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Amphibians and reptiles from northern Departamento Loreto, Peru: Taxonomy and biogeography

January 1995
The University of Kansas science bulletin 55(10):329-376

DOI:10.5962/bhl.part.779

Authors:

W. E. Duellman

University of Kansas

Joseph R Mendelson III

Zoo Atlanta

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Ectothermic inhabitants of tropical forests are subjected to constant environmental temperatures, which determine their passive thermoregulatory strategies. We observe these trends during the summer of 2017, in the anurans Dendropsophus bifurcus, Rhinella marina, and Scinax ruber, in a tropical rainforest from the Upper Amazon Basin of Ecuador. D. bifurcus and S. ruber showed a tendency to tigmothermy, whereas R. marina presented tendencies towards heliothermy. Body temperatures (Tbs) did not differ between D. bifurcus and R. marina, but S. ruber presented a lower Tb. Our results suggest that thermal environment is influencing different thermoregulatory strategies as tigmothermy and heliothermy of frogs and toads distributed in tropical environments at low elevation.

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A preliminary survey of amphibians and reptiles on the Nassau and Lely plateaus Eastern Suriname

Chapter

Apr 2007

INTRODUCTION Amphibians and reptiles are a species-rich and often conspicuous component of many neotropical forests. Three aspects of amphibians and reptile biology make them a valuable focal group for biological surveys: (1) the small body size of many species often results in high population densities, making it possible to collect a large amount of data in a relatively short period of time; (2) they perceive their environment on relatively small scales and many species show strict habitat requirements, making it possible to compare diversity patterns across finely-defined habitats; (3) their intermediate role in food webs ties them to both primary and secondary consumers. Amphibians are of particular interest because their moist, permeable skin makes them more sensitive to changes in their environment (e.g., contamination, climate change) than other vertebrate groups, and the biphasic lifestyle of many species exposes them to changes in both aquatic and terrestrial environments. Widespread reports of enigmatic amphibian declines in seemingly pristine locations are of urgent conservation concern (Lips 1998), and it appears that amphibians as a group are more threatened than other terrestrial vertebrates (Stuart et al. 2004, Beebee and Griffiths 2005). As part of the CI RAP survey in eastern Suriname, we surveyed the herpetofauna of Nassau and Lely mountains for six days each. Here we compare three response metrics (species richness, species composition, and an estimate of density) between the two mountains, and place these preliminary observations in a regional context by making comparisons with other sites in the Guayana Shield and the Amazon Basin. We also describe the distribution of species at a regional scale and among macrohabitats at the two sites, and discuss the conservation implications of our observations. METHODS We surveyed amphibians and reptiles for six days each at the Nassau mountain (25 – 30 October 2005) and Lely mountain (1 – 6 November 2005) using a combination of opportunistic surveys and time-constrained Visual Encounter Surveys (VES). Opportunistic surveys require actively searching for animals over large areas (i.e., up to several square kilometers) in order to increase the probability of encountering as many different species as possible. This method is effective for sampling species richness (Donnelly et al. 2004), but because not all individuals encountered are recorded, and cryptic or inactive individuals may be easily overlooked, the method is inappropriate for comparing density. In contrast, VES involve intensive sampling over small areas (i.e., a few hundred square meters) and all individuals encountered are recorded, making it possible to calculate an index of density by comparing the number of individuals encountered per unit time (Crump and Scott 1994). We conducted opportunistic surveys throughout the range of habitats available at each site, walking trails, forest creeks, and searching in natural and anthropogenic clearings both day and night throughout our stay. We conducted ten VES (eight nocturnal and 2 diurnal) at each site, concentrating effort in forest and forest stream habitats (Table 10.1). Table 10.1. Schedule of herpetofaunal sampling at Nassau and Lely, October–November, 2005. Species richness Because observed species richness is almost always an underestimate of true species richness (Colwell and Coddington 1994, Hellmann and Fowler 1999), we used program EstimateS (Colwell 1997) to calculate extrapolated estimates of amphibian and reptile richness of each site. There is considerable debate as to which of the many species richness estimators provides the most robust results (Hellman and Fowler 1999, Herzog et al. 2002), so we included four of the most commonly used estimators (ACE, ICE, Chao1 and MMMean). We included observations from both VES and opportunistic encounters in our analysis because few reptiles were encountered during VES surveys at Lely, making it impractical to estimate species richness from only that data set. Although not all individuals of all species were recorded during opportunistic surveys, the combined data set accurately reflects observed species abundances (i.e., rare species only occur once or twice in the entire data set, whereas the most commonly encountered species appear frequently), so we assume that the combined data set provides a reasonable basis for comparing species richness between sites. Species composition In addition to comparing species richness on the two mountains, we also wanted to describe overlap in species identity. We began simply by comparing the number of species occurring at only one of the sites with the number occurring at both sites. We conducted a formal test of the compositional difference between the two mountains using analysis of similarity (ANOSIM) based on the Bray-Curtis dissimilarity index (Clarke and Warwick 2001), and present these results in a non-metric multidimensional scaling nMDS) graph. Species composition is known to vary with geographic distance (Steinitz et al. 2005), so in addition to establishing that a compositional difference between the two sites exists, we also wanted to determine whether the difference in species composition between Nassau and Lely was more or less than would be expected given the distance between the two mountains. We compiled data on amphibian and reptile surveys from five sites in the Guayana Shield: Nouragues and Arataye, French Guiana (Born and Gaucher 2001); Petit Saut, French Guiana (Duellman 1997), Piste Ste. Elie, French Guiana (Born and Gaucher 2001); and Iwokrama, Guyana (Donnelly et al. 2005). Distance between these sites ranged from 64 – 713 kilometers. Herpetofaunal survey data are available for eight sites separated by a maximum distance of 263 kilometers in the Madre de Dios region of southeastern Peru (Duellman and Thomas 1996, Morales and McDiarmid 1996, Doan and Arriaga 2002), and because distances among sites in Madre de Dios are more similar to those of interest here (straight-line distance between Nassau and Lely is approximately 63 kilometers), we include a comparison with those sites as well. Although some species certainly remain undetected at sites in the Guayana Shield and Peru, those sites are relatively well-sampled compared with Nassau and Lely. For all pairs of sites, we calculated straight-line distance based on coordinates included in the primary literature cited above. We compiled a species by site presence/absence matrix for all sites and calculated dissimilarity among all pairs of sites using the Bray-Curtis dissimilarity index in Program Primer (Clarke and Warwick 2001). Density In order to have maximal flexibility and not be constrained to surveying fixed transects that may have resulted in the observation of few individuals, we opted to constrain our VES by time rather than area. At each site we conducted ten VES, eight nocturnal and two diurnal. As an index of density, we calculated the number of individuals encountered per survey minute (# individuals/# minutes surveyed), averaged this value across all ten surveys, and multiplied this average by 60 to provide an average number of individuals encountered per hour of survey at each site. Species-specific data When individuals of new species were encountered during opportunistic surveys, we noted the habitat in which the observation occurred. Similarly, we noted the habitat where VES occurred. Thus, we are able to assign species occurrences to one or more habitat categories: forest, forest stream, clearing, berg forest (Nassau only), swamp forest (Nassau only), savannah forest (Lely only), or forest clearing (Lely only). Because of the complex interdigitation of savannah forest and high forest around the camp at Lely, we refer to both as ‘forest’. We did survey a discreet patch of berg forest east of the main camp at Nassau, and a patch of savannah forest with many bromeliads near the northeastern corner of the airstrip at Lely. We assigned each species to one of two regional distribution patterns: Guayana Shield for those species endemic (or nearly so) to the Guayana Shield, and Widespread for species that also occur beyond the boundaries of the Guayana Shield. Distributional data were taken from Ceñaris and MacCulloch (2005) for amphibians and Ávila Pires (2005) for reptiles. Threat status for each species was established based on IUCN Red List guidelines (www.iucnredlist.org). Data for amphibians were taken from the Global Amphibian Assessment online database (www.globalamphibians.org). Data on the crocodilian were extracted from the IUCN website. For the lizards and snakes (for which no IUCN specialist group currently exists) we used our knowledge of probable distributions and potential threats to assign a threat status based on IUCN criteria. We did not include threat status for unidentified species thought to represent new species for science, because determination of threat status will require more survey work to establish the geographic range of those species. RESULTS We observed a total of 49 species in 12 days of sampling at the two sites (Table 10.2). The data presented herein include only species observed by the two authors; species observed by other members of the RAP team and on a herpetological expedition to Lely in 1979 are included in Appendix 16. Comparison with other well-studied sites in the Guayana Shield indicate that many species remain undetected on the two mountains, and that reptiles were undersampled on the RAP relative to amphibians (because they represent a smaller percentage of the total herpetofauna at RAP sites than at more well-sampled sites, Table 10.2). Despite the fact that many species remain to be detected on both mountains, preliminary observations indicate that Lely appears to be the richer of the two mountains; we observed 36 species there and 29 at Nassau (Figure 10.1). Extrapolated species richness estimates were largely consistent with the notion of higher richness at Lely than Nassau (Table 10.3). However see Chapter 11 for additional data from Nassau. Table 10.2. Herpetofaunal richness at nine sites in the Guayana Shield, including Nassau and Lely mountains. In each column, data are presented as raw species number/percentage of total herpetofauna. Table 10.3. Observed and estimated species richness for amphibians and reptiles at Nassau and Lely. A simple review of the species list for the two sites indicates that species composition differs between Nassau and Lely, with only 15/49 = 31% of all species occurring on both mountains. Forty-eight percent of the species at Nassau were unique to Nassau, whereas the percentage was 57% at Lely. As expected, the species occurring at the two sites represented a mix of widespread species that occur throughout lowland portions of much of the Amazon Basin, in addition to species known from lowland forest of the Guayana Shield (Appendix 16). Five records are particularly noteworthy because they represent taxa that could not be assigned to any known species. Four of these records were species of the genus Eleutherodactylus; one species was encountered at both Lely and Nassau, whereas the other three new species of Eleutherodactylus were found at Lely. We also collected what appears to be an undescribed species of Adenomera at Lely. Bray-Curtis dissimilarity between Nassau and Lely is 51.7% for reptiles and 44.4% for amphibians, and the two mountains are compositionally distinct (Global R = 0.669, P = 0.002; Figure 10.2). Comparison of the regression lines describing the relationship between compositional dissimilarity and geographic distance among sites in the Guayana Shield and Tambopata, Peru reveal that observed dissimilarity between Nassau and Lely is greater than would be expected based on observations from the reference sites (Figure 10.3). Like richness, herpetofaunal density was higher at Lely (mean = 7.4 individuals/hour) than at Nassau (4.5 individuals/hour). At Nassau, the highest density of individuals occurred in transects running through the IJskreek and forest adjacent to the stream, and lowest in savannah forest. At Lely, density was greatest in forest streams, slightly lower in forest, and lowest in savannah forest. Habitat use, distribution, and threat status for each species are presented in Appendix 16. We draw particular attention to the observation that forest streams are important habitat for many species encountered during our surveys. Just under half of the species occurring at each site made use of forest streams, and one quarter of the species encountered at Lely and one third of the species encountered at Nassau were only found in or along forest streams. In addition, two of the five new species encountered during our surveys were associated with forest streams. At Lely, density was higher in forest streams than in any other habitat, whereas at Nassau, density was broadly similar between forest and streams, but higher there than in other habitats. Overall, we suggest that forest streams be considered keystone habitat structures (Tews et al. 2004) of paramount biological and conservation value at the two sites because (1) they cover a small proportion of the total habitat at each site, (2) house a substantial fraction of overall herpetofaunal richness at the two sites, (3) are an important habitat for undescribed and probably narrowly endemic taxa recorded during our surveys, and (4) at Lely, have a greater herpetofaunal density than any habitat surveyed. DISCUSSION In our short surveys we sampled only a fraction of what is likely a rich herpetofauna on both mountains. Estimated species richness estimates for amphibians and reptiles were not much greater than observed richness (Table 10.3). Our impression is that low estimated richness is a function of seasonal fluctuations in activity (for amphibians) and small sample sizes (for both amphibians and reptiles), rather than being indicative of a depauperate herpetofauna on the two mountains. The rapid accumulation of species during a dry period during which many amphibians and reptiles were likely inactive is suggestive of potentially high richness, as is the geographic proximity to sites in western French Guiana with the highest known richness of amphibians and reptiles in the Guayana Shield (Petit Saut and the Nouragues reserve; Table 10.2). Comparison of our species accumulation curves with those from other sites indicate that species accumulated faster at Lely and Nassau than at individual camps in the Iwokrama reserve in Guyana (Donnelly et al. 2004), and were more similar to species-rich sites in the western Amazon (Duellman and Mendelson 1995, Cadle et al. 2002, Moravec and Aparicio 2005). Although sampling on the two mountains is far from complete, available evidence suggests that Lely is likely to be the richer of the two sites. Our observation that compositional dissimilarity between the two mountains is greater than expected given their geographic distance suggests that conservation of both areas is not redundant, but necessary in order to conserve a representative regional fauna. Beyond acting as reservoirs of a rich herpetofauna, the two mountains are home to a suite of endemic taxa that is of great regional importance. Particularly striking was the four Eleutherodactylus species encountered during our surveys. Previous to our surveys, five species of Eleutherodactylus were known from Suriname; our work on the two mountains has almost doubled the representation of the genus in the country. Two of the new species encountered during our surveys (Adenomera sp. and Eleutherodactylus sp. 1) utilized both forest and forest stream habitats and were abundant where they occurred. The three other new species (Eleutherodactylus sp. 2, Eleutherodactylus sp. 3, and Eleutherodactylus sp. 4) were found in the forest at Lely and were represented by only one or two individuals each. Although the forest-inhabiting Eleutherodactylus appeared to be rare, they likely occur throughout the forest and because they do not require standing water for breeding, their persistence is not as dependent on particular habitat requirements as the other frogs. Therefore, we consider the species associated with forest streams to be the most in need of conservation attention. Amphibians tend to have limited dispersal abilities, often moving less than 500 m (Smith and Green 2005). Because body size of the remaining four new species is small (< 40 mm, implying relatively limited dispersal abilities; Etienne and Olff 2004) and they appear to be reliant on a habitat type that is relatively scarce in the landscape, it may be unlikely that individuals can move to more suitable habitat (i.e., another stream) if they are disturbed. Amphibians tend to be dietary generalists, feeding on a variety of arthropods (Duellman 1978, Parmelee 1999), so it is unlikely that distributions of any of these species are limited by the availability of food resources. Protection of streams where they are known to occur should be considered the best conservation action for these new species, as well as the other species that utilize forest stream habitat on the two mountains. Streams are a keystone habitat feature of critical importance for amphibians at Nassau and Lely. Almost half of the species encountered during our surveys made at least some use of streamside habitat. Stream-associated amphibians are of paramount conservation significance because many species in this guild have experienced precipitous population declines (Lips et al. 2003). Like virtually all other taxonomic groups, amphibians have been affected by habitat loss and fragmentation, overharvest, and other anthropogenic disturbances. More alarming are population declines, many to the point of extinction, of amphibians in protected areas where the agent of decline is not so obvious. These enigmatic declines have resulted in the loss of many moderate- to high-elevation anurofaunas (Young et al. 2001), so the presence of abundant, diverse, stream-associated amphibian assemblages at Nassau and Lely is of significant conservation value. The densities we observed at Nassau and Lely are comparable to pre-decline data from forest streams and adjacent forest in Panama (Lips 1999), suggesting that the stream-associated fauna of Nassau and Lely have not experienced the dramatic declines that have occurred in other parts of the Neotropics (Young et al. 2001). This provides an excellent opportunity to protect an intact, upland stream-associated herpetofaunal assemblage. CONSERVATION RECOMMENDATIONS Our first and foremost conservation recommendation is to maintain the integrity of forest streams at both Lely and Nassau. Anthropogenic activity at Lely is minimal, so there are no current threats, but every attempt should be made to ensure that future activity at Lely be kept away from stream habitats. The stream at Nassau probably has been impacted and will continue to be impacted by the higher level of human activity. Of most concern is the presence of the camp clearing and a dirt path used by motorized vehicles that crosses the Ijskreek through the clearing. Because of the possibility that human activity may negatively impact stream quality at Nassau, we make the following recommendations: • (1) Because sedimentation and runoff from the clearing and the road have the potential to impact water quality in the stream, we recommend that no further expansion of the existing camp take place, and that vehicular traffic across the stream be reduced to an absolute minimum. • (2) The immediate initiation of a water-quality monitoring project in conjunction with herpetofaunal surveys. We suggest twice yearly surveys of the stream-associated herpetofauna at Nassau using fixed monitoring points established throughout the watershed. Species may be located visually and/or acoustically, but we recommend the utilization of a visual method (i.e., VES) in order to estimate population density as accurately as possible. Because interspecific variation in detection probabilities may compromise results (Mackenzie and Royle 2005), it will be necessary to incorporate methods that will allow for robust density estimation (discussed in Schmidt 2004). Concomittant with the faunal surveys, we recommend the collection of basic water quality data (dissolved oxygen, conductance, temperature, pH, and turbidity) at the beginning of each transect or monitoring point. • (3) An ongoing monitoring project to detect the presence of Batrachochytrium dendrobatidis in adult frogs along forest streams. Batrachochytrium dendrobatidis is a chytrid fungus that has been linked to amphibian declines in many parts of the Neotropics (Lips et al. 2005), and although we are not aware of reports of amphibian declines from the Guianas, conditions favorable for the occurrence of B. dendrobatidis are predicted to occur in the vicinity of Nassau and Lely mountains (Ron 2005). The presence of B. dendrobatidis can be detected via analysis of dermal swabs from live animals. We recommend collecting 300 swabs/visit (i.e., one swab per individual from the first 300 individuals encountered). To detect the presence of B. dendobatidis, analysis may be conducted on pooled samples of 10 swabs. If the fungus is detected, individual analysis of all swabs will be necessary to identify infected species. Should B. dendrobatidis be detected, the Declining Amphibian Population Task Force (http://www.open.ac.uk/daptf/index.htm) may be contacted for recommended action. • (4) We recommended expanded surveys of streams on the two mountains and in adjacent lowlands in order to more accurately quantify abundance and extent of occurrence of stream-associated frogs, particularly new species whose distributions are unknown. Determining the IUCN red list status of these five species will hinge on estimating the geographic range of these species, so a special effort should be made to determine their extent of occurrence. • (5) It is difficult to provide meaningful guidelines for the area required to effectively protect amphibian populations because the availability of breeding habitat is probably more important than area per se (Zimmerman and Bierregaard 1986). Reptiles, on the other hand, probably benefit more from larger areas, though relative to endothermic vertebrates their energetic needs (and therefore area required to sustain populations; Pough 1980) are low. It has been suggested that the 1500 ha of the La Selva reserve in Costa Rica is sufficient to protect the herpetofauna at that site (Guyer 1994), although population declines of both amphibians and reptiles have occurred there (S. Whitfield pers. com.). We therefore regard 1500 ha as the ‘minimum critical area’ necessary to protect a reasonably intact sample of the local herpetofauna, and suggest that at least this amount be preserved within the concessions at Lely and Nassau. Additionally, because we have identified streams as keystone habitat whose importance is disproportionate to their area, we recommend a forest buffer of at least 50 m (Lee et al. 2004) on both sides of all creeks running through the concessions. Authors' note: As this chapter was going to press, we became aware of a record of the toad Atelopus cf. spumarius from the forest near the basecamp at Nassau. Atelopus spumarius is a polymorphic taxon, and it is possible that more than one species is included under the name (some authors recognize the Guayana Shield taxon to be a distinct species, A. hoogmoedi). Although A. spumarius (as either A. spumarius sensu stricto or A. hoogmoedi) has a larger geographic range than many species of Atelopus, these toads have experienced precipitous population declines in much of Latin America, most likely due to infection by B. dendrobatidis, and A. spumarius senso lato is classified as vulnerable by the IUCN. A population of Atelopus at Nassau would therefore be of significant conservation concern. We recommend that efforts to establish the extent of occurrence of the new taxa encountered during our surveys include A. cf. spumarius. REFERENCES 1 Ávila PiresT. C S. 2005Reptiles.In HollowellT. R. P.Reynolds (eds.)Checklist of the terrestrial vertebrates of the Guiana ShieldBulletin of the Biological Society of WashingtonUSANumber 13. Pp25–40Google Scholar 2 3 BornM. P.Gaucher 2001Amphibian and reptile species at the Nouragues Nature Reserve.In BongersF. P.Charles-Dominique P-M.Forget M.Théry (eds.)Nouragues. Dynamics and plant-animal interactions in a neotropical rainforestKluwer Academic PublishersPp371381Google Scholar 4 CadleJ. J.Icochea J. P.Zúniga A.Portilla C.Rivera 2002La herpetofauna encontrada en el Refugio Juliaca y en el Puesto de vigilancia Enahuipa del Santuario Nacional Pampas del Heath.In MontambaultJ. R. (ed.)Informes de las evaluaciones biológicas Pampas del Heath, Perú Alto Madidi, Bolivia y Pando, BoliviaRAP Working Papers Number 24Washington, D.CConservation InternationalPp5257Google Scholar 5 CeñarisJ. C. R.MacCulloch 2005Amphibians.In HollowellT. R. P.Reynolds (eds.)Checklist of the terrestrial vertebrates of the Guiana ShieldBulletin of the Biological Society of WashingtonUSANumber 13. Pp923Google Scholar 6 ClarkeK. R. R. M.Warwick 2001Change in marine communities: an approach to statistical analysis and interpretationSecond editionPRIMER-EPlymouth, U.KGoogle Scholar 7 ColwellR. K. 1999EstimateS version 5.0.1. Statistical Estimation of species richness and shared species from samplesWeb site: viceroy.eeb.uconn.edu/estimatesGoogle Scholar 8 9 CrumpM. L. N. J.ScottJr. 1994Visual Encounter Surveys.In HeyerW. R. M. A.Donnelly R. W.McDiarmid L. C.Hayek M. S.Foster (eds.)Measuring and Monitoring Biological Diversity Standard Methods for AmphibiansWashington D.CSmithsonian Institution PressPp8491Google Scholar 10 11 12 DonnellyM. A. M. H.Chen G. G.Watkins 2005The Iwokrama herpetofauna: an exploration of diversity in a Guayanan rainforest.In DonnellyM. A. B. I.Crother C.Guyer M. H.Wake M. E.White (eds.)Ecology and Evolution in the TropicsChicago, IllinoisThe University of Chicago PressPp428460Google Scholar 13 14 15 16 17 18 19 GuyerC. 1994The reptile fauna: diversity and ecology.In McDadeL. A. K. S.Bawa H. A.Hespenheide G. S.Hartshorn (eds.)La SelvaChicago, IllinoisUniversity of Chicago PressPp210216Google Scholar 20 21 22 23 24 25 26 27 28 MoralesV. R. R. W.McDiarmid 1996Annotated checklist of the amphibians and reptiles of Pakitza, Manu national park Reserve Zone, with comments on the herpetofauna of Madre de Dios, Perú.In WilsonD. E. A.Sandoval (eds.)ManuWashington, D. CSmithsonian Institution PressPp503522Google Scholar 29 30 31 32 33 34 35 36 37 38 39 ZimmermanB. L. R. O.Bierregaard 1986Relevance of the equilibrium theory of island biogeography andGoogle Scholar Appendices Appendix 16 List of Reptiles and Amphibians recorded on the Nassau and Lely plateaus James I. Watling and Lucille F. Ngadino Data include habitat use, Distribution (W=Widespread Amazonian, GS=Guayana Shield), and IUCN Threat Status (LC = least concern, NE = not evaluated). ‘X’ indicates presence of species not observed by the authors.

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Jun 2023

The white-sand ecosystems in the Solimões-Negro Interfluve are among the less studied in Amazonia. Recent herpetological surveys conducted west of Manaus, Brazil (central Amazonia) indicate that white-sand forests host a unique anuran fauna comprising habitat specialized and endemic species. In the present study we describe a new species of rain frog belonging to the Pristimantis unistrigatus species group from the white-sand forest locally called ''campinarana'' (thin-trunked forests with canopy height below 20 m). The new species is phylogenetically close to rain frogs from western Amazonian lowlands (P. delius, P. librarius, P. matidiktyo and P. ockendeni). It differs from its closest relatives mainly by its size (male SVL of 17.3-20.1 mm, n = 16; female SVL of 23.2-26.5 mm, n = 6), presence of tympanum, tarsal tubercles and dentigerous processes of vomers, its translucent groin without bright colored blotches or marks, and by its advertisement call (composed of 5-10 notes, call duration of 550-1,061 ms, dominant frequency of 3,295-3,919 Hz). Like other anuran species recently discovered in the white-sand forests west of Manaus, the new species seems to be restricted to this peculiar ecosystem.

Pfeilgiftfrösche der Gattung Epipedobates mit rot granuliertem Rücken aus dem Oriente von Ecuador und Peru

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Aug 1989

Karl-Heinz Jungfer

A review of red-backed granulated dendrobatid frogs from the Amazonian lowlands (Oriente) of Ecuador and Peru has resulted in the description of two new species. Epipedobates bilinguis sp. n. has conspicuous yellow flecks on upper arms and thighs and has previously been confused wich E. parvulus. The latter species is redefined. E. ardens sp. n. from the Departamento San Martin, Peru, has been confused wich E. zaparo, but is different in ventral coloration and finger lengths. lt might be conspecific with an unidentified frog known only from a color photo from the upper Rio Manu, Departamento Madre de Dios, Peru.

Herpetofaunas in neotropical rainforests: comparative composition, history, and resource use

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