolo, R.D. Rndriniin, F.M. Rtsovin, nd D.R. Vieites. Referred specimens: ZSM 3224/2012 (ZCMV 12159) nd UADBA-A (ZCMV 12160), collected on 7 Jnury 2012 t Ankrfntsik Ntionl Prk by A. Rkotorison. KUZA 0001 (2010-F002), collected on 2 December 2010 t Ampijoro in Ankrfntsik Ntionl Prk (S , E , 274 m.s.l.) by R. Ito; KUZA 0002 (2010-F003), collected on 2 December 2010 t Ampijoro by T. Jono; KUZA 0003 (2010-F004), collected on 5 December 2010 t Ampijoro by R. Ito; KUZA 0004 (2010-F005), collected on 6 December 2010 t Ampijoro by T. Jono; KUZA 0005 (2011-Ad- 049), collected on 25 Februry 2011 t Ampondrbe (S , E46.9) by A. Mori; KUZA 0006 (2011-Ad-050), collected on 25 Februry 2011 t Ampondrbe by U. Kwi; KUZA 0007 (2011-Ad-059), collected on 25 Februry 2011 t Ampondrbe by A. Mori; KUZA ( ), collected on 19 Jnury 2013 by pitfll trp t Ampijoro by T. Jono; KUZA 0010 (2013-NoCode), collected on 12 Mrch 2013 t Ampijoro by R. Ito; KUZA 0011 (2010-F001), collected on 25 November 2010 t Ampijoro by T. Jono. Distribution. This species is known from two loclities, (1) its type loclity (27 km northest of Antsohihy) nd (2) Ankrfntsik Ntionl Prk. Popultions of the ltter loclity re here llocted to S. nlmin bsed on morphology nd moleculr genetics with Ankrfntsik specimens being only slightly differentited geneticlly from topotypic ones (Fig. 3). Nturl history. Frogs sty on or under leves on the forest floor when they cll. The hbitt consists of dense shrub nd lef litter in the reltively humid prcels of 304

35 VERTEBRATE ZOOLOGY 67 (3) 2017 b c d Fig. 22. Spectrogrms (bove) nd oscillogrms (below) of the dvertisement clls of Stumpffi nlmin (1 s nd 6 s durtion sections ech). ( b) Clls recorded in cptivity from topotypes t 23 C ir temperture; (c d) cll recorded in nturl hbitt by T. Jono t Ampijoro t 28.5 C ir temperture. Note tht d shows the sounds produced by two individuls intermittently clling, of which one is louder (i.e., closer to the microphone). dry deciduous forests. During field surveys t Ankrfntsik ( ; dt in Fig. 23), S. nlmin clled throughout the study period, from erly December to mid Februry. Clling ctivity ppered to be rther independent from rinfll intensity nd from ir temperture (Fig. 23), lthough clling ctivity decresed for few weeks in erly Jnury 2013 when rinfll cesed. Although this species usully clled in the evening nd t night, they lso emitted clls on severl riny mornings. In the observtion times, clls peked round 17:00. Cll. The dvertisement cll of S. nlmin consists of single short note repeted in series t regulr intervls. Clls of topotypic mle recorded by J. Klges in cptivity (Klges et l., 2013) t 23 C ir temperture hd the following cll prmeters (Fig. 22): cll durtion (= note durtion) ms (260 ± 23.6 ms; N = 5), inter-cll intervls ms (2242 ± 136 ms; N = 4), nd dominnt frequency t Hz (6210 ± 80.4 Hz, N = 5). Clls recorded by T. Jono on 5 December 2010, t Ampijoro, Ankrfntsik Ntionl Prk, hd the following prmeters (ir temperture 28.5 C): cll durtion ms (153 ± 13.5 ms; N = 10), inter-cll intervls ms (1120 ± 30 ms; N = 9), nd dominnt frequency t Hz (5399 ± 91 Hz, N = 10). Generlly, chrcteristics of clls from both loclities re in greement, but slight differences re obvious, with the Ampijoro clls being more tonl in chrcter, wheres clls from the topotype re more pulstile. However, the ltter could likely represent n rtifct, becuse recordings were obtined from individuls kept in terrri (see Klges et l., 2013). Cll repetition rte is more drmticlly different, with clls recorded from Ankrfntsik being lmost twice s frequently repeted s those record- 305

36 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Fig. 23. Grphic scheme showing clling ctivity of Stumpffi nlmin t Ankrfntsik Ntionl Prk, recorded in by T. Jono. The level of cll ctivity 0 indictes no vocliztions were herd; level 1: single or few clls were herd, but did not mke chorus; level 2: clls or choruses were herd intermittently, but in most of the time no vocliztions occurred; level 3: clls or choruses were frequently herd, but silent intervls were still obvious; level 4: intensive choruses were herd during most of the census, but severl occsionl cesstion of the chorus ws recognized; level 5: choruses were intensive nd never cesed. Air temperture is given s line grph nd precipittion s br chrt. ed from terrrium (inter-cll intervls ms vs ms). This might be explined by differences in ir temperture (5.5 C wrmer in Ankrfntsik thn in the terrrium), or individul motivtion. Stumpffi gimmeli Glw & Vences, 1992 (Figures 24 nd S93) Nme-bering type. Holotype ZFMK from primry forest ner Benvony (environments of Ambnj, NW-Mdgscr) ccording to the originl description. Identity nd dignosis. As currently defined, S. gimmeli is species from the Smbirno Region in northern Mdgscr. It is known from vrious loclities nd includes substntil mount of genetic vrition. Becuse t present, insufficient dt re vilble bout the vrious popultions, we follow Klges et l. (2013) nd consider them s deep conspecific lineges in preliminry wy. This implies tht S. gimmeli s defined here includes Stumpffi sp. 3 nd sp. 4 in Vieites et l. (2009) (not included in Glw & Vences, 2007; Wollenberg et l., 2008; Köhler et l., 2010; Klges et l., 2013; only Stumpffi sp. 4 included in Scherz et l., 2016). (1) Smll- to modertely-sized (up to 18.1 mm; dult mle SVL 14.5 mm); (2) mnus with four fingers (first finger slightly reduced in length) nd pes with five toes (first toe lmost not reduced in length); (3) terminl phlnges of fingers nd toes with slightly enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum slightly to modertely tuberculr; (6) gryish or brownish colortion with vrible but typiclly indistinct pttern nd without contrsted ventrl colortion, red color elements on ventrl side, or shrp color border between dorsum nd flnks, typiclly with hint of yellow on the bdomen; (7) regulrly repeted short single-note chirp cll. Distinguished from S. psologloss by shorter nd unpulsed dvertisement cll, lrger size, nd color pttern nd skin texture in most specimens; nd from S. nlmin by distinctly lrger size nd lower degree of reduction of first toe. Specimens exmined. Holotype ZFMK from Benvony, collected by F. Glw nd J. Müller on 16 Februry 1992; prtypes ZFMK , collected by F. Glw nd M. Vences on 27 Mrch 1991 t Ambnj (ZFMK clered nd stined); prtypes ZFMK , collected on 17 Februry 1992 by F. Glw & J. Müller t Ambnj; prtype ZFMK nd , collection dt s holotype. Referred specimens: ZSM 412/2000, collected on 18 Februry 2000 in Berr (Anbohzo) by M. Vences, F. Andreone, nd J.E. Rndrinirin; ZSM 597/2001 (MV ), collected on 30 Jnury 2001 t Antsirsir by F. Andreone, F. Mttioli, J.E. Rndrinirin & M. Vences; ZSM 833/2003 (FG/MV ), collected on 2 Jnury 2003 in Mnongrivo, Cmp 0 (S , E , 688 m.s.l.) by F. Glw, R.D. Rndriniin & M. Vences; ZSM 3231/2012 (ZCMV 13538), collected on 31 Jnury 2012 t 27 km from Ambnj by A. Rkotorison nd A. Rzfimmntso; ZSM 3240/2012 (ZCMV 13552), UADBA-A (ZCMV 13550) nd UADBA-A (ZCMV 13551), ll collected on 4 Februry 2012 in Antsirsir by A. Rkotorison nd A. Rzfimnntso; ZSM 631/2014 (DRV 6450), collected on 30 June 2010 in Angorony forest frgment ner Mromndi (S , E , 115 m.s.l.); ZSM 614/2014 (DRV 6475) nd ZSM 615/2014 (DRV 6481), collected on 30 June 2010 in Mromindr forest frgment ner Ankrmy (S , E , 283 m.s.l.) by F.M. Rtsovin, S. Rsmison, T. Rjofirison, nd F. Rndrinsolo. 306

37 VERTEBRATE ZOOLOGY 67 (3) 2017 b c d e f g h i j Fig. 24. Stumpffi gimmeli: () ZSM 412/2000 from Berr; (b) Specimen from Ambnj (photo not ssignble to collected voucher specimens); (c d) UADBA-A (ZCMV 13550) from Antsirsir; (e f) UADBA-A (ZCMV 13551) from Antsirsir; (g h) ZSM 3240/2012 (ZCMV 13552) nd (i j) ZSM 3231/2012 (ZCMV 13538) collected t 27 km from Ambnj. Distribution. This species is known from eight loclities in northestern Mdgscr, including (1) Benvony (type loclity), (2) Antsirsir, (3) Ambnj nd vicinity, (4) Angorony, (5) Mnongrivo, (6) Mromindr, (7) Berr, nd (8) Tsrtnn (moleculr dt only; Mnrikob Forest, Andmpy cmpsite). All these popultions were llocted to S. gimmeli minly bsed on their moleculr phylogenetic reltionships, lthough substntil degree of genetic differentition mong popultions is present (see Fig. 3). Nturl history. Typiclly found in the lef litter of rinforest nd trnsitionry forest, s well s cco plnttions. Specimens cll in the evening nd t night (Glw & Vences, 1992; 1994). The kryotype of specimen ZMA (FG/MV ) from Mnongrivo hs been described by Apre et l. (2007). Cll. The dvertisement cll of S. gimmeli consists of single short note repeted in series t regulr intervls. Recordings re vilble from two popultions tht re here ssigned to this species (Fig. 25): (1) Clls recorded by F. Glw on 16 Februry 1992 t Benvony (26 C ir temperture; Vences et l., 2006: CD3, trck 40), with the following prmeters: cll durtion (= note durtion) ms (103.5 ± 6 ms; N = 10), inter-cll intervls ms (1892 ± 106 ms; N = 9), nd dominnt frequency t Hz (4584 ± 31.5 Hz, N = 10). (2) Clls recorded by M. Vences on 2 Februry 2001 t Andmpy Cmpsite, Mnrikob forest, Tsrtnn (25 C ir temperture), nd here tenttively ssigned to this species, with the following prmeters: cll durtion ms (160 ± 16 ms; N = 7), inter-cll intervls ms (2990 ± 616 ms; N = 6), nd dominnt frequency t Hz (5167 ± 25 Hz, N = 7). Although detiled prmeters vry slightly mong popultions, these differences re likely explined by differences in motivtion, temperture nd body size of the mles recorded (frequency). The clls reported here ll sound very similr to the humn er. Stumpffi mdgscriensis Mocqurd, 1895 (Figures 26, S94 S95) Nme-bering type. Holotype MNHN collected sur l montgne d Ambre ccording to the originl description. 307

38 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b c d Fig. 25. Spectrogrms (bove) nd oscillogrms (below) of the dvertisement clls of Stumpffi gimmeli from two loclities (1 s nd 6 s durtion section ech): ( b) cll from Benvony; (c d) cll from Mnrikob forest, Tsrtnn (bndpss filter pplied). Identity nd dignosis. The identity of S. mdgsc riensis hs been difficult to scertin due to the very poor stte of preservtion of the holotype. However, bsed on the type loclity (Montgne d Ambre), color ptterns s vilble from the originl description, s well s body size nd digitl reduction, Köhler et l. (2010) ssigned this nomen to one of the severl species occurring on Montgne d Ambre. (1) Miniture to smll-sized (SVL mm); (2) mnus with four fingers (first finger modertely reduced in length) nd pes with five toes (first toe strongly reduced in length); (3) terminl phlnges of fingers not enlrged, those of toes with slightly enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum slightly tuberculr; (6) gryish or brownish dorslly, often with drker centrl ptch covering most of the posterior dorsum, this ptch either distinct or fintly recognizble only. Typiclly with shrp color border between lighter dorsum nd drker flnks. No contrsted ventrl colortion nd no red color elements on ventrl side; (7) regulrly repeted short single-note chirp cll. Distinguished from S. psologloss by shorter unpulsed dvertisement cll, nd by color pttern nd skin texture in most specimens; from S. nlmin by color pttern, possibly by longer inter-cll intervl, nd possibly by proportionlly longer hnd nd feet; nd from S. gimmeli by smller body size, color pttern, nd longer inter-cll intervl. Specimens exmined. Holotype MNHN , collected between My nd July 1893 on Montgne d Ambre by C. Alluud & Mr. Belly. Referred specimens: ZSM 201/2004 (FGZC 389) nd ZSM 202/2004 (FGZC 390), collected on 19 Februry 2004 on Montgne d Ambre (S , E , 1052 m.s.l.) by F. Glw, M. Puente, R.D. Rndriniin & A. Rzfimnntso; ZSM 2168/2007 (FGZC 1241), collected on 12 Mrch 2007 on Montgne d Ambre (S E , 1050 m.s.l.) by F. Glw, P. Bor, H. Enting, J. Köhler, A. Knoll; ZSM 3242/2012 (ZCMV 12185), collected on 16 Jnury 2012 on Montgne d Ambre by A. Rkotorison nd A. Rzfimnntso. 308

39 VERTEBRATE ZOOLOGY 67 (3) 2017 b c d Fig. 26. Stumpffi mdgscriensis in life: ( b) Specimen from Forêt d Ambre Specil Reserve (not sequenced; identifiction tenttive); (c d) ZSM 3242/2012 (ZCMV 12185) from Montgne d Ambre Ntionl Prk. b Fig. 27. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi mdgscriensis from Montgne d Ambre Ntionl Prk, recorded from specimen ZSM 3242/2012 (ZCMV 12185). () 1 s durtion section; (b) 6 s durtion section. Note tht the initil pek in mplitude t beginning of ech note is the result of n rtifct produced by dmged recording device. 309

40 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Distribution. Only known from Montgne d Ambre Ntionl Prk, lthough probbly lso occurring in Forêt d Ambre Specil Reserve (specimens tenttively identified from photos by color pttern; Fig. 26). Specimens from Montgne des Frnçis re considered to represent closely relted cndidte species (S. cf. mdgscriensis C25; see remrk below). Nturl history. Specimens were observed clling from the lef litter of rinforest in the evening. Cll. Advertisement clls were recorded by A. Rkotorison on 16 Februry 2012 in Montgne d Ambre Ntionl Prk from specimen ZSM 3242/2012 (ZCMV 12185) nd consist of single short note repeted in series t regulr intervls (Fig. 27). Numericl prmeters re s follows: cll durtion (= note durtion) ms (195 ± 5 ms; N = 5), inter-cll intervls ms (4224 ± 153 ms; N = 4), nd dominnt frequency t Hz (4231 ± 197 Hz, N = 5). It is importnt to mention tht recordings contin rtifcts such s initil peks of mplitude t beginning of ech note, which were produced by dmged recording device. Therefore the cll dt presented herein must be treted with cution. Remrk. We here discuss long with S. mdgscriensis geneticlly divergent form found t Montgne des Frnçis nd nmed Stumpffi sp. C25 ccording to Vieites et l. (2009), Köhler et l. (2010), Klges et l. (2013), nd Scherz et l. (2016). This linege is resolved with high support s sister to S. mdgscriensis from Montgne d Ambre bsed on genetic informtion from severl specimens. Despite substntil genetic divergence ( % in the 16S gene) the vilble dt do not llow cler conclusion s to whether this linege is better considered deep conspecific linege of S. mdgscriensis, or distinct species. No informtion on cll nd nturl history re vilble. Referred specimens of S. sp. C25: ZSM 2108/2007 (FGZC 1103), ZSM 2109/ 2007 (FGZC 1105) collected on 27 Februry 2007 on Montgne des Frnçis (S , E ) by F. Glw, P. Bor, H. Enting, J. Köhler nd A. Knoll. Stumpffi pygme Vences & Glw, 1991 (Figures 28 nd S96) Nme-bering type. Holotype ZFMK from m Strßenrnd, c. 1 km nördlich von Andony, Nosy Be, Nordwest-Mdgskr, trnslted: t the edge of the rod, c. 1 km north of Andony (= Hellville), Nosy Be, northwest Mdgscr, from the originl description. Identity nd dignosis. Stumpffi pygme is smllsized species of the north-western linege, without strong digitl reduction, tht so fr hs only been found on the offshore islnd Nosy Be nd Nosy Komb. (1) Miniturized species (SVL mm); (2) mnus with four fingers (first finger slightly reduced in length) nd pes with five toes (first toe slightly reduced in length); (3) terminl phlnges of fingers nd toes without enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum smooth or very slightly tuberculr; (6) light brownish colortion with typiclly indistinct or bsent pttern nd without contrsted ventrl colortion, red color elements on ventrl side, or shrp color border between dorsum nd flnks; (7) regulrly repeted short singlenote chirp cll. Distinguished from S. psologloss by shorter unpulsed dvertisement cll, nd by color pttern nd skin texture in most specimens; from S. nlmin by longer inter-cll intervls, nd possibly by proportionlly longer hnd nd feet; from S. gimmeli by smller body size, nd higher dominnt frequency of clls; nd from S. mdgscriensis by lck of shrp color border between dorsum nd flnks. Morphologiclly similr to S. nlmin but phylogeneticlly not closely relted to tht species. Sister group to S. mdgscriensis, which lso is morphologiclly similr but differs in color pttern, cll, nd concordnt nucler nd mitochondril DNA divergence. Specimens exmined. Holotype ZFMK 52541, collected on Mrch 1991 ner Andony, Nosy Be by F. Glw nd M. Vences; prtypes ZFMK 52542, ZFMK nd ZSM 557/1999 (formerly ZFMK 52543), sme collection dt s holotype. Referred specimens: ZFMK , 53772, 53773, ll collected in Jnury Februry 1992 on Nosy Be by F. Glw nd J. Müller; ZSM 430/2010 nd ZSM 431/2010, collected on 13 Februry 2010 from forest ner Ampsipohy, Nosy Be (S , E , 39 m.s.l.) by F. Glw. Distribution. This species is known from severl different loclities on Nosy Be, nd the nerby islnd of Nosy Komb (Andreone et l., 2003; Hyde Roberts & Dly, 2014; Blumgrt et l., 2017). Nturl history. A loclly quite common species in Nosy Be, in lef litter of secondry forest nd ylngylng plnttions. Clling ctivity mostly observed 1 2 hours before dusk, from the lef litter, with rin lso during the dy (Vences & Glw, 1991). Cll. Advertisement clls were recorded by F. Glw nd M. Vences in 1991 t the type loclity Nosy Be (Vences et l., 2006: CD3, trck 36) nd consists of single highpitched, tonl note (Fig. 29). Numericl prmeters re s follows: cll durtion (= note durtion) ms (196 ± 10 ms; N = 5), inter-cll intervls ms (3447 ± 419 ms; N = 4), nd dominnt frequency t Hz (6054 ± 49 Hz, N = 5). Stumpffi ngeluci sp. nov. (Figures 30 nd S97) Holotype. ZSM 3241/2012 (ZCMV 12184) (Figs. 12, 30 d e), n dult mle, collected on 16 Jnury 2012, t the prk entrnce to Montgne d Ambre Ntionl Prk (S , E , c. 975 m bove se level), Antsirnn Province, Mdgscr, by A. Rkotorison nd A. Rzfimnntso. 310

41 VERTEBRATE ZOOLOGY 67 (3) 2017 b c d Fig. 28. Stumpffi pygme. Specimens in life from Nosy Be (photos not ssignble to voucher specimens). () neometmorph specimen on mtchstick; (b c) dults; (d) fom nest with embryos nd endotrophic tdpoles. b Fig. 29. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi pygme from Nosy Be: () 1 s durtion section; (b) 6 s durtion section. Prtypes. ZSM 219/2004 (FGZC 428), ZSM 223/2004 (FGZC 433), ZSM 224/2004 (FGZC 434), UADBA-A (FGZC 436) nd UADBA-A (FGZC 435), collected on Februry 2004 in Montgne d Ambre Ntionl Prk by F. Glw, M. Puente, R.D. Rndriniin nd A. Rzfimnntso; ZSM 303/2004 (FGZC 586), ZSM 300/2004 (FGZC 580), ZSM 301/2004, (FGZC 581), ZSM 302/2004 (FGZC 583), UADBA-A (FGZC 582) nd UADBA-A (FGZC 585), collected on 28 Februry 2004 in Montgne des Frnçis (S12.333, E49.35, m.s.l.) by F. Glw, M. Puente, R.D. Rndriniin nd the employees of Kings Lodge; ZSM 1671/2008 (FGZC 1901), collected on 2 Mrch 2008 in Montgne des Frnçis (S , E , 140 m.s.l.) by F. Glw; ZSM 1639/2012 (FGZC 4915), collected on 10 December 2012 in Forêt d Ambre Specil Reserve, c. 5 km SW Skrmy 311

42 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs (S , E , 487 m.s.l.), by F. Glw, O. Hwlitschek, T. Rjofirison, A. Rkotorison, FM. Rtsovin, A. Rzfimnntso; ZSM 3243/2012 (ZCMV 12197), collected on 17 Jnury 2012 round the prk entrnce to Montgne d Ambre Ntionl Prk (S S, E , c. 975 m.s.l.) by A. Rkotorison nd A. Rzfimnntso; ZSM 229/2016 (ZCMV 13608), collected in Joffreville between by A. Rkotorison nd A. Rzfimnntso; UADBA-A (ZCMV 13612), collected in Montgne d Ambre Ntionl Prk between by A. Rkotorison nd A. Rzfimnntso. Dignosis. One of severl morphologiclly similr but prtly symptric Stumpffi species from northern Mdgscr. This species hs been previously listed s Stumpffi sp. 27 in Vieites et l. (2009), Köhler et l. (2010), Klges et l. (2013), Scherz et l. (2016) nd Peloso et l. (2017). (1) Smll- to modertely-sized species (dult mle SVL mm); (2) mnus with four fingers (first finger not obviously reduced in length) nd pes with five toes (first toe slightly to modertely reduced in length); (3) terminl phlnges of fingers without, those of toes with slightly enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum slightly tuberculr; (6) dorslly uniformly pricot with some smll, indistinct nd poorly contrsted drk pttern, tht cn be more intense. Ventrlly trnslucent pech with some indistinct drk nd light mottling especilly on the chest, nd sometimes drk color on throt. No red color ventrlly, no shrp border between dorsl nd lterl color; (7) regulrly repeted single-note tonl cll. Distinguished from S. be, S. grndis, S. hr, S. kibomen, S. megsoni, S. roseifemorlis, nd S. stffordi by smller body size; from S. tridctyl by lower degree of digitl reduction; from S. miery nd S. gimmeli by longer cll durtion in dvertisement cll nd less expressed length reduction of fingers; from S. tetrdctyl by less expressed length reduction of first finger nd especilly first toe; from S. psologloss by slightly more enlrged toe discs, colortion, nd shorter, unpulsed dvertisement cll; from S. pygme, S. mdgscriensis, nd S. nlmin by lrger body size. Phylogeneticlly in clde A1, which contins only two nominl species (S. psologloss nd S. gimmeli), which re distinguished by biocoustic chrcters. Description of the holotype. Specimen in good stte of preservtion, piece of left thigh removed s tissue smple for DNA extrction. Body round; hed slightly wider thn long, nrrower thn body; snout slightly pointed in dorsl view, pointed in lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis distinct, stright; lorel region stright, slightly oblique; tympnum distinct, bout 62% of eye dimeter; suprtympnic fold not visible; tongue long, brodening posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone ovl. Forelimbs slender; subrticulr tubercles single, distinct; outer metcrpl tubercle distinct, single, ovl; plmr tubercle distinct, single, round, equl in size to outer metcrpl tubercle; prepollex either smll or inner metcrpl tubercle, bout the sme size s other crpl tubercles; fingers without webbing; no fingers reduced; reltive length of fingers 1 < 2 = 4 < 3, fourth finger subequl in length to second; finger tips not expnded into discs. Hind limbs slender; TIBL 48% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle distinct, smll, nd ovl; outer mettrsl tubercle bsent; no webbing between toes; toes not reduced; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe slightly shorter thn third. Skin on dorsum smooth, without distinct dorsolterl folds; ventrl skin smooth; in life, dorsl skin ws grnulr. Colortion of the holotype. After three yers in 70% ethnol, the dorsum is pinkish with crem flnks. Two drker dots re present over the nterior of the scpulr region, forming the nterior ends of fint X-like mrking bove the scpule. A brown spot is present bove the insertion of ech rm, nd in the inguinl region. A brown M-like line runs between the posteriors of the eyes. Nostril surrounded by brown; drk suprtympnic stripe ending just beyond the tympnum; lterl hed below this stripe is beige. Ventrlly beige flecked with crem, slightly trnslucent nterior to the legs; ventrl surfce of thigh s trunk but slightly browner, shnk drk beige; sole of foot brown; vent region brown. Dorsl thigh pinkish s dorsum, with fint brown crossbnd; posterodorsl surfce of thigh brown towrds knee. Dorsl shnk pinkish, with perpendiculr brown crossbnd midwy long its length. Foot dorslly pinkish, with drk spot on its outer edge; toes mottled brown. Arms dorslly pinkish; lower rm with single brown crossbnd; hnds speckled. Color in life (holotype nd vrition). Dorsum pricot with subtle mrkings mde of dense speckles of ebony, with one forming concve interoculr br, wek nterior chevron round the suprscpulr region, nd nother wek nterior chevron from the inguinl region to the mid-bck. These mrkings cn be more intense (Fig. 30). The flnk becomes incresingly crem ventrlly. A lrge ebony spot is present bove the insertion of the rm (cn be weker, cf. Fig 29d, or stronger with more spots on the flnk, cf. Fig. 30). Another ebony mrking runs from the posterior mrgin of the eye, curving towrd the nterior insertion of the rm over the tympnum. The dorsl forelimb is s the bck, with n ebony crossbnd on the forerm. The fingers re mottled gry nd pricot with whitish nnulus before the terminl phlnge of ech finger. The dorsl legs re s the bck, with one wek crossbnd t the mid-thigh nd one on the mid-shnk. The toes re mottled ebony nd tn, with white nnulus before the terminl phlnge of ech toe. The ventrl skin is trnslucent pech cross the whole body, with crem flecks in firly dense bnd t nd just posterior to the pectorl girdle. The iris of the holotype is drk copper reticulted with blck, but it cn lso be gold with blck reticultions. Vrition. For vrition in mesurements mong specimens, see Tble 3. In generl, ll exmined specimens 312

43 VERTEBRATE ZOOLOGY 67 (3) 2017 b c d e f g Fig. 30. Stumpffi ngeluci sp. nov. from Montgne d Ambre Ntionl Prk in life: ( c) ZSM 3243/2012 (ZCMV 12197); (d e) holotype ZSM 3241/2012 (ZCMV 12184); (f g) ZSM 3243/2012 (ZCMV 13608). b Fig. 31. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi ngeluci sp. nov. from Joffreville recorded on 17 Jnury 2016 from prtype UADBA-A (ZCMV 13612): () 1 s durtion section, (b) 6 s durtion section. 313

44 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs gree strongly with the holotype in morphology. Tubercles of the hnd vry from the condition of the holotype by plmr nd outer metcrpl tubercles being fused. An intermedite degree of vribility ws noted in the colortion of specimens; the M- nd X-like mrkings of the holotype re lcking from ll other specimens. The drk dots bove the insertion of the rm re wekly present in ZSM 3243/2012 nd bsent in ll other prtypes. In one specimem (ZSM 300/2004), two pirs of white dots re present on the dorsum, one behind the scpulr region, nd one in front of the inguinl region. However, the inguinl spots, surptympnic stripe, brown clocl region, nd brown nostril re present in ll specimens (degree of visibility vries). The degree of visibility of leg crossbnds vries strongly, but they re present to some degree in ll specimens. Etymology. The species nme is ptronym honoring Angeluc Rzfimnntso, in recognition of his invluble help in herpetologicl explortion of Montgne d Ambre nd neighbouring regions. Distribution. This species is known from northern Mdgscr nd occurs in nd round Montgne d Ambre Ntionl Prk, t Joffreville, in Forêt d Ambre Specil Reserve s well s Montgne des Frnçis. The rnge encompsses elevtions from m bove se level. Nturl history. The clling mle ws observed hiding between tree roots on the forest floor, nd other ctive individuls were observed sitting on plnts, bout 5 cm bove the ground. Cll. Advertisement clls were recorded t Montgne d Ambre Ntionl Prk by A. Rkotorison on 17 Jnury 2016 from prtype UADBA-A (ZCMV 13612) nd consist of single, short, tonl note repeted in series t regulr intervls (Fig. 31). Numericl prmeters re s follows: cll durtion (= note durtion) ms (182 ± 2.7 ms; N = 6), inter-cll intervls ms (3596 ± 254 ms; N = 5), nd dominnt frequency t Hz (4651 ± 47 Hz; N = 6). Stumpffi huwei sp. nov. (Figures 32 nd S98) Holotype. ZSM 221/2016 (ZCMV 13618) (Figs. 12 b, 32 nd S98 b), n dult mle, collected between in Montgne d Ambre Ntionl Prk (S , E , 1018 m bove se level), Antsirnn Province, Mdgscr by A. Rkotorison nd A. Rzfimnntso. Prtypes. ZSM 3245/2012 (ZCMV 13514) nd UADBA-A (ZCMV 13515), UADBA-A (ZCMV 13619), ZSM 222/2016 (ZCMV 13620), UADBA-A (ZCMV 13621) nd ZSM 223/2016 (ZCMV 13622), ll collected between in Montgne d Ambre Ntionl Prk, by A. Rkotorison nd A. Rzfimnntso; ZSM 224/2016 (ZCMV 13611), collected in December 2011 in Montgne d Ambre Ntionl Prk, by A. Rkotorison nd A. Rzfimnntso; ZSM 904/2003 (FG/MV ), collected on 18 Februry 2003 in Montgne d Ambre by F. Glw, R.D. Rndriniin nd A. Rzfimnntso. Dignosis. One of the severl morphologiclly similr but prtly symptric Stumpffi species from northern Mdgscr. This species hs not been previously listed s cndidte species in ny publiction. (1) Smll-sized species (mle SVL mm); (2) mnus with four fingers (first finger not or very wekly reduced in length) nd pes with five toes (first toe slightly reduced in length); (3) terminl phlnges of fingers without, those of toes with slightly enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum smooth; (6) dorslly gryish to reddish brown, sometimes with lrge centrl drk teddyber shped ptch bordered with lemon on the dorsl flnks. Ventrlly some specimens uniformly gryish with some yellowish color on belly, other specimens with more intensive yellowish color on belly nd especilly on throt. No red color ventrlly, no shrp border between dorsl nd lterl color; (7) regulrly repeted single-note tonl cll with cll durtion > 100 ms. Distinguished from S. be, S. grndis, S. hr, S. kibomen, S. megsoni, S. roseifemorlis, nd S. stffordi by smller body size; from S. tridctyl by lower degree of digitl reduction; from S. miery by lower frequency rnge of dvertisement clls, nd less expressed length reduction of fingers; from S. tetrdctyl by less expressed length reduction of first finger nd especilly first toe; from S. psologloss by slightly less reduced first finger, skin texture, colortion, nd shorter, unpulsed dvertisement cll; from S. gimmeli by shorter cll durtion; from S. mdgscriensis by the lck of shrp border between dorsl nd lterl color nd slightly lrger body size; from S. pygme nd S. nlmin by lrger body size; from S. ngeluci by distinctly shorter cll durtion. The species is included in clde A1 but is not the direct sister group of ny of the nominl species described so fr (see below for its distinction from its sister species, S. mmitik sp. nov.). Description of the holotype. Specimen in good stte of preservtion, left thigh muscle removed s tissue smple. Body elongte; hed slightly longer thn width, nrrower thn body; snout rounded in dorsl view, rounded in lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis slightly distinct, stright; lorel region concve, oblique; tympnum slightly distinct, bout 60% of eye dimeter; suprtympnic fold not recognizble; tongue long, broding posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone rounded. Forelimbs slender; subrticulr tu bercles single, distinct; outer metcrpl tubercle not recognizble; inner metcrpl tubercle smll, ovl; fingers without webbing; reltive length of fingers 1 < 4 < 2 < 3, fourth finger slightly shorter thn second; finger tips not expnded into discs. Hind limbs slender; TIBL 47% of SVL; lterl mettrsli strongly con nected; inner mettrsl 314

45 VERTEBRATE ZOOLOGY 67 (3) 2017 b c d e f g Fig. 32. Stumpffi huwei sp. nov. specimen from Montgne d Ambre Ntionl Prk in life: () Holotype ZSM 221/2016 (ZCMV 13618); (b c) ZSM 3245/2012 (ZCMV 13514); (d e) UADBA-A (ZCMV 13621); (f g) ZSM 223/2016 (ZCMV 13622). tubercle smll, ovl; outer mettrsl tubercle bsent; no webbing between toes; toe tips slightly expnded; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe shorter thn third; subrticulr tubercles distinct, single. Skin on dorsum reltively smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After four yers in 70% ethnol, the dorsum is light beige, bnded with brown from the middle of the eyes to the inguinl region. The dorsl surfce of the hed is light beige flecked with brown. The nostril is beige surrounded with brown. The lterl surfce of the hed is s the dorsl surfce of the hed. The flnks re lterlly nd ventrlly beige flecked with brown. The flnk colortion merges with the ventrl colortion. The ventrl trunk is uniformly beige. The chin is beige vermiculted with brown. The ventrl thigh is beige spotted with brown. The shnk is ventrlly beige vermiculted with light brown. The trsus is ventrlly beige spotted with brown. The sole of the foot is s the trsus but drker. Dorslly, the thigh is beige spotted with brown nd with one brown crossbnd. The posterodorsl surfce of the thigh is beige mottled with drk brown. The shnk is s the thigh but without crossbnds. The trsus, foot, nd toes re s the shnk. The clocl region is drk brown. The rms re beige spotted with brown. The dorsomedil surfce of the hnd is beige. The fingers re s the rms. The underside of the rm is s the chin. Color in life (holotype nd vrition). Dorsl color mottled copper nd burnt umber, bordered by lemon yellow on the dorsolterl hed, spreding down to the flnks (Fig. 32). The lterl hed to behind the insertion of the forelimb is s the dorsum. The forelimb is lso s the bck, with wek lighter bnds before ech terminl phlnge of the hnd. The dorsl leg is s the bck, with one burnt umber crossbnd on the mid-thigh nd mid-shnk, nd distinct whitish bnds before the terminl phlnge of ech toe. The venter of the holotype ws not photogrphed in life, but the venters of prtypes vry from yellowish crem dppled with whitish spots, to lemon yellow with whitish dppling in the posterior body nd burnt umber reticultions under the chin (Fig. 32 d, e, g). The ventrl legs re tupe. The iris is brown to copper. Vrition. For vrition in mesurements mong spe cimens, see Tble 3. In generl, ll exmined specimens gree strongly with the holotype in morphology, with the exception of ZSM 224/2016 (ZCMV 13611), which is the lrgest known specimen. A high degree of vribility 315

46 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b c d Fig. 33. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi huwei sp. nov. from Montgne d Ambre Ntionl Prk, from ( b) holotype ZSM 221/2016 (ZCMV 13618) nd (c d) prtype UADBA-A (ZCMV 13621), recorded between : (, c) 1 s durtion section; (b, d) 6 s durtion section. ws noted in the colortion of specimens in preservtive: UADBA-A (ZCMV 13621) is dorslly pinkish with brown X-shped mrking strting from the eyes to the scpulr region nd brown spot in the inguinl region; ZSM 223/2016 (ZCMV 13622) is uniformly drk brown with some metllic spots on dorsum; UADBA-A (ZCMV 13619) is beige with brown X shped mrking from the eyes to the inguinl re, nd shnk nd trsus with brown crossbnd; ZSM 904/2003 (FG/ MV ) is dorslly light brown with brown V-shped mrking in the scpulr region. Etymology. The nme is ptronym honoring Reinhrd Huwe, dedicted nturlist nd ornithologist in Brunschweig nd technicin t the Zoologicl Institute of the Technicl University of Brunschweig, in recognition of his immense nd cretive contributions to reserch nd teching in zoology. Distribution. The species is known only from Montgne d Ambre Ntionl Prk. The rnge encompsses elevtions from m bove se level. Nturl history. The species occurs in rinforest in Montgne d Ambre Ntionl Prk. Active individuls were found in dense lef litter, however the clling mles were found t dusk t sites with less dense lef litter cover s well. Cll. The dvertisement cll consists of single, very short note emitted in series t regulr intervls (Fig. 33). Two recordings re vilble from Montgne d Ambre Ntionl Prk: (1) Clls recorded by A. Rkotorison between from specimen ZCMV hd the following numericl prmeters: cll durtion (= note durtion) ms (70 ± 2.4 ms; N = 6), inter-cll intervls ms ( ± 267 ms; N = 6), nd dominnt 316

47 VERTEBRATE ZOOLOGY 67 (3) 2017 b Fig. 34. Stumpffi ihrn sp. nov. from Fnmbn forest in life. Prtype, ZSM 1652/2012 (FGZC 3927) in () dorsolterl nd (b) ventrl view. frequency t Hz (5020 ± 39 Hz, N = 6). (2). Clls recorded by A. Rkotorison between from specimen ZCMV 13621: cll durtion ms (66.7 ± 2.75 ms; N = 7), durtion of inter-cll intervls ms ( ± 424 ms; N = 6), nd dominnt frequency t Hz (5081 ± 60.8 Hz, N = 7). Stumpffi ihrn sp. nov. (Figures 34 nd S99) Holotype. ZSM 1651/2012 (FGZC 3895) (Figs. 12 c, S99 c d), collected on 4 December 2012 in gllery forest southest of Andrfinkon (S , E , 820 m bove se level), Antsirnn Province, Mdgscr, by F. Glw, O. Hwlitschek, F.M. Rtsovin, A. Rkotorison, T. Rjofirison, nd A. Rzfimnntso. Prtypes. ZSM 1650/2012 (FGZC 3893) nd UADBA-A (FGZC 3894), with sme collecting dt s holotype; ZSM 1642/2012 (FGZC 3800) collected on 3 December 2012 ner Ambodimndresy (c. S , E , 778 m.s.l.) by F. Glw, O. Hwlitschek, F.M. Rtsovin, A. Rkotorison, T. Rjofirison, nd A. Rzfimnntso; ZSM 1652/2012 (FGZC 3927) nd UADBA-A (FGZC 3926), collected on 6 December 2012 in Fnmbn forest (S , E , 53 m.s.l.) by F. Glw, O. Hwlitschek, F.M. Rtsovin, A. Rkotorison, T. Rjofirison, nd A. Rzfimnntso. Dignosis. One of severl morphologiclly similr but prtly symptric Stumpffi species from northern Mdgscr. This species hs not been previously listed s cndidte species in ny publiction. (1) Smll-sized species (dult SVL mm; femle SVL 14.7 mm); (2) mnus with four fingers (first finger not or very wekly reduced in length) nd pes with five toes (first toe slightly reduced in length); (3) terminl phlnges of fingers without, those of toes with slightly to modertely enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum smooth with few scttered tubercles; (6) dorslly snd brown with poorly defined drker mrkings. Ventrl skin trnslucent with dense crem flecks over the pectorl girdle, becoming fine speckling posteriorly. No red color ventrlly, no shrp border between dorsl nd lterl color. Distinguished from S. be, S. grndis, S. hr, S. kibomen, S. megsoni, S. roseifemorlis, nd S. stffordi by smller body size; from S. tridctyl by lower degree of digitl reduction; from S. miery by less expressed length reduction of fingers; from S. tetrdctyl by less expressed length reduction of first finger nd especilly first toe; from S. psologloss, S. nlmin, S. mdgscriensis, nd S. pygme by lrger body size; from S. mdgscriensis lso by the lck of shrp border between dorsl nd lterl color nd slightly lrger body size. Most similr to S. gimmeli, S. ngeluci, nd S. huwei. These three species re ll included with S. ihrn in clde A1. The new species cn be distinguished from S. ngeluci nd S. huwei becuse these do not constitute its direct phylogenetic sister groups, nd do not shre mitochondril DNA or Rg-1 hplotypes. Phylogeneticlly the new species is sister to S. gimmeli. As discussed bove, S. gimmeli contins numerous deep genetic lineges nd might represent species complex; we here consider nd describe s distinct species S. ihrn the most divergent of these lineges which differs concordntly in mtdna nd by not shring Rg-1 hplotypes, nd by substntil geogrphic distnce from its known distribution re. This species hypothesis requires future confirmtion by thorough revision of the S. gimmeli complex with the inclusion of biocoustic dt nd finescle geogrphic smpling. Description of the holotype. Specimen in good stte of preservtion, tongue removed s tissue smple for DNA extrction. Body round; hed longer thn wide, wider thn body; snout rounded in dorsl view, rounded in lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis slightly distinct, 317

48 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs concve; lorel region stright, oblique; tympnum distinct, bout 60% of eye dimeter; suprtympnic fold not distinct; tongue bsent, mxillry teeth nd vomerine teeth bsent, chone ovl. Forelimbs slender; subrticulr tubercles distinct, single, rounded; outer metcrpl tubercle smll, elongte; inner metcrpl tubercle distinct; fingers without webbing; first finger slightly reduced; reltive length of fingers 1 < 4 < 2 < 3, fourth finger shorter thn second; finger tips slightly expnded into discs. Hind limbs slender; TIBL 47% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle prcticlly indistinguishble; outer mettrsl tubercle bsent; no webbing between toes; toe tips not expnded; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe shorter thn third; subrticulr tubercles distinct, single. Skin on dorsum smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After four yers in 70% ethnol, the dorsum is slightly pinkish beige with brod brown vertebrl bnd. The dorsl surfce of the hed is s the bck. The nostril is brown. The lterl surfce of the hed is beige mottled with brown. The flnks re crem mottled with beige. The flnk colortion merges with the ventrl colortion. The ventrl trunk is s the flnk. The chin is s the trunk. The ventrl thigh is s the belly. The shnk is ventrlly s the thigh. The trsus is ventrlly s the shnk. The sole of the foot is beige mottled with crem. Dorslly, the thigh is s the bck with brown crossbnd. The posterodorsl surfce of the thigh is s the inguinl region. The shnk, trsus, nd feet re s the thigh. The toes re s the feet with crem crossbnds. The clocl region is not drk brown. The rms re s the dorsum, brown crossbnd present on the lower rm. The dorsomedil surfce of the hnd is crem. The fingers hve smll crem crossbnds. The underside of the rm is s the ventrl trunk. Color in life (prtypes). Dorsum sndy brown with subtle, grnulr coffee-brown mrkings nd ptternings. The dorsl hed is clen tn nterior to the level of the mid-eye. The flnks re s the dorsum, becoming incresingly flecked with crem nd decresingly brown ventrlly. The lterl hed is s the dorsum, though the nostril is surrounded by drk brown, s is the posterodorsl edge of the tympnum. The dorsl forelimb is s the trunk, with sooty crossbnd on the forerm. The fingers re mottled with brown, with light nnulus before ech terminl phlnge. The dorsl hindlimb is s the dorsum, with one grnulr coffeebrown crossbnd on the thigh, one on the shnk, nd one on the trsus. The toes re mottled s the dorsum, with light nnulus before ech terminl phlnge. The ventrl skin is trnslucent crem, immculte on the centrl chin but with crem speckles below the lower jw. Dense crem flecks cover the pectorl girdle nd decrese in density posteriorly, becoming fine speckles tht cover the posterior bdomen nd ventrl legs. The iris is golden with blck reticultions. Vrition. For vrition in mesurements mong specimens, see Tble 3. In generl, ll exmined specimens gree strongly with the holotype in morphology, except: in ZSM 1642/2012 (FGZC 3800) snout slightly pointed dorslly nd lterlly, big elongte inner metcrpl tubercle, smll elongte outer metcrpl tubercle, smll elongte inner mettrsl tubercle, dorslly browner thn the holotype, brown spot beyond the rm; in ZSM 1650/2012 (FGZC 3893) outer metcrpl tubercle indistinguishble, dorsl colortion gryish, brown X shpe in the scpulr region, reversed V in the inguinl region; in ZSM 1652/2012 (FGZC 3927) smll elongte outer metcrpl tubercle, smll inner mettrsl tubercle, dorsl colortion s the holotype, except for the presence of beige tringle on the hed. Etymology. The species nme is noun in pposition to the genus nme, nd refers to the distribution of the new species, ner Ihrn (Vohemr in French), in northestern Mdgscr. Distribution. Currently known from four loclities: (1) south-est of Andrfinkon (type loclity), (2) Fnmbn forest, (3) ner Ambodimndresy, nd (4) Bezvon. The ltter loclity Bezvon is ssigned to this species bsed on DNA sequence deposited in GenBnk (ccession number DQ283411) corresponding to the specimen AMNH A (not exmined by us). This species occurs in remrkbly brod ltitudinl rnge ( m.s.l.). Nturl history. This species inhbits the lef litter of costl nd mid-ltitude rinforest. Cll. Unknown. Stumpffi lrinki sp. nov. (Figures 35 nd S100) Holotype. ZSM 3236/2012 (ZCMV 13525) (Fig. 12 d, 35), collected on 26 Jnury 2012 t Cmpement des Princes (pp. S , E , 90 m bove se level), Ankrn Ntionl Prk, Antsirnn Province, Mdgscr, by A. Rkotorison nd A. Rzfimnntso. Prtypes. UADBA (FG/MV ), UADBA (FG/MV ), ZSM 779/2003 (FG/MV ), nd ZSM 861/2003 (FG/MV ), ll collected on 11 Februry 2003 in Ankrn Ntionl Prk by F. Glw, R.D. Rndriniin, nd A. Rzfimnntso; ZSM 1669/2008 (FGZC 1620), collected on 13 Februry 2008 in Ankrn Ntionl Prk, Petit Tsingy (= loclity Cmpement des Princes; S , E , 90 m.s.l.), by F. Glw, M. Frnzen, J. Köhler, N. D Cruze; ZSM 205/2016 (ZCMV 13617), clling mle, collected t the type loclity by A. Rkotorison nd A. Rzfimnntso. Dignosis. One of severl morphologiclly similr nd prtly symptric Stumpffi species from northern Md gscr. This species hs been previously listed s Stumpffi sp. 26 in Vieites et l. (2009), Köhler et l. (2010), Klges et l. (2013), Scherz et l. (2016) nd 318

49 VERTEBRATE ZOOLOGY 67 (3) 2017 b c Fig. 35. Stumpffi lrinki sp. nov. from Ankrn Ntionl Prk in life: ( c) Holotype ZSM 3236/2012 (ZCMV 13525). Peloso et l. (2017). (1) Smll-sized species (SVL mm); (2) mnus with four fingers (first finger not or very wekly reduced in length) nd pes with five toes (first toe very slightly reduced in length); (3) terminl phlnges of fingers without, those of toes with slightly to modertely enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum smooth; (6) dorslly iridescent copper with fint burnt umber ptch from the eyes to the mid-dorsum or vent. Ventrlly trnslucent gry or light plum. No red color ventrlly, no shrp border between dorsl nd lterl color; (7) regulrly repeted pulstile single-note cll with cll durtion > 100 ms. Distinguished from S. be, S. grndis, S. hr, S. kibomen, S. megsoni, S. roseifemorlis, nd S. stffordi by smller body size; from S. tridctyl by lower degree of digitl reduction; from S. miery by longer cll durtion nd less expressed length reduction of fingers; from S. tetrdctyl by less expressed length reduction of first finger nd especilly first toe; from S. psologloss by pulstile but not clerly pulsed cll; from S. gimmeli by lower dominnt frequency in clls nd slightly smller body size; from S. mdgscriensis by the lck of shrp border between dorsl nd lterl color nd slightly lrger body size. Morphologiclly most similr to S. ngeluci, S. ihrn, S. pygme, nd S. nlmin, but not the direct sister group of ny of those species, strongly divergent in mitochondril nd nucler DNA, nd distinguished from S. pygme nd S. nlmin by slightly lrger body size, from S. ngeluci by slightly 319

50 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b Fig. 36. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi lrinki sp. nov. from Ankrn Ntionl Prk, recorded from prtype ZSM 205/2016 (ZCMV 13617): () 1 s durtion section; (b) 6 s durtion section (including some microphone noise). shorter cll durtion, nd from S. huwei by longer cll durtion. The species is included in clde A1 but is not the direct sister group of ny nominl species, nd is chrcterized by comprtively long brnch seprting it from its closest reltives. Description of the holotype. Specimen in good stte of preservtion, left leg prtilly removed s tissue smple, skin of the right leg slightly dmged. Body round; hed wider thn long, nrrower thn body; snout rounded in dorsl view, somewht truncte in lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis stright or slightly concve; lorel region concve, oblique; tympnum visible, bout 55% of eye dimeter; suprtympnic fold not visible; tongue long, slightly brodening posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone rounded. Forelimbs slender; subrticulr tubercles not visible; outer metcrpl tubercle distinct, elongte; plmr metcrpl distinct, tringulr, prepollex visible, elongte; fingers without webbing; first finger reduced; reltive length of fingers 1 < 2 < 4 < 3, fourth finger slightly longer thn second; finger tips not expnded into discs. Hind limbs slender, TIBL 47% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle smll, ovl; outer me t trsl tubercle bsent; no webbing between toes; no toes reduced; toe tips slightly expnded into discs; rel tive length of toes 1 < 2 < 5 < 3 < 4; fifth toe slightly shorter thn third; subrticulr tubercles distinct, single. Skin on dorsum reltively smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After three yers in 70% ethnol, the dorsum is beige in the lterl regions with brod light brown bnd from the hed to the nterior inguinl region. No inguinl spots. The dorsl surfce of the hed is s the bck with blck ovl spot in the scpulr region. The nostril is light brown. The lterl surfce of the hed nd the flnks re beige spotted with brown. The flnk colortion merges with the ventrl colortion. The ventrl body is uniformly crem on the bdomen nd spotted with brown lterlly. The chin is s the ventrl bdomen, with crem ptches. The ventrl thigh is s the belly. The shnk nd trsus re ventrlly s the thigh. The sole of the foot is brown mottled with crem. Dorslly, the thigh is beige spotted with brown. The posterodorsl surfce of the thigh is s the dorsl surfce. The shnk is s the thigh with wekly distinct crossbnd. The trsus nd foot re s the thigh, without crossbnds. The toes re mottled with drk brown. The clocl region is brown. The rms re s the dorsum. A wekly contrsted brown crossbnd is present on the lower rm. The dorsomedil surfce of the hnd nd the fingers re beige spotted with brown. The underside of the rm is s the ventrl trunk. In life, dorsum iridescent copper with fint burnt umber ptch running from between the eyes to the middorsum. A wek lterl color border divides the dorsl colortion from the burnt umber flnk. Whitish flecks re present on the flnk. The dorsl rm is s the bck, with the fingers being burnt umber with light nnulus before ech terminl phlnge. The dorsl hindlimb is s the bck with burnt umber flecks. The toes hve light nnulus before ech terminl phlnge. The ventrl skin is trnslucent, light gry over the vocl sc, light plum over the pectorl region, nd tupe on the posterior trunk nd hindlimbs. The iris is golden reticulted with blck. Vrition. For vrition in mesurements mong specimens, see Tble 3. In generl, ll exmined specimens gree strongly with the holotype in morphology, ex- 320

51 VERTEBRATE ZOOLOGY 67 (3) 2017 cept: ZSM 779/2003 (FG/MV ), inguinl spot present, brod indistinct drk ptch beginning in the middle of the eyes region nd running over most of the dorsum; in ZSM 861/2003 (FG/MV ), dorsum with brown X-shped mrking strting in the scpulr region nd ending in the middle of the body. Etymology. The species nme is ptronym honoring Otto Lrink, Professor Emeritus t Technicl University of Brunschweig, in recognition to his chievements in zoology nd his continued ssistnce for specilized reserch on the Mlgsy fun. Distribution. This species is only known from Ankrn Ntionl Prk. Nturl history. The species occurs in dry forest in Ankrn Ntionl Prk. One clling mle ws found t night, clling from limestone rock nd second clling mle ws found during the dy hiding in smll limestone cve. Cll. Advertisement clls were recorded by A. Rkotorison on 8 Jnury 2016 in Ankrn Ntionl Prk from prtype ZSM 205/2016 (ZCMV 13617). The cll consists of single pulstile note repeted in series t regulr intervls (Fig. 36). Numericl cll prmeters re s follows: cll durtion (= note durtion) ms (138 ± 21 ms; N = 3), inter-cll intervls ms (2216 ± 103 ms; N = 2), nd dominnt frequency t Hz (2914 ± 124 Hz, N = 3). The pulstile nture of notes is evident from expnded oscillogrms s well s recognizble by the humn er. However, pulses re not clerly seprted or countble. Stumpffi mledict sp. nov. (Figures 37 nd S101) Holotype. ZSM 2079/2007 (FGZC 1049) (Figs. 12 e, S101 b), n dult, collected on 25 Februry 2007 ner the Gite d etpe cmpsite (S , E , c m bove se level), Montgne d Ambre Ntionl Prk, Antsirnn Province, Mdgscr by F. Glw, P. Bor, H. Enting, J. Köhler, nd A. Knoll. Prtypes. ZSM 2169/2007 (FGZC 1244) nd UADBA (FGZC 1243), collected on 12 Mrch 2007 t Voie des Milles Arbres (S E, , 1050 m.s.l.) in Montgne d Ambre Ntionl Prk by F. Glw, P. Bor, H. Enting, J. Köhler, nd A. Knoll; ZSM 3244/2012 (ZCMV 13504), collected on 22 Jnury 2012 t Lc Mudit, Montgne d Ambre Ntionl Prk by A. Rkotorison nd A. Rzfimnntso. Dignosis. One of severl morphologiclly similr but prtly symptric Stumpffi species from northern Mdgscr, with probbly very long inter-cll intervl durtion. This species hs been previously listed s Stumpffi sp. 28 in Köhler et l. (2010), Klges et l. (2013), Scherz et l. (2016) nd Peloso et l. (2017). (1) Smllto modertely-sized species (SVL up to 16.8 mm; one dult mle SVL 13.8 mm); (2) mnus with four fingers (first finger wekly reduced in length) nd pes with five toes (first toe distinctly reduced in length); (3) terminl phlnges of fingers without, those of toes with slightly enlrged discs; (4) reltive hnd nd foot length, HAL/ SVL , FOTL/SVL ; (5) dorsum slightly to modertely tuberculr; (6) dorslly uniformly snd brown with some smll, indistinct nd poorly contrsted drk pttern. Ventrlly trnslucent lemon yellow with gry lip nd nterior chin, with off-white speckles posterior to the pectorl region. No red color ventrlly, no shrp border between dorsl nd lterl color; (7) regulrly repeted single-note tonl cll of c. 100 ms durtion; if vilble cll recordings refer to dequtely motivted specimens, this species hs the longest intercll intervl durtion of ll known Stumpffi species ( ms). Distinguished from S. be, S. grndis, S. hr, S. kibomen, S. megsoni, S. roseifemorlis, nd S. stffordi by smller body size; from S. tridctyl by lower degree of digitl reduction; from S. miery nd S. huwei by longer cll durtion nd less expressed length reduction of fingers; from S. tetrdctyl, by less expressed length reduction of first finger nd especilly first toe; from S. psologloss by tonl cll (vs pulsed); from S. pygme, S. mdgscriensis, nd S. nlmin, by lrger body size; from S. ngeluci by shorter cll durtion nd longer inter-cll intervls; from S. ihrn by seprte phylogenetic position nd lck of hplotype shring in Rg-1. The species is recovered in clde A1. It is the sister group of S. ngeluci nd occurs syntopiclly with its sister species. However, the two species re concordntly differentited in mitochondril nd nucler genes despite symptry, nd pper to differ biocousticlly, thus confirming their species sttus. Description of the holotype. Specimen in good stte of preservtion, its right rm removed s tissue smple. Body round; hed wider thn long, nrrower thn body; snout pointed in dorsl view, pointed in lterl view; nostrils directed lterlly, not protubernt, nerer to the tip of snout thn to eye; cnthus rostrlis distinct, concve; lorel region stright, slightly oblique; tympnum distinct, bout 80% of eye dimeter; suprtympnic fold not visible; tongue dmged, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone rounded. Forelimbs slender; subrticulr tubercles single, distinct; outer metcrpl tubercle lrge, ovl; plmr tubercle round, butting outer tubercle; inner metcrpl tubercle distinct, ovl; fingers without webbing; first finger reduced; reltive length of fingers 1 < 2 = 4 < 3, fourth finger subequl in length to second; finger tips not expnded into discs. Hind limbs robust; TIBL 49% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle indistinct, smll, ovl; outer mettrsl tubercle bsent; no webbing between toes; first toe reduced; toe tips slightly enlrged; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe distinctly shorter thn third; subrticulr tubercles distinct, single. Skin on 321

52 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b c Fig. 37. Stumpffi mledict sp. nov. in life: ( c) prtype ZSM 3244/2012 (ZCMV 13504) from Lc Mudit, Montgne d Ambre Ntionl Prk. dorsum smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After eight yers in 70% ethnol, the dorsum is beige, with no pttern. No spots re present in the inguinl region. The dorsl surfce of the hed is s the bck. The color of the nostril is not distinct from the lterl hed colortion. The lterl surfce of the hed is grnulr brown, with irregulr crem dots. A crem stripe runs from the tympnum towrd the insertion of the rm. The color of flnks is continution of the lterl hed colortion, forming distinct color border with the dorsum. The flnk colortion is distinct from the ventrl colortion. The ventrl bdomen is crem with trces of brown speckling. The chin hs more intense brown flecking thn the bdomen. The ventrl thigh is crem. The shnk is ventrlly crem mottled with light brown. The trsus is ventrlly s the shnk. The sole of the foot is beige. Dorslly, the thigh is beige s the bck. The nterodorsl thigh is brown with irregulr crem dots, like the flnk. The posterodorsl surfce of the thigh is s the nterodorsl side. The shnk is dorslly beige. The trsus is s the shnk. The foot is lterlly beige nd medilly crem t its bse, fding to become crem distlly. The toes re mottled crem nd beige. The clocl region is not drker thn the rest of the thighs. The rms re dorslly nd lterlly s the dorsum nd flnk, nd ventrlly clen crem. The dorsomedil surfce of the hnd is crem. The fingers re brown with crem flecks. Color in life (holotype nd prtypes). Dorsum snd brown with slightly drker midregion. The flnk fdes from the dorsl colortion to lemon yellow. The dorsl forelimb is s the bck, with sooty crossbnd on the 322

53 VERTEBRATE ZOOLOGY 67 (3) 2017 b Fig. 38. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi mledict sp. nov. from Montgne d Ambre Ntionl Prk, recorded from prtype ZSM 3244/2012: () 1 s durtion section; (b) 6 s durtion section. The initil pek in mplitude t the beginning of ech note most probbly represents n rtifct produced by dmged recorder. forerm. The fingers re mottled snd brown nd gry, with whitish nnulus before ech terminl phlnge. The dorsl hindlimbs re s the bck, with the toes s the fingers. The ventrl colortion of the holotype in life is not known, but one prtype hd trnslucent lemon yellow ventrl skin with gry lip nd nterior chin, nd off-white speckles from the pectorl region decresing in density to the legs (Fig. 37c). The iris is russet. Vrition. For vrition in mesurements mong specimens, see Tble 3. In generl, ll exmined specimens gree strongly with the holotype in morphology. ZSM 3244/2012 is overfixed, resulting in rtefctul suprtympnic folds nd trnsverse hed fold. This specimen hs more boxy snout thn the holotype nd prtype ZSM 2169/2007. A reltively low degree of vribility ws noted in the colortion of specimens in preservtive: in ZSM 3244/2012 nd ZSM 2169/2007 the dorsum is lightly flecked with brown, nd the distinction of the dorsum nd flnk is less strong thn in the holotype; in ZSM 2169/2007, n hour-glss shped mrking is present on the nterior dorsum. In this specimen the dorsl surfce of the thigh is lso reticulted with brown; ZSM 3244/2012 hs brown interoculr br. Etymology. The species nme is the feminine nomintive single form of the Ltin djective mledictus = cursed, referring both to one of the collecting loclities (Lc Mudit) nd to the difficulty in finding this species. Distribution. This species is only known from Montgne d Ambre Ntionl Prk. Nturl history. The species occurs in rinforest. The clling mle ws found t night, hiding between roots on the forest floor. Cll. Advertisement clls were recorded by A. Rkotorison on 16 Februry 2012 in Montgne d Ambre Ntionl Prk, ner Lc Mudit, from the specimen ZSM 3244/2012. The cll consists of single short tonl note repeted in series t regulr intervls (Fig. 38). Numericl prmeters re s follows: cll durtion (= note durtion) ms (102 ± 2.8 ms; N = 4), inter-cll intervls ms (5927 ± 735 ms; N = 3), nd dominnt frequency t Hz (4833 ± 21.5 Hz, N = 4). It is importnt to mention tht cll recordings of this species my suffer from certin rtifcts produced by dmged recording device. Results thus must be treted with cution. Stumpffi mmitik sp. nov. (Figures 39 nd S102) Holotype. ZSM 3237/2012 (ZCMV 13524) (Figs. 12 f, 39 e, j, k), n dult mle, collected on 26 Jnury 2012 ner the Cmpement des Princes (c. S , E , 90 m bove se level), Ankrn Ntionl Prk, Antsirnn Province, Mdgscr by A. Rkotorison nd A. Rzfimnntso. Prtypes. ZSM 862/2003 (FG/MV ), collected on 15 Februry 2003 round Mhmsin in Ankrn Ntionl Prk by F. Glw, R.D. Rndriniin, nd A. Rzfimnntso; ZSM 307/2004 (FGZC 592) colleted on 27 Februry 2004 in Ankrn Ntionl Prk, pitfll site (S , E , 121 m.s.l.) by F. Glw, M. Puente nd R. Rndriniin; ZSM 375/2005 (FGZC 2725), collected on 13 Februry 2005 between Andrkt nd Andp by F. Glw, M. Vences, nd R.D. Rndriniin; ZSM 3232/2012 (ZCMV 13528), UADBA-A (orig. ZSM 3233/2012, ZCMV 13529), UADBA-A (orig. ZSM 3234/2012, ZCMV 13530), nd UADBA-A (orig. ZSM 3235/2012, ZCMV 13531), ll collected on 29 Jnury 2012 from site 4 km from Vohemr by A. Rkotorison nd A. Rzfimnntso; ZSM 228/2016 (ZCMV 13616), collected between in Ankrn Ntionl Prk (S , E ) by A. Rkotorison nd A. Rzfimnntso. 323

54 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Dignosis. One of severl morphologiclly similr but prtly symptric Stumpffi species from northern Mdgscr. This species hs been previously listed s Stumpffi sp. 5 in Vieites et l. (2009), Köhler et l. (2010), Klges et l. (2013), Scherz et l. (2016) nd Peloso et l. (2017). (1) Smll-sized species (mle SVL mm); (2) mnus with four fingers (first finger slightly reduced in length) nd pes with five toes (first toe slightly reduced in length); (3) terminl phlnges of fingers without, those of toes with slightly enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum smooth with few scttered tubercles or slightly tuberculr; (6) dorslly russet to snd brown, often with lrge centrl drk teddyber shped ptch which cn however be indistinct. Ventrlly trnslucent gry to plum with flecks of iridescent crem scttered cross the bdomen nd/or chest. No red color ventrlly, no shrp border between dorsl nd lterl color; (7) regulrly repeted single-note tonl cll. Distinguished from S. be, S. grndis, S. hr, S. kibomen, S. megsoni, S. roseifemorlis, nd S. stffordi by smller body size; from S. tridctyl by lower degree of digitl reduction; from S. miery by longer cll durtion nd less expressed length reduction of fingers; from S. psologloss by n unpulsed cll structure; from S. tetrdctyl by less expressed length reduction of first finger nd especilly first toe; from S. pygme, S. mdgscriensis, nd S. nlmin by slightly lrger body size nd shorter durtion of clls. The species is plced in clde A1 nd within this clde it is distinguished from S. ngeluci by shorter durtion of clls nd of inter-cll intervls; from S. mledict by shorter durtion of inter-cll intervls; from S. lrinki by shorter inter-cll intervls. Most similr morphologiclly nd biocousticlly to S. gimmeli nd S. ihrn but not the direct sister group of these species, strongly divergent in mitochondril nd nucler DNA, nd possibly differing from S. gimmeli by slightly shorter durtion of inter-cll intervls. Direct sister txon of S. huwei, but distinguished from this species by longer cll durtion nd shorter inter-cll intervl durtion. Description of the holotype. Specimen in good stte of preservtion, piece of its inner thigh removed s tissue smple. Body round; hed wider thn long, nrrower thn body; snout slightly pointed in dorsl view, pointed in lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis distinct, stright; lorel region slightly concve, slightly oblique; tympnum indistinct, bout 70% of eye dimeter; suprtympnic fold not visible; tongue long, brodening posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone rounded. Forelimbs slender; subrticulr tubercles distinct, single; outer metcrpl tubercle distinct, ovl; inner metcrpl tubercle distinct, ovl; fingers without webbing; first finger slightly reduced; reltive length of fingers 1 < 2 < 4 < 3, fourth finger slightly longer thn second; finger tips not expnded into discs. Hind limbs slender; TIBL 44% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle smll, round; outer mettrsl tubercle bsent; no webbing between toes; first toe reduced; toe tips slightly expnded; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe slightly shorter thn third; subrticulr tubercles distinct, single. Skin on dorsum smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After three yers in 70% ethnol, the dorsum is light brown. The bck is flecked with drk brown, forming fint X-shped mrking over the scpulr region. Dorsolterl regions re gry. Drk spots re present in the inguinl region. The dorsl surfce of the hed is s the bck. The nostril is drk brown. The lterl surfce of the hed is mottled drk brown, with crem spots behind the ngle of the jw. An lmost blck suprtympnic stripe is present, extending towrd the insertion of the rm. The flnks re mottled gry nd drk brown with crem speckling. A drk brown spot is present bove the insertion of the rm. The flnk colortion fdes rpidly to the crem of the trunk. The ventrl bdomen is crem with brown flecks. The chin is browner, with some crem spots. The ventrl thigh is s the trunk. The shnk is ventrlly brown with crem spots. The trsus is ventrlly s the shnk. The sole of the foot is brown. Dorslly, the thigh is lighter brown thn the dorsum, with lrge brown flecks. The posterodorsl nd nterior surfces of the thigh re brown flecked with crem. The shnk, trsus, nd foot re s the thigh. The toes re flecked with light nd drk brown. The clocl region is drk brown. The rms re s the legs, without crossbnds. The dorsomedil surfce of the hnd is mottled crem nd brown. The fingers re brown with smll crem stripes. The underside of the rm is s the ventrl bdomen. Color in life (holotype nd vrition). Dorsum russet medilly, fding through copper to snd brown lterlly, nd over the flnks drkening to ebony. Lterl hed ebony with few copper flecks, including on the tympnum nd nterior to the eye. A fint burnt umber chevron is present in the suprscpulr region. A series of ebony spots re present on the flnks, with one bove the insertion of the forelimb, nd two on the mid-flnk t different heights. The flnk hs lso numerous bluish flecks, decresing in density dorslly. The dorsl forelimb is proximlly russet, becoming incresingly mottled with drker brown distlly. The fingers re mottled snd brown nd burnt umber, with light nnulus before ech terminl phlnge. The hindlimbs re s the dorsum with flecks of ebony, the toes re s the fingers. The ventrl skin is trnslucent, light gry over the chin, plum over the nterior trunk with flecks of iridescent crem, becoming tupe over the posterior body nd legs. The soles of the feet re burnt umber. The iris is dorslly copper, nteriorly nd posteriorly rust red. The prtypes hve rther different color pttern, with snd brown dorsum nd centrl burnt-umber-demrcted teddyber-shped mrking of vrying intensity, tn to tupe lterl colortion, nd cremy venter. 324

55 VERTEBRATE ZOOLOGY 67 (3) 2017 b c d e j f h k g i Fig. 39. Stumpffi mmitik sp. nov. from Ankrn Ntionl Prk nd Vohemr in life: ( b) UADBA-A (ZCMV 13530); (c d) UADBA-A (ZCMV 13531); (f g) ZSM 3232/2012 (ZCMV 13528); (h i) UABA-A (ZCMV 13529) from Vohemr; (e, j k) holotype ZSM 3237/2012 (ZCMV 13524) from Ankrn Ntionl Prk. Vrition. For vrition in mesurements mong specimens see Tble 3. In generl, ll exmined specimens gree strongly with the holotype in morphology. In some specimens the cnthus rostrlis is slightly concve, nd the snout cn be slightly rounded in lterl view. A high degree of vribility ws noted in the colortion of specimens in preservtive; in the oldest specimen, ZSM 862/2003, the color is lrgely obscured nd the whole dorsum is homogeneous brown; the flnks re still lighter; the lterl hed is s the dorsum; crem flecks re bsent from the posterior thigh. It lso lcks drk spots bove the tympnum, rms, nd in the inguinl region. This discolortion my be ssocited with strong fixtion or ge. In the other prtypes, ventrl colortion is generlly consistent with the holotype, lthough in some cses it hs more or less brown. The drk dorsl mrkings of the holotype re more pronounced in severl specimens. In these, border is formed between the lighter brown hed nterior to the eyes nd the drker brown dorsl colortion. In few specimens, the body is gry insted of brown. The inguinl spots re bsent in two specimens (ZSM 862/2003 nd ZSM 207/2004) but present in ll others. The suprtympnic mrking nd the spot bove the insertion of the rm re bsent from these specimens s well, nd lso from ZSM 3754/2005, but re present in ll other specimens. In ddition, drk flnk spot is 325

56 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b c d Fig. 40. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi mmitik sp. nov. (1 s nd 6 s durtion section ech): ( b) Clls from Ankrn Ntionl Prk recorded in 2016 from prtype ZSM 228/2016 (ZCMV 13616); (c d) clls from ner Andp recorded in 1995 nd tenttively ssigned to this species. present in ZSM 3234/2012 nd ZSM 3235/2012, which is lcking from the holotype. A light vertebrl stripe is present in ZSM 3235/2012. The forerm hs crossbnd in ZSM /2012. A crossbnd is present on the mid-shnk of some specimens. Etymology. The species nme is noun in pposition to the genus nme, derived from the Mlgsy word mmitik = sting, becuse the first uthor got stung severl times by wsp whilst she collected the specimens in Vohemr. Distribution. This species is known from three loclities: (1) Ankrn Ntionl Prk (type loclity), (2) Andrkt-Andp, nd (3) Vohemr. Specimens from Ankrn Ntionl Prk re geneticlly slightly divergent from those originting from Vohemr nd Andrkt-Andp. The rnge encompsses elevtions from m bove se level. Nturl history. The species occurs in dry forest in Ankrn NP s well s in mngo plnttion in Vohemr. In Ankrn NP, one clling mle ws found clling from bush bout 5 cm bove the ground, nd second one ws found hiding between brnches. The specimens collected in Vohemr were found in dense lef litter. All specimens were collected t night. Cll. The dvertisement cll of S. mmitik consists of single short tonl note repeted in series t regulr intervls (Fig. 40). Recordings re vilble for two different loclities: (1) Recorded by A. Rkotorison in Jnury 2016 t the type loclity, Ankrn Ntionl Prk, from prtype ZSM 228/2016 (ZCMV 13616), with the following numericl prmeters: cll durtion (= note durtion) ms (79 ± 7.3 ms; N = 4), inter-cll intervls ms (1569 ± 281 ms; N = 3), nd dominnt frequency t Hz (4619 ± 54 Hz, N = 4). (2) 326

57 VERTEBRATE ZOOLOGY 67 (3) 2017 b c d Fig. 41. Stumpffi sort sp. nov. from Sort forest in life: ( b) specimen not ssignble to voucher; (c d) ZSM 1643/2012 (FGZC 3618). Recorded by F. Glw on 19 Februry 1995 ner Andp (Vences et l., 2006: CD3, trck 44, s Stumpffi sp.) nd tenttively ssigned to this species, with the following numericl prmeters: cll durtion ms (103 ± 9 ms; N = 20), inter-cll intervls ms (955 ± 264 ms; N = 19), nd dominnt frequency t Hz (4749 ± 257 Hz, N = 20). Cll recordings obtined t Vohemr in 2012 suffer from severl rtifcts but re in principl identifible s belonging to this species. Stumpffi sort sp. nov. (Figures 41 nd S103) Holotype. ZSM 1644/2012 (FGZC 3621) (Figs. 12 g, S103 c d), collected on 26 November 2012 on the Sort Mssif (S , E , 1339 m bove se level), Antsirnn Province, Mdgscr by F. Glw, O. Hwlitschek, F.M. Rtsovin, A. Rkotorison, T. Rjofirison, nd A. Rzfimnntso. Prtypes. ZSM 1643/2012 (FGZC 3618), ZSM 1645/2012 (FGZC 3622), UADBA-A (FGZC 3617), UADBA-A (FGZC 3619), UADBA-A (FGZC 3620), nd UADBA-A (FGZC 3623), ll with sme collecting dt s holotype. Dignosis. One of severl morphologiclly similr but prtly symptric Stumpffi species from northern Mdgscr. This species hs not been listed s cndidte species in previous publictions. (1) Smll-sized species (SVL mm); (2) mnus with four fingers (first finger very slightly reduced in length) nd pes with five toes (first toe slightly reduced in length); (3) terminl phlnges of fingers without, those of toes with slightly enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum slightly to modertely grnulr; (6) dorslly tupe, occsionlly with horizontl bnds of tn or gry-brown, with numerous light spots. Severl ebony mrkings on the flnk, including one in the inguinl region. Ventrl skin trnslucent with dense pttern of flecks of crem on nd posterior to the chest; the throt occsionlly sooty or simply trnslucent. No red color ventrlly, no shrp border between dorsl nd lterl color. 327

58 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Distinguished from S. be, S. grndis, S. hr, S. kibomen, S. megsoni, S. roseifemorlis, nd S. stffordi by smller body size; from S. tridctyl by lower degree of digitl reduction; from S. miery by longer cll durtion nd less expressed length reduction of fingers; from S. tetrdctyl by less expressed length reduction of first finger nd especilly first toe; from S. nlmin, S. mdgscriensis, S. pygme, nd S. psologloss by lrger body size. The species is plced in clde A1 nd within this clde it is morphologiclly rther similr to the mjority of species, but is plced phylogeneticlly prt from most of them. It is prt of clde contining S. lrinki, nd is the direct sister txon to cndidte species (S. sp. C7) from Mrojejy. Although we lck biocoustic dt for this txon, we here suggest its sttus s new species S. sort due to its high genetic differentition from its closest reltive (pirwise 16S distnce from S. lrinki: 9.8%), s well s its grnulr dorsl skin (smooth in S. lrinki) nd ventrl crem flecks nd occsionlly drk throt (more uniform in S. lrinki). We re wre tht this species hypothesis requires further testing in the future. Description of the holotype. Specimen in good stte of preservtion, left thigh muscle removed s tissue smple. Body round; hed slightly wider thn long, nrrower thn body; snout rounded in dorsl view, rounded in lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis distinct, concve; lorel region slightly concve, slightly oblique; tympnum indistinct, bout 55% of eye dimeter; suprtympnic fold not visible; tongue long, expnding slightly posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone rounded. Forelimbs slender; subrticulr tubercles single, indistinct; outer metcrpl tubercle smll, ovl; plmr tubercle round; inner metcrpl tubercle smll, ovl; fingers without webbing; first finger slightly reduced; reltive length of fingers 1 < 2 = 4 < 3, fourth finger subequl in length to second; finger tips not or wekly expnded into discs. Hind limbs slender; TIBL 44% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle smll, round; outer mettrsl tubercle bsent; no webbing between toes; first toe reduced; toe tips slightly expnded; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe distinctly shorter thn third; subrticulr tubercles indistinct, single. Skin on dorsum smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After three yers in 70% ethnol, the dorsum is metllic silver. Drker silver res re present t the posterior end the dorsum, just behind the scpulr region, nd behind the eyes. Circulr diffuse blck mrkings re present behind the scpulr region. An oblong drk brown mrking is present in the inguinl region. The dorsl surfce of the hed is s the bck. The nostril is surrounded by brown. The lterl surfce of the hed is s the dorsum. A drk brown suprtympnic mrking is present, not exceeding the tympnum posteriorly. The flnks re s the dorsum. A lrge drk brown spot is present dorslly nd posteriorly to the insertion of the rm. This is followed by series of smller drk brown spots. These form border between the dorsl nd ventrl colortion. The ventrl trunk is crem flecked with brown, most intensely over the pectorl girdle. The chin is s the ventrl trunk. The ventrl thigh is brown spotted with crem. The shnk is ventrlly brown with few crem flecks. The trsus is ventrlly s the shnk. The sole of the foot is brown. Dorslly, the thigh is drk silver like the posterior dorsum. The posterodorsl surfce of the thigh is brown with crem spots. The shnk is s the thigh, with drk crossbnd midwy long its length. The trsus is s the shnk but lcks crossbnds. The exterior of the foot is s the trsus, with single drk spot; the medil foot is brown with crem flecks. The toes re mottled gry nd crem. The clocl region is drk brown. The rms re s the dorsum. A drk crossbnd is present on the lower rm. The dorsomedil surfce of the hnd is mottled gry nd crem. The fingers hve smll gry nd crem stripes. The underside of the rm is s the ventrl trunk. Color in life (prtype). Dorsl bse color tupe, with numerous crem to tn spots over the whole dorsum. Wide horizontl bnds of tn nd gry-brown cn be present on the dorsum. A lrge ebony mrking bordered with thin line of crem cn be present in the inguinl region. Further crem-bordered ebony mrkings re present round the nostril, below the eye, in the posterodorsl prt of the tympnum, nd in two spots of vrying shpe on the flnks, which re continuous with the dorsum. The dorsl forelimb is bisque, with sooty crossbnd on the forerm. The toes re bisque with whitish nnulus before ech terminl phlnge. The dorsl hindlimb is s the dorsum, without crossbnds. An ebony mrking cn be present on top of the foot. The throt skin cn be trnslucent nd mottled with lighter flecks, or sooty with wek lemon flecks. The ventrl trunk s skin is trnslucent, with crem flecks or speckles mostly concentrted on nd posterior to the pectorl girdle. The ventrl legs hve fint crem mottling. Vrition. For vrition in mesurements mong specimens, see Tble 3. In generl, ll exmined specimens gree strongly with the holotype in morphology, lthough two prtypes (ZSM 1643/2012 nd 1645/2012) hve pointed snouts in dorsl nd lterl view. An intermedite degree of vribility ws noted in the colortion of specimens; ZSM 1643/2012 is similr to the holotype in pttern, but brown colors previl insted of silver, with slightly fewer drk brown mrkings; ZSM 1645/2012 is dorslly pinkish, with the inguinl nd lterl spots nd brown clocl region nd shnk crossbnd of the other two specimens, more blck spots on the dorsum, one on the lower bck, nother lmost chevron shped over the scpulr region, nd few spots on the posterior hed. The legs of this specimen hve more drk brown spots thn the holotype, nd re lso pinkish in color. 328

59 VERTEBRATE ZOOLOGY 67 (3) 2017 Etymology. The species nme is noun in pposition to the genus nme, nd refers to the type loclity of the new species, the Sort forest in north-estern Mdgscr. Distribution. This species is known only from its type loclity in the Sort Mssif. Cll. Unknown. Stumpffi ynniki sp. nov. (Figures 42 nd S104) Holotype. ZSM 629/2014 (DRV 6413) (Fig. 12), collected on 29 June 2010 in forest frgment beside Route Ntionle 31, ner the border between the Belnn nd Antsohihy Districts (S , E , 1187 m bove se level), Mhjng Province, Mdgscr by M. Vences, D.R. Vieites, R.D. Rndriniin, F.M. Rtsovin, S. Rsmison, A. Rkotorison, E. Rjeririson, F. Rndrinsolo, nd T. Rjofirison. Prtypes. ZSM 1825/2010 (ZCMV 12600), UADBA-A (ZCMV 12601), UADBA (ZCMV 12599), nd UADBA (DRV 6412), ll with sme collecting dt s holotype. Dignosis. A poorly known species from north-western Mdgscr, previously listed s Stumpffi sp. 31 in Klges et l. (2013). (1) Smll-sized species (SVL mm); (2) mnus with four fingers (first finger modertely reduced in length) nd pes with five toes (first toe distinctly reduced in length); (3) terminl phlnges of fingers nd toes without enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum smooth; (6) dorslly with lrge nd well-contrsted centrl drk teddyber shped middorsl mrking, bordered on the hed nd dorsolterl body by lighter colortion. Ventrlly gryish with very little light mottling in preservtion. No red color ventrlly, no shrp border between dorsl nd lterl color. Distinguished from S. be, S. grndis, S. hr, S. kibomen, S. megsoni, S. roseifemorlis, nd S. stffordi by distinctly smller body size; from S. tridctyl by lower degree of digitl reduction; from S. miery by less expressed length reduction of fingers; from S. tetrdctyl by somewht lower degree of length reduction of first toe. The new species is phylogeneticlly plced in clde A2, nd differs from ll species in clde A1 (S. ngeluci, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. mmitik, S. mledict, S. sort) by slightly smller body size. Within clde A2 S. ynniki is distinguished from S. mdgscriensis by the lck of shrp border between dorsl nd lterl color, nd from S. psologloss by completely smooth dorsl skin. It is morphologiclly most similr to S. pygme nd S. nlmin, but strongly divergent in mitochondril nd nucler DNA from these species. Stumpffi ynniki is phylogeneticlly sister to S. nlmin, but ppers to differ slightly by hving reltively shorter hnds nd feet (Klges et l., 2013) nd, despite geogrphicl proximity of their rnges, these two species re geneticlly highly divergent. Fig. 42. Stumpffi ynniki sp. nov. from forest frgment beside Route Ntionle 31, ner the border between the Belnn nd Antsohihy Districts in life; prtype ZSM 1825/2010 (ZCMV 12600). Description of the holotype. Specimen in good stte of preservtion, left thigh muscle removed s tissue smple. Jw broken. Body stocky; hed wider thn long, nrrower thn body; snout rounded in dorsl view, rounded in lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis indistinct, stright; lorel region concve, oblique; tympnum indistinct, bout 80% of eye dimeter; suprtympnic fold bsent; tongue long, brodening posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone ovl. Forelimbs slender; subrticulr tubercles indistinct, single; outer metcrpl tubercle not recognizble; inner metcrpl tubercle smll, round; fingers without webbing; first finger reduced, second nd fourth fingers slightly reduced; reltive length of fingers 1 < 2 = 4 < 3, fourth finger subequl in length to second; finger tips not expnded into discs. Hindlimbs slender; TIBL 43% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle smll, round; outer mettrsl tubercle bsent; no webbing between toes; first toe strongly reduced; toe tips not expnded; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe slightly shorter thn third; subrticulr tubercles distinct, single. Skin on dorsum smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After five yers in 70% ethnol, the dorsum is light brown, with drk brown brod centrl stripe, nd crem vertebrl stripe. A drk spot is present in the inguinl region. The dorsl surfce of the hed is s the bck. The nostril is drk brown. The lterl surfce of the hed is drk brown. A drk stripe divides dorsl nd flnk colortion, running forwrd to the eye. Below it, the flnks re crem reticulted with brown. Flnk colortion extends onto the ventrl trunk. The chin is plin crem. The ventrl thigh is s s the chin. The shnk is ventrlly s the thigh. The trsus is ventrlly s the shnk. The sole of the foot is drk brown. Dor- 329

60 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs slly, the thigh is light brown with drk brown oblique crossbnd. The posterodorsl surfce of the thigh hs white line running its length, below which it is brown with white dots. The shnk is light brown with n oblique crossbnd. The trsus is s the shnk with n oblique crossbnd. The foot is s the shnk, with drk spot on its externl fce. The toes re brown with light brown flecks. The clocl region is drk brown. The rms re s the legs. A drk crossbnd is present on the forerm. The whole hnd is light brown. The underside of the rm is s the chin. Color in life (prtype). The only existing photo of specimen of this species in life (Fig. 42) hs distinct purple tinge to it. This my be result of chromtic berrtion, nd therefore colors re not nmed here in detil, but rther the overll pttern of the specimen is described: A centrl, drk, teddyber-like ptch is present on the mid-dorsum, bordered on the nterior hed nd dorsolterl body by lighter colortion. The lterl hed nd flnk re continuous, drker colortion, with blck mrking posterior to the insertion of the forelimb. The dorsl forelimb is brownish mottled with blck nd gry, s is the hnd. The dorsl hindlimb is continuous with the mid-dorsl colortion, but more mottled. The ventrl color in life is not known. Vrition. For vrition in mesurements mong specimens, see Tble 3. The prtype ZSM 1825/2010 is more or less identicl to the holotype in morphology, but differs strongly in colortion from the holotype. The dorsl color of this specimen is light brown, lterlly gry; the dorsum is flecked with drk brown; inguinl spots re present s in the holotype; the lterl surfce of the body is s in the holotype, but more solidly brown thn reticulted; the whole venter is more crem thn the holotype; the thighs lck the white stripe, but otherwise the leg colortion is similr, lbeit lighter; the rm is lso similr. Etymology. The species nme is ptronym, dedicted by the first uthor to her honorry nephew Ynnik Hill, to introduce him to the herpetofun of Mdgscr. Distribution. The species is known only from its type loclity, forest frgment beside Route Ntionle 31, ner the border between the Belnn nd Antsohihy Districts. Nturl history. Specimens were collected during the dy, from the lef litter in rinforest frgment, reltively close to strem. Cll. Unknown. 2. Lrge-sized species of clde A This section contins clde of four Stumpffi endemic to smll rnges in the North of Mdgscr tht form monophyletic group (subclde A3), sister to ll other species in clde A. Three of the species in subclde A3 re the lrgest known species of the genus. These lrgesized Stumpffi species hve no digitl reduction nd hve distinctly enlrged discs on terminl phlnges of fingers nd toes, nd they pper to be specilized to live in krstic environments, prticulrly in crevices nd cves. They my be superficilly confused with some species of Rhombophryne, but cn be distinguished from ll morphologiclly similr species of tht genus (e.g. R. minut) by the bsence of vomerine teeth (vs. presence in Rhombophryne). Stumpffi be Köhler, Vences, D Cruze & Glw, 2010 (Figure 43) Nme-bering type. Holotype ZSM 1668/2008 (FGZC 1612) (Fig. 43 b), from the edge of the river below the Point de Vue Petit Tsingy, S 2.95, E , c. 90 m.s.l., Ankrn Specil Reserve, Antsirnn Province, northern Mdgscr ccording to the originl description. Identity nd dignosis. This species hs been listed s Stumpffi sp. 23 in Vieites et l. (2009) (not included in Glw & Vences, 2007 nd Wollenberg et l., 2008). A lrge species of Stumpffi known only from the krstic Ankrn Mssif. (1) Lrge-sized species (femle SVL 25.2 mm); (2) mnus with four fingers (none of them reduced in length) nd pes with five toes (none of them reduced in length); (3) terminl phlnges of fingers nd toes with enlrged discs; (4) reltive hnd nd foot length, HAL/SVL 0.3, FOTL/SVL 0.6; (5) dorsum smooth; (6) light brown with indistinct pttern. Conceled surfces of hindlimbs with ornge-red color. Ventrlly uniformly gryish. Juveniles pprently with different colortion, blckish with mny smll white dots nd brown ptches on dorsum; (7) tonl single-note cll of ms durtion, emitted in regulr series (inter-cll intervls ms). Distinguished from S. psologloss, S. nlmin, S. ngeluci, S. gimmeli, S. grndis, S. huwei, S. ihrn, S. lrinki, S. mdgscriensis, S. mledict, S. mmitik, S. miery, S. pygme, S. sort, S. tetrdctyl, S. tridctyl, nd S. ynniki by distinct red color on conceled prts of hindlimbs. From the species listed, nd from S. kibomen nd S. roseifemorlis by lrger discs on fingers, nd lck of digitl length reduction. From ll these species except for S. grndis, S. kibomen, nd S. roseifemorlis by lrger body size. Specimens exmined. Holotype ZSM 1668/2008 (FGZC 1612), collected on 12 Februry 2008 from the edge of the river below the Point de Vue Petit Tsingy in Ankrn Ntionl Prk (S12.95 E , 90 m.s.l.) by Jcques, N. D Cruze, M. Frnzen, F. Glw, nd J. Köhler; prtype ZSM 1664/2008 (FGZC 1643), juvenile, collected on 13 Februry 2008, 20 m west of holotype loclity by N. D Cruze, M. Frnzen, F. Glw, nd J. Köhler. 330

61 VERTEBRATE ZOOLOGY 67 (3) 2017 b c Fig. 43. Stumpffi be from Ankrn Ntionl Prk in life: ( b) holotype ZSM 1668/2008 (FGZC 1612); (c) juvenile prtype, ZSM 1664/2008 (FGZC 1643). b Fig. 44. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi be, bsed on the recording of Lttenkmp et l. (2016). () 1 s durtion section; (b) 6 s durtion section. Distribution. This species is known from its type loclity t the estern side of Ankrn Ntionl Prk. Nturl history. Specimens were found on rocks nd vegettion 50 cm bove the ground in krstic limestone res; one specimen contined remins of lrge blck nts in its stomch (Köhler et l., 2010). A mle ws observed clling from crck in limestone rock in riverbed (Lttenkmp et l., 2016). Cll. The dvertisement cll hs been described by Lttenkmp et l. (2016). It consists of single short tonl note repeted in series t regulr intervls (Fig. 44). Cll prmeters s tken from this publiction were s follows: cll durtion (= note durtion) ms (174 ± 3 ms; N = 7), inter-cll intervl ms (919 ± 84 ms; N = 6), dominnt frequency Hz (3912 ± 11 Hz; N = 7). Stumpffi hr Köhler, Vences, D Cruze & Glw, 2010 (Figure 45) Nme-bering type. Holotype ZSM 1666/2008 (FGZC 1813) (Fig. 45 b), from smll creek on the western flnk of Nosy Hr, S, E [S , E ], 20 m.s.l., Antsirnn Province, northern Mdgscr ccording to the originl description. 331

62 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b c d Fig. 45. Stumpffi hr from Nosy Hr in life: ( b) holotype ZSM 1666/2008 (FGZC 1813); (c d) prtype ZSM 2195/2007 (FGZC 1293). Identity nd dignosis. This species hs been listed s Stumpffi sp. 22 in Vieites et l. (2009) (not included in Glw & Vences, 2007 nd Wollenberg et l., 2008). A lrge-sized species of Stumpffi known only from the islnd Nosy Hr. (1) Lrge-sized species (femle SVL mm); (2) mnus with four fingers (none of them reduced in length) nd pes with five toes (none of them reduced in length); (3) terminl phlnges of fingers nd toes with enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum smooth; (6) light brown with indistinct pttern. Ventrlly gryish with numerous smll whitish dots. Distinguished from S. psologloss, S. nlmin, S. ngeluci, S. gimmeli, S. grndis, S. huwei, S. ihrn, S. kibomen, S. lrinki, S. mdgscriensis, S. mledict, S. mmitik, S. miery, S. pygme, S. roseifemorlis, S. sort, S. tetrdctyl, S. tridctyl, nd S. ynniki by lrger discs on fingers, nd lck of digitl length reduction; nd from ll these species except for S. grndis, S. kibomen nd S. roseifemorlis by lrger body size. The species is included in clde A3, nd therefore not closely relted to ny of the previously mentioned species. Distinguished from S. be by the lck of red color on conceled surfces of hindlimbs. Specimens exmined. Holotype ZSM 1666/2008 (FGZC 1813), collected on 22 Februry 2008 from smll creek on the western flnk of Nosy Hr, Antsirnn (S , E49.175, 20 m.s.l.), by F. Glw nd J. Köhler; prtype ZSM 1665/2008 (FGZC 1801), sme collecting dt s holotype; prtypes ZSM 2195/2007 (FGZC 1293), ZSM 2196/2007 (FGZC 1296), nd UADBA-A (FGZC ) collected on 7 Mrch 2007 from cve entrnce on the western flnk of Nosy Hr, Antsirnn (S , E , 9 m.s.l., by H. Enting, F. Glw, nd J. Köhler. Distribution. This species is known only from its type loclity t the western slope of Nosy Hr. Nturl History. Found in forest long smll creek nd inside nd t the entrnce of cve. A lrge number of rthropods of considerble size were found in the stomch of one specimen (Köhler et l., 2010). Cll. Unknown. Stumpffi megsoni Köhler, Vences, D Cruze & Glw, 2010 (Figure 46) Nme-bering type. Holotype ZSM 1663/2008 (FGZC 1951) (Fig. 46), from Ampombofofo, from smll cve, S, E, 20 m.s.l., Antsirnn Province, northern Mdgscr ccording to the originl description. 332

63 VERTEBRATE ZOOLOGY 67 (3) 2017 b Fig. 46. Stumpffi megsoni holotype ZSM 1663/2008 (FGZC 1951) from Ampombofofo in life: () dorsolterl nd (b) ventrl view. Identity nd dignosis. This species hs been listed s Stumpffi sp. 21 in Vieites et l. (2009) (not included in Glw & Vences, 2007 nd Wollenberg et l., 2008). A modertely lrge-sized species of Stumpffi, possibly less specilized to krstic evironments nd cves thn the other three species in this subclde. (1) Modertely lrgesized species (femle SVL mm); (2) mnus with four fingers (none of them reduced in length) nd pes with five toes (none of them reduced in length); (3) terminl phlnges of fingers nd toes with only very slightly enlrged discs; (4) reltive hnd nd foot length, HAL/SVL 0.2 3, FOTL/SVL 0.7; (5) dorsum smooth with smll tubercles; (6) light brown with indistinct pttern. Ventrlly uniformly gryish without pttern. Distinguished from S. psologloss, S. nlmin, S. ngeluci, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. mdgscriensis, S. mledict, S. mmitik, S. miery, S. pygme, S. sort, S. tetrdctyl, S. tridctyl, nd S. ynniki by lrger body size, nd from mny of these species by lower degree of digitl length reduction; from S. kibomen, nd S. roseifemorlis by the lck of red color on ventrl surfces; from S. grndis by lck of distinct blck-light gry/blue mrbling ventrlly. The species is included in clde A3, nd therefore not closely relted to ny of the previously mentioned species. Distinguished from S. be by the lck of red color on conceled surfces of hindlimbs; nd from S. hr nd S. be by smller terminl discs on fingers nd toes. Specimens exmined. Holotype ZSM 1663/2008 (FGZC 1951), collected on 23 Februry 2007 t Ampombofofo from smll cve (S , E , 20 m.s.l.) by S. Megson; prtype ZSM 1662/2008 (FGZC 1951), collected on 21 November 2006 t the sme loclity s holotype, by S. Megson. Distribution. This species is known from its type loclity Ampombofofo in northernmost Mdgscr. Nturl history. Specimens were found on wlls in smll limestone cve on steep slope, close to primry forest nd strem, but lso in secondry hbitt. Body infltion nd deth-feigning were observed s probble ntipredtor mechnisms (Köhler et l., 2010). Cll. Unknown. Stumpffi stffordi Köhler, Vences, D Cruze & Glw, 2010 (Figure 47) Nme-bering type. Holotype ZSM 1667/2008 (FGZC 1674) (Fig. 47, b, c), from lrge cve, S, 49 20, c. 260 m.s.l., Montgne des Frnçis, Antsirnn Province, northern Mdgscr ccording to the originl description. Identity nd dignosis. This species hs been listed s Stumpffi sp. 24 in Vieites et l. (2009) (not included in Glw & Vences, 2007 nd Wollenberg et l., 2008). The lrgest of ll Stumpffi, known only from the krstic mssif of Montgne des Frnçis. (1) Lrge-sized species (mle SVL mm); (2) mnus with four fingers (none of them reduced in length) nd pes with five toes (none of them reduced in length); (3) terminl phlnges of fingers nd toes with enlrged discs; (4) reltive hnd nd foot length, HAL/SVL 0.3, FOTL/SVL 0.6; (5) dorsum smooth with few wekly expressed tubercles; (6) reddish brown or gryish brown with indistinct pttern. Ventrlly gryish with some drk mottling prticulrly on the throt. Distinguished from S. psologloss, S. nlmin, S. ngeluci, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. mdgscriensis, S. mledict, S. mmitik, S. miery, S. pygme, S. sort, S. tetrdctyl, S. tridctyl, nd S. ynniki by lrger body size, lrger discs on fingers, nd from most of these species by the lck of digitl length reduction. The species is included in clde A3, nd thus not closely relted to ny of the previously mentioned species. Distinguished from S. be by the lck of red color on conceled surfces of hindlimbs, from S. megsoni by 333

64 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b c d e Fig. 47. Stumpffi stffordi from Montgne des Frnçis in life: ( c) holotype, ZSM 1667/2008 (FGZC 1674); (d e) prtype UADBA (FGZC 1601). lrger body size, nd from S. hr by the lck of whitish spotting on venter. Specimens exmined. Holotype ZSM 1667/2008 (FGZC 1674), collected on 16 Februry 2008 on Montgne des Frnçis (S , E49.333, 260 m.s.l.) by N. D Cruze, M. Frnzen, F. Glw, J. Köhler, nd Z. Ngy; prtype UADBA (FGZC 1601), collected on 11 Februry 2008 t sme loclity s holotype by N. D Cruze. Distribution. This species is known only from its type loclity on Montgne des Frnçis. Nturl history. Specimens were found deep inside krstic cve, with the holotype specimen perching t the edge of smll puddle (Fig. 47c) (Köhler et l., 2010). Cll. Advertisement cll unknown. The distress cll hs been described by Köhler et l. (2010). 3. Smll-sized or miniturized species of cldes B nd D For convenience, we include in this cluster severl species of smll-sized Stumpffi tht belong to two distinct cldes, B nd D (Fig. 3). Most of these species re chrcterized by distinct digitl reduction, yet cldes B nd D do not pper to be sister groups, nd clde B lso includes one lrge-sized species described in the next section. Stumpffi miery Ndrintso, Riemnn, Vences, Klges, Rminoso, Rödel & Glos, 2013 (Figure 48) Nme-bering type. Holotype ZMB (#_611) (Fig. 48, d) from Mdgscr, Rnomfn region, Ambolo forest frgment, S , E (site nme: FFst04), pp. 660 m.s.l., ner smll strem ccording to the originl description. Identity nd dignosis. Stumpffi miery is smll-sized species included in clde B. Its occurrence in the southern centrl est t comprtively low elevtions is striking, s only one other species (S. nigrorubr sp. nov.) is known from this re. (1) Smll-sized species (SVL mm); (2) mnus with four fingers (first finger very strongly reduced in length; lso second nd fourth fingers short) nd pes with five toes (first toe strongly reduced in length, lmost rudimentry); (3) terminl phlnges of fingers nd toes without enlrged discs; (4) reltive hnd nd foot length, HAL/SVL 0.15, FOTL/SVL ; (5) dorsum smooth or slightly grnulr; (6) light brown to reddish brown dorsl color, typiclly with distinct drk brown teddyber shped centrl mrking, nd often dditionl drk ptches lterlly. Ventrlly gry with drk gry mottling but without contrsted ventrl colortion, red color elements on ventrl side, or shrp continuous color border between dorsum nd flnks; (7) regulrly repeted short single-note chirp cll. Distinguished from S. be, S. grndis, S. hr, S. kibomen, S. megsoni, S. roseifemorlis, nd S. stffordi by distinctly smller body size; from S. tridctyl by lower degree of digitl reduction; from S. tetrdctyl by more expressed length reduction of first finger nd somewht less expressed length reduction of first toe. The stronger first toe reduction lso distinguishes S. miery from ll species in clde A, including the roughly similr-sized or smller sized S. psologloss, S. nlmin, S. ngeluci, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. mdgscriensis, S. mledict, S. mmitik, S. miery, S. pygme, S. sort, S. tetrdctyl, S. tridctyl, nd S. ynniki. Further distinguished from most of the species in clde A (ll for which clls re known) by shorter cll durtion. Specimens exmined. Holotype ZMB (#_611) collected on 1 Mrch 2011 t Ambolo forest frgment (S , E , 660 m.s.l.) by J.C. Riemnn nd A. Telo; prtype ZMB (#_677), collected on 10 Mrch 2011 t Andlngin, ner Ifndin (S , E , 515 m.s.l.) by J.C. Riemnn nd A. Telo; prtype ZMB (#_768) collected on 4 April 2011 t 334

65 VERTEBRATE ZOOLOGY 67 (3) 2017 b c d e Fig. 48. Dorsolterl nd ventrl views of Stumpffi miery from the Rnomfn region, southestern Mdgscr, in life: (, d) mle holotype ZMB 77453; (c) mle prtype ZMB 77454; (e) mle prtype ZSM 2447/2007. Ambolo forest frgment (S , E47.509, 640 m.s.l.) by J.C. Riemnn nd A. Telo; prtype ZSM 121/2011 (#_804) collected on 6 April 2011 in coffee nd bnn plnttion in Andlngin (S S, E , 450 m.s.l.) by J.C. Riemnn nd A. Telo; prtype ZMB ( #_09) collected on 9 Mrch 2010 t Ambolo forest frgment (S , E , 670 m.s.l.) by S.H. Ndrintso, J.C. Riemnn, J. Solo, A. Telo, nd M.-O. Rödel; prtype UADBA-A (#_621) collected on 2 Mrch 2011 in forest c. 335

66 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b Fig. 49. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi miery from Ambohitsr (prtype ZSM 244/2007), recorded in () 1 s durtion section; (b) 6 s durtion section. 150 m to next strem (S , E , 680 m.s.l.) by J.C. Riemnn nd A. Telo; prtype ZSM 2447/2007 (ZCMV 5868) collected on 3 Mrch 2007 t Ambohitsr (S E , 294 m.s.l.) by M. Vences; prtype UADBA-A (#_770) collected on 4 April 2011 t Ambolo forest frgment (S , E , 700 m.s.l.) by J.C. Riemnn nd A. Telo; prtype UADBA-A (#_1393) collected on 2 Februry 2011 t Andlngin (S , E , 490 m.s.l.) by S.H. Ndrintso nd J. Solo; prtype UADBA-A (#_1738) collected on 10 Mrch 2011 in coffee nd bnn plnttion in Andlngin (S , E , 450 m.s.l.) by J.C. Riemnn nd A. Telo. Vrition. Although our 16S tree (Fig. 3) only contins single specimen of S. miery, DNA sequences of nother segment of the 16S gene re vilble from three dditionl specimens (see Ndrintso et l., 2013). Distribution. This species is known from three loclities in the Rnomfn region: (1) Ambolo forest frgment (type loclity), (2) Andlngin, ner Ifndin, nd (3) Ambohitsr. Nturl history. Summrized from Ndrintso et l. (2013). Clling mles were found in the lef litter of rinforest. Specimens were lso collected in degrded forest frgments nd mixed coffee bnn plnttion. Clling mles were herd in the fternoon but lso t night, usully from rther hidden positions in the lef litter. Cll. The dvertisement cll of S. miery consists of very short single note repeted in series t regulr intervls (Fig. 49). Numericl cll prmeters for the mle holotype (ZMB 77453) recorded on 1 Mrch 2011, t 20.5 C, in Ambolo forest frgment re s follows (ccording to Ndrintso et l., 2013): cll durtion (= note durtion) ms (73 ± 12, N = 10), inter-cll intervls ms (3102 ± 456, N = 10), nd dominnt frequency t Hz (8057 ± 136.9, N = 10). For the prtype (ZSM 2447/2007) recorded on 3 Mrch 2007 in the rinforest ner Ambohitsr, t n estimted temperture of 25 C, prmeters re s follows: cll durtion ms (47 ± 5, N = 10), inter-cll intervls ms (2105 ± 243, N = 10), nd dominnt frequency t Hz (7877 ± 109, N = 10). Stumpffi tridctyl Guibé, 1975 (Figures 50 nd S105) Nme-bering type. Holotype MNHN from Mssif du Mrojézy ccording to the originl description. Identity nd dignosis. Stumpffi tridctyl is highly miniturized species with strong digitl reduction, known only from the Mrojejy Mssif in north-estern Mdgscr, tht forms with its sister species (described below s S. contumeli sp. nov.) clde D, which is phylogeneticlly plced sister to ll other Stumpffi. (1) Miniturized species (SVL mm); (2) mnus with one finger only (first finger not visible; second nd fourth fingers reduced to smll knobs; third finger brodened to tringulr shpe) nd pes with three toes (first toe not recognizble, second toe reduced to smll knob); (3) terminl phlnges of fingers nd toes without enlrged discs; (4) reltive hnd nd foot length HAL/SVL , FOTL/SVL ; (5) dorsum smooth; (6) reddish brown dorslly, with distinct color border to the drk brown flnks. Ventrlly without contrsted ventrl colortion or red color elements; (7) regulrly repeted short single-note frequency-modulted whistle cll. Distinguished from ll nominl species of Stumpffi by stronger digitl reduction with only one finger nd three toes recognizble (shred with the new spe- 336

67 VERTEBRATE ZOOLOGY 67 (3) 2017 b c e f d g Fig. 50. Stumpffi tridctyl from Mrojejy Ntionl Prk in life: ( b) ZSM 527/2016 (ZCMV 15282) from Mrojejy, high ltitude; (c d) unctlogued specimen from Mrojejy, high ltitude; (e) ZSM 382/2005 (FGZC 2844) from Mrojejy, Cmp Simpon; (f g) femle specimen ZSM 526/2016 (ZCMV 15284) from Mrojejy, Cmp Simpon. cies S. contumeli sp. nov. nd S. obscoen sp. nov. described below). Also distinguished from most other Stumpffi by its frequency-modulted cll. Phylogeneticlly the species is sister to the new species S. contumeli sp. nov., the only other known representtive of clde D; for its differentition from tht species, see its description below. Specimens exmined. Holotype MNHN , nd prtypes MNHN , ll collected from the Mrojejy Mssif by C. Blnc. Referred specimens: ZSM 382/2005 (FGZC 2844) collected on 17 Februry 2005 from Cmp Simpon, Mrojejy Ntionl Prk (S E , 1326 m.s.l.), by F. Glw, R.D. Rndriniin nd M. Vences; ZSM 526/2016 (ZCMV 15284) nd ZCMV collected November 2016 from Cmp Simpon, Mrojejy Ntionl Prk (S , E , 1326 m.s.l.) by A. Rkotorison, M.D. Scherz, M.C. Bletz, J.H. Rzfindribe, A.Rzfimnntso, nd M. Vences. ZSM 528/2016 (ZCMV 15290) collected 19 November 2016 from high-elevtion site in Mrojejy Ntionl Prk (S , E , 1573 m.s.l.) by M.D. Scherz, M.C. Bletz, J.H. Rzfindribe, A.Rzfimnntso, nd M. Vences. ZCMV 15227, ZSM 527/2016 (ZCMV 15282), nd ZSM 525/2016 (ZCMV 15283) collected 19 November 2016 from high-elevtion site in Mrojejy Ntionl Prk (S , E , 2026 m.s.l.) by M.D. Scherz, M.C. Bletz, J.H. Rzfindribe, A.Rzfimnntso, nd M. Vences. Distribution. This species is known only from Mrojejy Ntionl Prk. Elevtionl rnge bsed on voucher specimens extends from m.s.l., but ccording to our observtions in 2016, clls cn be herd from distinctly lower nd higher elevtions, probbly rnging t lest from c m.s.l. Nturl history. At Mrojejy, clling specimens were herd from the lef litter in primry rinforest, during the dy. The specimens were deeply hidden in the lef litter nd exceedingly difficult to find. In 2016, clls were herd from rinforest s well s from shrubs nd vegettion mts including grsses bove the treeline. Cll. The dvertisement cll very probbly emitted by ZSM 382/2005 (specimen not seen clling) consists of single, short, pulstile, high-pitched, nd frequency- 337

68 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b Fig. 51. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi tridctyl (probbly from ZSM 382/2005) from Mrojejy Ntionl Prk: () 1 s durtion section; (b) 6 s durtion section. modulted note repeted in series t regulr intervls (Fig. 51). Clls recorded by M. Vences nd F. Glw on 16 Februry 2005 t Mrojejy Ntionl Prk, ner cmp Simpon, t 21.5 C ir temperture (Vences et l., 2006: CD3, trck 38, cut 1) re s follows: cll durtion (= note durtion) ms (132 ± 23 ms; N = 18), inter-cll intervls ms (1012 ± 39 ms; N = 17), nd dominnt frequency t Hz (7244 ± 200Hz, N = 18). Within notes, distinct frequency modultion is evident, with steep increse in dominnt frequency from the beginning to the end. Stumpffi contumeli sp. nov. (Figures 52 nd S106) Holotype. ZSM 443/2010 (FGZC 4258), n dult mle, collected on 2 April 2010 in Ambodivongy (S , E , 287 m bove se level), Tomsin Province, Mdgscr, by F. Glw, J. Köhler, P.-S. Gehring, M. Pbijn, nd F.M. Rtsovin. Prtypes. ZSM 441/2010 (FGZC 4248) nd ZSM 442/2010 (FGZC 4252) with the sme collection dt s the holotype. Dignosis. A very smll species of Stumpffi with strong digitl reduction in clde D, relted nd similr to S. tridctyl. Not included in previous studies. (1) Miniturized species (dult mle SVL mm); (2) mnus with one finger only (first finger not visible; second nd fourth fingers reduced to smll knobs; third finger brodened to tringulr shpe) nd pes with three toes (first toe not recognizble, second toe reduced to smll knob); (3) terminl phlnges of fingers nd toes without enlrged discs; (4) reltive hnd nd foot length, HAL/ SVL , FOTL/SVL ; (5) dorsum smooth with few scttered tubercles; (6) iridescent copper with irregulr drk flecks, with distinct color border to the drk flnks in t lest some specimens nd in the nterior hlf of the body. Ventrlly slte gry with whitish flecks posterior to the throt. Without contrsted ventrl colortion or red color elements on ventrl side; (7) regulrly repeted short single-note frequency-modulted whistle cll. Distinguished from ll nominl species of Stumpffi except S. tridctyl (nd S. obscoen sp. nov., described below) by stronger digitl reduction with only one finger nd three toes recognizble. Also distinguished from most other Stumpffi by its frequency-modulted cll. The new species is plced in clde D s the sister species of S. tridctyl, but differs from tht species by (1) strong genetic divergence in both mitochondril nd nucler genes, (2) more grnulr dorsum, (3) probbly shorter cll durtion, higher frequency nd steeper frequency modultion, lthough these trits re difficult to ssess due to differences in recording conditions (especilly temperture). Description of the holotype. Specimen in good stte of preservtion, right hindlimb removed s tissue smple. The right forelimb is frctured. Body stocky; hed slightly longer thn wide, nrrower thn body; snout rounded in dorsl view, pointed in lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis stright; lorel region stright, verticl; tympnum distinct, bout 52% of eye dimeter; suprtympnic fold indistinct; tongue long, brodening posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone smll, rounded. Forelimbs slender; subrticulr tubercles single, indistinct; outer metcrpl tubercle tringulr, indistinct; inner metcrpl tubercle distinct, oblong; fingers without webbing; ll fingers except third reduced; finger tips not expnded into discs. Hind limbs reltively thick; TIBL 338

69 VERTEBRATE ZOOLOGY 67 (3) 2017 b c Fig. 52. Stumpffi contumeli sp. nov. from Ambodivongy in life: ( c) prtype ZSM 441/2010 (FGZC 4248). 49% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle elongted, thin; outer mettrsl tubercle bsent; no webbing between toes; first toe bsent, second strongly reduced; reltive length of toes 2 < 5 < 3 < 4; fifth toe slightly shorter thn third. Skin on dorsum smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After five yers in 70% ethnol, the dorsum is silver with irregulr blck mrkings, nd occsionl circulr white spots; blck spots in the inguinl region. The suproculr regions contin severl smll blck spots. Dorsl surfce of the hed s the bck. Nostril brown, the lterl surfce of the hed is brown mottled with crem. A strk color border is present between the silver dorsl colortion nd the drk brown lterl colortion, running over the tympnum to the rostrum. White spots permete the drk brown color long the lower lip. The ventrl trunk is vermiculted white nd drk brown. The chin hs crem bse color spotted with tiny drk brown spots. Vermicultions extend onto the ventrl surfce of the thighs to become brown spotted with white. Interior surfce of the shnk crem. Dorslly, the thigh is bronze, with irregulr drk mrkings. The shnk hs strong drk crossbnd midwy down its length. The feet lso hve strong drk crossbnd, bordered either side by silver. The toes re striped silver nd drk brown. The rms re bronze nd silver flecked with drk mrkings. A crossbnd is present in the middle of the lower rm. Color in life (bsed on prtype ZSM 441/2010). Dorsum iridescent copper with irregulr drk flecks nd lighter spots. Superoculr region with drk ptches. Inguinl spots present, blck. Strk lterl color border trversing the middle of the eye; flnk blck to gry with smll irregulr iridescent light flecks, t high concentrtion posterior to the insertion of the forelimb. Forelimbs dorslly s the dorsum, with single blck crossbnd on the distl hlf of the forerm. Hnds mottled gry. Hindlimbs s the dorsum, with one drk crossbnd on the shnk, nd one on the dorsl surfce of the foot. Fourth toe dorslly gry with light golden spot just posterior to the terminl phlnge. Ventrl body slte gry in bse color. Dense concentrtion of iridescent whitish flecks directly posterior to the pectorl girdle, decresing in density posteriorly. Light gry spots present on the posteroventrl thighs. Iris rust red round the pupil, drkening externlly (Fig. 52). Vrition. For vrition in mesurements mong specimens, see Tble 3. In generl, ll exmined specimens strongly gree with the holotype in morphology. A high degree of vribility ws noted in the colortion of specimens: ZSM 441/2010 grees well with the holotype, except in lcking shnk crossbnd, hving weker inguinl spots, nd lcking superoculr blck spots; ZSM 442/2010 differs strongly, with the ventrl colortion of trunk extending onto the chin, the ventrl colortion being less vermiculted thn spotted, dorslly with n X- like mrking in the scpulr region, nd fint brown chevron extending nteriorly to the mid-dorsum from the lrge drk inguinl spots. These chevron nd inguinl spots extend onto the thigh s n oblique crossbnd. The shnk hs thick drk crossbnd. The dorsl surfce of the eye lso lcks blck spots. Etymology. The species nme is noun in pposition to the generic nme, derived from Ltin contumēli = insult. It is in reference to the offensive shpe of the hnd cused by digitl reduction. Distribution. Known only from its type loclity Ambodivongy. 339

70 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b Fig. 53. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls ttributed to Stumpffi contumeli sp. nov. from Ambodivongy: () 1 s durtion section; (b) 6 s durtion section. Nturl history. At Ambodivongy, mles clled from the forest floor from hidden positions under lef litter during the dy nd t dwn. Cll. The dvertisement cll of S. contumeli consists of single, very short, high-pitched nd pulstile note repted in series t regulr intervls (Fig. 53). Clls recorded by J. Köhler on 2 April 2010 t Ambodivongy very likely corresponding to this species (mles not observed clling, but found in lef litter from where clls were emitted) hd the following numericl prmeters: cll durtion (= note durtion) ms (42 ± 4 ms; N = 7), inter-cll intervls ms (543 ± 27 ms; N = 6), nd dominnt frequency t Hz (7493 ± 50 Hz, N = 7). Within notes, distinct frequency modultion is evident, with n increse in dominnt frequency from the beginning to the end. Fig. 54. Stumpffi obscoen sp. nov. from Andsibe in life, photogrphed 1991 (not ssignble to voucher specimen). Stumpffi obscoen sp. nov. (Figures 54 nd S107) Holotype. ZSM 381/2005 (FGZC 2666), sex undetermined, collected on 9 Februry 2005 round Andsibe (S , E , 900 m bove se level), Tomsin Province, Mdgscr, by F. Glw, R.D. Rndriniin, nd R. Dolch. Prtypes. ZSM 380/2005 (FGZC 2664), UADBA (FGZC 2663), nd UADBA (FGZC 2665), sme collecting dt s holotype; ZFMK collected in Februry 1991 round Andsibe by F. Glw nd M. Vences; ZFMK collected on 11 nd 13 Jnury 1992 round Andsibe by F. Glw nd J. Müller; ZFMK collected in Jnury or Februry 1994 in Andsibe by F. Glw nd M. Vences. Dignosis. A very smll species of Stumpffi with strong digitl reduction, phenotypiclly similr to S. tridctyl but plced in clde B. This species hs previously been listed s Stumpffi sp. 6 (Vieites et l., 2009; Köhler et l., 2010; Peloso et l nd Scherz et l., 2016), s Stumpffi sp. 7 in Wollenberg et l. (2008), nd s prt of Stumpffi tridctyl in Blommers-Schlösser & Blnc (1991) nd Glw & Vences (2007: ). (1) Miniturized species (SVL mm); (2) mnus with one finger only (first, second nd fourth finger not visible; third finger brodened to tringulr shpe by extended lterl fringes) nd pes with three toes (first toe not recognizble, second toe reduced to vestigil knob); (3) terminl phlnges of fingers nd toes without enlrged discs; (4) reltive hnd nd foot length, HAL/ SVL , FOTL/SVL ; (5) dorsum smooth with few scttered tubercles; (6) chocolte dorslly, with distinct color border to the ebony flnks in t lest some specimens nd in the nterior hlf of the body. Ventrlly gryish with drk mottling, throt drk. With- 340

71 VERTEBRATE ZOOLOGY 67 (3) 2017 b Fig. 55. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi obscoen sp. nov. from Andsibe: () 1 s durtion section; (b) 6 s durtion section. out contrsted ventrl colortion or red color elements on ventrl side; (7) regulrly repeted short single-note frequency-modulted whistle cll. Distinguished from ll nominl species of Stumpffi except S. tridctyl nd S. contumeli by stronger digitl reduction with only one finger nd three toes recognizble. Also distinguished from most other Stumpffi by its frequency-modulted cll. The new species is very similr t first glnce to S. tridctyl nd S. contumeli, but ccording to the moleculr phylogeny is not closely relted to those species, being plced in clde B rther thn in clde D. It differs from S. tridctyl nd S. obscoen by (1) strong genetic divergence in both mitochondril nd nucler genes nd distnt phylogenetic position, (3) distinctly longer inter-cll intervl, nd (3) different externl morphology of the hnd, with the enlrgement of the third finger mde up by lterl fringes rther thn by more uniform brodening of the entire finger into conicl shpe. Description of the holotype. Specimen in good stte of preservtion, its left forelimb removed s tissue smple. Body ovl; hed slightly wider thn long, nrrower thn body width; snout slightly pointed in dorsl view, somewht truncte in lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis indistinct, stright; lorel region stright, verticl; tympnum distinct, bout 70% of eye dimeter; suprtympnic fold not visible; tongue long, brodening posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone ovl. Forelimbs slender; subrticulr tubercles single, distinct; outer metcrpl tubercle distinct, brod, flt, nd round; inner metcrpl tubercle distinct, oblong; fingers without webbing, ll fingers except third reduced, finger tips not expnded into discs. Hind limbs reltively thick; TIBL 46% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle elongted, thin; outer mettrsl tubercle bsent; no webbing between toes; first toe prcticlly bsent, second toe strongly reduced; reltive length of toes 2 < 5 < 3 < 4; fifth toe distinctly shorter thn third. Skin on dorsum smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After 10 yers in 70% ethnol, the dorsum is brown with two oblong beige ptches over the hip region, nd further two spots of beige over the scpulr region. A drk brown oblique fleck is present in the inguinl region. The nostril is beige surrounded with brown. The lterl sides of the hed re drker brown thn the dorsum. The ventrl surfce is brown speckled with beige; speckling extends up lterl sides of body; chin slightly drker thn trunk. Ventrl surfce of thighs speckled s trunk, becoming lighter on ventrl surfces of shnks nd feet. Anterodorsl surfce of thigh lso speckled with beige, posterodorsl region beige without speckling; proximl hlf of shnk brown, distl hlf beige, with beige nd brown hlves seprted by drk crossbnd. Trsus beige. Feet dorslly beige medilly, drk brown lterlly. Clocl region drk brown. Upper rm dorslly beige, ventrlly brown, elbow brown, lower rm brown; hnd brown dorsolterlly, beige dorsomedilly, ventrlly beige. In life, dorsum chocolte with irregulr blck or drk brown flecks. Superoculr region with drk ptches. Inguinl spots present, blck. Strk lterl color border trversing the middle of the eye; flnk ebony with smll irregulr iridescent light flecks, t high concentrtion posterior to the insertion of the forelimb. Forelimbs dorslly s the dorsum, without crossbnds. Hnds dorslly brown. Hindlimbs s the dorsum, with one drk crossbnd on the proximl thigh, nd one on the distl 341

72 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs shnk, which line up when the leg is bent. Foot without crossbnds. Fourth toe dorslly brown with light iridescent flecks. Ventrl color in life not known. Iris rust red round the pupil, drkening externlly (Fig. 54). Vrition. For vrition in mesurements mong specimens, see Tble 3. In generl, ll exmined specimens strongly gree with the holotype in morphology. A high degree of vribility ws noted in the colortion of specimens: The dorsum of FGZC 2664 is light brown (lighter thn the holotype), with beige regions on the dorsum s in the holotype but more fint, but between hip nd shoulder spots n lmost tringulr drk brown mrking is present. Inguinl spots re present s in the holotype. A strong color border is present between the light brown dorsum nd the drk brown lterl region extending to the lterl side of the hed. Vent region is drk s in the holotype. The dorsl surfce of the whole leg is beige insted of brown with beige flecks; drk crossbnd is present on the dorsl surfce of the proximl end of the thigh. A discontinuous crossbnd is lso present in the middle of the tibi s in the holotype. The foot is dorsolterlly drk surrounded by light beige res, nd dorsomedilly beige. The ventrl surfce is s in the holotype except tht speckling of beige is bsent from the chin. ZFMK nd ZFMK re strongly discolored (orngish), but the former clerly grees with the holotype, while the ltter grees with FGZC 2664 in colortion. In ll specimens, the drk vent region, inguinl spots, leg crossbnds, nd ventrl flecking re distinct. The specimens ZFMK nd gree with holotype in color nd morphology. Etymology. The species nme is the feminine nomintive singulr form of the Ltin djective obscēn/obscoen = obscene. As for S. contumeli bove, it is in reference to the offensive shpe of the hnd cused by digitl reduction. Distribution. Known from forests round Andsibe. Nturl history. Specimens typiclly cll during the dy from the lef litter in primry rinforest but lso from secondry (mture euclypt) forest, from hidden positions. Sometimes mny clling mles cn be herd, suggesting tht these frogs occur ll over the forest. Cll. The dvertisement cll of S. obscoen consists of single pulstile note repeted in series t regulr intervls (Fig. 55). Clls recorded by F. Glw on 9 Jnury 1992 t Andsibe (Vences et l., 2006: CD3, trck 38, cut 2) hd the following numericl prmeters: cll durtion (= note durtion) ms (144 ± 8 ms; N = 8), intercll intervls ms (4619 ± 990 ms; N = 5), nd dominnt frequency t Hz (8361 ± 68Hz, N = 6). Within notes, distinct frequency modultion is evident, with n increse in dominnt frequency from the beginning to the end. Stumpffi dvidttenboroughi sp. nov. (Figures 56 nd S123) Holotype. ZSM 204/2016 (ACZCV 0106), Figs. 56 nd S123, dult mle, collected on 6 November 2013 in Shmbendrn (S , E , 558 m bove se level), Betmpon Strict Nture Reserve, Tomsin Province, Mdgscr, by G.M. Ros, D.J. Hrris, M.Rndrimiliso, nd H. Lv. Dignosis. A smll species of Stumpffi with digitl reduction. This species hs not been listed s cndidte species in previous publictions. (1) Miniturized species (dult mle SVL mm); (2) mnus with four fingers (first finger very strongly reduced in length; lso second nd fourth finger short) nd pes with four toes (first toe bsent, second nd fifth toes reduced in length); (3) terminl phlnges of fingers nd toes without enlrged discs; (4) reltive hnd nd foot length, HAL/SVL 0.17, FOTL/SVL 0.55; (5) dorsum smooth; (6) dorslly moch bordered round the lterl edges with chmpgne, forming teddyber shpe. Flnks moch, with shrp continuous color border between dorsum nd flnks. Distinguished from ll nominl species of Stumpffi by the pttern of digitl reduction, with completely reduced first toe nd length-reduced second toe (otherwise only found in S. tetrdctyl, but with different degree of finger reduction), nd strongly length-reduced first finger (only found in S. miery). Phylogeneticlly plced in clde B, forming the sister txon of S. obscoen, which presents much stronger digitl reduction. Description of the holotype. Specimen in good stte of preservtion, left foot removed s tissue smple. Body slender; hed longer thn width, slightly nrrower thn body; snout slightly pointed in dorsl view, rounded in lterl view; nostrils directed lterlly, not protubernt, slightly nerer to tip of snout thn to eye; cnthus rostrlis not distinct; lorel region concve, oblique; tympnum distinct, bout 80% of eye dimeter; suprtympnic fold not distinct; tongue long, brodening posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone ovl. Forelimbs slender; subrticulr tubercles single, distinct; outer metcrpl tubercle not recognizble; inner metcrpl tubercle distinct, ovl, fused with the prepollicl tubercule; fingers without webbing; first finger very strongly reduced, second nd fourth fingers short; reltive length of fingers 1 < 2 < 4 < 3, fourth finger slightly longer thn second; finger tips not expnded into discs. Hind limbs slender; TIBL 30% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle smll, round; outer mettrsl tubercle bsent; no webbing between toes; toe tips slightly expnded; first toe bsent, second nd fifth toes reduced in length; reltive length of toes 2 < 5 < 3 < 4; fifth toe shorter thn third; subrticulr tubercles distinct, single. Skin on dorsum reltively smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After three yers in 70% ethnol, the dorsum is metllic brown with brown ted- 342

73 VERTEBRATE ZOOLOGY 67 (3) 2017 b c Fig. 56. Stumpffi dvidttenboroughi sp. nov. from Betmpon Strict Nture Reserve in life: ( b) holotype ZSM 204/2016 (ACZCV 0106); (c) egg clutch found close to the specimen; note tht ssignment of the clutch to this species is tenttive nd requires confirmtion by further field observtion. dyber shpe strting from the middle of the eyes to the inguinl region. A drk brown spot, bordered with light brown line is present in the inguinl region. The dorsl surfce of the hed is metllic nd brown in color with beige line connecting the eyes nd defining the teddyber shpe long the dorsum. The nostril is surrounded with beige. The lterl surfce is homogenously drk brown flecked with smll beige dots. The flnk colortion merges with the ventrl colortion. The ventrl trunk is brown flecked with beige. The chin is brown spotted with beige but with less nd smller flecks thn the other ventrl prts. The ventrl thigh is s the belly. The shnk, trsus, nd foot re ventrlly s the thigh. Dorslly, the thigh is beige spotted with brown with drk crossbnd bordered by whitish line. The posterodorsl surfce of the thigh is beige spotted with brown nd with drk brown ovl shpe bordered with withish line close the clocl region. The shnk is beige spotted with brown with drk crossbnd bordered by whitish line nd in the distl portion is pinkish. The colortion of the distl portion of the shnk merges with the colortion of the trsus. The foot including the toes is s the trsus but drker. The rms re beige with pinkish color. The dorsomedil surfce of the hnd is beige. The underside of the rm is s the ventrl trunk. In life, dorslly dichromtic: internlly moch in teddyber shpe, bordered round the lterl edges nd the nterior hed by stripe of chmpgne. The flnk is drker, with smll iridescent light flecks. Dorsl forelimbs drk brown proximlly, becoming copper before the elbow, fding gin to brown distlly. Smll iridescent light flecks re present on the hnd. The hindlimbs re trichromtic: from the hip to the mid-shnk is moch. At the mid-shnk, drk crossbnd bordered with chmpgne line is present. Beyond this point, the leg becomes coppery ornge. A wek drk crossbnd is present on the trsus. The dorsl foot hs irregulr light iridescent flecks. The ventrl colortion in life is unknown (Fig. 56). Etymology. We dedicte this species to Sir Dvid Attenborough, who celebrted his 91 st birthdy on the 8 th of My 2017, in recognition of his services to science communiction nd nturl history brodcsting with the BBC for the lst 65 yers. Sir Dvid first visited Mdgscr in the 1960s, nd hs been n mbssdor for the islnd s wildlife, nd for thretened wildlife round the globe, throughout his long creer. Distribution. This species is so fr known only from its type loclity in the Betmpon Strict Nture Reserve. Nturl history. This specimen ws found during the dy in the lef litter in close proximity with jelly nest. The individul seemed to be gurding the egg clutch inside this curled dry lef forming sort of hlf pipe. This lef hd yet nother lef on top providing sfer microhbitt, both by keeping higher level of humidity nd wy from potentil predtors. Unfortuntely, the developing embryos were not smpled nd it is therefore not possible to unequivoclly ssign these eggs to this species. However, if they were indeed the eggs of this species, they would represent the first evidence of jelly- insted of fom-nests in Stumpffi, more similr to those of Rhombophryne nd other cophylines (see lso other observtions on S. chillei sp. nov. below). This species ppers to be uncommon, or hve extremely elusive hbits. At lest four mphibin surveys coupled with moleculr txonomic identifiction were crried out in the reserve (from 27 Februry to 23 Mrch 2004; from 3 Februry to 2 April 2007, from 25 October to 16 December 2007, nd from 3 to 25 November 2013) nd only 343

74 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs one of the mny collected Stumpffi individuls hs been ssigned to this new species. Cll. Unknown. 4. Lrge-sized species of clde B Clde B contins three smll-sized to miniturized species (S. dvidttenboroughi, S. miery, S. obscoen, discussed bove), ll with t lest certin degree of digitl reduction. Surprisingly, this clde contins with high support lso one undescribed, geogrphiclly disprte lrge-sized species without obvious digitl reduction, which we describe in this section. Stumpffi meikee sp. nov. (Figures 57 nd S108) Holotype. ZSM 1823/2010 (ZCMV 12374) (Fig. 57 d e), n dult mle, collected on 13 June 2010 t Mtsbory Miky Cmpsite, Tsrtnn Strict Nture Reserve (S , E , 2021 m bove se level), Antsirnn/Mhjng Province, Mdgscr, by M.Vences, D.R. Vieites, R.D. Rndriniin, F.M. Rtsovin, S. Rsmison, A. Rkotorison, E. Rjeririson, F. Rndrinsolo, nd T. Rjofirison. Prtypes. UADBA-A (DRV 6178), UADBA-A (DRV 6172), UADBA-A (DRV 6169), UADBA-A (DRV 6176), UADBA-A (DRV06173), UADBA-A (DRV06174), UADBA-A (ZCMV 12378), UADBA-A (ZCMV 12375), ZSM 1821/2010 (ZCMV 12372), ZSM 1822/2010 (ZCMV 12373), ZSM 617/2014 (DRV 6163), ZSM 618/2014 (DRV 6170), ZSM 619/2014 (DRV 6171), ZSM 620/2014 (DRV 6175), ZSM 621/2014 (DRV 6177), ZSM 622/2014 (DRV 6179), ZSM 623/2014 (DRV 6184), ZSM 624/2014 (DRV 6185), ZSM 625/2014 (DRV 6188), nd ZSM 626/2014 (DRV 6189), ll with the sme collecting dt s the holotype. Dignosis. A firly lrge, conspicuouly colored species from high elevtions on the Tsrtnn Mssif. Previously listed s S. sp. C33 in Scherz et l. (2016) nd Peloso et l. (2017). (1) Modertely lrge-sized species (SVL mm); (2) mnus with four fingers (first finger slightly reduced in length) nd pes with five toes (first toe distinctly reduced in length); (3) terminl phlnges of fingers nd toes with slightly enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum smooth; (6) dorslly gry to chocolte brown, with drker ptterns tht re highly vrible mong specimens but generlly consist of n interoculr-nterior dorsl tringle, nd severl smller ptches bordered with crem. Ventrlly, the chin nd bdomen is chmpgne to slmon colored, while the hindlimbs re more intense slmon or ornge colortion. Distinguished from S. nlmin, S. ngeluci, S. contumeli, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. m dgscriensis, S. mmitik, S. mledict, S. miery, S. ob scoen, S. psologloss, S. pygme, S. so r t, S. te tr dctyl, S. tridctyl, nd S. ynniki by lrger body size; from S. grndis by the lck of contrsted light blue-blck ventrl pttern; from S. be, S. hr, S. megsoni, nd S. stffordi by stronger length reduction of first toe; from S. grndis, S. hr, S. megsoni, nd S. stffordi by chmpgne to slmon-colored belly nd ventrl slmon to ornge color on hindlimbs (vs. no or very little reddish ventrl color); from S. kibomen by different dorsl pttern, nd by lck of dense blckish pigmenttion of the throt. Morphologiclly most similr to S. roseifemorlis, which hs similr ventrl nd dorsl color pttern nd lso occurs t comprtively high elevtions, but occupies very distnt position in the Stumpffi phylogeny, nd is concordntly differentited in mitochondril nd nucler genes. A probble morphologicl difference between these two species is reltive toe length, with the third toe being disctinctly longer thn the fifth toe in S. roseifemorlis, but only slightly longer thn the fifth toe in the new species. Description of the holotype. Specimen in good stte of preservtion, thigh muscle on right leg prtly removed s tissue smple. Body per shped; hed slightly longer thn wide, nrrower thn body; snout pointed in dorsl view, pointed in lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis distinct concve; lorel region concve, oblique; tympnum distinct, bout 59% of eye dimeter; suprtympnic fold not disctinct; tongue long, brodening posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone ovl. Forelimbs slightly thick; subrticulr tubercles single, round; outer metcrpl tubercle distinct, smll, plmr metcrpl visible, close to the outer metcrpl, round; prepollex present, ovl; fingers without webbing; first finger reduced, second finger reduced; reltive length of fingers 1 < 2 < 4 < 3, fourth finger distinctly longer thn second; finger tips expnded into discs. Hind limbs slender, 52% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle smll, ovl; outer mettrsl tubercle bsent; no webbing between toes; first toe strongly reduced; toe tips slightly expnded; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe distinctly shorter thn third; subrticulr tubercles distinct, single, elongte. Skin on dorsum reltively smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After five yers in 70% ethnol, the dorsum is brown with drk brown ptches. Drk brown reversed V in the inguinl region. The dorsl surfce of the hed is s the bck, with reversed tringulr drk brown ptch strting in the middle of the eye to the forelimb, drk brown ptches in the lterl side close to the forelimb. The nostril is beige. The lterl surfce of the hed is brown mottled with beige. The flnks re beige spotted with brown. The flnk colortion merges with the ventrl colortion. The ventrl bdomen is s the flnks. The chin is beige flecked with brown. The ventrl thigh is s the belly. The shnk nd trsus re ventrlly s the thigh. The sole of the foot is drk brown mottled 344

75 VERTEBRATE ZOOLOGY 67 (3) 2017 b c Fig. 57. Stumpffi meikee sp. nov. from Mtsbory Miky cmpsite, Tsrtnn Strict Nture Reserve, in life: () prtype ZSM 1821/2010 (ZCMV 12372); (b c) prtype ZSM 1822/2010 (ZCMV 12373); (d e) holotype ZSM 1823/2010 (ZCMV 12374). d e with beige. Dorslly, the thigh is drk brown with perpendiculr brown crossbnd. The posterodorsl surfce of the thigh is beige flecked with brown. The shnk is s the thigh with n oblique drk brown crossbnd. The trsus is lighter thn the thigh with perpendiculr crossbnd. The foot is s the thigh without crossbnd. The toes re striped with beige ptches. The clocl region is drk brown. The rms re drk brown. A drk crossbnd is present on the lower rm. The dorsomedil surfce of the hnd is beige. The fingers hve smll white nnulus before ech terminl phlnge. The underside of the rm is s the trunk. Color in life (holotype nd vrition). Dorsum with distinctive pttern formed of drk brown mrkings bordered by crem: n isoscelese tringulr mrking strts between the eyes nd nrrows to point round the suprscpulr region (cn be irregulr in some specimens, compre Fig. 57b with 57 nd 57d). A second mrking converges nteromedilly from the inguinl region to form second semi-tringulr mrking. A further two drk mrkings re present behind the suprscpulr region. These together with the second semi-tringulr mrking cn form n symmetricl X-like mrking; compre Fig. 57d with 57b nd 57. A further smll mrking cn be present on the snout. Dorsl bse color burnt umber (lighter in some prtypes, even rnging to gry; Fig. 57). No dorsolterl color border is present. The flnks re mostly s the dorsum, fding to the ventrl chmpgne, with smll irregulr whitish flecks. A crem mrking runs from the tympnum to the insertion of the rm. The dorsl forelimb is s the bck. A crossbnd is present on the forerm. The hnd is dorsomedilly crem, dorsolterlly s the dorsum, with irregulr whitish flecks, including one before ech terminl phlnge. Dorsl hindlimb s the bck, with crem-bordered drk brown crossbnd t the mid-thigh, mid-shnk, nd on the foot. The dorsl foot nd toes hve severl whitish flecks, including one before ech terminl phlnge. The ventrl trunk is chmpgne to slmon. The chin is chmpgne with brown flecks, decresing in density posteriorly (mostly bsent in some prtypes, Fig. 57). The ventrl legs nd inguinl region re slmon. Ventrl hnds nd feet rosy brown. The iris is russet (lighter browns in some prtypes, Fig. 57). Vrition. For vrition in mesurements mong specimens, see Tble 3. In generl, ll exmined specimens gree strongly with the holotype in morphology, except ZSM 1822/2010 (ZCMV 12373): first toe slightly bigger thn in the holotype, inner metcrpl smller, colortion uniformly brown; ZSM 624/2014 (DRV 06185): snout dorslly slightly round, tongue brodening posteriorly, inner mettrsl smll ovl, colortion in dorsl view is s the holotype but with white spots cross ll prts of the body; ZSM 619/2014 (DRV 6171): dorslly on the hed present, drk brown spot in the chone region; ZSM 622/2014 (DRV 6179) nd 626/2014 (DRV 6189): outer metcrpl present nd ovl, colortion s ZSM 624/2014; ZSM 621/2014 (DRV 6177): outer metcrpl smll, ovl, colortion dorslly lighter thn the holotype with crem spot; ZSM 620/2014 (DRV 6175): plmr metcrpl close to the outer metcrpl, brown brod mid-dorsl stripe on the bck strting between the eyes nd running to the inguinl region; ZSM 618/2014 (DRV 6170): lrge, elongte inner metcrpl tubercle, plmr metcrpl close to the outer metcrpl, both elongte. Colortion in dorsl view similr to the holotype except for the presence of crem spots cross the whole body. In ZSM 617/2014 (DRV 6163) colortion uniformly brown, outer metcrpl lrge, elongte, plmr metcrpl smll elongte. In ZSM 1821/2010 (ZCMV 12372) the thigh und trsus lck crossbnds, nd the plmr metcrpl is smll. For five prtypes, no moleculr dt re vilble: ZSM 617/2014 (DRV 6163), ZSM 623/2014 (DRV 6184), ZSM 624/2014 (DRV 6185); ZSM 625/2014 (DRV 6188), ZSM 626/2014 (DRV 6189). However, since these specimens were collected t exctly the sme site s the rest of the type series, nd they gree with the holotype in morphology, we here ssign them to S. meikee. 345

76 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b Fig. 58. Stumpffi grndis from bove Cmp Simpon, Mrojejy Ntionl Prk, in life (not ssignble to voucher specimen): () dorsolterl view; (b) ventrl view. Etymology. The species nme is mtronym honoring Meike Kondermnn, to whom we re plesed to dedicte this ttrctively colored species in recognition for the invluble help, supervision nd friendship she provided to the first uthor nd mny other students during moleculr lbwork t TU Brunschweig. Distribution. Known only from its type loclity, Mtsbory Miky, Tsrtnn Strict Nture Reserve, 2021 m bove se level. Nturl history. Specimens were found fter the min riny seson (in June), inctive under stones, boulders nd roots in the dry bed of tiny strem. Multiple individuls were found under the sme stones, suggesting they might hve been estivting t this plce. Cll. Unknown. 5. Moderte- to lrge-sized species of clde C Three nominl species of Stumpffi re comprtively lrge-sized, distributed in estern nd north-estern Mdgscr, nd belong to clde C (Fig. 3). These three species plus the smll-sized S. tetrdctyl re the only described species in this clde. A lrge number of dditionl clde members exist nd will be described s new species in the following. In this section, we review nd describe first the species of moderte to lrge size, nd in the subsequent section we focus on the smll-sized species of clde C. Stumpffi grndis Guibé, 1974 (Figure 58) Nme-bering type. Holotype MNHN from Mssif du Mrojezy (1300 m) ccording to the originl description. Identity nd dignosis. A comprtively lrge-sized species only known from the Mrojejy Mssif in northestern Mdgscr, distinguished from ll other species of the genus by its highly contrsted blck-bluish ventrl color pttern. (1) Modertely lrge-sized species (SVL mm); (2) mnus with four fingers (first finger slightly reduced in length) nd pes with five toes (first toe slightly reduced in length); (3) terminl phlnges of fingers without, those of nd toes with slightly enlrged discs; (4) reltive hnd nd foot length not mesured HAL/SVL , FOTL/SVL ; (5) dorsum smooth; (6) reddish brown dorslly, with distinct drk bown to blck ptches. Ventrlly light bluish with distinct nd strongly contrsted blck pttern, especilly in the throt, which cn be entirely blck, without red color elements on ventrl side. Distinguished from S. nlmin, S. ngeluci, S. con tumeli, S. dvidttenboroughi, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. mledict, S. mdgscrien sis, S. mmitik, S. miery, S. obscoen, S. psologloss, S. pygme, S. sort, S. tridctyl, nd S. ynniki by distinctly lrger body size. From ll these species nd S. be, S. hr, S. meikee, S. megsoni nd S. stffordi by distinct nd contrsted ventrl color pttern nd lrger body size, furthermore from S. contumeli, S. dvidttenboroughi, S. miery, S. obscoen, S. tetrdctyl, nd S. tridctyl by distinctly lower degree of digitl length reduction. Specimens exmined. Holotype MNHN collected in July 1972 from Mrojejy (1300 m.s.l.) by C.P. Blnc; prtypes MNHN with sme collection dt s holotype. Referred specimens: ZSM 372/2005 (FGZC 2820) collected on 16 Februry 2005 from bove cmp Simpon, Mrojejy Ntionl Prk (coordintes not tken) by F. Glw, M. Vences, nd R.D. Rndriniin (note: this niml unfortuntely disintegrted in preservtive). Distribution. Known only from Mrojejy Ntionl Prk t round m.s.l. Cll nd nturl history. Unknown. 346

77 VERTEBRATE ZOOLOGY 67 (3) 2017 b c d e f Fig. 59. Stumpffi kibomen in life: ( b) specimen from Andsibe (not collected); (c d) femle specimen from An Al; (e f) mle prtype NMBE from ner Andsibe. Plte from Glw et l. (2015). Stumpffi kibomen Glw, Vlln, Andreone, Edmonds, Dolch & Vences, 2015 (Figures 59 nd S109) Nme-bering type. Holotype ZFMK (Fig. S109) from ner Andsibe (S , E ), c. 900 m elevtion, centrl-estern Mdgscr ccording to the originl description. Identity nd dignosis. A recently described modertely lrge-sized species of Stumpffi from the northern centrl est of Mdgscr, with distinct red color on belly nd limbs. (1) Modertely lrge-sized (dult mle SVL mm); (2) mnus with four fingers (first finger slightly reduced in length) nd pes with five toes (first toe slightly reduced in length); (3) Terminl phlnges of fingers nd toes without enlrged discs; (4) reltive hnd nd foot length, HAL/SVL 0.2, FOTL/SVL 0.7 (ccording to the originl description); (5) dorsum smooth, sometimes with few scttered tubercles; (6) reddish brown dorslly, with distinct drk brown ptches usully forming longitudinl bnds, nd one tringulr pth between the eyes nd running long the nterior third of the dorsum. Ventrlly light reddish to deep red on belly; throt with dense blck pttern, often lmost completely blck; (7) regulrly repeted short single-note tonl cll. 347

78 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b Fig. 60. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi kibomen recorded ner Andsibe: () 1 s durtion section; (b) 6 s durtion section. Distinguished from S. nlmin, S. ngeluci, S. contumeli, S. dvidttenboroughi, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. mledict, S. mdgsc riensis, S. mmitik, S. miery, S. obscoen, S. psologloss, S. pygme, S. sort, S. tridctyl, nd S. ynniki by distinctly lrger body size. From ll these species nd S. be, S. hr, S. megsoni, nd S. stffordi, distinguished by distinct red color on belly (vs. red color bsent from ventrl surfce, or restricted to limbs in S. be). Furthermore distinguished from S. contumeli, S. dvidttenboroughi, S. miery, S. obscoen, S. tetrdctyl, nd S. tridctyl by distinctly lower degree of digitl length reduction. Stumpffi meikee hs reddish color on belly similr to S. kibomen, but in S. meikee this color is typiclly more fint (chmpgne to slmon), the throt hs less expressed blck pttern, the dorsl pttern is different, nd the species belongs to completely different mjor clde of Stumpffi (clde B rther thn clde C). Specimens exmined. Holotype ZFMK collected on 17 Jnury 1995 ner Andsibe (S , E , c. 900 m.s.l.) by F. Glw; prtype NMBE collected on 14 Februry 1997 in An Al forest (S , E , c. 850 m.s.l.) by D. Vlln; prtype NMBE collected on 13 Februry 1995 ner Andsibe (S , E , c. 950 m.s.l.) by D. Vlln. Distribution. Known from estern rinforests t Andsibe nd An Al. Nturl history. Specimens were found during the dy in lef litter of primry rinforest, euclypt forest, nd fern scrub. Clls were herd in the morning nd fternoon (Glw et l., 2015). Cll. The dvertisement cll of S. kibomen consists of single short nd slightly pulstile note repeted in series t regulr intervls (Fig. 60). Clls recorded on 20 Mrch 1995 t 15:15 h ner Andsibe nd t 24 C ir temperture hd the following numericl prmeters (ccording to Glw et l., 2015): cll durtion (= note durtion) ms (73 ± 2 ms, N = 9), inter-cll intervls ms (797 ± 15 ms, N = 9), nd dominnt frequency t Hz. Within notes, distinct frequency modultion is evident, with n increse in dominnt frequency from the beginning to the end. Remrk. We here list long with S. kibomen two lineges corresponding to the cndidte species here nmed S. sp. C11 nd S. sp. C34 (note: the cndidte species C34 s used in Scherz et l did not refer to this species). These re resolved in clde with S. kibomen but seprted by substntil genetic divergence. Ech is known from single individul not vilble to us for morphologicl exmintion in the frmework of the present study, nd bsed on these deficient dt, their sttus must remin unresolved. Stumpffi sp. C11 is known from the specimen MRSN A2583 from Ambolokoptrik, wheres S. sp. C34 is represented by UADBA (ZCMV 3319) from Rnomen forest in Rnomfn Ntionl Prk, collected by M. Vences nd collbortors. According to field notes, this specimen is juvenile with red belly in life. Stumpffi roseifemorlis Guibé, 1974 (Figures 61 nd S110) Nme-bering type. Holotype MNHN from Mssif du Mrojezy (1300 m) ccording to the originl description. Identity nd dignosis. A modertely lrge-sized species of Stumpffi described from the Mrojejy Mssif in 348

79 VERTEBRATE ZOOLOGY 67 (3) 2017 b c d e Fig. 61. Stumpffi roseifemorlis from Mrojejy Ntionl Prk in life: ( b) ZSM 374/2005 (FGZC 2883); (c) ZSM 487/2005 (FGZC 2047); (d e) ZSM 529/2016 (ZCMV 15172). the north est of Mdgscr, chrcterized by distinct reddish color ventrlly on limbs. (1) Modertely lrgesized species (SVL mm); (2) mnus with four fingers (first finger slightly reduced in length) nd pes with five toes (first toe distinctly reduced in length); (3) terminl phlnges of fingers without, those of toes with slightly enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) Dorsum smooth; (6) gryish brown to reddish brown dorslly, with poorly contrsted drk brown teddyber shped mrking centrlly on dorsum, without shrp color border between color of dorsum nd flnks. Ventrlly with reddish-ornge color on belly nd chest, nd more intense reddish color on ventrl surfce on hindlimbs, without obvious drk pttern on ventrl surfce; (7) regulrly repeted long tonl single-note dvertisement cll. Distinguished from S. nlmin, S. ngeluci, S. contumeli, S. dvidttenboroughi, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. mledict, S. mdgscriensis, S. mmitik, S. miery, S. obscoen, S. psologloss, S. pyg me, S. sort, S. tridctyl, nd S. ynniki by distinctly lrger body size. From ll these species nd S. hr, S. megsoni, nd S. stffordi, distinguished by the distinct red color on belly. Furthermore distinguished from S. contumeli, S. dvidttenboroughi, S. miery, S. obscoen, S. tetrdctyl, nd S. tridctyl by distinctly lower degree of digitl length reduction. Among species with red ventrl color, distinguished from S. be by smller body size nd smller terminl discs on fingers; from S. kibomen by less expressed blck pttern on throt nd different dorsl pttern. Most similr to S. meikee but this species belongs to completely different mjor clde of Stumpffi (clde B rther thn clde C). Specimens exmined. Holotype MNHN collected in July 1972 from the Mrojejy Mssif by C.P. Blnc; prtypes MNHN with sme collection dt s holotype. Referred specimens: ZSM 373/2005 (FGZC 2808) collected on 14 Februry 2005 from Mrojejy Ntionl Prk, probbly Cmp Mntell (S E , 481 m.s.l.) by F. Glw, R.D. Rndriniin, nd M. Vences; ZSM 374/2005 (FGZC 2883), nd ZSM 349

80 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b Fig. 62. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi roseifemorlis from Mrojejy Ntionl Prk (cll recorded from ZSM 529/2016, ZCMV 15172): () 1 s durtion section; (b) 6 s durtion section. 487/2005 (ZCMV 2047), collected on 16 nd 17 Februry 2005 t Cmp Simpon, Mrojejy Ntionl Prk (S E , 1326 m.s.l.) by F. Glw, R.D. Rndriniin, nd M. Vences. ZCMV , ZSM 529/2016 (ZCMV 15172, ZSM 530/2016 (ZCMV 15206), ZCMV 15207, ZCMV 15209, ZSM 531/2016 (ZCMV 15274), ZSM 532/2016 (ZCMV 15277), ZCMV , ll collected November 2016 from Cmp Simpon, Mrojejy Ntionl Prk (S , E , 1326 m.s.l.) by A. Rkotorison, M.D. Scherz, M.C. Bletz, J.H. Rzfindribe, A. Rzfimnntso, nd M. Vences. Distribution. Known from Mrojejy Ntionl Prk between m.s.l. (occurrence t low elevtion t Cmp Mntell indicted by single voucher specimen only nd in need of confirmtion). Nturl history. In November 2016, specimens were very common in the lef litter round Cmp Simpon in Mrojejy Ntionl Prk, in prticulr in moist nd dense lef litter long smll strems. However, clls puttively belonging to this species were rrely herd, from only limited number of specimens, in the evening nd t night. Cll. The dvertisement cll here preliminrily ssigned to S. roseifemorlis consists of single, modertely long note emitted in series t regulr intervls (Fig. 62). Clls were recorded by M. Vences on 18 November 2016 (21:00) in Mrojejy Ntionl Prk, probbly from specimen ZSM 529/2016 (ZCMV 15172; not seen clling, but found t the spot of cll emission with prtly extended vocl sc; ir temperture estimted t 17 C), nd hd the following prmeters: cll durtion (= note durtion) ms (278 ± 1.6 ms; N = 6), inter-cll intervls ms ( ± 152 ms; N = 6, nd dominnt frequency t Hz ( ± 35.1 Hz, N = 6). Remrk. Geneticlly divergent specimens tht phylogeneticlly re sister to S. roseifemorlis were recorded from the lowlnd loclity of Ambodivongy (specimens FGZC 4238 nd 4250; both deposited in the UADBA collection nd not vilble for morphologicl exmintion in this study). They re here referred to s new unconfirmed cndidte species, Stumpffi sp. C57, with strong genetic divergence (9.6% in the 16S rrna) to S. roseifemorlis. Stumpffi chillei sp. nov. (Figures 63 nd S111) Holotype. ZSM 536/2016 (ZCMV 15149), n dult mle, collected on 16 November 2016 ner Cmp Mntell, Mrojejy Ntionl Prk (c. S14.438, E49.776, 450 m bove se level), Antsirnn Province, Mdgscr, by M.D. Scherz, A. Rkotorison, M. Vences, M. Bletz, J.H. Rzfindribe, nd A. Rzfimnntso. Prtypes. ZSM 493/2005 (ZCMV 860) nd 494/2005 (ZCMV 861) collected on 19 Februry 2005, nd UADBA (FGZC 2729), UADBA (FGZC 2765), nd ZSM 377/2005 (FGZC 2750), collected on 14 Februry 2005, in Cmp Mntell, Mrojejy Ntionl Prk (S14.421, E , 481 m bove se level), Antsirnn Province, Mdgscr, by F. Glw, M. Vences, nd R.D. Rndriniin. UADBA (ZCMV 2095) collected on 19 Februry 2005 from site below Cmp Mntell (coordintes not tken), by F. Glw, M. Vences, nd R.D. Rndriniin. UADBA (ZCMV 866) collected on 19 Februry 2005 in Cmp Mrojeji, Mrojejy Ntionl Prk by F. Glw, M. Vences nd R.D. Rndriniin; ZFMK nd ZFMK (Fig. 63) collected on Mrch 1994 from Mrojejy Ntionl Prk below 500 m.s.l. by F. Glw, N. Rbibiso, nd O. Rmilison; ZFMK collected between 25 Februry to 1 Mrch 1995 from Mrojejy Ntionl Prk (coordintes not tken) by F. Glw nd O. Rmilison. ZSM 534/2016 (ZCMV 15141) nd ZSM 535/2016 (ZCMV 15143), collected on 16 November 2016, nd ZSM 533/2016 (ZCMV 15308), mle, nd ZCMV 15307, eggs collected long with ZSM 533/2016, collected on 21 November 2016, ll from Cmp Mntell, Mrojejy Ntionl Prk (S , E , 456 m.s.l.) by A. Rkotorison, M.D. Scherz, M.C. Bletz, J.H. Rzfindribe, A. Rzfimnntso, nd M. Vences. 350

81 VERTEBRATE ZOOLOGY 67 (3) 2017 b e c d Fig. 63. Stumpffi chillei sp. nov. from Mrojejy Ntionl Prk in life: () prtype ZFMK 57461; (b) mle, ZSM 535/2016 (ZCMV 15143); (c) ZCMV 15307, egg clutch of S. chillei; (d) clling holotype. Dignosis. A species from the north-est of Mdgscr chrcterized by the fstest cll repetition rte mong those species with known dvertisement clls. It hs been previously listed s Stumpffi sp. 10 in Vieites et l. (2009), Köhler et l. (2010), Scherz et l. (2016) nd Peloso et l. (2017), s Stumpffi sp. 4 in Wollenberg et l. (2008), s Stumpffi sp. Mrojejy 2 in Glw & Vences (2007), nd s Stumpffi sp. b in Glw & Vences (1994). (1) Modertely lrge-sized species (SVL mm); (2) mnus with four fingers (first finger slightly reduced in length) nd pes with five toes (first toe distinctly reduced in length); (3) terminl phlnges of fingers nd toes with slightly enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum smooth with wek dorsolterl ridges or rows of tubercles; (6) dorslly typiclly with snd brown mid-dorsl stripe bordered by vrious drker nd ligher brown mrkings over the flnks. Ventrlly with some drk pttern especilly on the throt, without contrsted ventrl colortion or red color elements on ventrl side; (7) regulrly repeted short single-note tonl cll with fst repetition rte. Distinguished from S. nlmin, S. ngeluci, S. contumeli, S. dvidttenboroughi, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. mledict, S. mdgscriensis, S. mmitik, S. miery, S. obscoen, S. psologloss, S. pygme, S. sort, S. tridctyl, nd S. ynniki by distinctly lrger body size; from S. be, S. kibomen, S. meikee, nd S. roseifemorlis by the lck of red color ventrlly or on limbs; from S. grndis by the lck of contrsted light blue-blck ventrl pttern. Furthermore distinguished from S. contumeli, S. dvidttenboroughi, S. miery, S. obscoen, S. tetrdctyl, nd S. tridctyl by distinctly lower degree of digitl length reduction; from S. hr, S. megsoni, nd S. stffordi by smller body size nd stronger length reduction of first toe; from ll other species by the presence, in mny individuls, of brod light ptch or bnd covering most of the dorsum; nd from ll Stumpffi for which dvertisement clls re known by shorter inter-cll intervls. Description of the holotype. Specimen in good stte of preservtion, left thigh muscle removed s tissue smple. Body slightly rounded; hed slightly longer thn wide, nrrower thn body; snout rounded in dorsl view, rounded in lterl view; nostrils directed lterlly, not protubernt, nerer to the tip of snout thn to the eye; cnthus rostrlis distinct, concve; lorel region slightly concve, verticl; tympnum distinct, bout 53% of eye dimeter; suprtympnic fold not distinct; tongue long, stright, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone ovl. Forelimbs slender; subrticulr tubercles single, wekly visible; outer metcrpl tubercle visible, single; inner metcrpl tubercle visible, ovl; fingers without webbing; no reduced fingers; reltive length of fingers 1 < 2 < 4 < 3, fourth finger longer thn second; finger tips expnded into smll discs. Hind limbs slender; TIBL 57% of SVL; 351

82 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs lterl mettrsli strongly connected; inner mettrsl tubercle smll, ovl; outer mettrsl tubercle bsent; no webbing between toes; first toes slightly reduced; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe distinctly shorter thn third. Skin on dorsum reltively smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After one yer in 70% ethnol, the dorsum is gry with brod light gry dorsl stripe covering the whole hed, nrrowing to the suprscpulr region, nd then running to the vent, with smll drker gry hert-shped mrking over the posterior hed, nd light dorsl snout. The gry dorsl stripe is bordered by fine blckish irregulr lines, nd then drker gry ptches on the flnks; further, rounded light gry res re present in the inguinl region. Blck spots re present bove the tympnum nd over the suprscpulr region, the insertion of the eye, nd nterior to the inguinl region. Inguinl spots re present, smll nd irregulr nd blck. Tympnum white, suprtympnic stripe ending behind the tympnum. The nostril is blckish. The lterl surfce of the hed is drk gry with blck spots, especilly lrge blckish re below the eye. The flnk colortion merges with the ventrl colortion. The ventrl trunk is immculte crem, with brown mottling in the lterl region nd on the chin. The ventrl thigh is brown with crem mottling. The shnk is ventrlly crem. The trsus is ventrlly brown. The sole of the foot is brown. Dorslly, the thigh is beige with one perpendiculr brown crossbnd. The posterodorsl surfce of the thigh is beige. The shnk is beige with one crossbnd. The trsus is s the shnk but with two crossbnds. The foot is crem speckled with beige with one crossbnd. The toes re speckled with beige. The clocl region is drk brown. The rms re s the dorsum. A drk crossbnd is present on the forerm. The dorsomedil surfce of the hnd is crem. The fingers hve smll drk crossbnd. The underside of the rm is s the ventrl trunk. Color in life (holotype nd vrition). A dorsl stripe of snd brown to light rust brown covers the whole hed, nrrows to the suprscpulr region, nd then runs to the vent. The lterl body is gry-brown or snd-brown in bse color with light rust mrkings rising from the inguinl region running to contct the dorsl stripe, which cn be bordered with crem. Drk mrkings re present posterodorsl to the tympnum, below the eye, nd round the nostril, bove the insertion of the rm, sometimes on the middle of the flnk, on the suprscpulr region, nd in the inguinl region. The dorsl forelimb is light pech to beige, with drk crossbnd on the forerm. The hnd is internlly crem, lterlly s the rest of the forelimb, with smll whitish nnulus before ech terminl phlnge. The clocl region is blck. The legs re snd brown to light pech to beige in dorsl colortion, with one gry-brown to drk brown crossbnd on the thigh, two on the shnk, nd one ech on the tibiofibulr nd mettrsl regions of the foot. The toes re mottled crem nd drk brown. Ventrlly muve in bse color, fding to pech over the chin, with light lemon flecks on the bdomen. The ventrl thighs nd feet smoke colored, trnslucent over the nterior portion of the thigh. The iris is golden with blck reticultions. Vrition. For vrition in mesurements mong specimens, see Tble 3. All exmined specimens gree strongly with the holotype in morphology, nd hve highly similr colortion, except ZSM 494/2005, which hs no distinct dorsl stripe, nd hs drk spot centrlly over the pectorl region nd lcks inguinl spots. A drk spot is lwys present bove the insertion for the rm nd over the tympnum. The legs re s the holotype. Ventrl colortion in preservtive cn be mottled with beige, brown, or gry, except in ZSM 493/2005, where the venter is flecked with brown nteriorly nd under the chin. Etymology. The species nme is ptronym honoring Achille P. Rselimnn, in recognition for his substntil contributions to the study nd conservtion of the herpetofun of Mdgscr. Distribution. The species is known only from Mrojejy Ntionl Prk, between m bove se level. Nturl history. Clling specimens were herd during the dy from lef litter in primry rinforest, in the lte fternoon nd in the night shortly fter dusk. In November, one mle specimen (ZSM 533/2016) ws found in wter-filled empty snil shell long with 86 unpigmented (white) erly embryos of 2.5 mm in dimeter, surrounded by jelly cpsule of 5.1 mm in dimeter (verge vlues). The embryos were identified s S. chillei by DNA brcoding. They were in different embryonl stges (75 eggs in stge 11, nd 11 eggs in stge 13, ccording to Gosner, 1960); therefore, nd becuse of their high number, it is probble tht they originted from different femles. This discovery constitutes further evidence tht fom nest building is not the rule mong Stumpffi species, nd their reproductive hbits re more diverse thn previously thought. Cll. The dvertisement cll of S. chillei consists of single, short nd high-pitched note repeted in series t fst succession. Recordings re vilble from three different clling mle tht re here ssigned to this species (Fig. 64): (1) Clls recorded by F. Glw on 29 Mrch 1994 (18:00 h) in Mrojejy Ntionl Prk t n ir temperture of 23.4 C (Vences et l., 2006: CD3, trck 46) Fig. 64. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi chillei sp. nov. from Mrojejy (1 s nd 6 s durtion section ech): ( d) cll recorded from specimen ZFMK 59893; (b e) cll recorded from holotype ZSM 536/2016 (ZCMV 15149); (c f) cll recorded from specimen ZSM 535/2016 (ZCMV 15143). 352

83 VERTEBRATE ZOOLOGY 67 (3) 2017 b c d e f 353

84 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs hd the following numericl prmeters: cll durtion (= note durtion) ms (60 ± 10 ms; N = 13), intercll intervls ms (422 ± 21 ms; N = 12), nd dominnt frequency t Hz (5873 ± 60 Hz, N = 13). (2) Clls recorded by M. Vences on 16 November 2016 (19:00 h; c. 22 C ir temperture) from specimen ZSM 535/2016 (ZCMV 15143) in Mrojejy Ntionl Prk with following prmeters: cll durtion (= note durtion) ms (44.3 ± 3.4 ms; N = 10), inter-cll intervls ms (545 ± 16 ms; N = 10), nd dominnt frequency t Hz (5917 ± 95.7 Hz, N = 10). (3) Clls recorded by M. D. Scherz on 16 November 2016 (c. 22 C ir temperture) from holotype ZSM 536/2016 (ZCMV 15149) in Mrojejy Ntionl Prk with following prmeters: cll durtion (= note durtion) ms (41.8 ± 3.1 ms; N = 10), inter-cll intervls ms (515.8 ± 7.9 ms; N = 10), nd dominnt frequency t Hz ( ± 58.2 Hz, N = 10). Stumpffi nlnjirofo sp. nov. (Figures 65 nd S112) Holotype. ZSM 489/2005 (ZCMV 873) (Fig. 65 b), dult femle, collected on 22 Februry 2005 in Nosy Mngbe Specil Reserve (c. S15.500, E49.766, c m bove se level), Tomsin Province, Mdgscr, by F. Glw, M. Vences, nd R.D. Rndriniin. Prtypes. ZSM 492/2005 (ZCMV 2178), ZSM 491/2005 (ZCMV 2104), UADBA (ZCMV 2161), UADBA (ZCMV 2105), nd UAD- BA (ZCMV 2142), ll with sme collecting dt s holotype; UAD- BA (ZCMV 7232), collected on n unknown dte in Ambtorom by J.E. Rndrinirin; ZSM 225/2016 (ACZCV 0121), collected on 8 November 2013 t Shmbendrn (S , E , 447 m.s.l.), Betmpon Strict Nture Reserve by G.M. Ros, D.J. Hrris, M. Rndrimiliso, nd H. Lv; ZSM 226/2016 (AC- ZCV 0224) nd ZSM 227/2016 (ACZCV 0225), collected on 16 November 2013 t Vohitsivln (S , E , 487 m.s.l.), Betmpon Strict Nture Reserve by A. Crottini, D. Slvi, E. Scnrini, nd George. Referred specimen. ZSM 490/2005 (ZCMV 889), with sme collecting dt s holotype, is not included in the type series due to the lck of relible moleculr dt. Dignosis. A species from north estern lowlnds of Mdgscr which hs not previously been included in ny study s seprte cndidte species; listed under Stumpffi tetrdctyl in Glw & Vences (2007), Wollenberg et l. (2008), nd Vieites et l. (2009). (1) Modertely lrge-sized species (femle SVL mm); (2) mnus with four fingers (first finger slightly reduced in length) nd pes with five toes (first toe distinctly reduced in length); (3) terminl phlnges of fingers with very slightly enlrged discs, those of toes with modertely enlrged discs; (4) reltive hnd nd foot length, HAL/ SVL , FOTL/SVL ; (5) dorsum modertely tuberculr, with numerous distinct scttered tubercles which sometimes re lrge nd even form smll ridges; (6) dorslly tn, with lrge indistinct symmetricl drk mrkings. Ventrl skin trnsprent to trnslucent in life, with few crem nd drk gry flecks behind the pectorl girdle, on the chin, nd on the ventrl legs, without ny red color elements. Distinguished from S. nlmin, S. ngeluci, S. contumeli, S. dvidttenboroughi, S. gimmeli, S. huwei, S. ih rn, S. lrinki, S. mledict, S. mdgscriensis, S. mmitik, S. miery, S. obscoen, S. psologloss, S. pyg me, S. sort, S. tridctyl, nd S. ynniki by distinctly lrger body size; from S. be, S. kibomen, S. mei kee, nd S. roseifemorlis by the lck of red color ventrlly or on limbs; from S. grndis by the lck of contrsted light blue-blck ventrl pttern. Furthermore, distinguished from S. contumeli, S. dvidttenboroughi, S. miery, S. obscoen, S. tetrdctyl, nd S. tridctyl by distinctly lower degree of digitl length reduction; from S. be, S. hr, nd S. stffordi by smller body size, nd from them nd S. megsoni lso by stronger length reduction of first toe. Within clde C2 the species is the direct sister group to S. chillei, which lso occurs in the estern lowlnds nd which is rther closely relted (16S divergence 3.0%). A biocoustic comprison of the two species is currently impossible due to the lck of cll recordings of S. nlnjirofo; however, due to concordnt differentition in mitochondril nd nucler genes, the bsence of cler brod dorsl light brown stripe, nd the strikingly tuberculr dorsum of S. nlnjirofo, we here describe it s distinct species, hypothesis tht requires confirmtion from future studies. Description of the holotype. Specimen in good stte of preservtion, prt of right thigh removed s tissue smple. Body elongte; hed slightly wider thn long, nrrower thn body width, snout slightly pointed in dorsl view, slightly pointed in lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis not distinct, concve; lorel region concve, oblique; tympnum distinct, bout 68% of eye dimeter; suprtympnic fold not visible; tongue long, slightly brodening posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone rounded. Forelimbs slender; subrticulr tubercles single, round; outer metcrpl tubercle distinct, smll, ovl; inner metcrpl tubercle distinct, elongte; fingers without webbing; no reduction of fingers; reltive length of fingers 1 < 2 < 4 < 3, fourth finger longer thn second; finger tips expnded into discs. Hind limbs slender; TIBL 49% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle smll, ovl; outer mettrsl tubercle bsent; no webbing between toes; first toe strongly reduced; toe tips expnded; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe shorter thn third; subrticulr tubercles indistinct. Skin on dorsum reltively smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After 10 yers in 70% ethnol, the dorsum is beige mottled with brown. A brown chevron rises nteriorly from the inguinl region. Inguinl spots re present. The dorsl surfce of the hed is beige mottled with brown. A brown hert shpe is present 354

85 VERTEBRATE ZOOLOGY 67 (3) 2017 b c d Fig. 65. Stumpffi nlnjirofo sp. nov. in life: ( b) holotype ZSM 489/2005 (ZCMV 873) from Nosy Mngbe; (c) ZSM 226/2016 (ACZCV 0224) from Betmpon; (d) ZSM 227/2016 (ACZCV 00225) from Betmpon. in the scpulr region. The nostril is light brown. The lterl surfce of the hed is beige mottled with brown. The flnks re s the lterl surfce of the hed. The flnk colortion merges with the ventrl colortion. The ventrl trunk is uniformly beige in the middle nd lterlly beige mottled with brown. The chin is more mottled with brown thn the trunk. The ventrl thigh is s the chin. The shnk ventrlly is more mottled with brown thn the thigh. The trsus nd foot re ventrlly s the shnk. Dorslly, the thigh is beige mottled with brown, with drk brown crossbnd. The posterodorsl surfce of the thigh is beige mottled with brown. The shnk is s the trsus with two fine drk brown crossbnd. The trsus is s the shnk without crossbnds. The foot nd toes re s the shnk. The clocl region is brown. The rms re beige mottled with brown. A drk brown crossbnd is present on the forerm. The dorsomedil surfce of the hnd is beige mottled with brown. The fingers hve smll white crossbnds. The underside of the rm is s the lterl prt of the ventrl trunk. Color in life (holotype nd vrition). Dorsl bse color tn. Chocolte mrkings re present over the suprscpulr region (common lso to ll of the prtypes) nd in chevron from the inguinl region to the mid-bck (more or less complete in ll prtypes). A drk spot is present on the dorsl hed between the eyes of the holotype (possibly scr). The flnk is flecked with irregulr ebony mrkings, including one round the nostril, one over the tympnum (present in ll prtypes), one over the insertion of the forelimb (present in ll prtypes), nd one on the midbody (present in ll specimens). The dorsl forelimb is s the bck, with one incomplete ebony crossbnd on the forerm. The hnd is mottled tn nd ebony. A light nnulus is present on ech finger before the terminl phlnge. The dorsl hindlimb is s the bck, with two chocolte crossbnds on the thigh nd three on the shnk. Three crossbnds re present on the feet of the prtypes, nd presumbly lso present on the holotype, but not visible from imges. The toes re tn, with white nnulus before ech terminl phlnge. The ventrl skin is mostly trnsprent. Mottled crem nd brown pigment is present on the bdomen, chin, nd legs. The iris is golden reticulted with blck. Vrition. For vrition in mesurements mong specimens, see Tble 3. In generl, ll exmined specimens gree strongly with the holotype in morphology. In the prtypes, brown interoculr br is present. Prtypes from Betmpon Strict Nture Reserve hve tubercles on 355

86 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs the bck, which cn be prticulrly numerous, such s in ZSM 227/2016 (ACZCV 0225). ZSM 226/2016 (AC- ZCV 0224), nd ZSM 227/2016 (ACZCV 0225) present shde of copper colortion on top of the overll tnned color. The tn/ebony pttern in some specimens is strongly contrsted s in ZSM 225/2016 (ACZCV 0121), but in others quite muted. Etymology. The species nme is noun in pposition to the genus nme nd refers to the known rnge of the new species with loclities in the so-clled Anlnjirofo region in estern Mdgscr. Distribution. Known from three loclities in estern Mdgscr: (1) Nosy Mngbe Specil Reserve (type loclity), (2) Betmpon Strict Nture Reserve, nd (3) Ambtorom. In Betmpon this species hs thus fr been found only round Shmbendrn nd Vohitsivln cmpsites, which re locted in the center nd northwest of the Reserve, respectively. Nturl history. In Betmpon this species seems to be uncommon (or hve secretive hbits) nd so fr restricted to intermedite elevtions within the protected re (records re thus fr vilble from m.s.l.). Cll. Unknown. Stumpffi diutissim sp. nov. (Figures 66 nd S113) Holotype. ZSM 378/2005 (FGZC 2751), n dult mle, collected on 14 Februry 2005 in Cmp Mntell, Mrojejy Ntionl Prk, (S14.421, E , 481 m bove se level), Antsirnn Province, Mdgscr by F. Glw, M. Vences, nd R.D. Rndriniin. Prtypes. ZSM 376/2005 (FGZC 2742) nd UADBA (FGZC 2730) collected on 14 nd 15 Februry 2005 in Cmp Mntell, Mrojejy Ntionl Prk (S14.421, E , 481 m.s.l.) by F. Glw, M. Vences, nd R.D. Rndriniin; ZSM 495/2005 (ZCMV 2067) collected on 18 Februry 2005 in Cmp Mrojeji, Mrojejy Ntionl Prk, (S , E , 746 m.s.l.) by F. Glw, M. Vences, nd R.D. Rndriniin; ZSM 496/2005 (ZCMV 2082) collected on 19 Februry 2005 in Mrojejy Ntionl Prk between Cmp Mrojeji nd Cmp Mntell by F. Glw, M. Vences, nd R.D. Rndriniin; UADBA-ZCMV 15057, ZSM 546/2016 (ZCMV 15064), ZSM 548/2016 (ZCMV 15077), UADBA-ZC- MV 15078, UADBA-ZCMV 15116, nd ZSM 547/2016 (ZCMV 15117), ll collected on 15 November 2016 t cmpsite clled Cmp 0 in Mrojejy Ntionl Prk (S , E , 310 m.s.l.) by A. Rkotorison, M.D. Scherz, M.C. Bletz, J.H. Rzfindribe, A.Rzfimnntso, nd M. Vences; ZSM 549/2016 (ZCMV 15144) nd ZSM 553/2016 (ZCMV 15153) collected on 16 November 2016 in Cmp Mntell, Mrojejy Ntionl Prk (S , E m.s.l.) by A. Rkotorison, M.D. Scherz, M.C. Bletz, J.H. Rzfindribe, A. Rzfimnntso, nd M. Vences. Dignosis. A modertely lrge-sized species from the north-est of Mdgscr. It hs been previously listed s Stumpffi sp. 13 in Vieites et l. (2009), Köhler et l. (2010), Scherz et l. (2016) nd Peloso et l. (2017), s Stumpffi sp. 3 in Wollenberg et l. (2008), s Stumpffi sp. Mrojejy 1 in Glw & Vences (2007). It might lso correspond to the species listed nd figured s Stumpffi sp. c in Glw & Vences (1994). (1) Modertely lrge-sized species (SVL up to 20.0 mm; dult mle SVL mm); (2) mnus with four fingers (first finger slightly reduced in length) nd pes with five toes (first toe distinctly reduced in length); (3) terminl phlnges of fingers with very slightly, those of toes with slightly enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum smooth; (6) dorslly highly vrible in colortion, brownish to reddish to yellowish with drk brown to blck pttern or mottling, with distinctly lighter ptch on the posterodorsl shnk. Ventrlly light with or without drk mottling, especilly in the throt region; (7) regulrly repeted single-note tonl cll with short note durtion. Distinguished from S. nlmin, S. ngeluci, S. contumeli, S. dvidttenboroughi, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. mledict, S. mdgscriensis, S. mmitik, S. miery, S. obscoen, S. psologloss, S. pygme, S. sort, S. tridctyl, nd S. ynniki by lrger body size of most specimens; from S. be, S. kibomen, S. meikee, nd S. roseifemorlis by the lck of red color ventrlly or on limbs; from S. grndis by the lck of contrsted light blue-blck ventrl pttern. Furthermore distinguished from S. contumeli, S. dvidttenboroughi, S. miery, S. obscoen, S. tetrdctyl, nd S. tridctyl by distinctly lower degree of digitl length reduction; from S. hr, S. megsoni, nd S. stffordi by smller body size nd stronger length reduction of first toe; from S. nlnjirofo by less tuberculr dorsum; from S. psologloss by n unpulsed cll; from S. nlmin, S. gimmeli, S. mdgscriensis, S. pygme, S. ngeluci, S. lrinki, S. mledict, S. mmitik, S. be, S. tridctyl, S. obscoen, nd probbly lso S. huwei nd S. kibomen, by shorter dvertisement cll durtion; nd from the symptric S. chillei by much longer inter-note intervl durtion. Description of the holotype. Specimen in good stte of preservtion, right thigh muscle removed s tissue smple. Body elongte; hed slightly wider thn long, nrrower thn body; snout pointed in dorsl view, pointed in lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis distinct, stright; lorel region stright, slightly oblique; tympnum distinct, bout 67% of eye dimeter; suprtympnic fold not visible; tongue long, brodening posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone rounded. Forelimbs slender; subrticulr tubercles single, indistinct; outer metcrpl tubercle indistict, rounded; prepollex present, lrge, ovl; fingers without webbing; first finger reduced; reltive length of fingers 1 < 2 < 4 < 3, fourth finger distinctly longer thn second; finger tips not expnded into discs. Hind limbs slender; tibiotrsl rticultion reching nterior eye corner; TIBL 52% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle smll, thin, ovl; outer mettrsl tubercle bsent; no 356

87 VERTEBRATE ZOOLOGY 67 (3) 2017 Fig. 66. Stumpffi diutissim sp. nov. from Mrojejy Ntionl Prk in life: () ZSM 546/2016 (ZCMV 15064); (b) ZSM 549/2016 (ZCMV 15144); (c) ZSM 547/2016 (ZCMV 15117). Smll inset photos showing ventrl surfces. webbing between toes; first toe strongly reduced; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe distinctly shorter thn third. Skin on dorsum smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After 10 yers in 70% ethnol, the dorsum is brown. Drk irregulr mottling is present on the bck. No distinct spot is present in the inguinl region. The dorsl surfce of the hed is s the bck. A drk horizontl interoculr br is present. The nostril is surrounded by brown. The lterl surfce of the hed is brown, with few crem flecks below the tympnic region. The flnks fde to lighter brown ventrlly, nd b c become incresingly spotted with crem. The flnk colortion merges with the ventrl colortion; the ventrl trunk is light brown mottled with crem. The chin is s the ventrl trunk but with less crem. The ventrl thigh is light brown with few crem dots. The shnk nd trsus re ventrlly s the thigh. The sole of the foot is light brown. Dorslly, the thigh is s the dorsum. The posterodorsl surfce of the thigh is light brown like the ventrl surfce. The shnk is dorslly lso s the thigh, except for light re from the heel to the mid-shnk on the internl surfce. The trsus nd foot re dorslly s the shnk; the foot is dorsomedilly crem. The toes re light brown with crem flecks. The clocl region is drk brown. The rms re s the dorsum. The dorsomedil surfce of the hnd is crem. The fingers hve smll white flecks. The underside of the rm is s the ventrl trunk. Color in life (vrition). Colortion is highly vrible in life. Dorslly cn be mostly ptternless brown with hed drk sooty ornge, lightening through burnt umber to brown posteriorly (unidentified specimen Fig. 66), or cn be yellowish, reddish, or brownish in bse color with series of blck to russet mrkings on the bck (Fig. 66b). Whtever the dorsl pttern, the legs nd rms mtch it, such tht the plin brown specimen hs plin brown rms nd legs, wheres ZSM 546/2016 hs yellow nd blck rms nd legs to mtch its dorsum, nd ZSM 549/2016 hs light brown rms with drk brown spots to mtch its dorsum, etc. In ll specimens, the posterior shnk is lighter color thn the rest of the legs, rnging from coppery to lemon yellow to cly (Fig. 66). In some specimens, there re hints of pttern elements tht might give rise to teddyber shpe if connected, such s hert-shped mrking over the posterior hed in ZSM 549/2016. A white nnulus is lwys present on ech toe before the terminl phlnge. The ventrl skin is trnslucent nd consequently plum to flesh in color, flecked with white over the bdomen nd ventrl legs (Fig. 66). Iris copper round the pupil, drkening externlly. Vrition. For vrition in mesurements mong specimens, see Tble 3. In generl, ll exmined specimens gree with the holotype in morphology. In ll prtypes, the cnthus rostrlis is concve nd distinct, the first toe is less reduced thn in the holotype, nd toes 2 5 hve slightly expnded terminl discs. Severl specimens lck prepollices, nd hve smll, ovl inner mettrsl tubercles. A high degree of vribility ws noted in the colortion of specimens in preservtive, tht reflects lso their vribility in life. In ll specimens, lighter re is present between the mid-shnk nd heel. This is most strongly visible in ZSM 495/2005, nd wekest in the holotype. Wholly brown chin ws noted only in the holotype nd ZSM 495/2005, which re lso the only specimens with distinct prepollices. Etymology. The species nme is derived from the Ltin dverb diutissime mening for long time nd refer- 357

88 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b Fig. 67. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi diutissim sp. nov. from Mrojejy Ntionl Prk (cll recorded from ZSM 549/2016, ZCMV 15144): () 1 s durtion section; (b) 6 s durtion section. ring to the considerble mount of time it took to describe this species fter its initil collection nd identifiction s new species in The species nme is used s n djective in the feminine nomintive singulr s lso done previously by some Ltin uthors. Distribution. The species is known only from Mrojejy Ntionl Prk t elevtions between m.s.l. Nturl history. Clling mles were found in the lef litter of primry rinforest in the fternoon nd in the evening. Cll. The dvertisement cll of S. diutissim consists of single, short, tonl note emitted in series t regulr intervls (Fig. 67). Clls recorded by A. Rkotorison on 16 November 2016 from specimen ZSM 549/2016 (ZCMV 15144) in Mrojejy Ntionl Prk t n ir temperture estimted round 22 C hd the following prmeters: cll durtion (= note durtion) ms (54.6 ± 1 ms; N = 10), inter-cll intervls ms ( ± ms; N = 10), nd dominnt frequency t Hz ( ± 51.8 Hz, N = 10). Stumpffi edmondsi sp. nov. (Figures 68 nd S114) Holotype. ZSM 371/2005 (FGZC 2677), collected on 9 Februry 2005 from Vohidrzn (S18.95, E48.5, c m bove se level), Tomsin Province, Mdgscr by F. Glw, R.D. Rndriniin, nd R. Dolch. Prtype. ZSM 1731/2012 (RDR 1065), collected on 6 November 2011 in the Andsibe region t the entrnce to the Anlmzotr Forest Sttion (sometimes clled the Mitsinjo Reserve), ner the rod leding to Andsibe, by M. Vences, D. Edmonds, R.D. Rndrininin, nd A. Rkotorison. Dignosis. A rther insconspicuous species from the northern centrl est which hs been previously listed s Stumpffi sp. 8 in Vieites et l. (2009), Scherz et l. (2016) nd Peloso et l. (2017), nd s Stumpffi sp. 2 in Wollenberg et l. (2008). (1) Modertely-sized species (dult mle SVL 17.4 mm); (2) mnus with four fingers (first finger modertely reduced in length) nd pes with five toes (first toe distinctly reduced in length); (3) terminl phlnges of fingers without, those of toes with very slightly enlrged discs; (4) reltive hnd nd foot length, HAL/SVL 0.25, FOTL/SVL ; (5) dorsum smooth; (6) dorslly ebony, with irregulr nd poorly contrsted blck ptches nd speckled with few smll iridescent whitish flecks. Posterodorsl shnk cn be burnt ornge. Ventrlly trnslucent skin, pricot on the posterior belly nd ventrl inguinl region; (7) regulrly repeted single-note tonl cll (herd in the wild but not recorded). Distinguished from S. nlmin, S. ngeluci, S. contumeli, S. dvidttenboroughi, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. mledict, S. mdgscriensis, S. mmitik, S. miery, S. obscoen, S. psologloss, S. pygme, S. sort, S. tridctyl, nd S. ynniki by lrger body size; from S. be, S. kibomen, S. meikee, nd S. roseifemorlis by the lck of lrge nd distinct res of red color ventrlly or on limbs (except for limited mount of burnt ornge on posterodorsl shnk); from S. grndis by the lck of contrsted light blueblck ventrl pttern; from S. be, S. hr, S. megsoni, nd S. stffordi by smller body size nd stronger length reduction of first toe; from S. chillei by dorsl color pttern; from S. nlnjirofo by less tuberculr dorsum. Furthermore distinguished from S. contumeli, S. dvidttenboroughi, S. miery, S. obscoen, S. tetrdctyl, nd S. tridctyl by distinctly lower degree of digitl length reduction. Stumpffi diutissim hs somewht 358

89 VERTEBRATE ZOOLOGY 67 (3) 2017 b Fig. 68. Stumpffi edmondsi sp. nov. from Andsibe in life: ( b) dorsl nd ventrl view of prtype ZSM 1731/2012 (RDR 1065). similr color pttern but belongs to different subclde of C1, nd, though vrible in colortion, seems to hve more trnsprent ventrl skin. Description of the holotype. Specimen in good stte of preservtion, right thigh muscle removed s tissue smple for DNA extrction. Body elongte; hed slightly longer thn wide, nrrower thn body width; snout pointed in dorsl nd lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis indistinct, concve, lorel region concve, oblique; tympnum distinct, bout 42% of eye dimeter; suprtympnic fold not visible; tongue long, slightly brodening posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone rounded. Forelimbs thick; subrticulr tubercles single, slightly distinct; outer metcrpl tubercle smll, ovl; prepollex distinct, thick, elongte; fingers without webbing; first finger reduced; reltive length of fingers 1 < 2 < 4 < 3, fourth finger longer thn second; finger tips slightly expnded into discs. Hind limbs slender; TIBL 53% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle smll, elongte; outer mettrsl tubercle bsent; no webbing between toes; first toes reduced; toe tips slightly expnded; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe shorter thn third; subrticulr tubercles distinct, single. Skin on dorsum reltively smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After 10 yers in 70% ethnol, the dorsum is beige with brod brown bnd running from the scpulr region to the inguinl region. Elongte brown ptches re present in the inguinl region. The dorsl surfce of the hed is drk brown. The nostril is brown. The lterl surfce of the hed is beige mottled with brown. The flnks re beige mottled with brown spots, which continues ventrlly onto the ventrl bdomen nd legs. The sole of the foot is brown mottled with beige. Dorslly, the thigh is brown mottled with beige, including its posterodorsl surfce. The shnk, trsus, nd foot re s the thigh, seemingly without distinctly different color to the posterodorsl shnk. The clocl region is brown. The rms re dorslly nd ventrlly s the dorsum. The dorsomedil surfce of the hnd is s the lower rm. The fingers re crem mottled with brown. Color in life (prtype ZSM 1731/2012). Dorsl bse colortion ebony. Irregulr, symmetricl blck mrkings re present, most notbly forming oblong spots in the inguinl region, nd n interoculr br on the posterior hlf of the eye. This br is nteriorly bordered by lighter brown. The dorsum is sprsely interspersed with iridescent whitish flecks. The dorsl forelimb is s the dorsum. The hnd is internlly rosy brown, lterlly s the rest of the forelimb, with smll whitish nnulus before ech terminl phlnge. The hindlimbs re s the dorsum except tht the posterior shnk is burnt ornge. The foot is dorslly s the shnk, with smll whitish nnulus before ech terminl phlnge. The ventrl skin is trnslucent. Whitish flecks re present on the lower jw. Silvery flecks re present t nd behind the pectorl gridle, becoming decresingly dense posteriorly. The lterl bdomen is pricot. The ventrl legs re tupe. The iris is drk. Vrition. For mesurements of the holotype nd prtype, see Tble 3. In generl, the prtype specimen grees strongly with the holotype in morphology, except tht the tongue brodens more strongly posteriorly; smll plmr metcrpl tubercle distinct, round; severl smll differences in color pttern exist, the most notble of which is the presence of distinct, burnt ornge posterodorsl shnk in life (beige in preservtive), which is bsent from the holotype. Etymology. The species nme is ptronym honoring Devin Edmonds to whom we re gld to dedicte this species in recognition for his efforts in mphibin conservtion in Mdgscr, nd for his help in collecting the prtype specimen of this new species. 359

90 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Description of the holotype. Specimen in good stte of preservtion, the right foot removed s tissue smple for DNA extrction. An incision ws mde in the right flnk to check the sex. Body elongted; hed slightly wider thn long, nrrower thn body; snout pointed in dorsl view, slightly pointed in lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis distinct, concve; lorel region concve, slightly oblique; tympnum distinct, bout 69% of eye dimeter; suprtympnic fold not visible; tongue long, brodening posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone rounded. Forelimbs slender; subrticulr tubercles single, indistinct; outer metcrpl tubercle subtringulr; plmr tubercle round; inner metcrpl tuber b Fig. 69. Stumpffi fusc sp. nov. from Ambodivongy in life: ( b) dorsolterl nd ventrl view of prtype ZSM 436/2010 (FGZC 4253). Distribution. The species is known from forests t Vohidrzn nd ner Andsibe, round m bove se level. Nturl history. One clling mle ws found in dense lef litter long the tril in Anlmzotr forest ner Andsibe. Cll. Unknown. Stumpffi fusc sp. nov. (Figures 69 nd S115) Holotype. ZSM 437/2010 (FGZC 4254), n dult ovigerous femle, collected on 2 April 2010 in Ambodivongy (S , E , c. 100 m bove se level), Tomsin Province, Mdgscr by F. Glw, J. Köhler, P.-S. Gehring, M. Pbijn, nd F.M. Rtsovin. Prtype. ZSM 436/2010 (FGZC 4253) with sme collecting dt s holotype. Referred specimens. UADBA (FGZC 4207) collected on 31 Mrch 2010 in Ambodivongy (S , E , c. 287 m.s.l.) by F. Glw, J. Köhler, P.-S. Gehring, M. Pbijn, nd F.M. Rtsovin (not included s prtype becuse no DNA sequence dt is vilble for this specimen). Dignosis. A rther insconspicuous species from northestern lowlnds of Mdgscr which hs been previously listed s Stumpffi sp. 33 in Perl et l. (2014) wheres the species ws not included in Scherz et l. (2016); in tht study, Stumpffi sp. C33 corresponds to S. meikee, described bove. (1) Modertely-sized species (femle SVL 17.7 mm); (2) mnus with four fingers (first finger slightly reduced in length) nd pes with five toes (first toe very strongly reduced in length); (3) terminl phlnges of fingers without, those of toes with modertely enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum smooth with numerous smll, distinct regulrly scttered tubercles; (6) dorslly twny, with lrge symmetricl blck copper-infused mrkings covering much or most of the dorsum. Ventrl skin trnslucent with mny lrge crem flecks, nd without ny red color elements. Distinguished from S. nlmin, S. ngeluci, S. contumeli, S. dvidttenboroughi, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. mledict, S. mdgscriensis, S. mmitik, S. miery, S. obscoen, S. psologloss, S. pygme, S. sort, S. tridctyl, nd S. ynniki by lrger body size in unmbiguously dult specimens; from S. be, S. kibomen, S. meikee, nd S. roseifemorlis by lck of red color ventrlly or on limbs; from S. grndis by the lck of contrsted light blue-blck ventrl pttern; from S. be, S. hr, S. megsoni, nd S. stffordi by smller body size. From S. chillei, S. diutissim, nd S. edmondsi, nd in generl from most other lrge-sized Stumpffi, by the more expressed length reduction of first toe. Furthermore distinguished from S. contumeli, S. dvidttenboroughi, S. miery, S. obscoen, S. tetrdctyl, nd S. tridctyl by lower degree of digitl length reduction (especilly on fingers). By color pttern reminiscent of the closely relted S. chillei nd S. nlnjirofo, but these species differ strongly in mitochondril nd nucler DNA sequences, nd the specimens exmined hve less expressed length reduction of first toe; S. nlnjirofo my lso hve somewht more tuberculr dorsum. 360

91 VERTEBRATE ZOOLOGY 67 (3) 2017 cle oblong, distinct; fingers without webbing; first finger slightly reduced; reltive length of fingers 1 < 2 < 4 < 3, fourth finger slightly shorter thn second; finger tips not expnded into discs. Hind limbs slender; TIBL 51% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle ovl, distinct; outer mettrsl tubercle bsent; no webbing between toes; first toe strongly reduced; toe tips expnded; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe distinctly shorter thn third; subrticulr tubercles distinct, single. Skin on dorsum smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After five yers in 70% ethnol, the dorsum is brown. A drk brown hert-like mrking is present on the hed, extending to the shoulder region. This is followed posteriorly by two blckish spots either side of the midline. These re followed by lrge, oblique, light brown mrkings extending down the flnks. A blck spot is present in the inguinl region (two smll spots on the right). Finlly, tringulr light brown ptch is present from between the inguinl regions to the vent. The nostril is surrounded by drk brown. The lterl surfce of the hed is beige speckled with blck. Drk flecks re present under the eye nd over the tympnum. The flnks fde from the dorsl mrkings to the crem of the venter. The ventrl trunk is crem. The chin is flecked with light brown. The ventrl thigh is light brown flecked with crem. The shnk is ventrlly light brown with few crem flecks. The trsus is ventrlly s the shnk. The sole of the foot is brown. Dorslly, the thigh is beige with two oblique brown crossbnds. The posterodorsl surfce of the thigh is crem with brown flecks. The shnk is light brown with three oblique crossbnds. The trsus is s the shnk with one crossbnd. The foot is light brown with drk brown flecks. The toes re crem speckled with brown. The clocl region is drk brown. The rms re beige speckled with drk brown. A drk crossbnd is present on the lower rm. The dorsomedil surfce of the hnd is crem. The fingers hve smll brown speckles. The underside of the rm is s the ventrl trunk. Color in life (prtype ZSM 436/2010). Dorsl bse colortion is twny, with semi-symmetricl blck mrkings with iridescent copper infusions: pir of brod lines converging on the midline from the inguinl region, pir of brod mrkings running from behind the suprscpulr region to the flnk, nd sub-tringulr mrking strting between the eyes nd nrrowing to point between the suprscpule. The flnk fdes to crem mottled with tupe, nd possesses two blck spots, one posterodorsl to the insertion of the rm, nd one posterodorsl to the tympnum. The lterl hed hs two further blck spots, one brod beneth the eye, nd one on the nostril. The dorsl rm is s the dorsum, interspersed with blck flecks. A whitish nnulus is present before the terminl phlnge of ech finger. The dorsl leg is s the bck, with one blck crossbnd with copper infusions on the mid-thigh, one on the mid-shnk, nd two on the foot. The toes re mottled, with light bnd present before ech terminl phlnge. The ventrl skin is trnslucent nd thus plum, densely flecked with crem mrkings on the chin nd posterior to the pectorl gridle, fding posteriorly. Two off-white mrkings re present on the bottom lip on either side of the hed, one beneth the eye, one beneth the nostril. The nterior point of the mndible is lso off-white. The ventrl legs re plum flecked with drk gry. The iris is copper. Vrition. For vrition in mesurements mong specimens, see Tble 3. In generl, ll exmined specimens gree strongly with the holotype in morphology. A low degree of vribility ws noted in the colortion of specimens; ZSM 436/2010 is lmost identicl to the holotype in dorsl colortion, but overll lighter. Ventrlly it is clen crem. Etymology. The species nme is the feminine nomintive singulr form of the Ltin djective fusc = brown. It refers to the vrious brown mrkings of the dorsum of this species. Distribution. The species is known only from its type loclity Ambodivongy, t c. 100 m bove se level. Nturl history. At the type loclity, specimens were collected from the lef litter on the forest floor during dytime. Nothing else is known of the species hbits. Cll. Unknown. Stumpffi jennoeli sp. nov. (Figures 70 nd S116) Holotype. ZSM 206/2016 (ACZCV 0056), dult mle, collected on 8 November 2013, t Shindrn (S , E , 344 m bove se level), Betmpon Strict Nture Reserve, Tomsin Province, Mdgscr by A. Crottini, D. Slvi, E. Scnrini, nd J.H. Velo. Prtypes. MRSN A6283 (FAZC 13682), collected on 28 Februry 2007 t Vohitsivln (S E , 517 m.s.l.) by G.M. Ros; MRSN A6386 (FAZC 13903), dult mle, collected on 19 November 2007 long Piste Principle (S , E , 514 m.s.l.) by G.M. Ros, F. Andreone, nd J. Nöel; MRSN A6281 (FAZC 13599), MRSN A6284 (FAZC 13587), MRSN A6245 (FAZC 13556) nd MRSN A6284 (FAZC 13587), ll collected between 7 nd 11 Februry 2007 t Shmbendrn (S , E , 458 m.s.l.) by G.M. Ros nd F. Andreone; MRSN A6261 (FAZC 13711) nd MRSN A6287 (FAZC 13729) collected on 6 nd 8 Mrch 2007 t Shbefoz (S , E , 349 m.s.l.) by G.M. Ros; MRSN A6307 (FAZC 13622), MRSN A6318 (FAZC 13656), nd MRSN A6277 (FAZC 13651) collected between 19 nd 22 Februry 2007 t Mintimbto (S E , 255 m.s.l.) by G.M. Ros; MRSN A6346 (FAZC 13900) collected on 18 November 2007 t Mintimbto (S , E , 277 m.s.l.) by G.M. Ros, F. Andreone nd J. Nöel; ZSM 208/2016 (ACZCV 0218) collected on 15 November 2013 t Vohitsivln (S , E , 481 m.s.l.) by A. Crottini, D. Slvi, E. Scnrini, nd George; ZSM 207/2016 (ACZCV 0167) collected on 22 November 2013 t Mintimbto (S , E , 274 m.s.l.) by G.M. Ros, D.J. Hrris, M. Rndrimiliso, nd H. Lv; ZSM 209/2016 (ACZCV 0118) 361

92 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b c d e f g h i j k l Fig. 70. Stumpffi jennoeli sp. nov. prtype specimens from Betmpon Strict Nture Reserve in life: ( b) ZSM 208/2016 (ACZCV 0218); (c) ZSM 211/2016 (ACZCV 0062); (d) ZSM 210/2016 (ACZCV 0070); (e) ZSM 209/2016 (ACZCV 0118); (f) ZSM 207/2016 (ACZCV 0167); (g) MRSN A6245; (h i) MRSN A6307; (j) MRSN A6346; (k l) MRSN A6386. collected on 07 November 2013 t Shmbendrn (S , E , 455 m.s.l.) by G.M. Ros, D.J. Hrris, M. Rndrimiliso, nd J.H. Lv; ZSM 210/2016 (ACZCV 0070) collected on 11 November 2013 t Rendrirendry (Piste Fotsimvo) (S , E , 204 m.s.l.) by A. Crottini, D. Slvi, E. Scnrini, George, G.M. Ros, D.J. Hrris, M. Rndrimiliso, nd H. Lv; ZSM 211/2016 (ACZCV 0062) nd ZSM 212/2016 (ACZCV 0063) collected on 7 nd 8 November 2013 t Shindrn (S , E , 344 m.s.l.) by A. Crottini, D. Slvi, E. Scnrini, nd J.H.Velo; ZSM 213/2016 (ACZCV 0247) collected on 7 nd 8 November 2013 t Shbefoz (S , E , 478 m.s.l.) by A. Crottini, D. Slvi, E. Scnrini, J.H.Velo, F. Andreone, J. Nöel, nd S. Frvelli. All these loclities refer to sites within Betmpon Strict Nture Reserve, Tomsin Province, Est Mdgscr. Dignosis. A modertely sized species from the northern centrl est which hs been previously listed s Stumpffi sp. 20 in Vieites et l. (2009), Scherz et l. (2016) nd Peloso et l. (2017) nd s Stumpffi sp. [C FJ559315] nd s Stumpffi sp. ff. grndis [C HM364799] in Ros et l. (2011, 2012). (1) Modertely-sized species (SVL mm, dult mle SVL 17 mm); (2) mnus with four fingers (first finger modertely reduced in length, especilly in mles) nd pes with five toes (first toe distinctly reduced in length); (3) terminl phlnges of fingers without, those of toes with very slightly enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum smooth with few scttered tubercles; (6) dorslly moch to tn, with vribly intense drker ptches of vrible size nd number. Ventrlly vrible, but lwys with pttern of light spots. The heel is often distinctly different in color to the rest of the leg. No red color ventrlly; (7) regulrly repeted single-note tonl cll. 362

93 VERTEBRATE ZOOLOGY 67 (3) 2017 b Fig. 71. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi jennoeli sp. nov. from Betmpon, MRSN A6386 (FJ559315): () 1 s durtion section; (b) 6 s durtion section. Distinguished from S. nlmin, S. ngeluci, S. contumeli, S. dvidttenboroughi, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. mledict, S. mdgscriensis, S. mmitik, S. miery, S. obscoen, S. psologloss, S. pygme, S. sort, S. tridctyl, nd S. ynniki by lrger body size; from S. be, S. kibomen, S. meikee, nd S. roseifemorlis by lck of red color ventrlly or on limbs; from S. grndis by the lck of contrsted light blue-blck ventrl pttern; from S. be, S. hr, S. megsoni, nd S. stffordi by smller body size; from S. chillei by distinctly longer inter-cll intervls; from S. nlnjirofo nd S. fusc by less tuberculr dorsum nd different dorsl nd ventrl pttern; from S. diutissim by slightly shorter cll durtion nd shorter inter-cll intervls; from S. edmondsi by the brownish rther thn reddish color on dorsl prt of shnk round tibiotrsl rticultion. Furthermore distinguished from S. contumeli, S. dvidttenboroughi, S. miery, S. obscoen, S. tetr dctyl, nd S. tridctyl by lower degree of digitl length reduction (especilly on fingers). Phylogeneticlly, the species is plced in subclde of clde C1 of unresolved internl nodes; within this subclde, morphologiclly similr species re only S. edmondsi nd S. nigrorubr sp. nov. (described below) but both of these species hve t lest some reddish color on hindlimbs nd t lest S. nigrorubr lso differs biocousticlly (see below). Description of the holotype. Specimen in good stte of preservtion, tongue removed s tissue smple for DNA nlysis. Body elongte; hed slightly wider thn long, nrrower thn body width; snout rounded in dorsl nd lterl view; nostrils directed lterlly, not protubernt, nerer to the tip of snout thn to eye; cnthus rostrlis slightly visible, concve; lorel region concve, oblique; tympnum distinct, bout 76% of eye dimeter; suprtympnic fold not distinct; mxillry teeth nd vomerine teeth bsent; chone rounded. Forelimbs slender; subrticulr tubercles single, distinct; outer metcrpl tubercle slightly visible, rounded; inner metcrpl tubercle visible, ovl, fused with the prepollicl tubercle; fingers without webbing; reltive length of fingers 1 < 2 < 4 < 3, fourth finger longer thn second; finger tips not expnded into discs. Hind limbs slender; TIBL 50% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle not visible; outer mettrsl tubercle bsent; no webbing between toes; toe tips slightly expnded; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe slightly shorter thn third; subrticulr tubercles distinct, single. Skin on dorsum reltively smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After three yers in 70% ethnol, the dorsum is uniformly pinkish beige. A brown hert shpe bordered with chmpgne line is present in the scpulr region. A brown ovl is present in the inguinl region. The dorsl surfce of the hed is s the bck with irregulr brown spots. The nostril is surrounded with beige. The lterl surfce of the hed is drk brown spotted with beige. The flnks re lterlly beige flecked with brown, ventrlly beige with white flecks. The lower flnk colortion merges with the ventrl colortion. The ventrl trunk is beige with white flecks bordered with brown line. The chin is s the bdomen. The ventrl thigh is s the bdomen. The shnk is ventrlly beige spotted with drk brown. The trsus nd foot re ventrlly s the shnk but drker. Dorslly, the thigh is pink-reddish nd beige with brown crossbnd. The posterodorsl surfce of the thigh is beige spotted with brown. The shnk nd trsus re s the bck, with reddish color on the posterodorsl shnk. The foot is beige with brown crossbnd. The toes re speckled with brown. The clocl region is drk 363

94 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs brown. The rms re s the dorsum. A drk crossbnd is present on the lower rm. The dorsomedil surfce of the hnd is beige. The fingers re beige spotted with brown. The underside of the rm is s the ventrl trunk. Color in life (vrition mong prtypes; see Fig 70). Dorsl bse colortion rnges from moch to tn. Drker mrkings, oftenly borderd by thin light line, re present, which rnge from fintly drker thn the bse color (e.g. Fig. 70k) to blck nd strongly contrsting (e.g. Fig. 70j). The mrkings follow firly consistent pttern: drk br is present posterior to the mid-oculr line nd is generlly delimited in front with thin chmpgne line; smll fleck is present on the snout; on the dorsum there cn be n X-like mrking between the suprscpule nd second X-like mrking on the middorsum (Fig. 70, c, h, k). The clocl region lso hs trpezoidl drk mrking (wekly visible in Fig. 70, c, d, j, k). The lterl fce is continuous with the dorsl hed, except for the drk nostril. Posterior to the eye, discontinuous, irregulr bnd of drk mrkings runs long the flnk, lwys incorporting the tympnum. The dorsl forelimb is s the dorsum, with severl drk mrkings, including crossbnd on the forerm. The hnd is internlly s the dorsum, nd lterlly drk, with white nnulus before the terminl phlnge of ech finger. The dorsl leg is s the dorsum, with one drk crossbnd t the mid-thigh, one on the shnk, nd up to four on the foot. The posterodorsl shnk tends to hve unique color, which cn be lmost yellow (Fig. 70d, f) or russet (Fig. 70g, j) the nterior forelimb usully hs the sme color s this shnk region, but slightly weker (Fig. 70g). The dorsl foot is s the rest of the leg, the toes being mottled, with white nnulus before ech terminl phlnge. The ventrl colortion is highly vrible, nd cn be lmost ebony brown with numerous whitish flecks (Fig. 70i), drk moch with white flecks nd with trnslucent beige chin nd posterior trunk (Fig. 70l), or beige with lrger white flecks bordered with brown line (Fig.70b). Density, size, nd position of whitish flecks re not consistent. The iris is golden. Vrition. For vrition in mesurements mong specimens, see Tble 3. In generl, ll exmined specimens gree strongly with the holotype in morphology. A low vribility ws noted in the colortion of specimens in preservtive; ACZCV 0218 hs pinkish colortion in the rm nd in the legs; in MRSN A6386 the dorsl pttern is less contrsted, resulting in n lmost uniform brown colortion. Etymology. The species nme is ptronym honoring Jen Nöel, tlented reserch guide nd pssionte herpetologist working t Betmpon Strict Nture Reserve, on behlf of Mdgscr Fun nd Flor Group in recognition of his ssistnce nd friendship. His interest hs contributed boosting herpetologicl reserch in Betmpon nd led to the identifiction of numerous new species of mphibins nd reptiles inhbiting this remnnt forest frgment of the est cost. Distribution. This species is so fr known only from Betmpon Strict Nture Reserve, where it is thought to hve brod distribution, being found in most of the surveyed res (Ros et l. 2012), from round m bove se level. Nturl history. The holotype ws found, together with other individuls of the sme species during the dy in the lef litter. Severl individuls were clling simultneously from hidden positions on the ground. However, the species is most often herd clling t dusk nd t night, prticulrly when it is drizzling or lightly rining. This species ppers to be quite bundnt cross the Reserve. Cll. The dvertisement cll consists of single, short tonl note emitted in series t regulr intervls (Fig. 71). Clls recorded by G.M. Ros (Ros et l., 2011: trck 51) on 6 November 2007 t Betmpon (21:45 h, ir temperture 21ºC) hd the following numericl prmeters: cll durtion (= note durtion) ms (44.2 ± 0.8 ms; N = 6), inter-cll intervls ms ( ± 17 ms; N = 5), nd dominnt frequency t Hz ( ± 76.9 Hz, N = 6). Stumpffi nigrorubr sp. nov. (Figures 72 nd S124) Holotype. ZSM 2448/2007 (ZCMV 5967) (Fig. S124), collected in Februry Mrch 2007 from loclity long the rod between Ifndin nd Tolongoin, Finrntso Province, Mdgscr (pproximtely t S21.35, E47.62) by M. Vences. Prtypes. ZSM 641/2003 (FG/MV ), n dult mle, collected on 15 Jnury 2003 in Rnomfn by F. Glw, M. Puente, L. Rhrivololoniin, M. Thoms, nd D.R. Vieites; ZSM 640/2003 (FG/MV ), dult mle, collected on 15 Jnury 2003 in Rnomfn villge by F. Glw, M. Puente, L. Rhrivololoniin, M. Thoms, nd D.R. Vieites; ZMA (ZCMV 17) collected on 18 Jnury 2004 t the type loclity by M. Vences; ZMA (ZCMV 98) collected on 21 Jnury 2004 in Ambohitsr (S E , 294 m.s.l.) by D.R. Vieites nd I. de l Riv; ZMB (JCR 5), clling mle, collected on 3 September 2010 in Ambolo forest frgment, Rnomfn (S , E , 643 m.s.l.) by J.C. Riemnn nd S.H. Ndrintso; UADBA-A (JCR 29) collected on 12 Mrch 2010 in Ambtolhidimy forest, Rnomfn (S , E , 1006 m.s.l.) by J.C. Riemnn nd S.H. Ndrintso; ZMB (JCR 562), clling mle, collected on 11 Februry 2011 in bnn plnttion, Ann- Fig. 72. Dorsolterl nd ventrl views of life specimens of Stumpffi nigrorubr sp. nov.: (, h) UADBA-A (JCR 29), mle from Ambtolhidimy, Rnomfn; (b) clling mle from Mnombo Reserve; (c) ZMB (NSH 2599), mle from Mnombo villge forest; (d, g) ZSM 640/2003 (FG/MV ) from Rnomfn villge; (e, i) JCR 3154, mle from Mnombo villge forest, see nuptil pds on right hnd; (f) ZMB (JCR 5), mle from Ambolo, Rnomfn, illustrting red colortion on heels nd outer prts of thighs. 364

95 VERTEBRATE ZOOLOGY 67 (3) 2017 b c d e g f h i 365

96 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs itehn, Rnomfn (S , E , 782 m.s.l.) by J.C. Riemnn nd S.H. Ndrintso; UADBA-A (JCR 678), clling mle, collected on 10 Mrch 2011 in bnn plnttion, Andlngin, ner Ifndin (S , E , 515 m.s.l.) by J.C. Riemnn nd S.H. Ndrintso; ZMB (JCR 855), clling mle, collected on 16 April 2011 in Shdikin forest frgment, Rnomfn (S , E , 643 m.s.l.) by J.C. Riemnn nd S.H. Ndrintso; UADBA-A (NSH 1322), collected on 27 Jnury 2011 in Ambolo forest frgment (S , E , 660 m.s.l.) by J.C. Riemnn nd S.H. Ndrintso; ZMB (NSH 1416), grvid femle, collected on 3 Februry 2011 in Andlngin forest frgment (S , E , 551 m.s.l.) by J.C. Riemnn nd S.H. Ndrintso; UADBA- A (NSH 2197) collected on 24 My 2011 in coffee nd bnn plnttion, Andlngin, ner Ifndin (S , E , 450 m.s.l.) by J.C. Riemnn nd S.H. Ndrintso; ZMB (JCR 3245), collected on 4 April 2012 in bnn plnttion, Andlngin, ner Ifndin (S , E , 515 m.s.l.) by J.C. Riemnn nd S.H. Ndrintso; UADBA-A (JCR 3246), clling mle, collected on 3 April 2012 in gllery forest ner Ambtolhy, Rnomfn (coordintes not tken) by J.C. Riemnn nd S.H. Ndrintso; ZMB (NSH 2599) nd JCR 3154, clling mles, collected on 29 Mrch 2012 in forest ner Mnombo villge (S , E , 46 m.s.l.) by J.C. Riemnn, S.H. Ndrintso, M.-O. Rödel, J. Glos, nd A. Rkotorison. Dignosis. A species from the southern centrl est of Mdgscr tht hs previously been listed s Stumpffi sp. 1 in Wollenberg et l. (2008), s Stumpffi sp. 9 in Vieites et l. (2009), Köhler et l. (2010), Scherz et l. (2016) nd Peloso et l. (2017), nd s Stumpffi sp. Rnomfn in Glw & Vences (2007). (1) Modertely-sized species (dult mle SVL mm); (2) mnus with four fingers (first finger distinctly reduced in length t lest in mles) nd pes with five toes (first toe distinctly reduced in length); (3) terminl phlnges of fingers with slightly enlrged discs, those of toes with very slightly enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum smooth to slightly grnulr; (6) dorslly mottled blck nd drk iridescent dirty golden with irregulr nd indistinct pttern of lighter nd drker elements, nd without shrp color border between color of dorsum nd flnks. Posterodorsl prt of shnk with rust red color. Ventrlly with rust red color on posterior belly nd hindlimbs, gry-brownish with white dotting on nterior belly, chest nd throt; (7) regulrly repeted short single-note tonl cll. Distinguished from S. nlmin, S. ngeluci, S. contumeli, S. dvidttenboroughi, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. mledict, S. mdgscriensis, S. mmitik, S. miery, S. obscoen, S. psologloss, S. pygme, S. sort, S. tridctyl, nd S. ynniki by lrger body size; from ll these species nd S. chillei, S. nlnjirofo, S. diutissim, S. fusc, S. grndis, S. hr, S. jennoeli, S. megsoni, nd S. stffordi by the presence of distinct rust red color ventrlly on belly nd hindlimbs, s well s posterodorslly on the shnks; furthermore distinguished from S. contumeli, S. dvidttenboroughi, S. miery, S. obscoen, S. tetrdctyl, nd S. tridctyl by lower degree of digitl length reduction (especilly on fingers); nd from S. nlnjirofo by less tuberculr dorsum. Distinguished from S. be by smller body size nd presence of red color on belly (vs. only on hindlimbs); from S. kibomen by indistinct dorsl pttern nd longer inter-note intervls; from S. roseifemorlis nd S. meikee by drker dorsl color nd presence of red color posterodorslly on shnks. Morphologiclly most similr to S. edmondsi, which is in the sme phylogenetic subclde but does not seem to be the direct sister species of S. nigrorubr, hs less red color on belly, nd does not shre hplotypes in Rg-1. Description of the holotype. Specimen in good stte of preservtion, left thigh muscle removed s tissue smple for DNA nlysis. Body elongte; hed slightly longer thn wide, nrrower thn body; snout pointed in dorsl nd lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis distinct, concve; lorel region concve, oblique; tympnum distinct, bout 61% of eye dimeter; suprtympnic fold not distinct; tongue long, brodening posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone rounded. Forelimbs slender; subrticulr tubercles single, slightly distinct; outer metcrpl tubercle not recognizble; inner metcrpl tubercle distinct, ovl; fingers without webbing; first finger reduced; reltive length of fingers 1 < 2 < 4 < 3, fourth finger longer thn second; finger tips slightly expnded into discs. Hind limbs slender; TIBL 56% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle smll, slightly visible, ovl; outer mettrsl tubercle bsent; no webbing between toes; first toe reduced; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe distinctly shorter thn third. Skin on dorsum reltively smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After eight yers in 70% ethnol, the dorsum is beige vermiculted with brown. The dorsl surfce of the hed is s the bck. The nostril is brown. The lterl surfce of the hed is beige with brown dots. The flnks re beige with brown stripe. The flnk colortion merges with the ventrl colortion. The ventrl trunk is crem mottled with brown. The chin is drker thn the bdomen. The ventrl thigh is crem speckled with brown. The shnk is ventrlly brown speckled with beige. The trsus is ventrlly beige with drker dots. The sole of the foot is beige. Dorslly, the thigh is beige with brown crossbnd. The posterodorsl surfce of the thigh is brown. The shnk nd the trsus re similr to the thigh. The foot is brown mottled with beige. The toes re brown mottled with beige. A crem Fig. 73. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi nigrorubr sp. nov. from Rnomfn: (, d) 1 s nd 6 s durtion sections of mle FG/MV ; (b, e) 1 s nd 6 s durtion sections of mle ZMB 81999; (c, f) 1 s nd 6 s durtion sections of mle ZMB

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98 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b Fig. 74. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi nigrorubr from Mnombo (ZMB 81999): () 1 s durtion section; (b) 6 s durtion section. mrking is present on the posterodorsl fce of the shnk. The clocl region is drk brown. The rms re drk brown. A beige crossbnd is present on the lower rm. The dorsomedil surfce of the hnd is drk brown. The fingers hve beige crossbnds. The underside of the rm is s the ventrl trunk. In life, dorsum mottled blck nd drk iridescent dirty gold. Flnks without color borders. A drk oblong mrking is present on the flnk, s well s fine irregulrly sized whitish spots. Dorsl forelimb proximlly rust red, fding to blck distlly. The hnd is internlly crem, lterlly s the dorsum, with irregulr whitish flecks, including n nnulus before ech terminl phlnge. Hindlimb proximlly blck, becoming rust red dorslly by the mid-shnk. The posterodorsl shnk is strongly rust red. The foot is dorslly blck, with irregulr whitish flecks, including n nnulus before ech terminl phlnge. The ventrl chin is drk plum with smll light flecks, becoming lrger posteriorly. The plum of the nterior trunk merges into strong rust red over the bdomen, which covers the whole ventrl legs. The ventrl forelimbs re plum. Vrition. For vrition in mesurements mong specimens, see Tble 3. In generl, ll exmined specimens gree strongly the holotype in morphology. ZSM 640/2003 (FG/MV ) is strongly fixed; colortion pttern uniformly brown with drk brown dots. Advertisement clls of specimens from Mnombo resemble those from Rnomfn in repetition rte, but differ in dominnt frequency ( Hz in Mnombo vs Hz in Rnomfn; see below). Etymology. The species nme is feminine nomintive singulr djective derived from the Ltin words niger = blck nd ruber = red, in reference to the blck bsl colortion nd red shnk, bdomen, nd ventrl legs of this species. Distribution. This species is known from vrious loclities in the Rnomfn region, including: (1) rod between Ifndin nd Tolongoin (type loclity), (2) Ambohitsr, (3) Ambolo forest frgment, (4) Ambtolhidimy forest, (5) Annitehn, (6) Andlngin, ner Ifndin, (7) Shdikin forest frgment, (8) ner Ambtolhy, nd (9) Rnomfn villge. Furthermore, it occurs (10) in forest ner Mnombo villge/mnombo Specil Reserve, > 120 km south of Rnomfn, constituting the southernmost record for the genus Stumpffi (16S sequences show 3.8% divergence to Rnomfn specimens; GenBnk ccession numbers MF MF768252, not included in Fig. 3). Nturl history. The species occurs in rinforest hbitt in Rnomfn Ntionl Prk nd is lso common in forest frgments nd bnn plnttions. Active individuls were observed in the lef litter during the dy nd t night, however clling mles were only found fter dusk. Grvid femles were found in the lef litter in Februry 2011 in forest frgment in Andlngin. The species hs rther wide ltitudinl rnge, from 46 m t Mnombo up to over 1000 m.s.l. in Ambtolhidimy forest. The kryotype hs been described under the nme S. sp. by Apre et l. (2007) for specimen ZMA (FG/MV ) from Rnomfn villge. Cll. The dvertisement cll consists of single short note emitted in series in slow regulr succession (Figs. 73 nd 74). Recordings re vilble from vrious loclities in the Rnomfn region. (1) Clls recorded by M. Vences on 15 Jnury 2003 t Rnomfn villge from specimen ZSM 640/2003 (Vences et l., 2006: CD3, trck 368

99 VERTEBRATE ZOOLOGY 67 (3) ) hd the following numericl prmeters: cll durtion (= note durtion) ms (67 ± 10 ms; N = 19), inter-cll intervls ms (1472 ± 104 ms; N = 18), nd dominnt frequency t Hz (5283 ± 70.5 Hz, N = 19); (2) Clls recorded by J.C. Riemnn nd S.H. Ndrintso on 11 Februry 2011 t 20:25 h in bnn plnttion ner Ambtolhy, Rnomfn (ir temperture 20.3 C) from specimen ZMB (JCR 562): cll durtion ms (72 ± 3 ms; N = 3), inter-cll intervls ms (1743 ± ms; N = 3), nd dominnt frequency t Hz (5440 ± 24.8 Hz, N = 3); (3) Clls recorded by J.C. Riemnn on 04 April 2012 t 21:15 h in gllery forest ner Ambtolhy (ir temperture 21.5 C) from specimen ZMB (JCR 3245): cll durtion ms (60 ± 3 ms; N = 4), inter-cll intervls ms (1464 ± 49.4 ms; N = 3), nd dominnt frequency t Hz ( ± 61 Hz, N = 4); (4) Clls recorded by J.C. Riemnn nd S.H. Ndrintso on 29 Mrch 2012 t night in forest ner Mnombo villge from specimen ZMB (NSH 2599): cll durtion ms (65 ± 3 ms; N = 5), inter-cll intervls ms (1416 ± 38 ms; N = 4), nd dominnt frequency t Hz (4573 ± 19 Hz, N = 5). Stumpffi prdus sp. nov. (Figures 75 nd S117) Holotype. ZSM 435/2010 (FGZC 4237) (Fig. 75 ), dult mle, collected on 4 April 2010 t Ambodivongy (S , E , c. 100 m bove se level), Tomsin Province, Mdgscr by F. Glw, J. Köhler, P.-S. Gehring, M. Pbijn, nd F.M. Rtsovin. Prtypes. Nine specimens collected t the sme site nd by the sme collectors s holotype: ZSM 432/2010 (FGZC 4210), ZSM 433/2010 (FGZC 4213), ZSM 434/2010 (FGZC 4214), UADBA (FGZC 4216) nd UADBA (FGZC 4217), ll collected on 31 Mrch 2010; ZSM 440/2010 (FGZC 4278) collected on 3 April 2010; ZSM 438/2010 (FGZC 4265), ZSM 439/2010 (FGZC 4268), nd UADBA (FGZC 4239) collected on 2 April 2010; ZSM 230/2016 (ACZCV 0147) collected on 13 November 2013 t Betkonn (S , E , 356 m.s.l.), Betmpon Strict Nture Reserve, by G.M. Ros, D.J. Hrris, M. Rndrimiliso, nd H. Lv; ZSM 770/2009 (ZCMV 11166) collected on 15 My 2009 from Melivinny-Mnompn by J.E. Rndrinirin; MRSN A2651 (FN 7021) collected on 2 December 1997 in Ambolokoptrik by F. Andreone nd J.E. Rndrinirin; MRSN A6204 collected on 17 Mrch 2007 long the Piste Principle in Betmpon Strict Nture Reserve (S , E , 438 m.s.l.) by G.M. Ros; MRSN A6276 (FAZC 13498) collected on 4 Februry 2007 in Rendrirendry (S , E , 325 m.s.l.), Betmpon Strict Nture Reserve, by G.M. Ros. Dignosis. A species from north-estern lowlnd sites with conspicuous nd contrsted color pttern in mny individuls, which hs been previously listed s Stumpffi sp. 12 in Vieites et l. (2009), s Stumpffi sp. ff. tetrdctyl [C HM364800] by Ros et l. (2012), nd s S. sp. CBetmpon2 in Scherz et l. (2016) nd Peloso et l. (2017). (1) Modertely lrge-sized species (dult mle SVL mm); (2) mnus with four fingers (first finger slightly reduced in length) nd pes with five toes (first toe slightly to distinctly reduced in length); (3) terminl phlnges of fingers with slightly, those of toes with modertely enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum smooth to very slightly grnulr; (6) dorslly vrible in color, most chrcteristiclly with reticultions of blck spots with pricot borders on bckground of copper, but cn lso be plum nd brown, rosy brown, or silver with drk mrkings. Mrkings often forming contrsted nd symmetricl series of lrge ptches on the centrl dorsum. The posterodorsl shnk cn be rust red to copper. Ventrlly light, with or without drk throt nd light spots or vermicultions; (7) regulrly repeted short single-note tonl cll repeted fter short intervls. Distinguished from S. nlmin, S. ngeluci, S. contumeli, S. dvidttenboroughi, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. mledict, S. mdgscriensis, S. mmitik, S. miery, S. obscoen, S. psologloss, S. pygme, S. sort, S. tridctyl, nd S. ynniki by lrger body size; from S. be, S. kibomen, S. meikee, S. nigrorubr, nd S. roseifemorlis by the lck of lrge nd distinct res of red color ventrlly or on limbs (only in some individuls, limited mount of red on shnk); from S. grndis by the lck of contrsted light blueblck ventrl pttern; from S. nlnjirofo nd S. fusc by less tuberculr dorsum, nd from these species nd S. chillei by different dorsl color pttern; from S. be, S. hr, S. megsoni, nd S. stffordi by smller body size nd stronger length reduction of first toe; from S. diutissim nd S. jennoeli by shorter inter-cll intervls nd lower dominnt frequency; from S. edmondsi by different dorsl color pttern in the mjority of specimens. Phylogeneticlly the sister species of S. diutissim, species tht differs quite strongly in biocoustics chrcters (longer inter-cll intervl, slightly longer cll durtion, higher dominnt frequency) nd is strongly divergent in mtdna, despite shring Rg-1 hplotypes. Description of the holotype. Specimen in good stte of preservtion, with its left rm removed s tissue smple. Body elongte; hed slightly longer thn wide, nrrower thn body; snout pointed in dorsl nd lterl view; nostrils directed lterlly, slightly protubernt, nerer to tip of snout thn to eye; cnthus rostrlis indistinct, concve; lorel region stright, lmost verticl; tympnum indistinct, bout 59% of eye dimeter; suprtympnic fold not visible; tongue long, brodening posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone rounded. Forelimb thick; subrticulr tubercles indistinguishble; outer metcrpl tubercle indistinct, ovl; prepollex present; fingers without webbing; first finger reduced; reltive length of fingers 1 < 2 < 4 < 3, fourth finger slightly longer thn second; finger tips not expnded into discs. Hind limbs slender; tibiotrsl rticultion reching center of eye; TIBL 49% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle indistinct, oblong; outer mettrsl tubercle bsent; no webbing between toes; first toe reduced; toe tips slightly expnded; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe distinctly shorter 369

100 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b e c d h i f g Fig. 75. Stumpffi prdus sp. nov. in life: () holotype ZSM 435/2010 (FGZC 4237); (b e) prtype ZSM 439/2010 (FGZC 4268); (c d) prtype ZSM 440/2010 (FGZC 4278), ll from Ambodivongy; (f g) prtype ZSM 230/2016 (ACZCV 0147); (h i) prtype MRSN A6204, both from Betmpon Strict Nture Reserve. thn third; subrticulr tubercles indistinguishble. Skin on dorsum smooth, without distinct dorsolterl folds. Ventrl skin smooth. The skin of the flnk nd especilly the nterior nd posterior thigh is perforted with holes, rnging in size from 0.1 to 0.7 mm dimeter. The cuse of these holes is unknown, but we suppose tht they my be cused by prsites. Colortion of the holotype. After five yers in 70% ethnol, the dorsum is drk brown. A few crem spots re present, but whether or not these re cused by prsites is uncler (see holotype description). No mrkings re present in the inguinl region. The dorsl surfce of the hed is s the bck but more gry. The nostril is surrounded by brown. The lterl surfce of the hed is brown s the dorsum, the lip being more crem. A light crem mrking extends from the posterior of the eye to the corner of the mouth in front of the tympnum, with crem spot directly below the tympnum. The flnks fde from the brown of the dorsum to the crem of the ventrl trunk, but re spotted with crem mrkings tht my be cused by prsites. The ventrl trunk is drk crem with light crem spots. The chin is s the ventrl trunk. The ventrl thigh is s the ventrl trunk. The shnk nd trsus re ventrlly beige. The sole of the foot is drk brown. Dorslly, the thigh is mottled drk nd light brown. The posterodorsl surfce of the thigh is light brown, with crem circlulr spots where skin is missing. The nterior surfce of the thigh is drk brown with mny of these spots where skin is missing. The shnk is light brown with drk brown oblique crossbnd nd other flecks of drk brown. The trsus is mottled light nd drk brown. The foot is externlly mottled light nd drk brown, fding to crem internlly. The toes re irregulrly striped light nd drk brown. The clocl region is drk brown. The rm is mottled light nd drk brown, without crossbnds. The dorsomedil surfce of the hnd is crem. The fingers hve smll drk nd light stripes. The underside of the rm is s the ventrl trunk. Color in life (totl vrition). Colortion extremely vrible. Most chrcteristic colortion is reticulted pttern of blck spots with pricot borders on bckground of reddish-copper (Fig.75b), but dorsl colortion cn lso be firly plin plum nd brown (Fig. 75) rosy brown with X-shped ebony mrkings (Fig. 75c), or reddish with 370

101 VERTEBRATE ZOOLOGY 67 (3) 2017 b Fig. 76. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi prdus sp. nov. recorded t Ambodivongy from the holotype (ZSM 435/2010): () 1 s durtion section; (b) 6 s durtion section. blck to burnt umber irregulr mrkings. Ventrl colortion is similrly vrible, nd cn be trnslucent plum (Fig. 75d), burnt umber with plum nd white spots (Fig. 75e), or brown with crem to white spots or vermicultions (Figs. 75g, i). Most specimens seem to hve rusty red to copper posterior shnk. All specimens hve white nnulus before ech terminl phlnge of their fingers nd toes. The iris rnges from drk brown to copper to gold. Vrition. For vrition in mesurements mong specimens, see Tble 3. In generl, ll exmined specimens gree strongly with the holotype in morphology. In lmost ll prtypes, the cnthus rostrlis is more distinct thn in the holotype. All specimens except the holotype hve slightly expnded finger tips, nd most lso hve distinctly longer fourth finger thn second. In specimens without prepollex, the first finger is still reduced. A high degree of vribility ws noted in the colortion of specimens in preservtive. Four distinct color morphs were noticeble in our eight type specimens. The first is tht of the holotype (N = 1). The second (N = 4; ZSM 440/2010, 439/2010, 770/2009, nd 438/2010) is figured in Fig. 75b-e. In preservtive, these specimens hve light bse color rnging from light brown to fuchsi with drk mrkings. These mrkings generlly consist of cross-like diffuse blotch on the bck surrounded by smller drk mrkings, but these rnge from strong to wek. In some specimens, drk dot is present on the rostrum. Drk flecks continue onto the legs nd rms, forming shnk crossbnd in some specimens. All of these specimens hve pink to fuchsi mrking on the posterodorsl shnk. The third (N = 1; ZSM 433/2010) is like the second, but the dorsl ptterning is more irregulr nd reminds of leoprd s spots. This lso hs pink mrking on the posterodorsl shnk. The fourth (N = 2; ZSM 432/2010 nd 434/2010) consists of gry bsecolor with the whole dorsum flecked with irregulr drk gry mrkings. The posterodorsl shnk lcks the distinct lighter re. Ventrl colortion of these four groups is inconsistent; ventrl colortion rnges from crem like the holotype to drk brown in ZSM 438/2010. The bottom lip of ll specimens is brown with irregulr crem spots. Etymology. The species nme is noun in pposition to the generic nme, derived from Greek πάρδος (párdos) = mle leoprd, in reference to the leoprd-like pttern of some specimens of this species. Distribution. This species hs wide distribution long the north-estern Mdgscn cost including four confirmed loclities: (1) Ambodivongy (type loclity), (2) Ambolokoptrik, (3) Melivinny-Mnompn, nd (4) Betmpon Strict Nture Reserve. In Betmpon this species hs thus fr been found only round the Rendrirendry nd Betkonn cmpsites, which re locted in the southern prt of the Reserve. The specimens from Ambodivongy nd Ambolokoptrik re slightly differentited geneticlly from those occurring t Betmpon nd Melivinny. The ltitude rnges from m.s.l. Nturl history. At both, Betmpon nd Ambodi vongy, specimens were found in lef litter t night. Mles clled during light rin from the ground not hiding under litter (Fig. 75). Cll. The dvertisement cll consists of single, very short note emitted in series nd fst succession (Fig. 76). Clls recorded by J. Köhler on 2 April 2010 t Ambodivongy from the holotype ZSM 435/2010 (FGZC 4237) hd the following numericl prmeters: cll durtion (= note durtion) ms (38.3 ± 2.3 ms; N = 12), durtion of inter-cll intervls ms (479.3 ± 7.6 ms; N = 12), nd dominnt frequency t Hz ( ± 73.3 Hz, N = 12). 371

102 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b c d e Fig. 77. Stumpffi tetrdctyl from Nosy Borh in life: () ZSM 593/2006 (ZCMV 3395); (d) ZSM 594/2006 (ZCMV 3396); the remining photos (b, c, e) cnnot be relibly ssigned to voucher specimens. 6. Smll-sized species of clde C While only single nominl species of smll-sized Stumpffi fell into clde C (S. tetrdctly), the moleculr dt suggest lrge diversity of new species in this clde, most of which re described in the following. Stumpffi tetrdctyl Vences & Glw, 1991 (Figures 77 nd S118) Nme-bering type. Holotype ZFMK from Strßenrnd, c. 1 km südlich von Mromndi n der Westküste der Insel Nosy Borh (= Sinte Mrie), Ost- Mdgskr ; trnslted: t the edge of the rod, c. 1 km south of Mromndi on the west cost of the islnd Nosy Borh (= Île Sinte Mrie), estern Mdgscr. Identity nd dignosis. Stumpffi tetrdctyl is smll-sized species phylogeneticlly embedded in clde C, nd so fr kown only from the offshore islnd Nosy Borh. (1) Smll-sized species (SVL mm); (2) mnus with four fingers (first finger modertely to strongly reduced in length) nd pes with four toes (first toe reduced to smll rudiment); (3) terminl phlnges of fingers nd toes without enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum smooth; (6) light brown to gryish dorsl color, with vrible drk dorsl mrkings, often with drk centrl teddyber shped ptch (e.g. Fig. 77), nd one individul known with light vertebrl line (Fig. 77d). Ventrlly gry with drk gry mottling nd sometimes drk throt, but without contrsted ventrl colortion, red color elements on ventrl side, or shrp continuous color border between dorsum nd flnks; (7) regulrly repeted short single-note chirp cll. Distinguished from S. chillei, S. nlnjirofo, S. be, S. diutissim, S. edmondsi, S. fusc, S. grndis, S. hr, S. jennoeli, S. kibomen, S. megsoni, S. meikee, S. nigro rubr, S. prdus, S. roseifemorlis, nd S. stffor di by smller body size; from the similr-sized species S. nlmin, S. ngeluci, S. gimmeli, S. huwei, S. ih r n, S. lrinki, S. mledict, S. mdgscriensis, S. mmi tik, S. psologloss, S. pygme, S. sort, nd S. ynni ki by more expressed digitl reduction (lmost complete reduction of first toe); from S. contumeli, S. obscoen, nd S. tridctyl by lower degree of digitl reduction; from S. miery nd S. dvidttenboroughi by less expressed digitl reduction on hnd. Furthermore distinguished from S. be, S. kibomen, S. meikee, S. nigrorubr, nd S. roseifemorlis by the lck of lrge nd distinct res of red color ventrlly or on limbs; from S. grndis by the lck of contrsted light blue-blck ventrl pttern; nd from mny species by dvertisement cll. Stumpffi tetrdctyl is currently the only smllsized nominl species of Stumpffi in clde C (numerous dditionl species will be described below). It belongs to highly supported subclde of clde C1 in which the only other smll-sized species re S. grrffoi sp. nov. nd S. spndei sp. nov. (described below). Specimens exmined. Holotype ZFMK collected on 4 Mrch 1991 from Nosy Borh by F. Glw nd M. Vences; prtype ZFMK with sme collecting dt s holoptype; prtype ZSM 558/1999 (previously ZFMK 52546) collected on 6 7 Mrch 1991 from Ambohiden forest, Nosy Borh, ner the 372

103 VERTEBRATE ZOOLOGY 67 (3) 2017 b Fig. 78. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi tetrdctyl from Nosy Borh (not ssocited with specimen): () 1 s durtion section; (b) 6 s durtion section. villge Ambohiden, by F. Glw nd M. Vences. ZSM 593/2006 (ZCMV 3395) nd ZSM 594/2006 (ZCMV 3396) collected on 7 8 Mrch 2006 in Nosy Borh, Mromndi villge (S , E , 20 m.s.l.) by M. Vences nd J. Rndrinirin. Distribution. The species is known from severl loclities on Nosy Borh. Nturl history. Specimens were observed both in primry rinforest nd in degrded, secondry vegettion nd plnttions, clling from the lef litter in the lte fternoon nd t night. At night, mles might cll from rther exposed positions (Vences & Glw, 1991). Cll. The dvertisement cll consists of single short note emitted in series t regulr intervls (Fig. 78). Clls recorded by M. Vences nd F. Glw on 4 9 Mrch 1991 t the type loclity Nosy Borh (Vences et l., 2006: CD3, trck 37, cut 1) hd the following numericl prmeters: cll durtion (= note durtion) ms (92 ± 9 ms; N = 11), inter-cll intervls ms (959± 244 ms; N = 10), nd dominnt frequency t Hz (5355 ± 138 Hz, N = 11). Stumpffi betmpon sp. nov. (Figures 79 nd S119) Holotype. ZSM 214/2016 (ACZCV 0210) (Figs. 79 b), dult femle, collected on 14 November 2013 t Vohitsivln (S , E , 481 m bove se level), Betmpon Strict Nture Reserve, Tomsin Province, Mdgscr by A. Crottini, D. Slvi, E. Scnrini, nd George. Prtypes. MRSN A6315 nd MRSN A6300, collected on 2 Mrch 2007 t Vohitsivln (S , E , 517 m.s.l.) by G.M. Ros; MRSN A6317, MRSN A6327, nd MRSN A6246, ll collected between 8 nd 11 Februry 2007 t Shmbendrn (S E , 458 m.s.l.) by G.M. Ros nd F. Andreone; MRSN A6313 (FAZC 13618), MRSN A6253, MRSN A6233, nd MRSN A6273, ll collected between 19 nd 22 Februry 2007 t Mintimbto (S17.894, E , 255 m.s.l.) by G.M. Ros; MRSN A6387 collected on 14 November 2007 t Shindrn (S , E , 341 m.s.l.) by G.M. Ros nd J. Nöel; MRSN A6255 collected on 8 Mrch 2007 t Shbefoz (S , E , 349 m.s.l.) by G.M. Ros; UADBA (ACZCV 0203) collected on 15 November 2013 t Vohitsivln (S , E , 514 m.s.l.) by A. Crottini, D. Slvi, E. Scnrini, nd George; ZSM 215/2016 (ACZCV 0039), UADBA (ACZCV 0040), nd ZSM 216/2016 (ACZCV 0041), ll collected on 7 November 2013 t Shindrn (S , E , 269 m.s.l.) by A. Crottini, D. Slvi, E. Scnrini, nd J.H. Velo; ZSM 217/2016 (ACZCV 0046) nd ZSM 218/2016 (ACZCV 0047) collected on 5 November 2013 t Shindrn (S , E , 360 m.s.l.) by A. Crottini, D. Slvi, E. Scnrini, nd J.H. Velo; ZSM 219/2016 (ACZCV 0103) collected on 5 November 2013 t Shmbendrn (S , E , 476 m.s.l.) by G.M. Ros, D.J. Hrris, M. Rndrimiliso, nd J.H. Lv. All these loclities refer to sites within Betmpon Strict Nture Reserve, Tomsin Province, Est Mdgscr. Dignosis. A smll sized species from the northern centrl est of Mdgscr, which hs been previously listed s Stumpffi sp. 17 in Vieites et l. (2009), Scherz et l. (2016) nd Peloso et l. (2017), nd s Stumpffi sp. [C FJ559312] in Ros et l. (2012). (1) Miniturized to smll-sized species (SVL mm); (2) mnus with four fingers (first finger modertely to strongly reduced in length, especilly in mles) nd pes with five toes (first toe very strongly reduced in length, rudimentry); (3) terminl phlnges of fingers nd toes without enlrged discs; (4) reltive hnd nd foot length, HAL/ SVL , FOTL/SVL ; (5) dorsum smooth with moderte number of scttered tubercles; (6) dorslly gry to sndy brown. Cn hve n intricte but indistinct pttern of smll drk brown ptches, but cn lso hve series of brod longitudinl stripes of color. Ventrlly with lrge crem flecks or reticultions, especilly on the chest. No red color ventrlly. 373

104 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Colortion of the holotype. After three yers in 70% ethnol, the dorsum is pinkish gry. Two brown spots re present over the scpulr region. Rounded drk brown spots re present in the inguinl region. The dorsl surfce of the hed is s the bck. Drk brown flecks pre b c d e f g h i j Fig. 79. Stumpffi betmpon sp. nov. from Betmpon Strict Nture Reserve in life: ( b) holotype ZSM 214/2016 (ACZCV 0210); (c d) ZSM 217/2016 (ACZCV 0046); (e f) ZSM 218/2016 (ACZCV 0047); (g) ZSM 219/2016 (ACZCV 0103); (h) FAZC (voucher not collected); (i) MRSN A6313; (j) MRSN A6315. Distinguished from S. chillei, S. nlnjirofo, S. be, S. diutissim, S. edmondsi, S. fusc, S. grndis, S. hr, S. jennoeli, S. kibomen, S. megsoni, S. meikee, S. nigrorubr, S. prdus, S. roseifemorlis, nd S. stffordi by smller body size; from the similr-sized species S. nlmin, S. ngeluci, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. mledict, S. mdgscriensis, S. mmitik, S. psologloss, S. pygme, S. sort, nd S. ynniki by more expressed digitl reduction (lmost complete reduction of first toe); from S. contumeli, S. obscoen, nd S. tridctyl by lower degree of digitl reduction; from S. miery nd S. dvidttenboroughi by less expressed digitl reduction on hnd. Furthermore, distinguished from S. be, S. kibomen, S. meikee, S. nigrorubr, nd S. roseifemorlis by the lck of lrge nd distinct res of red color ventrlly or on limbs; from S. grndis by the lck of contrsted light blue-blck ventrl pttern. Along with S. tetrdctyl nd severl other new species described below the species belongs to clde C; here it is plced sister to ll other clde C lineges, lbeit without support, suggesting it is not closely relted to ny other known species. It differs from S. tetrdctyl nd mny of the other smll-sized new species described below by the comprtively tuberculr dorsum. Description of the holotype. Specimen in good stte of preservtion, fourth toe removed s tissue smple. Body rounded; hed wider thn long, nrrower thn body; snout rounded in dorsl nd lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis slightly distinct, concve; lorel region distinct, concve, oblique; tympnum distinct, bout 80% of eye dimeter; suprtympnic fold not visible; tongue long, brodening posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone rounded. Forelimbs slender; subrticulr tubercles single, distinct; outer metcrpl tubercle not recognizble; inner metcrpl tubercle distinct, ovl; fingers without webbing; reltive length of fingers 1 < 4 < 2 < 3, fourth finger slightly shorter thn second; finger tips not expnded into discs. Hindlimbs slender; TIBL 48% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle not visible; outer mettrsl tubercle bsent; no webbing between toes; toe tips not expnded; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe shorter thn third; subrticulr tubercles distinct, single. Skin on dorsum reltively smooth, without distinct dorsolterl folds. Ventrl skin smooth. 374

105 VERTEBRATE ZOOLOGY 67 (3) 2017 b Fig. 80. Stumpffi dolchi sp. nov. from Nosy Mngbe in life: ( b) dorsolterl nd ventrl view of holotype ZSM 488/2005 (ZCMV 2143). sent bove the eyes. The nostril is surrounded by beige. The lterl surfce of the hed is drk brown striped with beige. The flnks re ventrlly beige mottled with brown nd lterlly beige. The flnk colortion merges with the ventrl colortion. The chin is brown with beige flecks. The ventrl trunk is beige mottled with brown. The ventrl thigh is brown with beige flecks. The shnk nd trsus re ventrlly s the thigh. The sole of the foot is s the thigh but drker. Dorslly, the thigh is pinkish spotted nd mottled with brown nd with brown crossbnd. The posterodorsl surfce of the thigh is s the dorsum. The shnk, trsus, nd foot re s the thigh. The toes re drk brown mottled with beige. The clocl region is drk brown. The rms re pinkish spotted with brown, mottled with drk brown on the lower rm. The dorsomedil surfce of the hnd is beige flecked with brown. The fingers re drk brown spotted with beige. The underside of the rm is s the ventrl trunk. Color in life (holotype nd vrition). The dorsum is dominted by brod medil stripe of light-brown running from the snout to the vent, nd irregulrly defined by beige lterl border. Along this brod bnd there re severl pinkish-beige tubercles. A thin beige line strts from the snout nd fdes behind the scpulr region. A stripe of dppled ebony nd gry runs from the lterl fce to the posterior trunk. The tympnum is gry. The dorsl forelimb is rosy tn, with two coppery-ebony crossbnds on the forerm. The hnd is mottled rosy tn nd ebony, with light nnulus before the terminl phlnge of the third finger. The dorsl leg is rosy tn speckled with ebony, with two light brown crossbnds on the thigh nd shnk. The foot nd toes re light slmon flecked with ebony. The chin is brown with crem reticultions. The ventrl skin of the trunk is trnslucent in bse color, nd thus plum over the pectorl girdle becoming incresingly yellow towrd the posterior bdomen, reticulted with splotches of crem. The ventrl legs re brown with crem spots. The iris is coppery with blck reticultions. Vrition. For vrition in mesurements mong specimens, see Tble 3. In generl, ll exmined specimens gree strongly with the holotype in externl morphology. Colortion in this species is quite vrible. Most individuls re sndy brown in dorsl colortion, sometimes mottled with brown flecks, nd severl do not hve dorsl stripes, or hve more intricte dorsl ptterning. The ventrl colortion generlly hs drk brown bckground colortion with dense crem mrkings, but is less reticulted thn the holotype. ACZCV 0103 is dorslly drk beige mottled with drk brown, the shnk in dorsl view is beige with one brown crossbnd, nd the lower rm hs brown crossbnd. ACZCV 0047 is dorslly beige mrbled with brown, hs brown tringulr shpe from the middle of the eye until the scpulr re, nd brown crossbnd on the thigh nd shnk. ACZCV 0046 is overll pinkish-beige nd lcks brod medin bnd. Etymology. The species nme is noun in pposition to the genus nme, nd refers to the type loclity of the new species, the forest of Betmpon Strict Nture Reserve in estern Mdgscr. Distribution. This species is so fr known only from Betmpon Strict Nture Reserve, where it is found cross the entire Reserve with, thus fr, the exception of Betkonn, lthough the species is expected to be present there s well. Within the Betmpon, it rnges from m.s.l. Nturl history. The holotype ws found t night in the lef litter. This species ppers to be quite bundnt nd distributed cross the Betmpon Strict Nture Reserve, therefore in res completely overlpping with the lrger species S. jennoeli described bove. Cll. Unknown. 375

106 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Stumpffi dolchi sp. nov. (Figure 80) Holotype. ZSM 488/2005 (ZCMV 2143) (Fig. 80) collected on 22 Februry 2005 in Nosy Mngbe Specil Reserve, ner Mrontsetr (S15.5, E49.8, c m bove se level), Tomsin Province, Mdgscr, by F. Glw, M. Vences, nd R.D. Rndriniin. Dignosis. A smll sized species from the north est of Mdgscr, which hs been previously listed s deep conspecific linege of S. tetrdctyl in Vieites et l. (2009), nd s S. sp. C32 in Scherz et l. (2016) nd Peloso et l. (2017). (1) Miniturized species (SVL 11.6 mm); (2) mnus with four fingers (first finger strongly reduced in length) nd pes with four toes (first toe completely reduced); (3) terminl phlnges of fingers nd toes without enlrged discs; (4) reltive hnd nd foot length, HAL/SVL 0.18, FOTL/SVL 0.75; (5) dorsum smooth; (6) dorslly chocolte brown with indistinct nd poorly contrsted drk speckling, or with some lrger drk symmetricl mrkings. Flnks drk brown, with distinct color border between dorsl nd lterl color. Ventrlly ebony with distinct pttern of bluish white flecks, especilly on chest. No red color ventrlly. Distinguished from S. chillei, S. nlnjirofo, S. be, S. diutissim, S. edmondsi, S. fusc, S. grndis, S. hr, S. jennoeli, S. kibomen, S. megsoni, S. meikee, S. nigrorubr, S. prdus, S. roseifemorlis, nd S. stffordi by smller body size; from the similr-sized species S. nlmin, S. ngeluci, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. mledict, S. mdgscriensis, S. mmitik, S. psologloss, S. pygme, S. sort, nd S. ynniki by more expressed digitl reduction (complete reduction of first toe); from S. contumeli, S. obscoen, nd S. tridctyl by lower degree of digitl reduction; from S. miery nd S. dvidttenboroughi by less expressed digitl reduction on the hnd, especilly of the fourth finger. Furthermore, distinguished from S. be, S. kibomen, S. meikee, S. nigrorubr, nd S. roseifemorlis by the lck of lrge nd distinct res of red color ventrlly or on limbs; from S. grndis by the lck of contrsted light blue-blck ventrl pttern; from S. betmpon by smooth dorsum; very similr morphologiclly to S. tetrdctyl but differs by more strongly expressed color border between flnks nd dorsum, nd strongly in mitochondril nd nucler DNA. Along with S. tetrdctyl nd severl other new species described herein the species belongs into clde C; phylogeneticlly it is plced s sister species to S. mkir sp. nov. (described below) which is morphologiclly similr but differs by more stout body nd more expressed digitl reduction on the hnd, especilly of the fourth finger, nd by concordnt differentition in mitochondril nd nucler DNA. Description of the holotype. Specimen in good stte of preservtion. The right thigh muscle removed s tissue smple. Body elongte; hed longer thn width, nrrower thn body; snout pointed in dorsl view, pointed in lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis not distinct; lorel region slightly concve, oblique; tympnum distinct; bout 40% of eye dimeter; suprtympnic fold not visible; tongue destroyed, ttched nteriorly; mxillry teeth nd vomerine teeth bsent; chone rounded. Forelimbs slender; subrticulr tubercles not recognizble; outer metcrpl tubercle not distinct; inner metcrpl tubercle distinct, rounded; fingers without webbing; reltive length of fingers 1 < 2 < 4 < 3, fourth finger slightly longer thn second; finger tips not expnded into discs. Hind limbs slender; TIBL 48% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle smll, ovl; outer mettrsl tubercle bsent; no webbing between toes; toe tips not expnded; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe shorter thn third; subrticulr tubercles distinct, single. Skin on dorsum reltively smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After 10 yers in 70% ethnol, the dorsum is beige vermiculted with brown. The dorsl surfce of the hed is s the bck. The nostril is beige. The lterl surfce of the hed is brown. The flnks re lterlly brown spotted with beige; ventrlly beige spotted with brown. The flnk colortion merges with the ventrl colortion. The ventrl trunk is beige mottled with brown. The chin is s the tunk. The thigh, shnk, trsus, nd foot re ventrlly s the bdomen. Dorslly, the thigh is beige spotted with brown. The posterodorsl surfce of the thigh is s the dorsl spect. The shnk is s the thigh. The trsus is s the shnk. The foot is s the trsus. The toes re s the foot. The clocl region is brown. The rms re beige spotted with brown. The dorsomedil surfce of the hnd is s the rm. The underside of the rm is s the ventrl trunk. In life, dorsum chocolte brown with drker brown speckles. A strong lterl color border is present between the chocolte dorsum nd the ruddy brown flnk. Iridescent bluish flecks re present long the ventrl edge of the flnk. The dorsl forelimb is s the bck, with some bluish flecks on the hnd, including n nnulus before the terminl phlnge of ech finger. The dorsl hindlimb is s the bck, without crossbnds, but with smll drk brown speckles. The toes re drk brown with smll light blue flecks, including n nnulus before the terminl phlnge of ech finger. The venter is ebony with firly lrge bluish white flecks throughout, t their highest density t the mid-trunk, decresing in density on the thigh nd chin. The ventrl thigh is lighter thn the trunk. The iris is copper. Etymology. The species nme is ptronym honoring Riner Dolch, in recognition of his extrordinry chievements in the study nd conservtion of Mdgscr s biot. Distribution. The species is known only from its type loclity on Nosy Mngbe t c m.s.l. Nturl history. The holotype ws collected in the lef litter of low-ltitude rinforest. 376

107 VERTEBRATE ZOOLOGY 67 (3) 2017 Cll. Not relibly known. Remrk. Two specimens from Nosy Mngbe ssigned to S. tetrdctyl in Vences & Glw (1991) probbly belong to S. dolchi but re here not included in the type series becuse of the lck of genetic dt. These specimens (ZFMK ) emitted clls differing from S. tetrdctyl from the type loclity by much longer internote intervls nd higher frequency of bout 6000 Hz. Stumpffi grrffoi sp. nov. (Figures 81 nd S120) Holotype. ZSM 1752/2008 (ZCMV 8803) (Figs. 81 b), dult specimen, collected on 14 Februry 2008 in Mhso Forest ner Ambodisko villge (S , E , 1032 m bove se level), NE Vohimen/Lke Alotr, Tomsin Province, Mdgscr, by D.R. Vieites, J.L. Ptton, P. Bor, nd M. Vences. Prtypes. ZSM 631/2009 (ZCMV 8684), ZSM 632/2009 (ZCMV 8685), ZSM 633/2009 (ZCMV 8687), ll collected August 2009 in Tmpolo Forest by H. Rsolonjtovo nd R.D. Rndriniin; MRSN A6278 (FAZC 13489) collected on 4 Februry 2007 t Rendryrendry, Betmpon Strict Nture Reserve (S , E , 325 m.s.l.) by G.M. Ros; ZSM 220/2016 (ACZCV 0006) nd UADBA (ACZCV 0007) collected on 4 November 2013 long Piste Fotsimvo t Rendrirendry, Betmpon Strict Nture Reserve (S , E , 287 m.s.l.) by A. Crottini, D. Slvi, E. Scnrini, George, G.M. Ros, D.J. Hrris, M. Rndrimiliso, nd H. Lv; UADBA (ACZCV 0071) collected on 11 November 2013 long Piste Fotsimvo t Rendrirendry, Betmpon Strict Nture Reserve (S , E , 239 m.s.l.) by A. Crottini, D. Slvi, E. Scnrini, Clude, G.M. Ros, D.J. Hrris, M. Rndrimiliso, nd H. Lv; DRV 5698, DRV 5628, DRV 5622, DRV 5621 (to be ctlogued in MNCN) with sme collection dt s holotype. Fig. 81. Stumpffi grrffoi sp. nov. in life: ( b) dorsolterl nd ventrl views of holotype, ZSM 1752/2008 (ZCMV 8803) from M hso Forest; (c) MRSN A6278 from Betmpon Strict Nture Re serve. Dignosis. A smll Stumpffi from vrious loclities long the northern Mlgsy est cost which hs been previously listed s Stumpffi sp. 18 in Vieites et l. (2009), Scherz et l. (2016) nd Peloso et l. (2017) nd Stumpffi sp. [C FJ559313] in Ros et l. (2012). (1) Smll-sized species (dult mle SVL mm); (2) mnus with four fingers (first finger distinctly reduced in length) nd pes with five toes (first toe very strongly reduced in length, externlly recognizble s rudiment only); (3) terminl phlnges of fingers without, those of toes with slightly enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum smooth; (6) dorslly with brod medil stripe of burnt umber in teddyber-like shpe, lterlly bordered by copper. A strong dorsolterl color border cn be present. Ventrlly rosy brown to plum with drk mottling on the throt, nd lrge crem flecks on the chest; (7) regulrly repeted single-note tonl cll. Distinguished from S. chillei, S. nlnjirofo, S. be, S. diutissim, S. edmondsi, S. fusc, S. grndis, S. hr, S. jennoeli, S. kibomen, S. megsoni, S. meikee, S. nigrorubr, S. prdus, S. roseifemorlis, nd S. stffordi by smller body size; from the similr-sized species S. nlmin, S. ngeluci, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. mledict, S. mdgscriensis, S. mmitik, S. psologloss, S. pygme, S. sort, nd S. ynniki by more expressed digitl reduction (lmost complete length reduction of first toe); from S. contumeli, S. obscoen, nd S. tridctyl by lower degree of digitl reduction; from S. miery nd S. dvidttenboroughi by less expressed digitl reduction on hnd, especilly on fourth finger. Furthermore distinguished from S. be, S. kibomen, S. meikee, S. nigrorubr, nd S. roseifemorlis by the lck of lrge nd distinct res of red color ventrlly or on limbs; from S. grndis by the lck of contrsted light blue-blck ventrl pttern; from S. betmpon by smooth dorsum; from S. dolchi by lck of b c 377

108 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b Fig. 82. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi grrffoi sp. nov. from Mhso Forest, recorded from holotype ZSM 1752/2008 (ZCMV 8803): () 1 s durtion section; (b) 6 s durtion section. strongly expressed color border between flnks nd dorsum; from S. tetrdctyl by longer durtion of inter-cll intervls. Phylogeneticlly, plced within subclde of clde C1 tht otherwise contins only lrge-sized species, except for S. tetrdctyl nd S. spndei sp. nov. (described below) which both differ biocousticlly nd in mitochondril nd nucler DNA. Description of the holotype. Specimen in firly good stte of preservtion, right thigh muscle removed s tissue smple. Body round; hed slightly longer thn wide, nrrower thn the body; snout pointed in dorsl nd lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis distinct, concve; lorel region stright, lmost verticl; tympnum indistinct, bout 41% of eye dimeter; suprtympnic fold not visible; tongue long, brodening slightly posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone rounded. Forelimbs slender; subrticulr tubercles distinct, single; outer metcrpl tubercle not recognizble; inner metcrpl tubercle distinct, ovl; fingers without webbing; first finger strongly reduced, second nd fourth fingers reduced; reltive length of fingers 1 < 2 < 4 < 3, fourth finger slightly longer thn second; tip of finger three expnded into disc. Hind limbs slender; TIBL 55% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle oblong; outer mettrsl tubercle bsent; no webbing between toes; first toe strongly reduced, recognizble only s rudiment, second toe reduced; toe tips expnded; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe distinctly shorter thn third; subrticulr tubercles indistinct, single. Skin on dorsum smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After lmost eight yers in 70% ethnol, the dorsum is light gry. Almost symmetricl drk brown nd crem lines extend from the inguinl region to the bck of the hed on either side of the midline. A drk brown spot is present in the inguinl region. The dorsl surfce of the hed is gry with drk gry mrkings beside the eyes. The nostril is surrounded by brown. The lterl surfce of the hed is mottled gry nd brown. A brown suprtympnic mrking is present. This is seprted from long drk mrking on the flnks. Otherwise the flnk is the color of the dorsum, fding ventrlly to the color of the ventrl trunk. The bdomen is crem with brown flecks, becoming incresingly brown nteriorly. The chin is brown with crem flecks. The ventrl thigh is s s the chin. The shnk nd trsus re ventrlly s the thigh. The sole of the foot is drk brown. Dorslly, the thigh is s the dorsum. The posterodorsl surfce of the thigh is drk brown with crem spots. The shnk is s the thigh, with brown perpendiculr crossbnd. The trsus is s the shnk, with similr crossbnd. The foot is externlly gry with brown spot on the exterior fce, nd brown interiorly. The toes re brown with crem flecks. The clocl region is drk brown. The rms re s the dorsum. A drk crossbnd is present on the lower rm. The dorsomedil surfce of the hnd is brown. The underside of the rm is s the ventrl trunk. In life, dorsl body with n irregulr brod medil stripe of burnt umber, with wvy border to copper colortion of the dorsolterl body nd hed (mking n lmost teddyber-like shpe). A lterl color border is present, the copper of the dorsolterl bck hs stright border, to the umber flnk, which fdes ventrlly to crem, merging with the venter. A high density of beige flecks is present on the flnk, decresing in density dorslly. The dorsl forelimb is copper, with burnt umber crossbnd on the forerm. The hnd is burnt umber with high concentrtion of beige flecks. The dorsl leg is umber with some copper shding nd hs high concen- 378

109 VERTEBRATE ZOOLOGY 67 (3) 2017 trtion of beige flecks. The chin is rosy brown nteriorly, becoming incresingly plum posteriorly, with numerous smll whitish flecks long the lips nd over the throt. The ventrl trunk is covered in lrge crem flecks between which the trnslucent skin is visible, which become smller on the ventrl legs. The iris is copper. Vrition. For vrition in mesurements mong specimens, see Tble 3. In generl, ll exmined specimens gree strongly with the holotype in morphology. A single exception is ZSM 631/2009, which hs expnded terminl discs on its toes. A reltively low degree of vribility ws noted in the colortion of specimens in preservtive; ZSM 632/2009 nd ZSM 631/2009 re similr to the holotype in pttern, but drk brown with drker brown mrkings, insted of light gry with drk brown mrkings. The mid-dorsl mrking is cross-shped in these specimens. Ventrlly, the bdomen is drker brown thn the holotype, nd the ventrl legs re brown s well. ZSM 631/2009 is similr to these specimens, but lighter brown. It lcks the long drk mrking on the flnk present in the other specimens, nd hs more vermiculted dorsl mrkings. Specimens from Betmpon Strict Nture Reserve: MRSN A6278 present mid-dorsl mrking which is more cross-shped thn teddyber-shpe, nd presents less evident copper colortion. In this specimen hindlimbs hve two crossbnds. ZSM 220/2016 (ACZCV 0006) hs poorly defined mid-dorsl mrking resulting in homogeneous dorsolterl colortion, which is copper in the bckground with irregulr umber mrkings. Etymology. The species nme is ptronym honoring H. Mrtin Grrffo, in recognition of his substntil contributions to the study of nurn lkloids, including numerous compounds of the Mlgsy genus Mntell. We emphsize tht Mrtin usully collborted with Tom Spnde, nd it is thus no coincidence tht Stumpffi grrffoi nd S. spndei sp. nov. (described below) occur syntopiclly in the Mhso Forest. Distribution. Known from three loclities in estern Mdgscr including: (1) Mhso Forest (type loclity), (2) Tmpolo Forest, nd (3) Betmpon Strict Nture Reserve. In Betmpon, lthough intensively smpled, the species is thus fr known to occur only round the villge of Rendrirendry (long Piste Fotsimvo). It occurs from m.s.l. Nturl history. At the type loclity, specimens were herd clling in the evening nd t night, from the lef litter in degrded rinforest, in mixed choruses with S. spndei sp. nov. (described below). The holotype ws found clling from the lef litter of Mhso Forest. In Betmpon this species seems to be uncommon (or with quite secretive hbits) nd restricted to low elevtions ( m.s.l.) in Rendryrendry nd the surroundings, living on the edge of secondry forest with some level of disturbnce. Cll. The dvertisement cll consists of single, short nd high-pitched note emitted in series t regulr intervls (Fig. 82). Clls recorded on 14 Februry 2008 by M. Vences t Mhso forest hd the following numericl prmeters: cll durtion (= note durtion) ms (103.4 ± 2.5 ms; N = 7), inter-cll intervls ms ( ± 150 ms; N = 6), nd dominnt frequency of Hz ( ± 84.2 Hz, N = 7). The wide rnge in inter-cll intervl durtion is presumbly cused by some clls being emitted fter typiclly long intervls due to disturbnce. Stumpffi mkir sp. nov. (Figures 83 nd S119) Holotype. ZSM 541/2009 (ZCMV 11257), dult mle (Fig. 83), collected on June 2009 on the Mkir Plteu, within the Mkir Nturl Prk, close to the source of the Fotsilnn River (S E , 1067 m bove se level), Mhjng Province, Mdgscr, by M. Vences, D.R. Vieites, F.M. Rtsovin, R.D. Rndriniin, E. Rjeririson, T. Rjofirison, nd J.L. Ptton. Dignosis. A smll sized species from the north est of Mdgscr, which hs previously been listed s S. sp. C38 in Scherz et l. (2016) nd Peloso et l. (2017). (1) Smll-sized species (SVL 12.1 mm); (2) mnus with four fingers (first finger strongly reduced in length, second nd fourth finger modertely reduced in length) nd pes with four toes (first toe externlly completely reduced); (3) terminl phlnges of fingers nd toes without enlrged discs; (4) reltive hnd nd foot length, HAL/SVL 0.16, FOTL/SVL 0.64; (5) dorsum smooth; (6) dorslly ebony bordered with gry-brown. Flnks ebony, with distinct color border between dorsl nd lterl color. Ventrlly with trnslucent plum skin, rnging from ebony on the throt to the lmost gry of the posterior bdomen, interspersed with incresingly dense crem flecks posteriorly. No red color ventrlly. Distinguished from S. chillei, S. nlnjirofo, S. be, S. diutissim, S. edmondsi, S. fusc, S. grndis, S. hr, S. jennoeli, S. kibomen, S. megsoni, S. meikee, S. nigrorubr, S. prdus, S. roseifemorlis, nd S. stffordi by smller body size; from the similr-sized species S. nlmin, S. ngeluci, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. mledict, S. mdgscriensis, S. mmitik, S. psologloss, S. pygme, S. sort, nd S. ynniki by more expressed digitl reduction (complete reduction of first toe); from S. contumeli, S. obscoen, nd S. tridctyl by lower degree of digitl reduction; from S. miery nd S. dvidttenboroughi by slightly lesser degree of digitl reduction on hnd, especilly of the fourth finger. Furthermore distinguished from S. be, S. kibomen, S. meikee, S. nigrorubr, nd S. roseifemorlis by the lck of lrge nd distinct res of red color ventrlly or on the limbs; from S. grndis by the lck of contrsted light blue-blck ventrl pttern; from S. betmpon by smooth dorsum; from S. dolchi nd S. tetrdctyl by stronger reduction of the fourth finger, nd by generlly more stout body shpe. Phylogeneticlly it 379

110 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b Fig. 83. Stumpffi mkir sp. nov. in life: ( b) dorsl nd ventrl views of holotype ZSM 541/2009 (ZCMV 11257) from the Mkir Plteu. is plced s sister species to S. dolchi, which is concordntly differentited in mitochondril nd nucler DNA. Description of the holotype. Specimen in good stte of preservtion, left thigh muscle removed s tissue smple. Body rounded; hed slightly wider thn long, nrrower thn body width; snout rounded in dorsl nd lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis concve; lorel region concve, oblique; tympnum slightly distinct, bout 57% of eye dimeter; suprtympnic fold not visible; tongue long, brodening posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone rounded. Forelimbs slender; subrticulr tubercles single, slightly distinct; outer metcrpl tubercle not recognizble; inner metcrpl tubercle distinct, ovl, fused with tubercule prepollicl; hnd without webbing; first finger strongly reduced, sound nd fourth fingers modertely reduced; reltive length of fingers 1 < 2 < 4 < 3, fourth finger longer thn second; finger tips not expnded into discs. Hind limbs slender; TIBL 45% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle smll, ovl; outer mettrsl tubercle bsent; no webbing between toes; toe tips not expnded; first toe totlly reduced, reltive length of toes 2 < 5 < 3 < 4; fifth toe shorter thn third; subrticulr tubercles distinct, single. Skin on dorsum reltively smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After seven yers in 70% ethnol, the dorsum is beige mottled with brown. A brown bnd crosses the dorsum from the middle of the eye to the inguinl re. A beige spot is present in the inguinl region. The dorsl surfce of the hed is s the bck. The nostril is beige. The lterl surfce of the hed is beige spotted with white dots. The flnks re s the lterl surfce of the hed. The flnk colortion merges with the ventrl colortion. The ventrl trunk is uniformly beige spotted with brown nd flecked with light beige. The chin is s the ventrl trunk. The ventrl thigh is s the bdomen. The shnk is ventrlly brown flecked with beige. The trsus nd foot re ventrlly drk brown flecked with beige. Dorslly, the thigh is drk brown flecked with drk beige. The posterodorsl surfce of the thigh is uniformly drk brown flecked with light beige. The shnk, trsus, nd foot re s the thigh. The toes re light brown speckled with light beige. The clocl region is drk brown. The rms re drker thn the dorsum, speckled with light beige. The dorsomedil surfce of the hnd is light beige, mottled with beige nd speckled with white. The fingers re uniformly beige. The underside of the rm is s the ventrl trunk. In life, dorsl bse color ebony, bordered in the dorsolterl regions nd on the hed by gry-brown. Numerous smll bluish flecks re present on the flnks. The dorsl forelimb is s the bck. The hnd hs severl bluish flecks, including n nnulus before the terminl phlnge of ech finger. The dorsl hindlimbs is s the bck, without crossbnds, with slightly more ornge brown on the posterodorsl shnk. The toes hve severl bluish flecks, including n nnulus before the terminl phlnge of ech toe. The ventrl skin is trnslucent, drk plum, especilly drk (resembling ebony) in the chin region, lightening posteriorly; interspersed with dense crem flecks in the posterior trunk, decresing in density nteriorly nd posteriorly on the legs. The iris is drk copper. Etymology. The species nme is noun in pposition to the genus nme, nd refers to the type loclity of the new species, the Mkir Nture Reserve in north-estern Mdgscr. Distribution. The species is known only from its type loclity on the Mkir plteu. Nturl history. The single known specimen ws collected during the dy, ctive in the lef litter of primry rinforest. Cll. Unknown. 380

111 VERTEBRATE ZOOLOGY 67 (3) 2017 Description of the holotype. Specimen in good stte of preservtion, left thigh muscle removed s tissue smple for DNA extrction. Body elongte; hed s wide s long; snout rounded in dorsl view, boxy in lterl view; nostrils directed lterlly, not protubernt, nerer to tip of snout thn to eye; cnthus rostrlis distinct, concve; lorel region concve, verticl; tympnum distinct, bout 52% of eye dimeter; suprtympnic fold not visible; tongue long, brodening posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone rounded. Forelimbs slender; subrticulr tubercles indistinguishble; outer metcrpl tubercle smll, indistinct, ovl; prepollex strong; fingers without webbing; first finger strongly reduced, second finger reduced; reltive length of fingers 1 < 2 = 4 < 3, fourth finger subequl in length to second; finger tips not expnded into discs. Hind limbs slender; TIBL 54% of SVL; lt b c d e f g h Fig. 84. Stumpffi miovov sp. nov. from Sort Forest in life: ( b) holotype ZSM 1649/2012 (FGZC 3656); (c d) prtype ZSM 1640/2012 (FGZC 3650); (e f) prtype ZSM 1647/2012 (FGZC 3675); (g h) prtype ZSM 1646/2012 (FGZC 3751). Stumpffi miovov sp. nov. (Figures 84 nd S121) Holotype. ZSM 1649/2012 (FGZC 3656) (Figs. 84 b), dult mle, collected on 28 November 2012 in Sort forest (c. S , E , 1398 m bove se level), Antsirnn Province, Mdgscr, by F. Glw, O. Hwlitschek, F.M. Rtsovin, A. Rkotorison, T. Rjofirison, nd A. Rzfimnntso. Prtypes. ZSM 1640/2012 (FGZC 3650), ZSM 1646/2012 (FGZC 3751), ZSM 1647/2012 (FGZC 3675), UADBA-A (originlly ZSM 1648/2012, FGZC 3676), UADBA-A (FGZC 3655) nd UADBA (FGZC 3663), ll collected between November 2012 on the Sort mssif (pproximtely between S , E , 1398 m nd S13.675, E , 1580 m.s.l.) by F. Glw, O. Hwlitschek, F.M. Rtsovin, A. Rkotorison, T. Rjofirison, nd A. Rzfimnntso. Dignosis. A modertely sized species with ornge-reddish ventrl side from the north est of Mdgscr, which hs not been listed s cndidte species in previous publictions. It is listed mong the smll-sized species of clde C, but it pproches the size of the lrger species in the clde. (1) Smll- to modertely-sized species (SVL mm, dult mle SVL 15.1 mm); (2) mnus with four fingers (first finger strongly to modertely reduced in length) nd pes with five toes (first toe distinctly reduced in length); (3) terminl phlnges of fingers nd toes without enlrged discs; (4) reltive hnd nd foot length, HAL/SVL , FOTL/SVL ; (5) dorsum smooth with number of distinct scttered tubercles; (6) dorslly copper, with copper-infused ebony mrkings. These mrkings nd dorsl color cn be vrible, nd cn include stripes or isolted color elements. Ventrl color with intense fiery ornge on the trunk, ventrl legs, nd sometimes the ventrl forelimbs. The ventrl chin cn hve light medin stripe. This species is esily distinguished from ll species except S. kibomen, S. be, S. roseifemorlis, S. edmondsi, S. nigrorubr, nd S. meikee by ornge to reddish ventrl colortion. Among these, from S. be by fiery ornge colortion on the ventrl hindlimbs (vs. only on the conceled portions of the hindlimbs); from S. kibo men, S. roseifemorlis, S. meikee, S. edmondsi, nd S. ni grorubr by more intense fiery ornge ventrl color, sometimes extending even onto the forelimbs (pler or more red in ll other tx). In ddition, it cn be distinguished from S. chillei, S. nlnjirofo, S. be, S. diutissim, S. edmon dsi, S. fusc, S. grndis, S. hr, S. jennoeli, S. kibo me n, S. megsoni, S. meikee, S. nigrorubr, S. prdus, S. roseifemorlis, nd S. stffordi by slightly sml ler body size; from numerous smll-sized species in cldes A, B, C nd D by lrger body size, i.e., from S. n lmin, S. betmpon, S. dolchi, S. grrffoi, S. m dg sc riensis, S. mkir, S. psologloss, S. pygme, S. te tr dctyl, nd S. ynniki. Furthermore, from S. contu meli, S. dvidttenboroughi, S. miery, S. obscoen, nd S. tridctyl by lower degree of digitl reduction of the toes; nd from S. grndis by the lck of contrsted light blue-blck ventrl pttern. Phylogeneticlly, S. miovov occupies rther isolted position nd possibly is the sister group of subclde C2 which exclusively contins Stumpffi species of reltively lrge body size. 381

112 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs erl mettrsli strongly connected; inner mettrsl tubercle smll, ovl; outer mettrsl tubercle bsent; no webbing between toes; first toe strongly reduced, second toe reduced; toe tips slightly expnded; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe distinctly shorter thn third; subrticulr tubercles distinct, single. Skin on dorsum smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After three yers in 70% ethnol, the dorsum is pinkish gry. Two drk brown spots re present over the scpulr region. Elongted drk brown mrkings re present in the inguinl region. The dorsl surfce of the hed hs lrge drk brown hert-shped mrking extending from the eyes to the bck of the hed. The tip of the rostrum hs drk brown spot s well. The nostril is surrounded by brown. The lterl surfce of the hed is s the dorsum. A drk brown mrking is present beneth the eye, nd suprtympnic mrking is present s well. The flnks re gry, with fint drk mrking bove the insertion of the rm. The flnk fdes to the crem of the ventrl trunk. The ventrl trunk is crem on the bdomen, nteriorly becoming flecked with incresing mounts of brown. The chin is speckled brown nd crem. The ventrl thigh, shnk, nd trsus re lso crem. The sole of the foot is light nd drk brown. Dorslly, the thigh is light brown. A fint oblique crossbnd is present. The posterodorsl surfce of the thigh is brown flecked with crem. The shnk is s the thigh with n oblique crossbnd nerer to the knee thn the heel. The trsus is s the shnk, without crossbnds. The foot is externlly s the shnk, internlly crem. The toes re flecked with crem nd brown. The clocl region is not different from the posterior thighs. The rms re s the dorsum. A drk crossbnd is present on the lower rm. The dorsomedil surfce of the hnd is crem. The fingers re brown with few crem flecks. The underside of the rm is s the thigh. Color in life (holotype nd vrition). Dorsl colortion copper, with copper-infused ebony tringulr mrking between the eyes converging to point between the suprscpule bordered by chmpgne. A pir of symmetricl suprscpulr ebony spots bordered with chmpgne behind the suprscpule, nd nother pir of lrger, oblong spots in the inguinl region. The flnk is spotted with whitish flecks. The nterior hlf of the tympnum is chmpgne. An ebony mrking is present below the eye nd round the nostril. The dorsl forelimb is s the bck, with n ebony crossbnd on the forerm. The medil hnd is beige, the lterl hnd burnt umber flecked with copper, with whitish nnulus before ech terminl phlnge. The dorsl hindlimb is redder copper, with one ebony-copper crossbnd on the mid-thigh nd midshnk. The medil foot is fiery ornge, the toes mottled fiery ornge, burnt umber, nd copper, with whitish bnd before ech terminl phlnge. The chin is grybrown spotted with pricot. From the pectorl girdle posteriorly the bse-color is fiery ornge, including the ventrl rms nd legs. Over the mid-trunk, whitish speckling nd flecking is present. The iris is copper dorslly, rust red nteriorly nd posteriorly. The dorsl nd ventrl colortion is highly vrible (Fig. 84). Dorsl ptterns include chmpgne to tn mid-dorsl stripe tht cn be brod or thin, n hourglss shped drk mrking, lterl drk spots, nd strong to wek hindlimb crossbnds. The chin spots cn be yellow, nd mid-chin chmpgne stripe bordered in burnt umber cn be present. The ornge of the rms nd legs cn be more yellow or more red. The iris cn be golden to copper, nd cn lck red colortion. Vrition. For vrition in mesurements mong specimens, see Tble 3. In generl, ll exmined specimens gree strongly with the holotype in morphology. The lorel region vries from verticl to slightly oblique. The first finger ppers less strongly reduced in specimens tht lck prepollex. A plmr tubercle is sometimes visible. A high degree of vribility ws noted in the colortion of specimens; ventrl colortion in preservtive ws more or less consistent, with some specimens hving medil brown chin stripe ending t the pectorl girdle (ZSM 1648/2012 nd 1640/2012), but most specimens hving more or less pure crem venter, nd immculte crem ventrl legs. Dorsl colortion cn hve vertebrl crem stripe (ZSM 1648/2012 nd 1640/2012), nd cn vry from pink (ZSM 1647/2012 nd 1648/2012) to gry (ZSM 1646/2012). A hert-shped mrking s in the holotype ws noted only in ZSM 1646/2012, but this ws posteriorly connected to tringulr drk gry mrking. This specimen lso hs the sme dorsl spots s the holotype. Thigh crossbnds vry from bsent (ZSM 1648/2012) to drk nd strong (ZSM 1646/2012). The tip of the rostrum is drk in ll specimens. Etymology. The species epithet miovov is Mlgsy word mening vrible, nd refers to the vribility of colortion in this species. It is used s noun in pposition to the genus nme. Distribution. This species is known only from the Sort Forest between c m bove se level. Nturl history. The species ws bundnt t mid-elvtions of the Sort mssif, nd observed jumping on the ground long tril within bmboo forest. Cll. Unknown. Stumpffi spndei sp. nov. (Figures 85 nd S122) Holotype. ZSM 1751/2008 (ZCMV 8802) (Figs. 85 nd S122), clling mle, collected on 14 Februry 2008 in Mhso Forest ner Ambodisko villge (S , E , 1032 m bove se level), NE Vohimen/Lke Alotr, Tomsin Province, Mdgscr, by D.R. Vieites, J.L. Ptton, P. Bor, nd M. Vences. 382

113 VERTEBRATE ZOOLOGY 67 (3) 2017 b Fig. 85. Stumpffi spndei sp. nov. from Mhso Forest in life: ( b) dorsolterl nd ventrl views of the holotype ZSM 1751/2008 (ZCMV 8802). b Fig. 86. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi spndei sp. nov. from Mhso Forest (recorded from the holotype ZSM 1751/2008): () 1 s durtion section; (b) 6 s durtion section. Dignosis. A smll Stumpffi from the northern centrl est, which hs been previously listed s Stumpffi sp. 19 in Vieites et l. (2009), Scherz et l. (2016) nd Peloso et l. (2017). (1) Smll-sized species (SVL 12.7 mm); (2) mnus with four fingers (first finger distinctly reduced in length) nd pes with five toes (first toe strongly reduced in length); (3) terminl phlnges of fingers without, those of toes with very slightly enlrged discs; (4) reltive hnd nd foot length, HAL/SVL 0.21, FOTL/ SVL 0.67; (5) dorsum smooth; (6) dorslly iridescent tupe with ebony to blck irregulr flecks, nd two pirs of semi-symmetricl beige mrkings bordered in blck. Ventrlly light gry on the chin, rosy posteriorly, flecked with tupe; (7) regulrly repeted single-note tonl cll. Distinguished from S. chillei, S. nlnjirofo, S. be, S. diutissim, S. edmondsi, S. fusc, S. grndis, S. hr, S. jennoeli, S. kibomen, S. megsoni, S. meikee, S. miovov, S. nigrorubr, S. prdus, S. roseifemorlis, nd S. stffordi by smller body size; from the similr-sized species S. nlmin, S. ngeluci, S. gimmeli, S. huwei, S. ihrn, S. lrinki, S. mledict, S. mdgscriensis, S. mmitik, S. psologloss, S. pygme, S. sort, nd S. ynniki by more expressed digitl reduction (lmost complete length reduction of first toe); from S. contumeli, S. obscoen, nd S. tridctyl by lower degree of digitl reduction; from S. miery by distinctly shorter inter-cll intervl length. Furthermore distinguished from S. be, S. kibomen, S. meikee, S. nigrorubr, S. roseifemorlis, nd S. miovov by the lck of lrge nd distinct res of red or ornge color ventrlly or on limbs; from S. grndis by the lck of contrsted light blueblck ventrl pttern; from S. betmpon by smooth dorsum; from S. dvidttenboroughi, S. dolchi, nd S. mkir by the lck of strongly expressed color border between flnks nd dorsum; from S. tetrdctyl by slightly shorter cll durtion (without overlp of vlues), nd concordnt divergence in mitochondril nd nucler genes. Phylogeneticlly, plced within subclde of 383

114 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs clde C1 tht contins only lrge-sized species, except for S. tetrdctyl nd S. grrffoi which both differ biocousticlly nd in mitochondril nd nucler DNA. Description of the holotype. Specimen in good stte of preservtion, left hindlimb removed s tissue smple for DNA extrction. Body round; hed slightly wider thn long, nrrower thn body width; snout slightly pointed in dorsl nd lterl view; nostrils directed lterlly, not protubernt, nerer to the tip of the snout thn to the eye; cnthus rostrlis distinct, slightly concve; lorel region slightly concve, oblique; tympnum slightly distinct, bout 52% of eye dimeter; suprtympnic fold not visible; tongue long, brodening posteriorly, ttched nteriorly, not notched; mxillry teeth nd vomerine teeth bsent; chone ovl. Forelimbs slender; subrticulr tubercles single, slightly distinct; outer metcrpl tubercle slightly visible, rounded; prepollex visible, elongte. Fingers without webbing; first finger reduced; reltive length of fingers 1 < 2 = 4 < 3, fourth finger subequl to second; finger tips not expnded into discs. Hind limbs slender; TIBL 49% of SVL; lterl mettrsli strongly connected; inner mettrsl tubercle indistinguishble from reduced first finger; outer mettrsl tubercle bsent; no webbing between toes; first toe strongly reduced, second toe reduced; toe tips not expnded; reltive length of toes 1 < 2 < 5 < 3 < 4; fifth toe shorter thn third; subrticulr tubercles single, distinct. Skin on dorsum reltively smooth, without distinct dorsolterl folds. Ventrl skin smooth. Colortion of the holotype. After seven yers in 70% ethnol, the dorsum is beige, with brod brown bnd strting in the middle of the eyes nd running to the inguinl region. Brown spots re present in the inguinl region. The dorsl surfce of the hed is s the bck with brown ptch. The nostril is brown. The lterl surfce of the hed is beige flecked with brown. The flnks re beige flecked with brown. The flnk colortion merges with the ventrl colortion. The ventrl trunk is beige mottled with brown. The chin is beige mottled with brown. The ventrl thigh is s the bdomen. The shnk is ventrlly brown slightly flecked with crem. The trsus is ventrlly beige with brown flecks. The sole of the foot s the trsus. Dorslly, the thigh is brown with drk brown crossbnd. The posterodorsl surfce of the thigh is brown flecked with crem. The shnk is similr to the thigh. The trsus is s the shnk with slightly brown crossbnd. The foot is beige flecked with brown. The toes re striped. The clocl region is brown with crem flecks. The forelimb is beige, with crossbnd on the forerm. The dorsomedil surfce of the hnd is beige spotted with brown. The fingers hve smll brown spots. The underside of the rm is s the ventrl trunk. In life, dorslly iridescent tupe in bse colortion, interspersed with ebony to blck irregulr flecks. Two pirs of beige mrkings bordered in blck re present, one in the suprscpulr region (symmetricl), nd one t the mid-bck (symmetricl). The hed is beige. The flnk hs row of lrge ebony spots, fusing posteriorly to colored bnd. A further ebony mrking is present below the eye nd round the nostril. The dorsl forelimb is s the bck; n ebony crossbnd is present on the forerm. The fingers re mottled, with light nnulus before ech terminl phlnge. The dorsl hindlimb is s the bck, with one ebony crossbnd with coppery inclusions on the midthigh, one on the midshnk, nd two on the foot. The posterodorsl shnk is beige. The toes re mottled tupe nd ebony, with smll whitish nnulus before ech terminl phlnge. The ventrl body is light gry on the chin, rosy posteriorly, flecked ll over with tupe. The iris is copper. Etymology. The species nme is ptronym honoring Thoms F. Spnde, in recognition of his substntil contributions to the study of nurn lkloids, including numerous compounds of the Mlgsy genus Mntell. Distribution. The species is known only from its type loclity in Mhso Forest, c m bove se level. Nturl history. At the type loclity, specimens were herd clling in the evening nd t night, from the lef litter in degrded rinforest in mixed choruses with S. grrffoi. Cll. The dvertisement cll consists of single, very short nd high-pitched note emitted in series t regulr intervls (Fig. 86). Clls recorded by M. Vences on 14 Februry 2008 from the holotype t Mhso forest hd the following numericl prmeters: cll durtion (= note durtion) ms (46 ± 1.8 ms; N = 11), inter-cll intervls ms (801.8 ± 48.6 ms; N = 10), nd dominnt frequency t Hz (5755 ± 69.8 Hz, N = 11). 7. Cndidte species not described herein Stumpffi sp. C7 (Figure 87) Remrk. This species is listed s Stumpffi sp. 7 in Vieites et l. (2009), Köhler et l. (2010), Klges et l. (2013), Scherz et l. (2016) nd Peloso et l. (2017), nd s Stumpffi sp. 6 in Wollenberg et l. (2008). It is chrcterized by its very long cll durtion; however, the moleculr phylogeny plces it sister to S. sort (Fig. 3), species of which dvertisement clls re unknown. Clrifiction of the sttus of S. sp. C7 will be possible in the future by more detiled comprisons of dvertisement clls nd nucler genes, once biocoustic dt for S. sort become vilble. Mteril exmined. ZSM 379/2005 (FGZC 2826), FGZC 2827, nd FGZC 2870, ll collected on 16 Februry 2005 from Cmp Simpon, Mrojejy Ntionl Prk (S , E , 1326 m.s.l.) by F. Glw, M. Vences, nd R.D. Rndriniin; FGZC 384

115 VERTEBRATE ZOOLOGY 67 (3) 2017 b c d Fig. 87. Stumpffi sp. C7 from Mrojejy Ntionl Prk in life: ( b) dult mle, ZSM 544/2016 (ZCMV 15181); (c) dult mle, ZSM 555/2016 (ZCMV 15182); (d) unctlogued specimen, ZCMV b Fig. 88. Spectrogrms (bove) nd oscillogrms (below) of dvertisement clls of Stumpffi sp. C7 from Mrojejy (cll recorded from specimen ZCMV 15181): () 1 s durtion section; (b) 6 s durtion section collected on 14 Februry 2005 from Cmp Mntell, Mrojejy Ntionl Prk (S E, E , 481 m.s.l.) by F. Glw, M. Vences, nd R.D. Rndriniin; ZSM /2016 (ZCMV ), ZCMV , 15056, 15061, ZSM 541/2016 (ZCMV 15069), ZSM 542/2016 (ZCMV 15070), ZCMV 15074, ZCMV 15079, nd ZCMV 15093, ll collected on 15 November 2016 from cmpsite clled Cmp 0 in Mrojejy Ntionl Prk (S , E , 310 m.s.l.), by A. Rkotorison, M.D. Scherz, M.C. Bletz, J.H. Rzfindribe, A.Rzfimnntso, nd M. Vences; ZCMV 15155, ZCMV 15166, 385

116 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs b Fig. 89. Preserved specimen of Stumpffi sp. C7, ZSM 379/2005 (FGZC 2826), from Mrojejy Ntionl Prk; scle br = 5 mm. ZSM 543/2016 (ZCMV 15168), ZCMV 15169, ZCMV 15179, ZSM /2016 (ZCMV ), ZSM 555/2016 (ZCMV 15187), ZSM 550/2016 (ZCMV 15212), ZCMV 15213, nd ZSM 554/2016 (ZCMV 15281), ll collected on November 2016, from Cmp Simpon, Mrojejy Ntionl Prk (S , E , 1326 m.s.l.) by A. Rkotorison, M.D. Scherz, M.C. Bletz, J.H. Rzfindribe, A.Rzfimnntso, nd M. Vences. Cll. The dvertisement cll of S. sp. C07 consists of single, modertely long, tonl note emitted in series t regulr intervls (Fig. 88). Clls were recorded by M.D. Scherz on 17 November 2017 from specimen ZCMV in Mrojejy Ntionl Prk, with the following prmeters: cll durtion (= note durtion) ms (293.8 ± 2.9 ms; N = 10), inter-cll intervls ms ( ± ms; N = 10), nd dominnt frequency t Hz ( ± 20.7 Hz, N = 10). Fig. 90. Stumpffi sp. C29 (DRV 6103) from Antevilmbzh, Tsrtnn Strict Nture Reserve, in life. Stumpffi sp. C11 Remrk. This species is listed s Stumpffi sp. 11 in Vieites et l. (2009). Mteril exmined. MRSN A2583, from Ambolokoptrik. Stumpffi sp. C29 (Figs. 90, S125) Remrk. This species hs been listed s Stumpffi sp. 29 in Klges et l. (2013). Mteril exmined. ZSM 1824/2010 (ZCMV 12387) collected on 13 June 2010 in Mtsbory Miky, Tsrtnn Strict Nture Reserve (S E , 2021 m.s.l.) by M. Vences, D.R. Vieites, R.D. Rndriniin, F.M. Rtsovin, S. Rsmison, A. Rkotorison, E. Rjeririson, F. Rndrinsolo, nd T. Rjofirison; ZSM 627/2014 (DRV 6103) collected on 10 June 2010 in Antevilmbzh, Tsrtnn Strict Nture Reserve (S , E , 1589 m.s.l.) by M. Vences, D.R. Vieites, R.D. Rndriniin, F.M. Rtsovin, S. Rsmison, A. Rkotorison, E. Rjeririson, F. Rndrinsolo, nd T. Rjofirison; MRSN A2653 collected in 2001 in the Tsrtnn Mssif by F. Andreone, J.E. Rndrinirin, nd M. Vences. Stumpffi sp. C30 Remrk. This species hs been listed s Stumpffi sp. 30 in Klges et l. (2013). Mteril exmined. UADBA-A (DRV 6459), UADBA-A (DRV 6458), UADBA-A (DRV 6457), UADBA-A (DRV 6456), UADBA-A (DRV 6455), ZSM 635/2014 (DRV 6454), ZSM 633/2014 (DRV 6452), ZSM 634/2014 (DRV 6453), ZSM 632/2014 (DRV 6451), nd UADBA-A (DRV 6487), ll collected on 30 June 2010 in Angorony forest frgment ner Mromndi (S , E , 115 m.s.l.) by F.M. Rtsovin, S. Rsmison, T. Rjofirison, nd F. Rndrinsolo. 386

117 VERTEBRATE ZOOLOGY 67 (3) 2017 Stumpffi sp. C42 Mteril exmined. ZMA (FG/MV ), collected in Februry 2002 in Fierenn by M. Vences nd D.R. Vieites. Discussion Fst trck txonomy The inventory of Mdgscr s mphibin fun hs seen steep progress over the pst three decdes (e.g., Köhler et l., 2005; Glw & Vences, 2007; Vieites et l., 2009; Glw et l. 2010; Perl et l., 2014; Scherz et l., 2016). Lrge numbers of new species hve been described, hundreds of dditionl, still undescribed cndidte species hve been discovered, phylogenetic reltionships mong tx hve been reveled by extensive moleculr dt sets, nd numerous field observtions hve considerbly improved our understnding of the nturl history of mny of these frogs. While Blommers-Schlösser & Blnc (1991) listed 133 described species of nurns in Mdgscr, this number hd risen to 244 species in Vieites et l. (2009), 292 in Perl et l. (2014), nd currently hs climbed up to bout 315 (AmphibiWeb, 2017; Frost, 2017). Even more cndidte species hve lredy been identified but require txonomic revisionry work before being confirmed, formlly recognized, nd nmed. While the discovery of new cndidte species becomes ever more efficient by routine ppliction of DNA brcoding pipelines in concert with more intense fieldwork in remote res, the forml species descriptions remin s mjor bottleneck in species inventory. Recent efforts to overcome this Linnen shortfll (Brown & Lomolino, 1998; Whittker et l., 2005; Hortl et l., 2015) include pproches of turbo-txonomy (Butcher et l., 2012) or fst-trck txonomy (Riedel et l., 2013) which chiefly combine DNA brcode sequences with concise morphologicl descriptions nd high-resolution digitl imging to stremline the forml description of lrger numbers of new species. This effort hs proven to be of extrordinry efficiency, yielding descriptions of hundreds of new species (Butcher et l., 2012; Riedel et l., 2013b, 2014, 2016). Other uthors (e.g., Dijkstr et l., 2015) hve lso succeeded in describing lrge numbers of species by reducing the extent of morphologicl descriptions, nd Renner (2016) rgues for n emphsis on DNA chrcters in the forml nming of species, complying with the essentil focus of Linneus on dignosis, not description. Although such simplified description pipeline is especilly useful for hyperdiverse orgnisms poorly covered by current txonomy, such s numerous groups of rthropods or nemtodes, we relized the potentil of n dpted nd simplified description procedure lso for vertebrtes such s Stumpffi, in order to chieve comprehensive revision of such species-rich nd poorly known gener within resonble time frme. Our txonomic pipeline, compred to the stndrds followed in numerous species descriptions of Mlgsy mphibins in the pst, is simplified in tht it relies on the following min components: (1) species delimittion is bsed on concordnt differentition in one mtdna nd one nucdna mrker, or one mtdna mrker nd biocoustics; (2) morphologicl dignoses re bbrevited nd simplified, only selected number of morphometric mesurements is tken, nd not ech species is dignosed ginst ech other species; dignoses focus especilly on differentition from those species tht were closely relted to the new species in the moleculr phylogeny, or tht occur syntopiclly (see Vijykumr et l., 2014 for distinction of linege dignosis nd field dignosis); (3) description of morphologicl vrition is somewht bbrevited, nd insted, s mny specimens s possible re documented by photogrphs in life nd in preservtive; nd (4) most importntly, we delibertely ccepted number of inconsistencies nd omissions in our pipeline, reflecting the lck of relible dt nd preferences of reserchers, s discussed in the following. The foremost qulity criterion for species-level txonomy is the robustness of the underlying species hypotheses. In generl, the genetic divergences of Stumpffi proved extrordinry, with levels of mtdna divergence in most cses much bove the usully defined thresholds of 3% or 5% uncorrected pirwise distnce in the 16S gene (e.g., Vences et l., 2005; Fouquet et l., 2007; Vieites et l., 2009). Allele shring in Rg-1 is furthermore exceedingly rre despite using only short frgment of this gene, with correspondingly low number of informtive sites. We re therefore convinced tht the plethor of new species described herein re robustly supported s independent evolutionry lineges, even in cses where biocoustic or morphologicl differences were fint or could not be ssessed due to missing dt. Nevertheless, in series of cses we opted not to describe species, even though these fulfilled t lest the criterion of very strong divergence in mtdna. This pplies especilly for the S. gimmeli complex, in which we include two deep lineges. In this species, we refrined from further splitting becuse (1) we do not hve clerly identified dult specimens nd clls from ll three lineges, (2) we do not hve genetic dt from the type loclity of the species, Ambnj, nd therefore cnnot scertin with certinty which of the lineges corresponds to S. gimmeli sensu stricto, nd (3) the S. gimmeli complex is one of the rre cses of nucler llele shring, thus necessitting more extensive nlyses of nucdna mrkers. Two relted lineges of S. kibomen re known from only single juvenile specimen (S. sp. C34 from Rnomfn) or hve no fresh tissue smples vilble to us (S. sp. C11 from Ambolokoptrik). We decided not to describe S. sp. C30 becuse we hve no cll recordings or photogrphs in life vilble, nd becuse we observed one highly divergent linege (specimen DRV 6487) suggesting tht two distinct species might be involved. Since the vilble smples nd specimens of S. sp. C30 were collected in the course of rpid field 387

118 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs survey trgeted t reptiles, we cnnot fully scertin ll metdt ccompnying the smples. For S. obscoen, we found similr sitution with few specimens from the type loclity Andsibe showing deeply divergent 16S sequences, but too little ssocited informtion (dvertisement clls, nucdna sequences) is vilble to txonomiclly interpret these differences. The sister linege of the highlnd species S. roseifemorlis is known from lowlnd loclity (S. sp. C57 from Ambodivongy) nd is therefore unlikely to represent the sme species; yet, with only two smples vilble for moleculr study, nd no voucher specimen vilble for morphologicl comprison, we refrined from txonomic conclusion in this cse. In generl, by not describing the lineges discussed bove we follow the ple of Mirlles et l. (2013) for conservtiveness in erecting new species, given tht under integrtive txonomic nd species concepts (Dyrt, 2005; De Queiroz, 2007; Pdil et l., 2010) it is epistemologiclly difficult to decisively reject two-species hypothesis nd thus synonymize described species-level tx. In only few cses did we decide to describe lineges tht re not fully supported by ll evidence s full species. This pplies to S. ngeluci nd S. mledict, two species tht re not seprted by the 4% threshold in the 16S gene. In this cse we consider their symptric occurrence nd lck of llele shring in Rg-1 s sufficient for description, especilly becuse we collected nd observed the respective specimens for this study. Furthermore, the recognition of both S. nlnjirofo nd S. chillei s species requires confirmtion, s they re sister species both occurring t lowlnd sites of the northest. We decided to describe them s distinct species becuse the moleculr species criteri did pply ( > 4% 16S divergence; no llele shring in Rg-1), nd becuse we hd sufficient mount of well-preserved mteril to scertin some fint morphologicl differences. Yet, we cknowledge tht the sttus of S. nlnjirofo requires confirmtion s it might in the future be rther considered DCL of S. chillei, once more dt on the genetic contct zone of the two tx become vilble. To conclude, our somewht relxed ppliction of the species criteri defined priori lrgely depends on our interprettion of dt qulity. In cses where DNA sequences were vilble from sufficient specimens nd could be ssigned to the vouchers with full relibility, nd other ssocited metdt were considered relible s well, we decided for recognition nd description of new species even if some species criteri were not or only brely fulfilled. On the other hnd, wherever we hd doubts s to the ssocition of DNA sequences, voucher specimens, color photos, nd collection metdt, we refrined from txonomic conclusions. Given the mssive mount of dt nlysed for this study, even in thoroughly compiled dt sets some errors will persist, nd it is importnt to crefully evlute the possibility of smple confusion or other errors before chnging the txonomic clssifiction of the orgnisms involved (Scherz et l., 2016; 2017). Deep genetic divergences within nd between species Besides the presence of deep Stumpffi lineges of still unclrified sttus, the surprisingly strong genetic divergence in these frogs wrrnts comment. Species in Stumpffi re in generl differentited by distinctly higher mitochondril differences thn other, co-distributed Mlgsy frogs, nd the lmost complete lck of llele shring in Rg-1 is exceptionl s well. For instnce, in species of the fmilies Mntellide nd Hyperoliide, nucler llele shring mong closely relted species hs been regulrly observed (e.g., Guibemntis: Vences et l., 2013; Boophis: Vences et l., 2012; Heterixlus: Gehring et l., 2012) despite the use of longer stretches of nucler genes in the respective nlyses. Most species of Stumpffi re only known from single loclities, or from very smll geogrphic rnges. In cses where smples were vilble from more thn one loclity, these lmost lwys hd distinct mitochondril hplotypes, e.g., S. nlmin from the type loclity vs. Ankrfntsik, S. gimmeli, S. mmitik, nd S. psologloss from vrious sites, or S. nlnjirofo from Betmpon, Nosy Mngbe, nd Ambtorom. An exception is found in S. ngeluci where specimens from Montgne d Ambre nd the nerby Montgne des Frnçis hve identicl 16S sequences. In generl, the strong genetic structuring in these frogs might be relted to their smll body size. Bsed on multi-species nlysis of mntellids, Pbijn et l. (2012) found n inverse ssocition between body size nd nucleotide divergence between popultions, nd hypothesized tht the lck of genetic cohesion mong popultions is excerbted in smll-sized species due to ecologicl nd physiologicl constrints, thus leding to regionl genetic frgmenttion. In principle, this process hs the potentil to ccelerte rtes of lloptric specition in smll frogs reltive to lrge species, nd it might hve led to the lrge number of microendemic species of Stumpffi uncovered in the current study. However, species diversifiction in Mdgscr is driven by multitude of fctors (Brown et l., 2014; Rodríguez et l., 2015). Stumpffi hve their center of species richness in northern Mdgscr where mny complementry lloptric nd symptric processes might be influencing specition (Vences et l., 2009; Brown et l., 2016), nd more detiled biogeogrphic nd phylogeogrphic nlyses re therefore necessry to understnd the processes driving their evolution. Reproductive modes Dt on the reproductive modes of Stumpffi re only vilble for few species, i.e., S. pygme, S. nlmin, S. chillei, nd possibly S. gimmeli nd S. dvidttenboroughi (pending confirmtion). Stumpffi pygme nd S. nlmin produce fom nests. In S. pygme, developing embryos nd non-feeding tdpoles were found in these fom nests in the wild (Glw & Vences, 1994), with 388

119 VERTEBRATE ZOOLOGY 67 (3) 2017 froglet mesuring < 3 mm snout-vent length. In cptivity, fom nests were found tht contined no eggs, but were gurded by mle frog. A fom nest of S. nlmin produced in cptivity contined no eggs but ws gurded by n dult frog (Klges et l., 2013). We lso found comprtively lrge fom nests contining numerous eggs, embryos, nd tdpoles in metmorphosis (ZFMK ), possibly of S. gimmeli, ttched to the inner wll of hollow tree in Mnongrivo Reserve in Februry Here we report tht the only known specimen of S. dvidttenboroughi, ws observed to gurd eggs in jelly nest, lthough unfortuntely those eggs were not collected nd it ws not possible to relibly scertin species identity of the embryos. Furthermore, we observed mle nd eggs of S. chillei in wter-filled snil shell, nd verified the identity of these eggs by DNA brcoding. This confirms tht the production of fom nests is not generl chrcteristic of breeding in Stumpffi. In the closely relted genus Rhombophryne, subterrnen or terrestril jelly nests re probbly the norm (lthough in this genus breeding ecology is lso poorly known). Other cophylines lso ly jelly-bound eggs, often in wter-filled tree holes or in burrows in the lef litter (e.g. Blommers-Schlösser, 1975). Given tht fom nests re otherwise unknown in the fmily Microhylide, it seems likely tht they re derived reproductive trit of Stumpffi tht might chrcterize minly or only the species in clde A, lthough more dt re needed to test this hypothesis. Furthermore, prentl cre seems to be distributed cross the mjority of cophyline species (e.g., Blommers-Schlösser, 1975; Köhler et l., 1997). This subfmily, nd the genus Stumpffi in prticulr, my therefore provide n excellent system for study of the evolution of breeding systems in nurns, especilly in light of the resolution of their txonomy nd systemtics. Biogeogrphic ptterns Our phylogenetic nlysis reveled tht the two most species-rich cldes of the genus (cldes A nd C) re predominntly distributed in two distinct res of Mdgscr, i.e., northern nd north-western Mdgscr vs. estern nd north-estern Mdgscr (Fig. 4). Within clde A, clde A1 nd A3 contin species locted only in northern Mdgscr, nd clde A2 contins species from northern nd north-western Mdgscr. In Clde C, both C1 nd C2, re composed of species from estern nd north-estern Mdgscr. Wollenberg et l. (2008) reveled tendency to microendemism in cophyline microhylid frogs. Indeed, mny Stumpffi re currently known only from smll res not lrger thn 50 to 100 km 2 (Köhler et l., 2010; Ndrintso et l., 2012). Rnge size in Stumpffi might be correlted to body size (Klges et l., 2013), in greement with generl pttern in Mdgscr s mphibins nd reptiles (Brown et l., 2016). Accordingly, one fctor responsible for this microendemism might be the overll smll size of Stumpffi species, s they re mong the smllest frogs in Mdgscr, nd some number mong the smllest frogs in the world (Glw & Vences, 2007). However, there re exceptions tht do not fit this pttern. Some species of clde A (i.e., smll-sized or miniturized frogs), S. mmitik, S. psologloss, nd S. gimmeli, hve reltively wide distribution, lthough more dt is necessry to support this especilly for S. psologloss nd S. gimmeli. Also, in clde C (i.e., smll- to modertesized species), S. grrffoi, S. prdus, nd S. nlnjirofo hve reltively wide distribution, nd the sme is true for S. nigrorubr lthough the southernmost known specimens from Mnombo re geneticlly strongly differentited. In totl, seven out of 41 nominl species hve wider distribution nd re not microendemics ccording to current knowledge. We suspect tht future reserch will led to rnge extensions of mny Stumpffi species but the generl pttern in the genus, i.e., overll smll rnges nd existence of true microendemic species, is unlikely to chnge. Only few Stumpffi were known to occur in the dry forests of Mdgscr (Klges et l., 2013). The results herein increse the number of these species occurring in dry forests. In ddition to S. stffordi, S. be, S. hr, nd S. nlmin, lso S. mmitik nd S. lrinki occur in Ankrn Ntionl Prk in northern Mdgscr. This new dtset lso reveled tht Stumpffi collected in 2010 nd 2012 by us from Ankrfntsik Ntionl Prk re indeed S. nlmin, s previously hypothesized by Klges et l. (2013), nd this probbly lso pplies to erlier collected mteril from this re tht previously ws ssigned to S. psologloss by Blommers-Schlösser & Blnc (1991). Ptterns of digitl reduction The loss of limbs is one of the most extreme morphologicl chnges in the history of tetrpods (Lnde, 1978). A chnge in the number of digits is process during the embryonic stge chrcterized by the process of construction followed by destruction (Glis et l., 2001). According to Alberch & Gle (1983, 1985), there is well-defined pttern in terms of wht elements re most likely to be reduced. In frogs, for exmple, the first toe is lwys reduced first. Severl digitl reductions re seen in Stumpffi. The order in which their fingers re reduced is consistently first finger first, followed by the fourth, then second, with the third remining s the only developed digit tht becomes brodened in S. tridctyl, S. contumeli, nd S. obscoen. Remrkbly, toe reduction does not seem to follow the scheme of Alberch & Gle (1983, 1985). In most tx, but especilly in S. tridctyl, S. contumeli, nd S. obscoen, the first nd second toes re more strongly reduced thn the fifth toe, t lest externlly (Figs. 9 11). If this trend is supported bsed on the phlngel development, then it would be significnt devition from wht hs been ccepted s typicl trend mong ll frogs. 389

120 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Severl sister tx re chrcterized by the sme type of digitl trnsformtions: the first toes re reduced in S. ngeluci nd S. mledict, the phlngel discs re slightly enlrged in S. gimmeli nd S. ihrn, the first finger nd toe re reduced in S. ynniki nd S. nlmin, nd in S. mdgscriensis nd S. pygme (ll of which re smll-sized or miniturized species from clde A), nd the first finger is very strongly reduced in length nd first toe bsent in S. obscoen nd S. dvidttenboroughi (smll-sized or miniturized species from clde B). In clde C, trend is seen to reduce in length especilly the first toe, even in some reltively lrge-sized species, nd with complete reduction in externl view in some smll-sized species (e.g., S. tetrdctyl). In clde D, both S. contumeli nd S. tridctyl hve digitl reductions, i.e., the first finger is not visible, the second nd fourth fingers re reduced to smll knobs, nd the third finger is brodened to tringulr shpe. Investigtions into the osteologicl bsis for these digitl reductions re on-going (Scherz et l., unpubl. dt). Body size ptterns The pttern of body size evolution is prticulrly expressed in Clde A. Cldes A1 nd A2 consist of only smll-sized nd miniturized species ( mm) distributed in the north nd northwest of Mdgscr, wheres A3 is composed of only lrge sized species (21 28 mm) from the north of Mdgscr. Clde B, in contrst, is vrible both in morphology nd distribution. It is composed of both estern (S. miery, S. obscoen, S. dvidttenboroughi) nd northwestern species (S. meikee), nd of smll-sized or miniturized species (S. miery, S. obscoen, S. dvidttenbouroughi) s well s lrger species (S. meikee). However, more dt re needed on this group, s it is only modertely supported (0.94 BPP) in the consensus tree. Within clde C there is considerble vrition in body size. However, pirs of sister species in clde C, bsed on dult mles, hve comprble body sizes, such s S. prdus nd S. diutissim (17 22 mm), S. mkir nd S. dolchi (12 mm), nd S. chillei nd S. nlnjirofo (19 20 mm). Clde D is formed by the two miniturized species S. tridctyl (10 11 mm) nd S. contumeli (8 9 mm). The ltter species is the smllest known frog species from Mdgscr nd mong the smllest frogs in the world: Rittmeyer et l. (2012) identified 29 of the world s smllest frogs, mong which 15 re microhylids. While this list would need to be djusted to ccount for recent descriptions, t present S. contumeli (mx. mle SVL 8.9 mm) would compete with Pedophryne mnuensis (mx. mle SVL 8 mm) nd P. swiftorum (mx. mle SVL 8.9 mm) for the title of the smllest frog in the world. To our knowledge no other species described since 2012 might lso be cndidte for this title. The history of the body size evolution of this group is thus clerly confounded by repeted shrinkge nd growth cross different cldes. However, there re severl indictions tht the group my hve hd very smll common ncestor, in prticulr digitl reduction in the mjority of species, the bsence of vomerine nd mxillry teeth in even the lrgest species ccording to currently vilble dt, nd the bsl reltionship of the diminutive species of clde D with the rest of the genus. Further reserch into the evolution of the size of these frogs, including ncestrl size reconstructions, my help us to understnd not just the vriety of sizes present in most cldes, but lso the dispersl nd specition history of the genus. Color ptterns Three of the new species, S. nigrorubr, S. miovov, nd S. meikee, show the red/ornge color on the belly tht ws previously known only from Stumpffi kibomen nd to some degree from S. roseifemorlis (Glw et l., 2015). While S. meikee is member of clde B, the other red-bellied species belong to clde C but do not form monophyletic group (Fig. 4). The function of the red/ ornge ventrl color is poorly known (Glw et l., 2015). Severl other color elements repet frequently cross mny unrelted tx, including the teddyber shped dorsl mrking, dorsl chevrons, crossbnds on the legs nd rms, light nnuli before the terminl phlnges of the fingers nd toes, nd lightened or bright res on the posterodorsl shnk. These my reflect some limittions or developmentl consistencies in these frogs, especilly s ll of these elements re known lso to some degree from other gener indeed, most microhylids hve light nnuli before their terminl phlnges. At lest mong Mlgsy frogs, the mrkings on the posterodorsl shnk seem to be more or less restricted to Stumpffi (possibly lso present in some species of Anodonthyl; Glw & Vences, 2007). Cll ptterns The clls of Stumpffi species re mostly composed of the repetition of single chirping notes, except for S. psologloss tht emits long note consisting of pulses (Glw & Vences, 1992). The clls of the newly described species re not fundmentlly different to most of the formerly recognized nominl species of Stumpffi. Interestingly, our dt revels similr pttern of cll differences between species occuring in symptry tht indictes prllel processes in cll evolution within Stumpffi: Species from Montgne d Ambre Ntionl Prk, S. ngeluci, S. huwei, nd S. mledict, differ in cll durtion, with S. ngeluci emitting longer clls ( ms) thn S. huwei (61 74 ms), nd S. mledict emitting shorter clls ( ms) thn S. ngeluci but longer thn S. huwei. Similrly, this is the cse for the species of the Mrojejy Mssif, with only some overlp in cll durtion between S. chillei (36 79 ms), S. diutissim (53 56 ms), S. tridctyl ( ms), nd S. roseifemorlis ( ms). 390

121 VERTEBRATE ZOOLOGY 67 (3) 2017 Also the durtion of inter-cll intervls often differs mong symptric species, e.g. in Ankrn Ntionl Prk, with S. mmitik ( ms; this is bsed on the longest cll series vilble for the species) clling fster thn S. lrinki ( ms). Similrly, this is the cse for the species from Rnomfn (inter-cll intervls for S. nigrorubr re ms nd for S. miery ms). In Ambodivongy, the dominnt frequencies of clls differ between symptric species with S. contumeli clls ( Hz) hving higher frequency thn those of S. prdus ( Hz). Conservtion sttus nd conservtion significnce of newly described species Most species of Stumpffi were previously ssessed s Dt Deficient, except S. nlmin, which ws considered Criticlly Endngered (Klges et l., 2013), S. gimmeli (Lest Concern; Andreone et l., 2005), nd S. pygme, S. be, S. hr, S. megsoni, nd S. stffordi (Vulnerble; Andreone et l., 2005, Köhler et l., 2010). All members of the genus were gin ressessed between 2014 nd 2016 by the IUCN Species Survivl Commission Amphibin Specilist Group for Mdgscr (long with ll other Mlgsy mphibins). This ressessment found three species to be Dt Deficient (S. megsoni, S. tetrdctyl, nd S. tridctyl), two Lest Concern (S. grndis nd S. gimmeli), one Vulnerble (S. stffordi), seven Endngered (S. be, S. kibomen, S. mdgscriensis, S. miery, S. psologloss, S. pyme, nd S. roseifemorlis), nd two Criticlly Endngered (S. nlmin nd S. hr). Our dt revels tht Stumpffi species occur both in protected s well s in unprotected res. Some of the species collected in unprotected res like S. mmitik in Vohemr, S. gimmeli in Antsirsir nd 27 km from Ambnj, nd S. ihrn in gllery forest ner Andrfinkon, were found in highly degrded res (i.e. mngo or vnill plnttions). Also, some species occurring in protected res (S. nlmin in Ankrfntsik NP, S. ngeluci in Montgne d Ambre NP, S. huwei in Montgne d Ambre NP, S. grrffoi in Betmpon Strict Nture Reserve) were found in rther disturbed res within these prks, i.e. either close to the prk office or in dense lef litter t the border of tril. We gree with the hypothesis of Ndrintso et l. (2013) tht most Stumpffi re ble to dwell in secondry forest s long s lyer of lef litter provides sufficient humidity nd some shde exists. On the other hnd, the strikingly distinct ptterns of distribution observed in some of the six different species of Stumpffi occurring t Betmpon Strict Nture Reserve might contrdict this hypothesis nd suggest subtle underlying hbitt speciliztion. However, studies from forest frgments in the Rnomfn region lso confirm tht Stumpffi spp. pprently persist in degrded hbitt (Riemnn et l., 2015). Accordingly, even if most species within the genus hve very smll rnge of distribution, we do not consider ll members of the genus to be under immedite thret of extinction. Nevertheless, strict ppliction of the IUCN Red List criteri (IUCN 2001) yields 2 species s Lest Concern, 6 species s Ner Thretened, 5 species s Vulnerble, 17 species s Endngered, 8 species s Criticlly Endngered, nd 3 species s Dt Deficient (Tble 6). The IUCN Red List criteri re however inherently nd intentionlly subjective; explicity they my tke into ccount the suspicions nd feelings of the ssessors, however vgue. This mkes them prcticlly impossible to pply consistently. To ttempt to overcome this, we hve mde concerted effort to keep these ssessments consistent with those recently published in which severl of the uthors were intensively involved nd in strict keeping with the criteri. Despite this, we wish to emphsise tht (1) strict ppliction of the criteri is often inpproprite; (2) even here there ws some dissent mong uthors s to which criteri should be pplied in which wy to ech species; nd (3) the most commonly used criterion, B, which ssesses the distribution of thret bsed on rnge size, occupncy, nd on-going hbitt nd species trends within tht rnge, is ill suited to hndling smll nd/or microendemic tx, but relisticlly only one other criterion (D2) cn be pplied to tx from which no popultion size dt re vilble. The current Red List system results in infltion of risk, nd emphsizes publiction of high thret ctegories with totlly indequte dt; the discovery of single individul tht expnds Criticlly Endngered species rnge by 40 km would instntly result in its thret sttus being downgrded. This is problem deeply nested within the IUCN s ssessment strtegy, nd one tht ought to be ddressed due to the extreme vlue plced on Red List rnkings by policy mkers, conservtion prctitioners, nd others. We therefore urge tht more species be listed s Dt Deficient insted of being given inflted sttuses, s this is n honest, scientific sttement of the current stte of knowledge on them, which is lrgely indequte. This includes not just species from which few individuls or few dt re vilble, but lso txonomic species complexes, for which ny ssessment bsed on the full distribution of the complex will be drstic overestimtion of ny individul member s rnge, nd n underestimtion of the threts ech species is fcing. Finl considertions With this study, we hve tken first big step towrd n integrtive resolution of the txonomy of the smller cophylines, which hve hd n enormous txonomic gp (Vieites et l., 2009; Perl et l., 2014). We hve incresed the nominl diversity of Stumpffi from 15 to 41, nd provided bseline informtion tht will llow the description of few more cndidte species once dditionl specimen mteril becomes vilble. This brings us to pproximtely 100 nmed cophyline species. However, in other gener, numerous cndidte species remin, nd we expect 391

122 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Tble 6. Evlution nd proposl of the IUCN Red List sttus for nominl species in the genus Stumpffi (in lphbeticl order) ccording to defined criteri (IUCN 2001). Plese note tht ppliction of criteri prtly remin subjective nd not lwys identicl rtionle ppers dequte (see column with comments). Abbrevitions used: TDL = Thret-defined loclitiy sensu IUCN 2001; AOO = Are of Occupncy; EOO = Extent of Occurrence; CR = Criticlly Endngered; EN = Endngered; VU = Vulnerble; NT = Ner Thretened; LC = Lest Concern; DD = Dt Deficient. Species Applied IUCN Criteri S. chillei B1b(iii) S. nlmin B1b(iii), B2b(iii) S. nlnjirofo B1b(iii) S. ngeluci B1b(iii) S. be B1b(iii) S. betmpon B1b(iii), B2b(iii) S. cotumeli B1b(iii) Comments Current Red List Sttus Proposed Red List Sttus One TDL, which is in protected re. On-going hbitt degrdtion due to illegl logging ctivity, especilly t low ltitudes. EOO nd AOO probbly > 10 km 2. In line with Rhombophryne svk from sme re. Two TDLs, one in protected re. Species proposed s CR in originl description becuse of single known loclity. Moleculr studies now reveled second loclity within protected re. New EOO still < 5000 km 2. Both known loclities under hevy nthropogenic pressure. Three TDLs including two protected res. Probbly reltively widespred in lowlnd rinforests long the northern est cost, but EOO still < 20,000 km 2, nd popultion extremely frgmented. Lowlnd rinforest hbitts re under severe humn pressure. Three TDLs in three protected res (Joffreville is ner to two of these protected res nd not considered its own TDL). EOO < 5000 km 2. Hbitt degrdtion is ongoing t known loclities. One TDL, which is protected re. Does not qulify for CR under criterion B s its speciliztion to krstic hbitt nd cves mens it is probbly not experiencing decline in extent or qulity of hbitt. Could rpidly become CR if nthropogenic ctivity in the prk increses drsticlly. One TDL, which is protected re. EOO ~26km 2. Lowlnd rinforest hbitt is under severe nthropogenic pressure. Probbly lso qulifies for CR under A2c. One TDL, which is in protected re. Current knowledge is poor, nd it my hve wider distribution, but ll specimens were collected t exctly the sme loclity, nd the hbitt is under nthropogenic pressure. Not yet evluted EN CR EN Not yet evluted VU Not yet evluted EN EN EN Not yet evluted CR Not yet evluted CR S. dvidttenboroughi B1b(iii), B2b(iii) See S. betmpon. Not yet evluted CR S. diutissim B1b(iii) See S. chillei. Not yet evluted EN S. dolchi D2 S. edmondsi n/ One TDL, which is protected re. AOO < 20 km 2. No on-going threts sensu criterion B. However, islnd popultion is vulnerble nd could esily be driven to CR or EX quickly. Currently one or two TDLs, in close proximity to severl protected res. Probbly widespred in Andsibe-Mntdi re, nd could extend north up to Zhmen through continuous forest; EOO is still certinly < 20,000 km 2. Apprently not immeditely thretened by declines in hbitt qulity, but if this chnges, will qulify for VU B1b(iii). Not yet evluted VU Not yet evluted NT S. fusc B1b(iii) See S. contumeli. Not yet evluted CR S. grrffoi B2b(iii) S. gimmeli None S. grndis B1b(iii) S. hr B1b(iii) S. huwei n/ S. ihrn B1 b(iii) Three TDLs, t lest one in protected re. Current EOO < 4000 km 2. Popultions severely frgmented. On-going hbitt decline, prticulrly of lowlnd rinforest. Roughly nine TDLs currently known, t lest two in protected res. Although current EOO is less thn 20,000 km 2, species is bundnt, hs pprently stble popultions, occurs in secondry nd disturbed hbitts, nd probbly is not significntly ffected by deforesttion. See S. chillei, but found t higher elevtion where threts re lower. EOO probbly < 100 km 2. Assessed s EN in line with Rhombophryne vventy nd R. serrtoplepbros recently ssessed from the sme loclity. One TDL. AOO < 10 km 2. Known from severl cves nd round smll creek on Nosy Hr, the ltter suffering from severe hbitt modifiction (construction of concrete stirs for tourism). One TDL, which is protected re. Current EOO is smll enough to qulify for CR, but currently there re no on-going declines or fluctutions in hbitt, popultions, or distribution t the reltively high ltitude of the known loclities. Three TDLs, none of which is protected. EOO < 5000 km 2. All loclities under hevy nthropogenic pressure, severe degrdtion of hbitt. Not yet evluted EN LC LC LC EN CR CR Not yet evluted NT Not yet evluted EN S. jennoeli B1b(iii), B2b(iii) See S. betmpon. Not yet evluted CR S. kibomen n/ See S. edmondsi. EN NT S. lrinki D2 See S. be. Less specilized to krstic hbitts thn tht species, but probbly lso not experiencing on-going declines in hbitt, rnge, or popultion size. Not yet evluted VU 392

123 VERTEBRATE ZOOLOGY 67 (3) 2017 Tble 6 continued. Species Applied IUCN Criteri S. mdgscriensis n/ S. mkir n/ Comments Current Red List Sttus Proposed Red List Sttus See S. huwei. Also found round Montgne des Frnçis (S. sp. C25), incresing the spred of risk, but still could become EN or CR. Recently ssessed s EN, but here shown to hve wider distribution nd is not known from edge res most ffected by deforesttion. Known from only one specimen. Species could be found throughout Mkir, or loclly restricted (could be CR, EN, or VU). No known immedite threts in the center of Mkir, but if found t the edges could be under significnt thret. More informtion is needed. EN NT Not yet evluted DD S. mledict n/ See S. huwei. Not yet evluted NT S. mmitik n/ S. megsoni n/ S. meikee n/ S. miery B1b(iii) S. miovov B1b(iii) S. nigrorubr B1b(iii) S. obscoen n/ S. prdus B1b(iii) S. psologloss B1b(iii) S. pygme B1b(iii) Three known TDLs, one in protected re. Current EOO < 20,000 km 2, but probbly more widespred. Occurs in secondry hbitts. No known on-going threts. Assessment of popultion trend is needed, but conservtively considered LC. One TDL, which is not protected nd is thretened. Apprently tolernt to some degree of hbitt modifiction, but certinly experiencing on-going decreses in qulity of hbitt. Difficult to ssess, here clssified s DD for consistency with current Red List. One confirmed TDL, but unpublished dt suggest t lest one more loclity southwest of Belnn (Scherz et l., unpubl. dt); currently in one protected re. EOO not currently clculble. Probbly thretened by on-going hbitt declines. Could qulify for CR, EN, or VU. More informtion is needed. At lst four TDLs, some inside protected re. EOO < 5000 km 2. On-going decline in extent nd qulity of hbitt, but pprently tolernt to some hbitt degrdtion. One TDL, which is not yet protected. EOO < 250 km 2. Continuing deforesttion nd hbitt degrdtion; high nthropogenic pressure. In line with Rhombophryne longicrus from the sme re. At lest six TDLs, some inside protected re. EOO < 5000 km 2. On-going decline in extent nd qulity of hbitt, but pprently tolernt to some hbitt degrdtion. One TDL, which is in protected re. Current EOO < 100 km 2, but probbly more widespred. On-going decline in extent nd qulity of hbitt outside of protected res, but overll probbly stble. Thus lmost qulifies for CR B1b(iii), but currently suspected to not stisfy ny subcriterion of Bb. Three TDLs, one of which is protected. EOO < 20,000 km 2. Popultions severely frgmented, nd lowlnd forest is under hevy nthropogenic pressure. Four TDLs, t lest one of which is protected. EOO < 5000 km 2, but probbly more widespred thn currently known. Anthropogenic pressure is high throughout rnge. One TDL, which is protected re. EOO < 100 km 2. High nthropogenic pressure, but somewht resilient to it. Could rpidly be driven to CR or EX by encrochment on its hbitt, fire, or introduction of invsive species. Not yet evluted LC DD DD Not yet evluted DD EN EN Not yet evluted EN Not yet evluted EN Not yet evluted NT Not yet evluted VU EN EN EN EN S. roseifemorlis B1b(iii) See S. chillei. EN EN S. sort B1b(iii) See S. miovov. Not yet evluted EN S. spndei B2b(iii) A single specimen is known. One TDL, which is not protected. Forest frgment where HT ws found hs n re of 6 km 2 (pprox. AOO), but species is probbly more widely distributed. Nerby forest is highly frgmented. Not yet evluted CR S. stffordi D2 AOO estimted to be < 10 km 2. Loclity is suffering from on-going hbitt degrdtion nd high nthropogenic pressure. VU VU S. tetrdctyl B1b(iii) One TDL, which is poorly protected. EOO < 300 km 2. Abundnt in forest nd secondry hbitt, but suspected to be declining from nthropogenic ctivity. DD EN S. tridctyl B1b(iii) See S. chillei. DD EN S. ynniki B1b(iii) One TDL, which is in new protected re. Known EOO is diminutive, but the species is probbly found more widely in the Tsrtnn region. On-going declines in extent nd qulity of hbitt throughout nerby res. More reserch is likely to lower the thret sttus of this species. Not yet evluted CR 393

124 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs to see similr pproches tken to resolving their txonomy in the coming yers. Yet, complete nd conclusive txonomy of Mdgscr s frogs is not on the horizon, nd will require both the clrifiction of old nmes, nd the collection of more mteril for txonomic purposes. Acknowledgments We re deeply indebted to R. Günther for numerous vluble comments on previous version of the mnuscript, s well s to R. Ernst, U. Fritz, nd A. Zrske for their editoril efforts nd to M.H. Fischer for the efficiency nd ptience in the typesetting nd proofing stges.. We re grteful to our collegues nd guides M.C. Bletz, P. Bor, H. Enting, R. Dolch, A. Knoll, N. D Cruze, P.-S. Gehring, M. Pbijn, D. Edmonds, J. Müller-Jung, M. Puente, L. Rhrivololoniin, M. Teschke, J.L. Ptton, H. Rsolonjtovo, R.D. Rndriniin, F.M. Rtsovin, S. Rsmison, E. Rjeririson T. Rjofirison, A. Rzfimnntso, J.H. Rzfindribe, J. Solo, A. Telo, F. Rndrinsolo, J. Nöel, D. Slvi, J.D. Hrris, E. Scnrini, S. Frvelli, M. Rndrimiliso, H. Lv, D. Vlln nd J.H.Velo for their help during the field work over mny yers, C. App nd H. Reuper for ssistnce with DNA sequencing nd preliminry phylogenetic nlysis. The Direction Générle des Forêts kindly grnted the reserch nd collection permits. This work ws crried out in frmework of collbortion ccords mong the Deprtment of Animl Biology Antnnrivo nd finncilly supported by the Volkswgen Foundtion, the Deutsche Forschungsgemeinschft (GL 665/1-1; RO 3064/2-1), the Europen Assocition of Zoos nd Aquri (EAZA), the Sint Louis Zoo s Field Reserch for Conservtion progrm (FRC# 12-12) of the Wildcre Institute, Museo Regionle di Scienze Nturli, Gondwn Conservtion nd Reserch, nd the Mohmed Bin Zyed Foundtion (project ,) nd by the Spnish Ministry of Economy under Grnts CGL nd CGL P to DRV. AR is supported by PhD fellowship of the Deutscher Akdemischer Austuschdienst. 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129 Senckenberg Gesellschft für Nturforschung, (3): I XXIV Electronic Supplement Appendix b c d Fig. S91. Stumpffi psologloss, preserved specimens from Nosy Be: ( b) ZSM 479/2000 (FG/MV ); (c d) ZSM 480/2000 (FG/MV ); Scle br = 5 mm. ISSN I

130 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Electronic Supplement b c d e f g h i j k l m n o p q r s t Fig. S92. Stumpffi nlmin, preserved specimens from Ankrfntsik Ntionl Prk: ( b) KUZA 0001 (2010-F002); (c d) KUZA 0002 (2010-F003); (e f) KUZA 0003 (2010-F004); (g h) KUZA 0004 (2010-F005); (i j) KUZA 0005 (2011-Ad-049), (k l) KUZA 0006 (2011-Ad-050); (m n) KUZA 0007 (2011-Ad-059); (o p) KUZA 0010 (2013-NoCode); (q r) KUZA 0008 ( ); (s t) KUZA 0009 ( ); Scle br = 5 mm. II

131 Electronic Supplement VERTEBRATE ZOOLOGY 67 (3) 2017 b c d Fig. S93. Stumpffi gimmeli, preserved specimens: ( b) ZSM 833/2003 (FG/MV ) from Mnongrivo; (c d) ZSM 597/2001 (MV ) from Antsirsir; Scle br = 5 mm. b b c d Fig. S94. Stumpffi mdgscriensis, preserved specimen from Montgne d Ambre (ZSM 201/2004); Scle br = 5 mm. Fig. S95. Stumpffi cf. mdgscriensis (= S. sp. C25), preserved specimens from Montgne des Frnçis: ( b) ZSM 2108/2007 (FGZC 1103); (c d) ZSM 2109/2007 (FGZC 1105); Scle br = 5 mm. III

132 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Electronic Supplement b c d e f Fig. S96. Stumpffi pygme, preserved specimens from Nosy Be: ( b) ZFMK 53769; (c d) ZFMK 53772; (e f) ZFMK 53773; Scle br = 5 mm. b c d e f g h i j k l Fig. S97. Stumpffi ngeluci sp. nov., preserved prtype specimens from Montgne des Frnçis: ( b) ZSM 302/2004 (FGZC 583); (c d) ZSM 301/2004 (FGZC 581); (e f) ZSM 224/2004 (FGZC 434); (g h) ZSM 223/2004 (FGZC 433); (i j) ZSM 303/2004 (FGZC 586); (k l) ZSM 300/2004 (FGZC 580); Scle br = 5 mm. IV

133 Electronic Supplement VERTEBRATE ZOOLOGY 67 (3) 2017 b c d e f g h Fig. S98. Stumpffi huwei sp. nov., preserved specimens from Montgne d Ambre Ntionl Prk: ( b) holotype ZSM 221/2016 (ZCMV 13618); (c d) prtype ZSM 222/2016 (ZCMV 13620); (e f) prtype UADBA-A (ZCMV 13621); (g h) prtype ZSM 223/2016 (ZCMV 13622); Scle br = 5 mm. b c d e f g h Fig. S99. Stumpffi ihrn sp. nov., preserved specimens from Sort: ( b) prtype ZSM 1642/2012 (FGZC 3800); (c d) prtype ZSM 1650/2012 (FGZC 3893); (e f) holotype ZSM 1651/2012 (FGZC 3895); (g h) prtype ZSM 1652/2012 (FGZC 3927); Scle br = 5 mm. V

134 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Electronic Supplement b c d e f Fig. S100. Stumpffi lrinki sp. nov., preserved prtype specimens from Ankrn Ntionl Prk: ( b) ZSM 779/2003 (FG/ MV ); (c d) ZSM 861/2003 (FG/MV ); (e f) ZSM 1669/2008 (FGZC 1620); Scle br = 5 mm. b c d e f Fig. S101. Stumpffi mledict sp. nov., preserved specimens from Mon tgne d Ambre: ( b) holotype ZSM 2079/2007 (FGZC 1049); (c d) prtype ZSM 2169/2007 (FGZC 1244); Scle br = 5 mm. VI

135 Electronic Supplement VERTEBRATE ZOOLOGY 67 (3) 2017 b c d e f g h i j Fig. S102. Stumpffi mmitik sp. nov., preserved prtype specimens: ( b) ZSM 375/2005 (FGZC 2725) from Andp; (c d) ZSM 862/2003 (FG/MV ) from Ankrn Ntionl Prk; (e-f) ZSM 3234/2012 (ZCMV 13531); (g h) ZSM 3233/2012 (ZCMV 13529); (i j) ZSM 3234/2012 (ZCMV 13530); Scle br = 5 mm. VII

136 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Electronic Supplement b c d e f Fig. S103. Stumpffi sort sp. nov., preserved specimens from Sort forest: ( b) prtype ZSM 1643/2012 (FGZC 3618); (c d) holotype ZSM 1644/2012 (FGZC 3621); (e f) prtype ZSM 1645/2012 (FGZC 3622); Scle br = 5 mm. b Fig. S104. Stumpffi ynniki sp. nov., preserved prtype specimen collected in forest frgment between Belnn nd Antsohihy, ZSM 1825/2010 (ZCMV 12600); Scle br = 5 mm. VIII

137 Electronic Supplement VERTEBRATE ZOOLOGY 67 (3) 2017 b c d e f g h Fig. S105. Stumpffi tridctyl, preserved prtypes from Mrojejy Ntionl Prk: ( b) MNHN ; (c d) MNHN ; (e f) MNHN ; (g h) MNHN ; Scle br = 5 mm. b c d Fig. S106. Stumpffi contumeli sp. nov., preserved prtype specimens: ( b) ZSM 441/2010 (FGZC 4248) from Mkir; (c d) ZSM 442/2010 (FGZC 4252) from Mkir; Scle br = 5 mm. b Fig. S107. Stumpffi obscoen sp. nov., preserved prtype specimen in dorsl nd ventrl view from Andsibe: ( b) ZSM 380/ 2005 (FGZC 2664); Scle br = 5 mm. IX

138 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Electronic Supplement b c d Fig. S108. Stumpffi meikee sp. nov., preserved prtype specimens from Mtsbory Miky, Tsrtnn Strict Nture Reserve: ( b) ZSM 1821/2010 (ZCMV 12372); (c d) ZSM 1822/2010 (ZCMV 12373); Scle br = 5 mm. Fig. S109. Stumpffi kibomen, preserved holotype from ner Andsibe (ZFMK 60007); Scle br = 5 mm. X

139 Electronic Supplement VERTEBRATE ZOOLOGY 67 (3) 2017 b c d e f Fig. S110. Stumpffi roseifemorlis, preserved specimens from Mrojejy Ntionl Prk: ( b) MNHN ; (c d) ZSM 373/2005 (FGZC 2808); (e f) ZSM 374/2005 (FGZC 2883); (g h) ZSM 487/2005 (ZCMV 2047); Scle br = 5 mm. XI

140 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Electronic Supplement b c d e f g h Fig. S111. Stumpffi chillei sp. nov., preserved specimens from Mrojejy: ( b) prtype ZFMK 57460; (c d) prtype ZFMK 57461; (e f) prtype ZFMK 59893; (g h) holotype ZSM 536/2016 (ZCMV 15149); Scle br = 5 mm. XII

141 Electronic Supplement VERTEBRATE ZOOLOGY 67 (3) 2017 c b d e f g h i j k l Fig. S112. Stumpffi nlnjirofo sp. nov., preserved prtype specimens from Nosy Mngbe Specil Reserve: b ZSM 491/2005 (ZCMV 2104); (c d) ZSM 488/2005 (ZCMV 2143); (e f) ZSM 492/2005 (ZCMV 2178); (g h) ZSM 225/2016 (ACZCV 0121); (i j) ZSM 226/2016 (ACZCV 0224); (k l) ZSM 227/2016 (ACZCV 0225); Scle br = 5 mm. XIII

142 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Electronic Supplement b c d e f Fig. S113. Stumpffi diutissim sp. nov., preserved prtype specimens from Mrojejy Ntionl Prk: ( b) ZSM 496/2005 (ZCMV 2082); (c d) ZSM 495/2005 (ZCMV 2067); (e f) ZSM 376/2005 (FGZC 2742); Scle br = 5 mm. XIV

143 Electronic Supplement VERTEBRATE ZOOLOGY 67 (3) 2017 b Fig. S114. Stumpffi edmondsi sp. nov., preserved prtype specimen : ( b) ZSM 1731/2012 (RDR 1065) from Andsibe; Scle br = 5 mm. b Fig. S115. Stumpffi fusc sp. nov., preserved prtype specimen from Ambodivongy, Mkir: ( b) ZSM 436/2010 (FGZC 4253); Scle br = 5 mm. XV

144 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Electronic Supplement b c d e f g h Fig. S116. Stumpffi jennoeli sp. nov., preserved prtype specimens from Betmpon: ( b) MRSN A6283; (c d) MRSN A6386; (e f) ZSM 207/2016 (ACZCV 0167); (g h) ZSM 208/2016 (ACZCV 0218); Scle br = 5 mm. XVI

145 Electronic Supplement VERTEBRATE ZOOLOGY 67 (3) 2017 e f g h i j k l Fig. S117. Stumpffi prdus sp. nov., preserved prtype specimens from Mkir: ( b) ZSM 432/2010 (FGZC 4210); (c d) ZSM 440/2010 (FGZC 4278); (e f) ZSM 438/2010 (FGZC 4265); (g h) ZSM 433/2010 (FGZC 4213); (i j) ZSM 434/2010 (FGZC 4214); (k l) ZSM 439/2010 (FGZC 4268); Scle br = 5 mm. XVII

146 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Electronic Supplement b c d Fig. S118. Stumpffi tetrdctyl, preserved specimens from Nosy Borh: ( b) ZSM 594/2006 (ZCMV 3396); (c d) ZSM 593/2006 (ZCMV 3395); Scle br = 5 mm. b c d e f g h Fig. S119. Stumpffi betmpon sp. nov., preserved prtype specimens from Betmpon: ( b) ZSM 216/2016 (ACZCV 0041); (c d) ZSM 217/2016 (ACZCV 0046); (e f) ZSM 218/2016 (ACZCV 0047); (g h) ZSM 219/2016 (ACZCV 0103); Scle br = 5 mm. XVIII

147 Electronic Supplement VERTEBRATE ZOOLOGY 67 (3) 2017 b c d e f g h i j Fig. S120. Stumpffi grrffoi sp. nov., preserved prtype specimens from Mhso Forest: ( b) ZSM 631/2009 (ZCMV 8684); (c d) ZSM 632/2009 (ZCMV 8685); (e f) ZSM 633/2009 (ZCMV 8687); (g h) ZSM 220/2016 (ACZCV 0006); (i j) MRSN A6278; Scle br = 5 mm. XIX

148 Rkotorison, A. et l.: Integrtive txonomy of Stumpffi frogs Electronic Supplement b c d e f g h Fig. S121. Stumpffi miovov sp. nov., preserved prtype specimens from Sort: ( b) ZSM 1640/2012 (FGZC 3650); (c d) ZSM 1647/2012 (FGZC 3675); (e f) ZSM 1648/2012 (FGZC 3676); (g h) ZSM 1646/2012 (FGZC 3751); Scle br = 5 mm. XX

149 Electronic Supplement VERTEBRATE ZOOLOGY 67 (3) 2017 b b Fig. S122. Stumpffi spndei sp. nov., preserved holotype from Mhso Forest: ( b) ZSM 1751/2008 (ZCMV 8802); Scle br = 5 mm. Fig. S123. Stumpffi dvidttenboroughi sp. nov., preserved holotype from Betmpon Strict Nture Reserve: ( b) ZSM 204/2016 (ACZCV 0106); Scle br = 5 mm. b Fig. S124. Stumpffi nigrorubr sp. nov., preserved holotype from long the rod between Ifndin nd Tolongoin: ( b) ZSM 2448/ 2007 (ZCMV 5967); Scle br = 5 mm. b Fig. S125. Stumpffi sp. C29, preserved specimen from Mtsbory Miky, Tsrtnn Strict Nture Reserve: ( b) ZSM 1824/2010 (ZCMV 12387); Scle br = 5 mm. XXI