A new cryptic species of poison frog from the Bolivian Yungas (Anura: Dcndrohati dae: E~?ipcdohutes)

Ein neuer kkrytischer Giftfrosch au's den bolivianischen Yungas (Ai~ura: Dendrobati dac: EpipednRu~~s)

Wir hrschrelben einen neuen Ciftfrosch dcr Ciattung Epspt,rkohufb~t.r aus den holi~ianischen Yungas Fr ahnclt E hnlmelr (B<IIJI t7-\(,I R , 1883) und E. p r r m (BLBRON nn ISCHUDI, 18381, untcrschcidct sich won ihncn abcr hinsichtlich des 1 arbmustcrs, Anzeigerul~ und in der Ra\mdblillye des I hC rRNA Gens

ABSTRACT

We describe a new poison frog oftf~c gcnuq Ep~prdr,hr~te\ him ihe Bolivian Yunga~ Ti is c ~ m ~ l a r 10 E hah- neia ( B o u ~ t ~ c ~ t a . 18x3) and E pzrtur (RIRRO\ in T s c ~ r nI. 1838) but differs from these in color pattern. adivcrt~sc- m a t call and base conlposltlon of thc I hS rRNh gene

KEY WORDS

Amphibia: Anura: Dendrohatldae: Ep~pedohate~yungicoIu spec. nol:; taxonomy; systcmitics; b~owoust~cs; RVA sequencing; Bollvia, Ncotmpics

INTRODUCTION

In recent years, numerous studies on systematics of neotropical poison frogs (Dendrobatidae) have become available, This incIudes higher taxa phylogeny (e.g. V ~ N C E S ct al. 2003) as well as poison frog taxonomy at Ihc spccies level (e.g. HADDAD & MKTINS 1994). Both incruased efforts in the field and a broader methodical away allowed for new insights.

Ncvcrtheless, the taxonomy of the cis-Andcan members of Epipedobafrs MYERS, 1087 remains poorly understood. Several of the recognised species havc been associated with E. picaus (BIBKOU in Tscrrunr, 1838). In contrast to the previous ticw (c.g. SILVERSTUNT; 1976), K. ppicfus is not a wide-ranged variable species. Actually, several, in part cryptic species,

' more or less resembling E. piutus, can be distinguished when life colors, tadpole morphology, bioacoustics andlor molecular geiletics are studicd. At present, the follow- ing species (at least phcnctically) related to E. ;.ictus are widely acccptcd (H~rlo4n &

MARTINS 1994; LOTTERS et al. 1997; Cim- z.4r.c~ et al. 1999; FROST 2004): (1) Epi- pedohate.~picm,~ sensu stricto from lowland la sub-Andean Bolivia and adjaucnt Brazil, (2) E. bulivian~~s (BOULEN~~LI~ , 1902) lrom sub-Andean Bolivia, (3) E. hruccatus ( S T B W D A C ~ E R , 1864) and (4) E. Juvopic- tus (A. L ~ r z , 1925) from the central-east- ern Brazilian highlands and south-eastern Bolivia. (5) E. huhneli (BOI,LLNC;~K, 1 883) froin the Guianas, (he Amazon basin and the adjacent Andean versant, (6) E. nrbri- ventris Lbrrn~s & DEBOLD & HENLE & GI.AW & KNELLER, 1997 from sub-Andean Peru. In addition to tlrese taxa, there is evi- dence for the existence of' somc unnamcd species (e.g. M D D ~ & MARTPIS 1994; VF.NCFS et al. 2003; own unpubl. data), while the status of a form originally named E. picfu.~ ~uaymen.~is (HEATM'OLL & SO- LAKO & HEATA'OLE, 1965) froin northern South America remains lo be resolved (cl: SILVERSTONE 1 976).

Fig. I . Map of Bolivia with political unils and !mown distribut~ons of Epa~wr!ohatt.r species (from data In GONZ~LES ct nl. 1999 and those in the present popcr):

w - E bolrvrunw. ;p - E. J!av(~picfrrs, A - E. hahnuli. 0 - E pautz<r and * - E. vwngicolu spec. nov.

Ahh. 1 : Karte von Bolivien mil politischen Einheiren lind dcn hekannten Vzrbrzitungw von .E>tpedohufr,s-Ar~m (nach Daten \,on C;O~%AI.ES el al. 1999 und aus der vorlicgcndcn Arbeil):

w - E, hoolil:iunwss, -41 - E..fiuvopicfu.~, L - E. hahncli, - E. piuwds and * - E y~n~qieof(~ SPCC. no^.

The purpose of this paper is to de- to E. piclacs and E. hahneli. Like most other scribe a new spccies of Epiyadobates li.orn E~~ipedobuies fiom Bolivia, it is only known sub-Andean Bolivia, i.e, the humid montanc from a relatively small gcagraphic range forest zone called Yungas, which is similar (fig. 1 ) .

MATERlALS AND METHODS

Specimens examined by the authors les "Noel Kempf'Mercado"', Santa Cruz de are deposited at RM (British Museuin, Lon- la Sierra). NMW (Naturhistorisches Mu- r don). UBF (Colccci6n Roliviana de Fauna, seum, Vienna), ZFMK (Zuologisches For- La Paz), NKA (Musco de Ciencias Natura- schungsmuseum Alexander Kocnig, Bonn)

N w L@ipedoErat~~ h i n Roll vin 117

and ZMB (Zoologisches Museum der Humboldt-Universitdt, Bcrlin) and include:

, Epipedohraars holivianus ( 5 specimens): BoEivia: BM 1947.2.13.84-90 (lecto- and paralectotype), San Carlos, Dcpartamento La Paz; BM 1947.2.1 3.91 (paralectotype), San Emesto, Departamcnlo La Paz; CBF 390 1, km 30, Caranavi-Yucumo road, Departamcnto La Paz; NKA 3707, Serrania Bcu, Departamento La Paz; Epipedohrate.~ ,fla~~opicrus (4 specimens): Bolivia: ZFMK 77442-44, Serrania dc Santiago, Departa- mento Santa Cruz; Brazil: BM 1988.144, Proximo ao Reciro ZC Correia-fa7enda Sallo, Estado Minas Gerais; F~~ipedohaaes huhneii (10 specimens): Bolivia: ZFMK 66809- 10, Cobija, Dcpartamento Pando; Peru: RM 1947.2.1 5 . 14-20 (lecto- and paralectotypcs), Yurimaguas, Departamento Lorcto; ZFMK 40742, Tarapoto, Departa- mento San Martin; Epipedo bates pictu.~ (5 1 specimens): Bolivia: ZFMK 72 153, Esta- cion Hiol6gica del Beni, Departamento del Bcni; ZFMK 66962-63, 66984, Parjacti- Cochabamba road, Departamento Cocha- bamba; La Paz; NJSA 1190, 1539, 1400, 15 10, Buenavista, Dcpartamento Santa Cruz; NKA 2 102-08, Concesion Forestal Oquiriquia, R i o San Martin, Departamento Santa C r u ~ ; ZFMK 66855-59, Parquc Nacional Amborh, Campamento Mataracu, Departamento Santa Cruz; NKA 1028- 1032, Proyecto Rios Blanco y Negro, Departamcnto Sanb Cmz; NKA 59-60, 98- 100, Pucrlo Almackn, Departamento Santa Cruz; NKA 166, Puerto Kico, Departamenlo S ~ L T Cmz; -NKA 686, 692, 703-708, 71 0,

776-84, San Ramhn, Departamento Santa Cruz; Brazil: NMW 19190: 3 (lectotypc of E. eucnemis), Rio MamorC, Estado Ron- dhnia; Epipedobute,~ pictus payanenszs (2 specimens): Guiana: ZMB 3001 9, Meamu Mouth; Ep+edohaies mbriventris (59 spec- imens): Pcru: ZFMK 30037-52, 37892-907, 39854-69, 30053, 37908-1 1 , 39870,64838- 42 (holo- and paratypes), Cordillera Azul. Depmmento Ucayali.

The availablc specimens of the new species arc adul t males which were ob- scrvcd calling in the field. Thcy were pre- served in 70 % ethanol. The holotype was photographed in life hd'ore preservation. For genetic analysis, tissue of one toe o f thc holotype of thc new and other species was stored in 98 % ethanol. The schcme of the description of the new specics Ibllows LB.I.ERS et al. ( 1 997). Measurements were taken with dial callipers to the nearest 0.1 mm; snout-vml-length is abbreviated SVL.

Bel'ore collecting the holotype, its advertisement calls were rccorded in the field with a Sonym WM D6C walkman, a Sennheisefi Me 66 dircclional microphone on metal cassetie tapes. Digital isation and analysis was performed with SyntrilliumR CoolEdit 2000$~ at FFT (Fast Fourier Trans- h-rnation) 1028: a spectrogram was elabo- rated at FPT 256. Terminology of calls fol- lows HEYER ct al. (1990).

Genetic analyses, focusing on the sequence of a 491 bp section of the rnito- chondrial 16s rKNA gene, followed the methods applied by V ~ u c t s et al. (2003).

DESCRIPTION

Epipedohates yupfgicola spec. nov. (ligs. 2,3)

Holotype: CBF3900(fieldnumbcr SR 89-31, adult male, km 10 on road liom Caranavi to Yolosa (15"53'17" S, 47'33' 09"' W, ca. 600 m above sea level), Yungas dc La Paz, Provincia Caranavi, Departa- mento La Paz, Bolivia, 2 October 1999, S. REICHLE leg.

Paratype: NKA 7908, adult male, same locality data as holotype, 29 October 2004, A. JOHN & S. REICHLE leg.

D i a g n o s i s : A spccies related to Epi- pedobarss pictus and allies (cf. SILVERSTON F 1976; HAD~IAIJ & M ~ T I N S 1994) with (1) adult male SVL ca. 22.8-23.5 rnm; (2) dor- sal skin slightly granular; ( 3 ) Finger I > Finger I1 when adprcsscd; (4) toes and fin- gers lacking webbing; (53 maxillary teeth present; (6) tympanum visible; (7) in lifc black with irregular grey dorsal flecks, yel- lowish cream labial line and dorsolateral

Fig. 2. Holorype af E w m yumgicoh spec. no. in life (WF 3W). Abb. 2. Holotrpug van Ep@&b&s yrrrgicoia M)V. im Lab (CBF 39W).

Ncw Epip~llohate s from Uolivitl 119

line from tip of snout via nostril and eye to groin, small red signal spots in axillary, thigh, lower femoral and calf'rcgions, venter dark grey with bluish grey marbling,

' extremities bronze with irregular dark brown stipples, iris black with goldcn stip- ples; (8) advcrtiscmcnt call exisling of rap- idly rcpcalcd notes (each of two pulses) last- ing ca. 3 1-34 rns at a dominant frequency > 3550 Hz and < 3750 Hz; (9) sequence of a 491 bp section of the mitochondria1 16s rRNA gene as listed below.

Thc new species is most similar to E. pictus and E. hahneli (see Comparisons).

Descr ipt ion o f holotype: Body slender, head narrower than body; snout in dorsal and lateral vicws rounded; maxillary teeth prescnt, vomerine teeth absent; choanac rounded; tongue twice as long as wide, free for about half its length; vocal slits and median subgular vocal sac present; nares slightly protuberant, almost not visi- ble from dorsal; canthus rostralis convex Ikom tip of snout to nostril. straight from nostril to eye; loreal region vertical; hori- zontal eye diameter larger than distancc from nostril to anterior corner of eye; lym- panum visible; skin slightly granular, most prominent on dorsum: Cool webbing absent; rclative length of toes: 1 < 11 < V i 111 < 1V; metatarsal tubercles well developed, round- ed. about the same size; rest of solc smooth; well developed subarticular tubercles at joints of all phalanges of foot; hand web- bing absent; relative length of fingers: 1V < I1 < 1 < 111, Finger 1 > Finger IT when adpressed; metacarpal tubercles well devcl- oped, rounded, outer about twice the size of inner; rest of palm smooth; well developed subarticular tubercles at joints of all pha- langes of hand.

In preservative. the holotype is dor- sally black, greyish on limbs, with white labial line from bclow nostril to arm inser- tion and whitc dorsolateral line from tip of snout above nostril and eye to groin; small pink signal spots in axillary, thigh, lower femoral and calf regions; ventrally grcyish, with white marbling on posterior belly and . limbs. Tn lifc, this specimen was black with irregular grey flecks on dorsum; labial and dorsolateral lines were yellowish cream; signal spots were red; ventrally it was dark

1

grey with bluish grey marbling; cxtremilies

werc bronze with irregular dark brown stip- ples; the iris was black with golden stip- ples.

Descr ipt ion o f pa ra type : The paratypc agrees well with the above descrip- tion of the holotype.

Measurements (in rnrn) and ra t ios of holotype (andparatype) : SVL,23.5 (22.8); hcad lcngth from tip of snout to angEc of jaws, 6.2 (5.6); head width at angles of jaws, 6.9 (6.8); interorbital dis- tance, 2.5 (2.4); distance between narcs, 1.7 (1.5); distance from nostril to antmior cor- ner of eye, 1.8 ( I .6): horizontal eye diame- ter, 3.3 (3.2); tibia length, 11.7 ( 1 1 . I) ; foot length from tip of longest toe to proximal outer metatarsal tubercle, 1 0.9 (1 0.1 ); hand length from tip of longest finger to proximal outer metacarpal tubcrcle, 6.3 (5.1); head IengthISVL, 0.26 (0.25); head width/SVL, 0.29 (0.30); tibia lengthlSVL, 0.50 (0.44); distance from nostril to eyeleye diamctcr, 0.55 ( O S O ) .

n i s t r i b u t i n n and ecology: Epi- pedohates ywngir,uiu is only known from two localities within a relatively small gco- graphic range in the Yungas dc La Paz, Bolivia (fig. 1). Apart from the type locali- ty, it was found and rccorded by JOHN (2003) at km 22 on the Caranavi-Yucurno road, Yungas de La Paz, Provincia Caranavi, Departamento La Paz (ca. 1,000 rn above sea level). The entire area is composed of humid mountain rain forest. At the type locality, howcver, human influence has convcrtcd allnost all forest to small scale banana plantations.

All specimens observed by the authors and JOHN (2003) were males, calling from exposed positions on he ground during the day. From thc same general area, E. bdi- vianus and E. picttls have been reported (scc GOKZALES et al. 1999). The lattcr was Ibund syntopically with E. jwngicola along the Caranavi-Yucumo road.

Advertiscment call: Vocalizations pro- duccd by the holotype of Epipedohate.~ yungico!a (fig. 4) consisted of numemus rapidly repeated notes. Twenty such notes were analysed (recordcd al a lemperature of 27.X°C). Thcsc were each composed of two pulses, showing a slight upward frequency swccp from around 3550 to 3740 H L . Dominant frequency was at about 3590-

5 Oms

Pig. 4. Oscillogrnm (abovc) and su~ind spectrogram (below) of advertivement call of Eplprdohcrte.~ j~uvgicoIu spec. UOV. (CRF 3900). Temperature during recording 27.X°C.

4hb. 4: Osj.rllugramm (oben) und Klungspcktrop~nrn (untm) des Anzeigerufes %on Fprpehhafcs jufigbut~la spec. no\ (CHI' 3900). Tempcralur wd~rcnd dcr Aufnahme 27,8OC.

57 19 Hz (mean 3660 * 49 Hz). Note length ranged frnm 3 1 to 34 ms (mean 32.4 * 0.9 ins), while note repetition ralc was 4.5-5.0 notcslsecond (mean 4.7 * 0.16 notes/sec- ond). The inter-note interval was 163-221 ms (mean 180.7 * 1 1.9 ms). The data coin- cide with those given by J U ~ I N (2003). The advertisement call o f E. ,vecngicola is not allocahle to any of the known calls defined for dcndrobatid frogs (cf. LOT~ERS et al. 2003).

Molecular genetics: The sequence of thc 491 bp fragment of the mitochondria1 DNA of the 16s rRNA gene of the holotype of Epipedobates yunxicola (GenHank Ac- cession Numbcr AY263239; I~ttp://www. ncbi.nlm.nih.gov/) is:

1 cccagtgact ttgttcaacg gccgcggtat cctaaccgtg cgaaggtagl: gtaatcactt

6 I gttctttaaa tgaggactag tatgaatggc cccacgaggg clgcactgtc tccmttct

12 1 aatcaatgaa actaatctcc ccglgaagaa gcgggaataa ccclataaga cgagaagacc

18 1 ctatggagct ttaaacaatt gaaacatttg cltttttctt gacctcttcc gagctcttta

24 1 tcttacttaa gcattcttat ttctagtttt aggttggggt gaccacggag caaaacttaa

301 cctccatgaa gaaatgaata tatttttaag ccacaaacta cccmtaag catcaacaaa

361 ttgaccttca ttgacccaat atattgalca acgaaccaag itaccctagg gataacagcg

421 caatctactt caagagctca htcgacaag . taggtttdcg acctcgatgt tggatcaggg

48 I tatcctagtg g

New Epprdobatt7s from Bollvia 121

Kemarks : Aftcr examination of the Iccqotype, we agree with previous authors

, (e.g. ITADDAD & MARTINS 19%) that rpi- pedo bates eucnemb (S~INDACHNEK, 1 S G 4 ) from Hrazil as a junior synonym of E. pic- fw.7.

Wc have not been able lo study the type material of E. yiclus flayanensis. From referable specimens from Guiana and according to the original description (HEAT-

WOLE ct al. 19651, this form is similar to E. pictu.s (scc Comparisons), although we sus- pect it to represent a distinct species rather than a disjunct northern population of the Ialtcr.

Elymolrrgy: The specific name means "inhabitant of the yungas" and refers to the eco-geographic region in which the new species occurs.

Fpipdohates yunzicola is morpho- logically similar to E. picbus and E. hahneli, both known from Bolivia (fig. 1). Tn adult male SVL, it matches both species, includ- ing large individuals of E. huhraeii. This lat- ter poison frog apparently comprises a com- plcx of species (cf. HAVDAU & MARTINS 1994; L~TTERS et al. 1997). Populations examined by HAIIDAT) & MARTIKS (1994) and us, including Bolivian and topotypic spccimens from Peru (see Material and Methods), are similar to thc new species in having (in life) black dorsal surl'aces with irregular grcy flecks and yellowish crcam lines. Howcvcr, they lack teeth ( v m u s present in E. yun~ir.o/a), have small yellow- ish orange (versus red in E. yztngicolu) sig- nal spots in axiIlary, thigh and calf regions excluding (versus including in E. yzsngicola) lower femoral region.

Epiprdubabe.~ pictu,~ (sensu HADDAD & MARTINS I9Y4), including specimens found by us near the type localily, shams the pres- ence or tceth and (in life) reddish signal spots with the new species. However, E. p ic tx~ has bright yellow (versus yellowish cream in E. yungicolu) dorsolateral lines and always lacks dorsal spots (versus grey dorsaI spots presenl in IT. )rungico/a). The red thigh signal is usual1 y larger in E. piths than in E. ywngi- cola, extending onto the upper femur (cf. HADDAD & MAWI'INS 1994: 292). A lower fcmoral signal spot is absent in E. picaus (ver-

, sus prcsent in E. jvingiuola). Characters of the colour pattern mentioned to distinguish the new species and I!: picaus are also amli- cable to distinguisll it and the form guava- ' nensis. Moreover, accordi ng to SILVERSTONE (1976, by implication), thc latter lacks teeth (versus prcsen t in E. pngicola).

Epipedohafc~ boliviunus from the same general arca as the new species (cf. GONZALES et a]. 1999) lacks teeth (versus present in E. yzlngicoda), has ye1 Iowish (ver- sus red in E. yuyuiuuia) axillary and thigh signal spot.% and lacks lower femoral and calf spots (versus prcsent in R yungicola).

Other Epipeddnbates resetnbling E. pictiw (see Introduction) diil'cr from E. ylmgicodn by the presence of yellowish dor- sal markings (E. E,braccarztls, E. f7uvopicruLs) or reddish vcntcr (E. mbriventris) (HADDAD & MARTINS I 994; L ~ T ~ E R S et al. 1997).

The advertisemrml call of E, yungicolu is similar to those ofE. piuhs and E. hahneli (cC data in L ~ T T ~ . R ~ & KNELLER 2000). However, vocalisations in these two poison frogs are unpulsed (vcrsus pulsed in E. ytingicolu) and their lu\vcr darn inant fre- quency range is higher (= 3750 Hz) than the upper range in E. ymgicolu (< 3750 Hz). The nok lcngth in E. yungicoiu is clearly longer than in Amazonian (including Bolivian) populations or clearly shorter than in a sub-Andean population (i.c. from Peru) o f tlic E. hahn~l i complex (temperature of E. hahneli recordings varied from 24.0-2X.0°C; cf. Lontns & KNELI.T.R 2000). In contrast, note length and number of notes per second overlap with the lower rangc known in E. pictus. Vocalization parameters in the latter mcnlioned species show great variation (cE HADDAD & MARTIYS 1994; DE LA R~VA et RI. 1996$, probably rclated to its large geo- graphic range (see fig. I ) or, more likely, the presence of unidentified additional cryptic bpccics. Recordings of E. piuh.s Imm ncar its type locality exhibit a note Ienglh of' abou~ 40-50 ms with 2.8-2.9 notes per sec- ond (according to H.4rlrl~n & MARTINS 1994

a

Table I (Ih~s and opposite P,~gc): Summary nl'thc uncorrected p-d~stanccs fur the 1 hS rRNA dilta set of den- drobatid frogs; value^ c I .X % are in bold (scc tcxt). For ongin uf specimens and GcnBank n u m k r ~ see V r ~ c r s et at. (2003).

Tab. 1 (diese und gcgcnuherliegende fcik): Zusarnrncnfas~cnde Darstellung i i k r die unkorr~p~crtcn p- ' Distanzcn dcr I hS rRNA Datcn von Ucndrobarldcn; Wer1.e 5 1 ,R% in Fettschift (slchc Text). Zur Aerki~nfi und dm GenRwk-Numnem dcr Exmplnre .;ithe C'EVCES et al. (2003).

Allobntrs cf. firnoruli,~ (B01 I .rNGtR, 1883) C[vpfop/~-i!ohafe.i azaircivrntris (Kvr.1.1 FR & ~IENLE, 1985) Dendmhuftlv auratus (GIR~RD, 1855) DendmBab~s imitator SCHL!LI'~:, 1986 Epiprckjbuf~~ aan f hnnyi (NOBLE, 192 1) Epiprdobdes Ps)casIt.si ( M F B - I N , 1941) Epipcrlohate.~ hzlinptr (JL'NGFER, 19x11) E~~~wcfohatesJ~n~opicfus .Epipedohrr~e.c hrzhnulz (T) Ej~ipedo bores huhneli (11) Epzp~dobotrs hnhneIi (11 I) Epipedohatcs pnmlus (BoLL~N~;~R. 1882) Lpip~do/)ute.t pirfz<,~ (1) Epiprdobcltes piclu,v ( I I) Epipedohafm silv~rstonri (MYEKS & DAI.Y, 1978) i7pipcdobn fes cf. triir.ok(~r ( B o u ~ .PNGER, 1899) Ljlipeciohatr.~ trivi:ilfuft~s (SPIX, 1824) Epip<doburr,s yupagiuola spec. nov.

[recorded at 22.5"CJ or own unpublished data [recorded at 23.0°C and 26.5OC]), thus di l'l'ering from E. pngicoln.

Wc used available 16s rRNA se- quences 01' species of the genus Ep*edo- haaes (cf. V~N( .ES et al. 2003) to compare them with the sequcnce gained from the holotype of E. ,win,yico!cr. The complete data matrix (1 6 samples; 591 bp including gaps; not shown) included the in-group species as listed as Epipedobutes in table 1. The matrix for ~ h c uncorrected p-distances for all nucleotide sites is also presented in this table.

Whcn applying both Most Parsimony and Baycsian analyses (cf. VLNCES et al.

2003) to the resulting data sel, wc received ovcrall coinciding topologies, strongly sup- porting thc same terminal clades (trees nnt shown). All analyses suggest E. jrir?~iuolu to be part of a wcll supported clade of cis- Andean species of Ep@ednhates.

Kegarding the uncorrected p-distances (bblc I), it becomes clear thal the cis- Andean species of Epipedohutes [i.e. all except E. unahonyi (No131.r~ 1 92 1 ) and E. cf.

A

rricolor (BOULEKGER, 1899)l are rather close to each other with comparatively low genetic variation. Epiprdohates yungicnla is separated from E. picrris by a genctic dif- ;

ference o f 1.8 % and E. hahncli by at least 2 % (table 1 j. Thcse values support its spe-

New Epiprdobnt~.~ from Bolivia 123

Table 1 (continued). Summary of the uncorrected p-diqances for the 1 6 s rRNA dab set of dmdrobatid frogs: values s I .R % are in bold (see text) I'or origin of spcclmcnf and GenBank numb- ree V r ~ c r s el at. (20031.

I Tab. 1 (I'ortsctzung). Zusamrnenfarsende Dilrsiellung iiher die unkorrigierten p-Distanzen dcr 16s rRNA Daten icon Dendrubatiden: Werte 5 1,8% in Pcttschrifi (sichc Text). Zur Herkui~li und den GenRank-Nummem der Exemplare s ~ c h c V t h ~ t s ct al. (2003).

cific distinctness, since the levcl of genetic difference is even lower among other Epipednhote~ that are well identifiable as distinct species on the basis of morphology and bioacoustics (c.g. SILVERSTO~T. 1 976; LOTTERS & KNELLER 2000), as Tor instance among E. biiinguis JIJNGFE~~, 1989 and E. trivittat~s (SPIX, 1824) or E. bussleri (ME- LIN, 1941) and E. fEmopictus (see table 1).

Comparing all available p-distances. 1 E. ,vungic~lu is most C ~ O S C to Eq flavopicfus

and LJ. wivitlaaaa (1.2 and 1.4 %; table 1). On the basis of morphology (differcnccs in SVL, color pattern, presencelabsence of teeth etc.) and advertisement call parame- ters (e.g. notc Icngth), the new species is

clearly distinct from both (cf. SILVERSTONE 1976; HAIIDAT) & MARTIUS 1994; LBTTFRS & KNELLEI~ 2000). Thus, the possibility that E. yungict~ia may be conspecific with anc of these has to be negated. In addition, a direct comparison of the sequences of E. Juvopic- [us, B, rrivirtufus and E. yungico/u, includ- ing all sequenced base pairs (an unambigu- ous alignment of the highly variable loop regions of the 16s rRNA gene is only possi- ble in very closely related species - these regions [77 bp] were consequently not used for the overall phylogeny), raises the true genetic diffcrcnces bctween them to about 3.1 % - 4.6 %.

Permits to conduct field work and to export cussionr abvut bioacoustlcs with our colleague JOrn matcrial were issued by CBF and NKA; special thnnks KOHLER (ZFMK) DVA sequcnccs wed ]>ere were ' i o 1,ucindo t i o n ~ ~ ~ c s and Claudia C O R T ~ ~ . Tlie IHP made available through running cooperat~orl with Programnit of the Luropcan Community kindly made Juachim Kos~:c I I (Universib of Mainz) and Miguel access to EM for ihe senior author through the SYS- Vr.uorr (University of Ams~erdilm). M i ~ u c l also pro- RESOURCE Major Research Infrastructure of the vidcd valrrahle co~nrnents or1 an earlier v m i w of tllc Natural History Museum (BM, London); he is cspe- manuscript. cially pratcful to Rarry T. CLARKE. We had fruitful dis-

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DATE OF SUBM ISSTON: Decembcr 2 1 HI, 2004 Correspollding editor: Hrinz Grillitsch

AUTHORS: Stefan LbTre~4, Institute of Zoology, Fcology L)cpartmeni, University of Mnin~, Saarstrde - 21, 55099 Mainz, Cierrnimy < loeCtcrs(@uni-nminz.de r; Andreas SCHMITZ, Muskurn d'Histuire Narurcllc, Deparl- mcnt of Herpetology and Ichihyology, C.P. 6434, 121 1 Geneva h, Srvit~erland; Stcffcn R t l r ~ 1I.r- Research Institute and Zoolo~iml Museum Alcxander Koenip, H w p e t o l o ~ Department, Adennauerallee 160, 531 13 Boiln, Germany.