F IG. l.

-Ho loty pe of Dendrobates sirensis, NHMW 31892, SVL

16.8 mm.

HERPETOLOGICA
VOL. 47
Herpetologica , 47(1), 1991, 1-5 1991 by The Herpetologists League, Inc.

MARCH 1991

NO.1

A NEW SPECIES OF POISON-DART FROG (ANURA: DENDROBATIDAE) FROM THE SERRANiA DE SIRA, PERU
MANFRED AICHINGER

Zoologisches lnstitut , Univ ersitiit Wien , Althanstraj3e 14 , A -1090 Wien, Austria

ABSTRACT: A new species of poison-dart frog, Dendrobates sirens is, is described from the Serrania de Sira , an isolated mountain range in east-central Peru, Departamento Huanuco, at an elevation from 750-1560 m . Adult specimens have a snout-vent length of 15-17 mm. The species is easily diagnosed from all dendrobatids by the uniform red coloration of the dorsum and the turquoise-green limbs.

Key words:

Anura; Dendrobatidae; Dendrobates sirensis; Peru

THE Serrania de Sira is an isolated mountain range in the upper Amazon basin between the Rio Pachitea and the Rio Ucayali in east-central Peru. The highest elevations are about 2400-2500 m (Terborgh and Weske, 1975) . For an approximate location of this mountain range, see Duellman and Toft (1979) . Our expedition followed the same route to the Serrania de Sira as described by Terborgh and Weske (1975) . Duellman and Toft (1979) first reported on a collection of anurans from this region, including the dendrobatids Col ostethus marchesianus, Phyllobates (now Epipedobates) petersi, and P. (now Epipedobates) pictus. During a herpetological survey from September 1987 to October 1988, five specimens of the poison-dart frog were captured. This new species was collected first in the Serrania de Sira in 1976 by Werner Hanagarth.

num was judged by dissection. Specimens are deposited in Naturhistorisches Museum Wien (NHMW) and Museo de Historia Natural "Javier Prado" in Lima (MHNJP). One paratype is in the collection of Staatliches Museum fiir Naturkunde Stuttgart (SMNS) . Museum abbreviations follow Frost (1985).

METHODS

Nineteen different measurements (to the nearest 0.1 mm) were made following Myers (1982). The condition of the om oster1

Dendrobates sirensis sp. nov . Fig. I-Frontispiece Holotype.-NHMW 31892 (Fig . 1), an adult female, collected on a horizontal tree 1.2 m above a stream in the Serrania de Sira, Rio Llullapichis drainage, 750 m, Departamento Huanuco, Peru (928' S, 7447' W), on 10 September 1987, at 0910 h by M. Aichinger. Paratypes.-MHNJP 11518 and MHNJP 11519, adult males, same location as holotype, 780 m, on the ground , on 20 March 1988, at 1530 h and at 1250 m, in September 1988, both by B. Wallnofer , SMNS 07092, adult male, same location , at 1560 m, on the ground , on 23 January 1976, by W. Hanagarth.

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Diagnosis.-A very small dendrobatid having an adult snout-vent length (SVL) of about 15-17 mm (three males 14.7-15.4 mm , 1 female 16.8 mm). Skin on back slightly granular, moderately granular on venter, and smooth on limbs; dorsum uniform dark red, arms and legs turquoisegreen, venter turquoise-green with a rectangular red patch on the belly, chin red; neither spots nor reticulation present dorsally; teeth absent; first finger much shorter than second; finger discs very conspicuous, enlarged to at least twice finger width; omosternum absent ; tympanum visibly concealed posterodorsally. D. sirensis is tentatively assigned to the quinquevittatus group as characterized by Schulte (1981) , Myers (1982), and Zimmermann and Zimmermann (1988). It differs from all members of this group (D. captivus, D . fantasticus, D . imitator, D . mysteriosus, D. quinquevittatus, D . reticulatus, D. vanzolinii, D. variabilis) by its uniform red dorsum, its unreticulated feet and venter, and its turquoise-green limbs. Dendrobates sirensis may be confused with other small poison-dart frogs of the genus Dendrobates or Minyobates, like D. granuliferus, D . pumilio, D . speciosus, M . opisthomelas, and M. steyermarki, which all have a bright red dorsal coloration. Dendrobates sirensis differs from D . speciosus, M. opisthomelas, and M. steyermarki in having turquoise-green limbs, from D . granuliferus in having only a slightly granular skin, and from D. pumilio in having a rectangular red patch on the turquoise-green belly . Description.-Size small, SVL of adults 14.7-16.8 mm; (three males 14.7-15.4 mm; one female 16.8 mm): head narrower than body; in all specimens, greatest head width 32-34% of SVL. Males lacking vocal slits and vocal sacs. Teeth absent, omosternum absent. Skin on the back is slightly granular in life, moderately granular on venter, and smooth on limbs. Snout subtruncate in dorsal aspect and sloping in lateral aspect (Fig. 2). Naris situated near tip of the snout and directed laterally; both nares visible from front and from below but not from above. Canthus rostralis rounded; loreal region

more or less flat. Interorbital area wider than upper eyelid. Center of naris to edge of eye/eye length = 0.75-0.86. Tympanum seemingly vertically elliptical, but concealed posterodorsally; its approximated diameter 56-64% of the eye length. Hand relatively large (Fig. 3); its length (proximal edge of metacarpal tubercle to tip of longest finger) 78-87% of greatest head width; relative length of adpressed fingers 3 > 4 > 2 > 1; adpressed first finger roughly three-fourths the length of second finger. Discs conspicuously expanded on all but first finger . Disc of third finger 22% times wider than distal end of adjacent phalanx; widths of discs without apparent sexual dimorphism. A circular metacarpal tubercle at base of palm, a smaller inner metacarpal tubercle at base of first finger , and one or two usually prominent subarticular tubercles on fingers (one each on fingers 1, 2, 4, two on finger 3). Distal subarticular tubercle on finger 3 small and inconspicuous, but all other tubercles on hand prominently raised; surfaces of discs rounded; lack of pigment contributing to prominent appearance of tubercles. Hind limbs of moderate length, with heel of adpressed limb reaching tympanum. Tibia 36-39% of SVL. Relative length of adpressed toes 4 > 5 > 3 > 2 > 1; first toe short, reaching bottom of subarticular tubercle on base of second toe, and with unexpanded disc; other toes with discs distinctly expanded (but smaller than finger discs) . Moderate-sized inner and small outer metatarsal tubercles, somewhat protuberant but with rounded surfaces. One to three subarticular tubercles (slightly protuberant tubercles, one each on toes 1, 2, two each on 3, 5; nonprotuberant tubercles, one each on toes 3, 5, three on toe 3). Hands and feet lack webbing, supernumerary tubercles, or a lateral fringe (Fig . 3).

Measurements (in mm) of holotype.The holotype is a female containing two mature ovarian eggs in both ovaries. Measurements of paratypes are given in Table l. Length from snout-vent 16.8; tibia length from heel to fold of skin on knee 6.4; greatest head width (between angles of jaws) 5.4; head width at level of eyes

March 1991]

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o
I

5m m

F IG . 3.-Palma r and plantar surfaces of right hand and foot of holot ype of Dendroba tes sire nsis , N H MW 3 1892.

5mm

FI G. 2.- Dorsal and lat er al view of the hea d of th e holotype of Dendroba tes sirensis, N HMW 3 1892.

5.3; approxim ate width of interorbital area 2.1; head length from tip of snout to angle of jaw 4.0 ; tip of snout to center of naris 0.8; center of naris to anterior corner of eye 1.5; distance between center of nares 2.0; eye length from anterior to posterior corner 1.8; approximated horizontal diameter of tympanum because tympanum posteriorl y indistinct 1.0 ; corner of mouth to lower edge of tympanic rin g 0.6. Length from pr oximal edge of large m edial palmar tubercle to tip of lon gest (third) finger 4.7; width of third finger di sc 1.1 ; width of third finger (pe n ultimate phalanx) below disc 0 .5. Width of di sc of third toe 0.7 ; width of third toe proximal to d isc 0 .5; width of disc of fourth toe 1.0; width of fourth toe (penultimate phalanx) pr oxim al to di sc 0.5.

Coloration in life.-Head and dorsum dark red , no di stinct spots or stripes ; tympanum dark brown ; iris black . Arms and legs turquoise-gr een ; turquoise colora tion lighter on th e ante rodorsal side of pro xim al half of upper a rm and in gro in anteroventrally on proxim al part of th igh extend ing onto th e bod y. Vente r turquoise-gr een with a rect an gular red pat ch on th e bell y (a ppr oxim at el y 5 x 3 mrn ): chin dark re d ; ve nter below rectangular pat ch and th e ve ntral sur faces of th e legs d ark turquoise . Surfaces of palms and soles black . Parat ype MH NJP 11519 had an indistinct brown spot in th e interorbital area. One individual which esca pe d from captivit y (sex unknown) showed a di spositi on of a rather wide , ill-defin ed re d lat er al st ripe exte nd ing horizontally fr om th e groin one -third of the dorsum to th e head (M . H enzl , personal com m unica tion) . A color tr ansparency of this spe ci men sug gests that this stripe is not homologou s with th e incomplet e oblique lat eral lin e that ch ar acterizes m any d endrobatid s. In preservati ve , th e red co loratio n a ppears pale gray; turquoise changes to blac k. The red co lora tio n of par at ype MHNJP 11519 cha nged to black. Parat ype SMNS 07092 becam e pale brown but th e coloration in life was as described above (W. H anagarth , per sonal com m un ica tion). Life history notes. -The holot ype con tained two we ll-develope d eggs in both ovaries (d iam eter 2.0, 2.1 mrn) as we ll as

HERPETOLOGICA T ABLE I.-Measurements (in mm) from paratypes.

MHNJP 11518 Male MH NJP 11519 Male

[Vol . 47 , No.1

Characteristic

SMNS 7092 Male

Length from snout to vent Tibia length from heel to fold of skin on knee Tibia/SVL Greatest head width (between angles of jaws) Head width/SVL Head width at level of eyes Width of interorbital area Head length from tip of snout to angle of jaw Tip of snout to center of naris Center of naris to anterior corner of eye Distance between center of nares Eye length from anterior to posterior corner Naris-edge of eye/eye length Horizontal diameter of tympanum (posteriorly indistinct) Diameter tympanum/eye length Corner of mouth to lower edge tympanic ring Length from proximal edge of large medial palmar tubercle to tip of longest (3rd) finger Length hand/greatest head width Width of third finger disc Width of third finger (penultimate phalanx proximal to disc) Third finger disc /width 3rd finger Width of disc of third toe Width of third toe (penultimate phalanx) below disc Width of disc of fourth toe Width of fourth toe below disc

14.7 5 .3 36 .1% 5.0 0.3 4.8 1.6 3.0 0.7 1.2 1.7 1.6 75.0% 0.9 56.3% 0.6 3.9 78 .0% 0.9 0.4 2.3 0.6 0.4 0.8 0.4

14.8 5.8 39 .2% 4.8 0.3 4.7 1.7 3.2 0.8 1.2 1.7 1.4 85.7% 0.9 64.3% 0.6 4.0 83 .3% 0.9 0.4 2.3 0.6 0.4 0.6 0.4

15.4 6.0 39 .0% 4.7 0.3 5 .1 2.0 3.8 0.8 1.4 2.0 1.4 100 .0% 0.9 64 .3% 0.6 3.9 83 .0% 1.0 0.5 2.0 0.7 0.4 0.9 0.4

three small eggs (diameter < 1 mm). An un preserved female laid a single egg on the surface of a leaf in a plastic box a few days after capture (M. Henzl, personal communication) . Individuals are very agile, jumping continuously. The species seems to be extremely rare or secretive. During one year of investigation, only five individuals were observed (two escaped from captivity). Etymology.- The species is named after the type locality, the Serrania de Sira.
DISCUSSION

Dendrobates sirensis is known only from the Serrania de Sira in east-central Peru at an elevation from 750-1560 m and seems to be endemic in this isolated mountain range. There it lives in sympatry with D. quinquevittatus, which was found from the lowland rainforest at 260 m up to 1080 m. Based on characters that were attributed to the genus by Silverstone (1975), the new species is assigned to Dendrobates. These

characters are first finger shorter than the second, well expanded finger discs, and a bright coloration in life. Dendrobates sirensis may be highly arboreal, judging from the relatively long hand with the well developed discs. One individual showed an indication of light red lateral stripes. A female contained only two mature ovarian eggs. Considering these facts, D. sirensis is tentatively assigned to the D. quinqueoittatus group. Based on characteristic behavioral parameters, Zimmermann and Zimmermann (1988) attributed the following characters to the quinquevittatus group: the arboreal frogs live in middle to upper strata in tropical rainforest; females lay only few eggs; males carry one or two tadpoles to bromeliads; larvae feed facultatively on unfertilized eggs laid by the female. Schulte (1981), Myers (1982), and Zimmermann and Zimmermann (1988) placed the following eight species in the quinquevittatus group: D. captivus, D. fantasticus, D . imitator, D . mysteriosus, D.

March 1991]

H ER P ETO LO GI CA

quinqueoittaius, D . reticulatus, D. vanzolinii, and D . variabilis. Myers (198 2) could not docum ent a relationship of D . captivus and D . mysteriosus to th e quinquevittatus group and left th em as possible close relatives to th at group. Dendrobat es quinquevittatus is widespread throughout the Ama zon Basin , inhabiting primaril y lowland rainforest (Le scure and Bechter, 1981; Silverstone , 197 5). All othe r spe cies of the quinquevittatus group occ ur onl y in com pa ra tively small ar eas in th e western Amazonian lowl ands (Kne lle r, 1983; Myers, 1982 ; Schulte , 1986; Zimmermann and Zimmermann , 1984, 1988). Dendrobates captivus, D . imitator, and D. mysteriosus may be restricted to sin gle localities in montane rainfor est. Ex cept for D . mysteriosus, which is known only from a single spec imen (Myers, 1982 ), D . quinquevittatus lives sym patrically with all other spe cies. Like Minyobates abditus and M . ste yermarki, which Myer s and Daly (197 6) assigned to th e minutus group as d efin ed by Silverstone (1975), and M . opisthomelas, Dendrobat es sirensis might be a terminal relict of an ancestor whose range wa s fragment ed by climatic vegetational change s that occur re d in th e Andes during Quaternar y time. F urthe r stud ies on skin al kaloids, knowledge of tadpoles, and th e reproductive biology of this species will be necessar y to define th e exact ge ne ric and group relationships of D . sirensis.
R ESUM E N

d a ta. I a m in debted to A. Schliiter (Stu tt gart ) and N. Carillo (Li m a ) for the loan of specimens. Than ks are a lso du e to M. H enzl, W . Hodl, K.-H. Jungf er , A. Schliite r, a nd E . Zimmermann for ca re fu llv review in g th e m anuscript a nd to th e Direccion Ge'ner al F ores ta l y d e F auna fo r permits (06 2-87-AG-DGFFDR FF-SDFF). F ina ncial su pport was de rived from "os terreich ische r F onds zur Fiirderung d er wissensc haftlic hen Forschung " (p rojec t P6399B ) and "Bundesm ini ster iu m Iu r W issensc haft u nd Forschung, osterre ich". LI T E RATURE CITED D UELLMA:-I, W . E., AND C. A. TOFT. 1979. Anurans fro m the Serrania de Sira , Amazon ian Peru: T a xonomy an d b iogeography. H er pet ologica 35 :60-70. F HOST, D. R. 1985. Am phib ian Species of the W orl d . Alle n Pr ess an d th e Associati on of Syste matics Co llecti on s, Lawrence, Kansas. KNELLER, M. 1983. Beobachtunge n an Dendrobates fantast icus im na tiirlic hen Lebe nsraum un d im T errariu m . Herpetofauna 5 :15-1 8. LESCURE, J., Al':D R. BECHTER. 1982. Le comportement de repro duction en captivite et Ie pol ymorph isme de Den drobates quinquevit tatus Steindachner. Revu e fro Aq ua rio!. 8 :107 - 118 . MYERS, C. W . 1982. Spo tted poison frogs: D escription of three new Dendrobates from weste rn Amazon ia , a nd resurrecti on of a lost species from "Chiriq u i" . Am. Mu s. Nov it . 2721 :1-23. MYEHS, C. W ., ANDJ. W . D ALY. 1976. A ne w species of poison fr og (De n drobates) fr om Ande a n Ecu ador includin g an a na lysis of its skin toxins . O cc. Pa p. Mu s. Nat. H ist. Un iv . Kan sas 59:1-1 2. SCHULTE, R. 1981. Dendrobat es qui nqueoittat us ; iiko logie, Halt ung und Zu cht. H er pet of auna 3 :2428. - - -. 1986. Eine neu e Den drobates-Art aus Ostpe ru (Am p hib ia ; Salientia ; Den drobatida e ). Sauria 8 :11-20. SILVERSTONE, P. A. 1975 . A revision of th e poisondart fro gs of th e ge nus Dendrobat es W agler. Nat. Hi st. Mu s. Los Angeles Co . Sci. Bull . 21:1-53. T EHBORGH, J., AND J. S. W ESK E. 1975. The ro le of co mpetit ion in th e d istributi on of Andean b ird s. Ec ology 56:5 62-576. ZIMMERMANN, H. , AND E . ZIMMERMANI':. 1984. D ur ch Nachzucht er ha lten : Baumsteigerfr iische Dendrobat es quinqu evittatus und D . reticula tus. Aq u. Mag. 18 :35-41. 1988. E tho- T axon om ie und zoogeogra ph ische Ar tengruppenbild ung bei Pfeilgift fr iischen (Anur a: Den drobatidae ). Salamandra 24 :125160.

Se describe Dendrobat es siren sis, una nueva especie de batracio ven en oso de la Serrania de Sira , Departamento Huanuco , Peru , a un a elevacion de 750 a 1560 m. Es una ran a diminuta (15- 17 mm ) que se puede facil d istinguir d e otros dendrob atidos por su color acion . La coloracion dorsal del cue r po es roja y la de las extre m idades, verde turqui.
Acknowledgments.-I thank B, W allnof er for co llecting spe cim ens an d M. Henzl for obse rva tiona l

Accepted : 26 Ap ri l 1990 Associate Edito r: D avid Cannate lla