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Zeitschrift/Journal: Herpetozoa

Jahr/Year: 1999

Band/Volume: 12_3_4

Autor(en)/Author(s): Gonzales Lucindo, Lötters Stefan, Reichle Steffen

Artikel/Article: On the dendrobatid from Bolivia: rediscovery of Epipedobates bolivianus (Boulenger, 1902), first record of Colostethus brunneus (Cope, 1887) and comments on other species (Anura: Dendrobatidae). 179-186 ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at

HERPETOZOA 12 (3/4) 179 -186 Wien, 30. Dezember 1999

On the dendrobatid frogs from Bolivia: rediscovery of Epipedobates bolivianus (BOULENGER, 1902), first record of Colostethus brunneus (COPE, 1887) and comments on other species (Anura: Dendrobatidae)

Zu den Dendrobatiden Boliviens: Wiederentdeckung von Epipedobates bolivianus (BOULENGER, 1902), Erstnachweis von Colostethus brunneus (COPE, 1887) und Bemerkungen zu anderen Arten (Anura: Dendrobatidae)

LUCINDO GONZALES & STEFAN LÖTTERS & STEFFEN REICHLE

KURZFASSUNG

Epipedobates bolivianus (BOULENGER, 1902) ist ein bisher nur von der Typusserie bekannter Pfeilgiftfrosch (Dendrobatidae) vom Ostabhang der Anden in Bolivien. Kürzlich wurde die Art unweit der Typuslokalität wiederent- deckt. Die Lebendfarbung wird erstmals beschrieben. Colostethus brunneus (COPE, 1887) wird erstmals und C. mcdiarmidi REYNOLDS & FOSTER, 1992 und C. trilineatus BOULENGER, 1883 werden an neuen Stellen nachgewie- sen. Die Verbreitung und Taxonomie der Pfeilgiftfrösche Boliviens wird zusammengefaßt und diskutiert

ABSTRACT

Epipedobates bolivianus (BOULENGER, 1902) is an uncommon poison (Dendrobatidae) from the eastern versant of the Bolivian Andes. It is only known from the type series. Recently, it was rediscovered close to the type lo- cality. Life coloration is described for the first time. Colostethus brunneus (COPE, 1887) is recorded for the first time, and C. mcdiarmidi REYNOLDS & FOSTER, 1992 and C. trilineatus BOULENGER, 1883 are reported from additional localities. The distribution and of the Bolivian poison frogs is summarized and discussed.

KEY WORDS

Anura, Dendrobatidae; Epipedobates bolivianus, Colostethus brunneus, first record; Bolivia.

South American poison frogs of the San Carlos (1200 m a.s.l.), both in the De- Epipedobates MYERS, 1987 (Dendro- partamento La Paz, Bolivia (FROST 1985). batidae) are known from low to moderate Since, the species was redescribed twice elevations both east and west of the Andes. (SILVERSTONE 1976; RODRÏGUEZ & MYERS Including Phobobates ZIMMERMANN & ZIM- 1993). Both contributions were based on MERMANN, 1988 as a junior synonym of Epi- the type specimens only as fresh material of pedobates, presently about 30 species are E. bolivianus has not been available, known (GLAW & al. 1998; MYERS & al. On 17 September 1998, LG collected 1998). Most species of Epipedobates occur a specimen of E. bolivianus, SVL 23.6 mm in the upper Amazon basin including the (fig. 1A), at Territorio Comunitario y Re- Andean foothills (FROST 1985). Although, serva de la Biosfera "Pilon Lajas" (15°06' in recent years, the knowledge on certain 36" S, 67°32' 33" W; ca. 1350 m a.s.l.), members of the genus has increased (e.g., Serrania Beu, Provincia Sudyungas, Depar- HADDAD & MARTINS 1994), several species tamento La Paz. This locality lies in the remain poorly known. For sure, E. bolivianus proximity of the above mentioned localities (BOULENGER, 1902) has to be included with (fig. 2). The specimen is deposited in the the least known species of Epipedobates collection of the Museo de His- (DE LA RIVA & al. 1996). The original de- toria Natural "Noel Kempff Mercado", scription was based on one specimen from Santa Cruz de la Sierra, Bolivia (NKA San Ernesto (800 m a.s.l.) and two from 3707). In addition, an adult female, SVL ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at

180 L. GONZALES & S. LÔTTERS & S. REICHLE

26.6 mm (figs. IB, 1C), was found by SR a.s.l. We summarise the Bolivian distribu- on 2 October 1999 at km 30 on the Yu- tions of both species mentioned in fig. 2 cumo-Quiquibey road (15°41'55" S, 67°28' based on records in REYNOLDS & FOSTER 52" W; ca. 1650 m a.s.l.), Provincia No- (1992), HADDAD & MARTINS (1994), DE ryungas, Departamento La Paz. This local- LA RIVA & al. (1996), KÖHLER & LÖTTERS ity is situated about 100 km south-east to (1999), and material at CBF, NKA and the former. The specimen is deposited at ZFMK (Zoologisches Forschungsinstitut und the Colección Boliviana de Fauna, La Paz, Museum Alexander Koenig, Bonn). Al- Bolivia (CBF 3901). We compared both though E. pictus and E. hahneli occur in specimens with the descriptions of E. bolivi- broad sympatry in south-eastern Peru anus by SILVERSTONE (1976) and ROD- (compare DUELLMAN & THOMAS 1996), there RiGUEZ & MYERS (1993) as well as b/w is presently no evidence of sympatry in photographs of the lectotype [BM (British Bolivia. However, E. pictus (NKA 3753- Museum, London) 1947.2.13.89]. Our com- 58, 3771, 3794) was also found at different parison revealed high coincidence in exter- sites within Territorio Comunitario y nal features except that NKA 3707 pos- Reserva de la Biosfera "Pilon Lajas" and sesses a small light thigh spot while in the thus occurs in sympatry with E. bolivianus lectotype and CBF 3901 the dorsolateral (see fig. 2); nevertheless, syntopic occur- line continues on the upper femur. Life rence cannot be confirmed. A record of E. coloration, previously unknown, was as pictus (NKA 2733-2734) from the northern follows (figs. 1A - 1C): ground colour black, Paraguay basin in extreme south-eastern dorsally turning into brownish orange (most Bolivia (see fig. 2) is remarkable since the intense at mid-dorsum); bright dorsolateral general area is semiarid and most probably line yellow (orange before eye in NKA does not meet the species' usual biotope re- 3707), turning into yellowish cream before quirements. groin; labial Une whitish cream; limbs dor- Other dendrobatid frogs known from sally brownish bronze, hind limb with black Bolivia belong to the genus Colostethus marbling in NKA 3707; ventral sides of COPE, 1866, a speciose group of apparently body and limbs greenish to bluish cream non-toxic and more cryptically coloured with black marbling; prominent light yel- species (FROST 1985; GLAW & al. 1998). low axillary and in NKA 3707 small light Colostethus mcdiarmidi REYNOLDS & FOS- yellow thigh spot (light calf spot absent). TER, 1992 was described from humid mon- Thus far known, this species is restricted to tane forest (ca. 1650-1700 m a.s.l.) in the humid montane forest (800-1650 m a.s.l.) Chapare region, Departamento Cocha- in the Yungas de La Paz. bamba. On 17 and 21 September 1998, LG Epipedobates bolivianus superficially found this species, two males (NKA 3708, resembles E. pictus (BIBRON in TSCHUDI, 3712) of 22.0 and 24.0 mm SVL (fig. 3), in 1838) from semi-humid and very humid humid montane forest at Territorio Comu- lowland rain forests of the south-western nitario y Reserva de la Biosfera "Pilon La- Amazon basin of Bolivia and adjacent Peru jas" (15°06'36" S, 67°32'33" W; ca. 1350 (HADDAD & MARTINS 1994). It can be dis- m a.s.l.), Serrania Beu, Provincia Sudyun- tinguished from E. bolivianus by having gas, Departamento La Paz. These speci- light reddish orange axillary, thigh and calf mens coincide exactly with the original de- spots as well as bluish marbled ventral scription. The records of C. mcdiarmidi are surfaces (HADDAD & MARTINS 1994). The all situated in the Yungas of Cochabamba third species of Epipedobates known to oc- and La Paz (fig. 2) separated by a distance cur in Bolivia is the Amazonian E. hahneli of more than 250 km. Due to insufficient (BOULENGER, 1883). It can be distinguished collecting, it remains to be resolved if the from E. bolivianus by having a whitish species is more or less continuously dis- dorsolateral line and bluish marbled ven- tributed (as we assume) or if it shows a tral surfaces (HADDAD & MARTINS 1994). disjunct distribution. In contrast to E. bolivianus, both E. pictus Colostethus trilineatus (BOULENGER, and E. hahneli are primarily lowland spe- 1883) was suggested to represent a valid cies, although, according to DE LA RIVA & species from the south-western Amazon al. (1996), E. pictus reaches up to 1300 m basin by MORALES (1994). According to ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at

Fig. 1 A: Epipedobates bolivianus (BOULENOER, 1902) in dorsal view (NKA 3707). Fig. IB: Epipedobates bolivianus (BOULENOER, 1902) in dorsal view (CBF 3901). Fig. IC: Epipedobates bolivianus (BOULENOER, 1902) in ventral view (CBF 3901).

Abb. 1A: Epipedobates bolivianus (BOULENOER, 1902) in Dorsalansicht (NKA 3707).

Abb. IB: Epipedobates bolivianus (BOULENOER, 1902) in Dorsalansicht (CBF 3901).

Abb. IC: Epipedobates bolivianus (BOULENOER, 1902) in Ventralansicht (CBF 3901). ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at

182 L. GONZALES & S. LOTTERS & S. RHCHLE

I 68e 64e 60e

— 10 10° — CBA = COCHABAMB A CHU = CHUQUISACA

14° —

— 22 22° —

e 68e 64 60e _L

Fig. 2: Map of Bolivia with Departamentos and localities where dendrobatid frogs have been found. A - Colostethus brunneus; • - C. mediartnidi; O - C. trilineatus; Y - Colostethus sp. from Cobija; "it - Epipedobates bolivianus; D - E. hahneli; M-E. pictus. Andean areas above 2S00 m a.s.l. are grey. Abb. 2: Bolivien mit seinen Departamentos und den Fundorten von Dendrobatiden. A - Colostethus brunneus; • - C. mediarmidi; O-C. trilineatus; Y - Colostethus sp. from Cobija; -Epipedobates bolivianus; Q-E. hahneli; M-E. pictus. Andine Bereiche über 2500 m ü. N. N. sind grau.

DE LA RIVA & al. (1996), REICHLE & KÖH- ow") from near Chipiri, Departamento Co- LER [1996 under the name Colostethus chabamba, and concluded that they are also marchesianus (MELIN, 1941)] and KÖHLER C. trilineatus. Recently, SR collected two & LOTTERS (1999), in Bolivia this species females, SVL 18.1 and 18.9 mm (NKA occurs in lowland rain forest (ca. 200-300 3842-43), referable to this dendrobatid frog m a.s.l.) of the Departamentos Cocha- species, at Mataracü (17°33(92" S, 63°52( bamba, Beni and Pando (fig. 2). DE LA 71" W; ca. 500 m a.s.l.), Parque Nacional RIVA & al. (1996) examined specimens re- Amboró, Provincia Ichilo, Departamento ported by FUGLER (1983 as Colostethus sp.) Santa Cruz (compare fig. 2). Due to the from Tumi Chucua, Departamento Beni, lack of physical barriers, we principally and by EDWARDS (1974 as C. "phaeocene- agree with DE LA RIVA & al. (1996) that C. ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at

Bolivian dendrobatid frogs 183

Fig. 4: Palmar surfaces of individuals from three populations of Colostethus from lowland Bolivia. Bar equals 4 mm. A - Colostethus trilineatus BOULENOER, 1883 [Pando, Cobija, ZFMK 66791 ] - right hand. B - Colostethus trilineatus BOULENOER, 1883 [Beni, Totaizal (EBB), ZFMK 62444] - left hand. C - Colostethus sp. [Pando, Cobija, ZFMK 66793] - left hand. Abb. 4: Handflächen von Individuen dreier Colostethuj-Populationen aus dem bolivianischen Tiefland Der Balken entspricht 4 nun. A - Colostethus trilineatus BOULENOER, 1883 [Pando, Cobija, ZFMK 66791] - rechte Hand B - Colostethus trilineatus BOULENOER, 1883 [Beni, Totaizal (EBB), ZFMK 62444] • linke Hand. C - Colostethus sp. [Pando, Cobija, ZFMK 66793] - linke Hand ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at

184 L. GONZALES & S. LÔTTERS & S. REICHLE

trilineatus is also widespread in the inter- Recently, the authors found members vening area of the known Bolivian locali- of the genus Colostethus (fig. 6) at Parque ties. Nevertheless, we noted differences in Nacional "Noel Kempff Mercado" (Cam- the size of the thenar tubercle in three Bo- pamento Pauserna, 13°31'56M S, 61°06'20" livian populations examined by us. In fe- W; ca. 140 m a.s.l.) in the north-eastern males from Mataracü (NKA 3842-43) and lowlands of the Departamento Santa Cruz, a male and female from Cobija, Departa- Provincia Velasco (fig. 2). They are de- mento Pando (ZFMK 66791-92), the thenar posited at NKA (3012, 3150, 3153, 3162, tubercle is about two thirds of the size of 3835-36) and ZFMK (69917-18). These the palmar tubercle (fig. 4A). In contrast, specimens, mean SVL 15.2 mm (range 14.6 - in ZFMK 62444, a male from Totaizal 16.0 mm, n=8) in which a dorsal rhombus- (EBB), Departamento Beni, the thenar tu- like pattern may be present or absent, coin- bercle is less than one third of the size of cide well with C. brunneus (COPE, 1887) as the palmar tubercle (fig. 4B). We will not redescribed by EDWARDS (1974) and LA exclude that our observation reflects inter- MARCA (1996). According to FROST (1985), specific variation in sibling species. With this species is broadly distributed over almost respect to this, it is noteworthy that, ac- the entire Amazon basin with extends into the cording to HANAGARTH & BECK (1996), the Guyanas (the type locality is in Brazil, Estado Beni savannahs apparently have little Mato Grosso). We here report C. brunneus for Zoogeographie relation to the Amazon ba- the first time from Bolivia. sin. In summary, three species of Epipe- KÖHLER & LÔTTERS (1999) recog- dobates and at least three, maybe five spe- nised another morphotype of Colostethus cies of Colostethus are known from Bo- from Cobija, Departamento Pando (com- livia. Due to distributions outside but close pare fig. 2). According to the authors, it to the Bolivian border (e.g., FROST 1985; differs from syntopic C. trilineatus (sensu MARTINS & HADDAD 1990; MORALES, 1992; MORALES 1994) in slightly larger size, RODRÌGUEZ & MYERS 1993; HADDAD & shape of dorsolateral stripes, the presence MARTINS 1994; MYERS & al. 1998), the fol- of white ventrolateral marks (fig. 5 A) and lowing dendrobatid frog species may be ex- having a yellowish ventral colour (fig. 5B). pected to occur in this country as well: Al- Dissections revealed that the two speci- lobates femoralis (BOULENGER, 1883), Colo- mens available (ZFMK 66793-94) are adult stethus sp. from south-eastern Peru (V. R. females. Colostethus trilineatus females are MORALES, pers. comm.), bio- otherwise known to have white ventral lat MORALES, 1992, D. vanzolinii MYERS, sides (e.g., NKA 3842-43, ZFMK 66792). 1982, D. quinquevittatus STEINDACHNER, In addition, the thenar tubercles of this 1864, Epipedobates braccatus (STEINDACH- morph and syntopic C. trilineatus (ZFMK NER, 1864), E. macero RODRÎGUEZ & MYERS, 66791-92, male and female) are different 1993, E. simulons MYERS, RODRÎGUEZ & in shape and size (figs. 4A, 4C). This sup- IcoCHEA, 1998 and E. trivittatus (SPK, ports the hypothesis of KÖHLER & LÔTTERS 1824). The presence of the latter was pre- (1999) that the two forms found at Cobija viously confirmed through call recordings are not conspecific. by APARICIO(1992).

Figs. 3, 5 A, 5B, 6 (opposite page) / Abb. 3, 5A, 5B, 6 (gegenüberliegende Seite)

Fig. 3: Colostethus mcdiarmidi (NKA 3708) illustrated for the first time in life. Abb. 3: Colostethus mcdiarmidi (NKA 3708). Erste Abbildung eines lebenden Exemplars. Fig. 5: Colostethus sp. from Cobija (ZFMK 66794), Departamento Pando; 5 A - dorsolateral view; 5B - ventral view. It is not clear if it is conspecific or not with C. trilineatus. Abb. 5: Colostethus sp. aus Cobija (ZFMK 66794), Departamento Pando; SA - Dorsolateralansicht; SB - Ventralansicht. Es ist unklar, ob die Form artgleich oder nicht mit C. trilineatus ist. Fig. 6: Colostethus brunneus (ZFMK 69917) from Parque Nacional "Noel Kempff Mercado". Abb. 6: Colostethus brunneus (ZFMK 69917) vom Parque Nacional "Noel Kempff Mercado". ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at

186 L. GONZALES & S. LÖTTERS & S. REICHLE

ACKNOWLEDGMENTS

We are grateful to DAMIAN RUMIZ for taking tario y Reserva de la Biosfera "Pilon Lajas". Field work photographs in the field of Colostethus mcdiarmidi and of two of the authors (SL and SR) in the Departamento Epipedobates bolivianus. JÖRN KÖHLER kindly pro- Santa Cruz was supported by Fundación Amigos de la vided photographs of the lectotype of E. bolivianus and Naturaleza (FAN), Santa Cruz de la Sierra. SL thanks of Colostethus sp. from Cobija. For help in the field we the Graduiertenforderung des Landes Nordrhein-West- thank MARTIN BEICHTER, ANGELIKA VON FOERSTER, fàlen and the German Academic Exchange Service (DAAD) ALEX WIDMER and GUSTAVO J. SCROCCHI. One of the for providing grants. Special thanks to TARAN GRANT, authors (LG) is indebted to the proyecto "Pilon Lajas" IONACIO DE LA RIVA, MICHAEL B. HARVEY, KARL- de Veterinarios sin Fronteras (V.S.F.), Rurrenabaque,* HEINZ JUNGFER, and VICTOR R. MORALES for com- which helped to realize field work at Territorio Comuni- ments on the matter and/or the manuscript

REFERENCES

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DATE OF SUBMISSION: April 13th, 1999 Corresponding editor: Heinz Grillitsch

AUTHORS: LUCINDO GONZALES, Museo de Historia Natural "Noel Kempff Mercado", Casula 2489, Santa Cruz de la Sierra, Bolivia; STEFAN LÖTTERS & STEFFEN REICHLE, Zoologisches Forschungsinstitut und Museum Al- exander Koenig, Adenauerallee 160, 53113 Bonn, Germany, [e-mail: [email protected]]