review and analysis of altitudinal distribution of the andean anurans in colombia

Accepted by M. Vences: 26 Jun. 2008; published: 21 Jul. 2008 1

ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2008 · Magnolia Press

Zootaxa 1826: 1–25 (2008) www.mapress.com/zootaxa/

Review and analysis of altitudinal distribution of the Andean anurans in Colombia

MANUEL HERNANDO BERNAL1, 2 & JOHN D. LYNCH1

1Laboratorio de Anfibios. Instituto de Ciencias Naturales, Universidad Nacional de Colombia. Bogotá, Colombia2Laboratorio de Herpetología & Eco-Fisiología, Grupo de Investigación en Zoologia. Universidad del Tolima, Ibagué, Colombia. E-mail: [email protected]

Abstract

In the following work we examine the richness and altitudinal distribution of Colombian Andean anurans trying toemphasize patterns of distribution. We also supply an updated checklist of Andean anurans in Colombia. At present,Colombian harbors about 396 Andean frogs: 153 species in the Cordillera Occidental, 187 species in the Cordillera Cen-tral, and 131 species in the Cordillera Oriental. Of these, the Cordillera Oriental presents the higher number and percent-age of endemic species. The frequency distribution of altitudinal ranges for Colombian Andean frogs shows that themajority of species have narrow altitudinal ranges, less than 500 m altitude, and only a few species have broad altitudinaldistributions. On the other hand, lowland species have broader altitudinal ranges than do highland species. The hypothe-sis of a wider altitudinal range of highland anurans is therefore not supported. Finally, the averages of the Jaccard simi-larity indices for the Andean anurans along altitudinal gradients in Colombian are approximately similar to those of othertropical anurans reported by Huey (1978), but notably lower than those of anurans of temperate localities. Thus, theseresults are in concordance with Janzen’s hypothesis (1967) about a broader altitudinal range for temperate species, likelybecause of their higher thermal tolerance.

Key words: Checklist, richness, altitude, temperature, Janzen’s hypothesis

Introduction

Tropical zone-species have been suggested to be more habitat specific and to have narrower thermal toleranceranges than do temperate zone-species, because of the relative uniformity of their local environmental condi-tions (Janzen 1967, Huey 1978). As a consequence, faunal turnover along altitudinal gradients should be rapidin the tropics, and tropical species should have relatively restricted altitudinal ranges, such that between altitu-dinal faunal and floral units overlaps would be reduced (Olson 1994, Ghalambor et al. 2005). Also, it is gen-erally acknowledged that biological diversity is higher in the tropics than in temperate zones (Janzen 1967,Huey 1978, Ghalambor et al. 2005) and that along elevational gradients, species richness decreases with alti-tude toward a faunal minimum at very high elevations, in both vertebrates and invertebrates (Bernal 2005,Navas 2003, Poynton 2003, Bruhl et al. 1999, Patterson et al. 1996, McCoy 1990). Several biotic and abioticfactors, as well as historical events, have been attributed to these differences in species richness and biodiver-sity along latitudinal and altitudinal gradients. One of these was Janzen’s (1967) hypothesis: that altitudinallyseparated populations in the tropics will experience reduced gene flow, because the “mountain passes shouldbe physiologically higher in the tropics”, leading to greater genetic divergence that favour the allopatric speci-ation and higher tropical biodiversity (Ghalambor et al. 2005). Navas (2003, 2005) reviewed factors influenc-ing herpetological diversity in Andean high-elevations. Navas (2003) argued that temperature should be the

TERM OF USEThis pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited.

BERNAL & LYNCH2 · Zootaxa 1826 © 2008 Magnolia Press

most important single factor limiting herpetological diversity, at least in tropical Andean high-elevations,although subsequently, Navas (2005), suggested that temperature must not be the most likely stressor limitinganuran colonization of high elevation in the tropics, and that other environmental factors may be more impor-tant than temperature, such as water pH, intensity of UV radiation and other variables that may impact embryodevelopment.

Here we examine the richness and altitudinal distribution of Colombian Andean anurans trying to empha-size patterns of their distribution. First, we compare data compiled on Andean frogs (Appendix 1) with themost current reports about frogs from the richest lowland places in Colombia, Amazonía (Lynch 2005) andChocó (Lynch & Suárez 2004), to determine the peaks in species richness among these Colombian geographicunits, and to contrast the results given by Lynch et al. (1997) about biogeographic patterns of Colombianfrogs. Second, we carry out an analysis of similarity of frog species along altitudinal ranges to evaluate if trop-ical anurans have relatively restricted altitudinal ranges, such that overlaps between altitudinal faunas wouldbe reduced, according with Janzen’s (1967) hypothesis. Third, we compare altitudinal ranges of highland andlowland frog species of the Andean mountains to test if highland anurans have wider altitudinal ranges thanlowland species because of the possibly broader range of environmental temperature (Ghalambor et al. 2005).Finally, we compare similarity indices for Colombian tropical frogs along altitudinal gradients with the simi-larity indices reported by Huey (1978) for temperate frogs, and with a current study on elevation pattern offrogs species in China (Fu et al. 2006), in order to test latitudinal pattern of between-altitude faunal similarity,an extension of Janzen’s hypothesis (Huey 1978).

Materials and methods

Data about altitudinal distribution of the Colombian Andean frogs were obtained from an extensive review onthe amphibian collection of the Instituto de Ciencias Naturales, Universidad Nacional de Colombia, and frompublished literature. In this work, Andean anurans are defined as species that live mainly above 1000 m alti-tude, and lowlands species as those that live below 1000 m altitude. Data analyses are described in the text.

Results and discussion

1. The richness of Andean frogs in Colombia.Lynch et al. (1997) compared the frog fauna of ten eco-geographic units in Colombia, and they reported

that the Andean Cordilleras (Figure 1) were the regions with the highest richness in anurans, so that highlandregions could explain the majority of Colombian frog diversity, more than the lowland regions. At the present,Colombia harbors about 754 reported species of amphibians: 700 species of anurans, 35 species of caeciliansand 19 of salamanders. Of these, 396 are Andean frogs: 153 species in the Cordillera Occidental, 187 speciesin the Cordillera Central, and 131 species in the Cordillera Oriental (Appendix 1), some species are found inmore than one cordillera. With respect to the richest lowland units in Colombia, 124 anurans species havebeen reported in the biogeographic Chocó (Lynch & Suárez 2004) and 97 species in a small section of theAmazonian region (Lynch 2005), although Lynch (2005) suggests a local fauna no fewer than 123 species.According to these recent data, the Andean mountains are still the richest geographic units in toads and frogsof Colombia, and particularly the Cordillera Central is the Andean mountain with the highest anuran richness.Thus, comparisons among ecogeographic units in Colombia show that highland regions are richer in frog spe-cies than lowland regions. Although, along an altitudinal transect, lowland sities harbor more species thanhighland places (Duellman 1988, Lynch 1998, Navas 2003, Poynton 2003, Bernal et al. 2005). Lynch et al.(1997) provided an explanation about this trend; it is that anuran diversity in Colombia is due to beta diversity(geographical replacement) rather than alpha diversity (ecological complexity).


Zootaxa 1826 © 2008 Magnolia Press · 3ALTITUDINAL DISTRIBUTION OF ANDEAN ANURANS

FIGURE 1. Map of Colombia showing the eco-geographic units sensu Lynch et al. (1997). Andean units: 1. CordilleraOccidental; 2. Cordillera Central; 3. Cordillera Oriental. A: Chocó; B: Caribean lowlands and interandean valleys; C:Amazonía.

With respect to endemic species, the Andean Cordilleras of Colombia harbor 217 species and of these theCordillera Oriental presents the higher number (76), and percentage of endemicity (Table 1). This value isnotably superior to the 27 endemic species of the Biogeographic Chocó (Lynch & Suárez 2004), which repre-sents the 19.4% of the total species for this ecogeografic unit and just the 0.036% of Colombian species. Con-trarily, the Andean endemic species represents the 28.8% of the Colombian anurans, and specifically theCordillera Oriental harbors the 19% of the Andean species and the 10.1% of the Colombian anurans. In con-clusion, comparing these update data about frog distribution in Colombia, the Andean mountains also harborthe highest number of endemic species.

Finally, a cluster analysis of anuran similarity for the three Colombian cordilleras shows that the Cordil-lera Central is closer to Cordillera Occidental (Jaccard Index = 0.17) than Cordillera Oriental (JI = 0.11).However, it is important to realize that anuran similarities among the three cordilleras are very low, such thatbetween the Cordillera Occidental and Oriental the Jaccard Index is only 0.038. It means that each Andean



mountain harbors almost its exclusive anuran fauna, or endemic anurans as mentioned above. Likely, the bio-geographic history of the Andean Cordilleras has played an important role in this particular diversity for eachAndean mountain chain. For example, when the glacial and interglacial periods produced cycles of contrac-tion and expansion of anuran habitats over long periods of time (Navas 2005, Duellman 1999), because itcould generate the scenario to speciation through vicariance, if gene flow across populations becamerestricted, and then the high anuran diversity. Cladistic analysis of some anurans inhabiting the Andes (Lynchet al. 1997, Lynch 1999) show that phylogenetically related species occur at about the same altitude, indepen-dent of their geographical distance, so Andean diversity is mainly explained through geographic replacementof species or vicariance.

TABLE 1. Number of anuran species found in the Andean mountains of Colombia. * Total: number of different Andeanspecies.

2. Altitudinal ranges and species similarity along Andean elevational gradients.Because of stability of environmental temperature in the tropics, in comparison with temperature zones, it

has been considered that tropical species should have relatively restricted altitudinal ranges, such thatbetween-altitudinal fauna overlaps would be reduced (Janzen’s 1967 hypothesis).

To test this hypothesis, here we estimated altitudinal ranges for Andean anuran species by measuring thedifference between the maximum and minimum values reported in the three Colombian Andean cordilleras(Appendix 1). Then, a bar graph was made to view the frequency of these altitudinal ranges. Finally, faunalturnover along altitudinal gradients was assessed by estimating the relative faunal similarity between twoadjacent altitudinal bands, each about 500 m wide. Jaccard index of species overlap (Krebs 1989), which usesonly presence/absence data, were calculated.

Figure 2A shows that the majority of Andean anurans have narrow altitudinal ranges, less than 500 m alti-tude, others have an altitudinal range to 1000 m, but only a few species have a broad altitudinal distribution.On the other hand, the Jaccard similarity index calculated for the Andean anurans of the three Colombian Cor-dilleras is low (Table 2), and consequently shows a sudden altitudinal change between species. This indicatesthat tropical mountain anurans are mainly altitude specialists (Navas 2005), or they are delimited by altitudeor temperature (Lynch 1999). In general, they have restricted altitudinal ranges, as Janzen (1967) hypothe-sized. Janzen’s hypothesis (1967) has been very influential because he derived a simple environmental-physi-ological model predicting that tropical mountain passes would be more effective barriers to organismal

Family Cordillera Occidental Cordillera Central Cordillera Oriental *Total

Aromobatidae 1 1 2 3

Brachycephalidae 74 88 50 185

Bufonidae 15 20 15 45

Centrolenidae 26 24 18 55

Dendrobatidae 17 19 8 37

Hemiphractidae 7 12 9 21

Hylidae 10 17 25 43

Leptodactylidae 1 5 2 5

Microhylidae 1 0 1 1

Ranidae 1 1 1 1

Total 153 187 131 396

Endemic species 67 74 76 217

% Endemic species 43.7 39.5 58.0 54.7%



FIGURE 2. Altitudinal ranges (maximum minus minimum reported altitude) for anurans of the: Andean mountains ofColombia (A), Colombian lowland species (< 1000 m elevation) (B), Colombian highland species (> 3000 m elevation)(C), Henduagan mountains of China (D).



dispersion than would temperate-zone passes of equivalent altitude (Ghalambor et al. 2005). It is mainly sup-ported because environmental temperature variation in the tropics is lower than in temperate zones, so thermaltolerances for tropical ectothermic species should be narrower than for temperate species, and consequentlythey should have narrow altitudinal ranges, as it has been confirmed in this paper.

TABLE 2. Jaccard Similarity Index (JSI) for the altitudinal comparisons between Andean anurans in the three Colom-bian cordilleras.

3. Altitudinal pattern of between highland and lowland Colombian Anurans. Ghalambor et al. (2005) suggested that at high altitudes in the tropics, organisms may experience sum-

mer-like conditions during the day and winter-like conditions at night during the year. Thus, with increasingaltitude, tropical ectotherm organisms should evolve relatively broad thermal tolerances to cope with the fluc-tuating diurnal changes in temperature. In this case, Janzen’s hypothesis (1967) could be applicable to altitudi-nal gradients in the tropics, such that altitudinal distributions of highland tropical anurans should be greaterthan lowland tropical species.

Because wide altitudinal ranges for frog species exist in the Andean mountains in Colombia, we com-pared the altitudinal range between highland Andean anurans (above 3000 m altitude) and lowland anurans(below 1000 m). For this, we used data from Appendix 1, and from Acosta (2000) for lowland species near theAndean mountains, specifically from the Caribbean ecobiogeographic unit (Lynch et al. 1997), which is con-sidered to extend to the interandean mountain valleys (Lynch 2006).

Figure 2B shows that lowland species have a broader altitudinal range than do highland species (Figure2C), mostly about 1000 and 2000 m, contrary to the predicted results. Hence the hypothesis about a broaderthermal tolerance and possible wider altitudinal range for highland anurans is not supported. Because ecto-therms typically restrict activity period in unfavourable environments (Stevenson 1985) and they can look forrefuges or simply escape adverse environmental temperature conditions, a selection for a broader thermal tol-erance may not be found in these highland organisms. On the other hand, wide environmental temperaturefluctuations attributed to high mountains are likely not comparable to body temperatures at which anurans areexposed or at temperatures of their microhabitat. In this case, thermal differences between highland and low-land anurans would not be as different as was initially hypothesized. On the contrary, lowland species couldhave a broader thermal tolerance and consequently the possibility of a wide range of altitudinal distributiondue to the high fluctuations of environmental temperature where they or their embryos develop, many of themin open areas without any forest protection on warm days and cold nights.

4. Latitudinal pattern of between-altitude anuran similarity.Following Janzen’s hypothesis (1967) according to which temperate species should have broader altitudi-

nal ranges than tropical species, it is expected that similarity indices between-altitudinal fauna bands shouldbe higher for temperate species than for tropical species. Huey (1978) gathered and analyzed data of amphibi-ans and reptiles from several altitudinal transects from Santa Marta (Colombia), Costa Rica, Guatemala, Mex-

Altitudinal bands (m) JSI Cordillera Occidental JSI Cordillera Central JSI Cordillera Oriental

(1001-1500) – (1501-2000) 0.54 0.23 0.55

(1501-2000) – (2001-2500) 0.40 0.43 0.45

(2001-2500) – (2501-3000) 0.27 0.49 0.31

(2501-3000) – (3001-3500) 0.03 0.39 0.40

(3001-3500) – (3501-4000) 0.42 0.32 0.62

(3501-4000) – (4001-4500) ------ 0.40 ------

AVERAGE 0.33 0.38 0.47



ico to California (USA), and tested the prediction that if Janzen was right, then “between-altitude faunalsimilarity should vary directly with latitude”. He effectively found a low faunal similarity in the tropics, and asignificant positive correlation between-latitude and faunal similarity, which was primarily a function of spe-cies replacements rather than species dropout.

Here, we compare data of Jaccard similarity indices for anurans from the three Colombian Cordilleras(Appendix 1) with Dice’s faunal similarity index used by Huey (1978), in order to test if similarity indices forAndean anurans agree with this report. That comparison is possible because these two indices are stronglycorrelated (Southwood 1966), and also the altitudinal comparison between species within a band is about1000 m wide. According to table 3, the averages of faunal similarity for these three Colombian cordilleras areapproximately similar to the anuran faunal similarity for the tropical countries studied by Huey (1978), butnotably lower than in the temperate localities. As Huey (1978) stated, these results can be in concordance withJanzen’s hypothesis about a broader altitudinal range for temperature species, likely due to a wider thermaltolerance. On the other hand, the low similarity index can mean high turnover species along altitudinal gradi-ents, as happens in the Colombian Cordilleras.

TABLE 3. Latitudinal comparison of between-altitude anuran similarity (AS).

We also gathered the altitudinal distribution of frogs of the three Colombian Cordilleras in just one dataset, to compare with a recent study on elevation patterns of frog species of the Hengduan mountains in China(Fu et al. 2006), a latitudinal different place. To establish the latitudinal comparison between-altitude anuransimilarity for Colombia and China, we estimated the Jaccard similarity index (JSI) between two altitudinalbands of 500 m wide, starting from 1000 m elevation in both cases, and then calculated the average of JSI foreach altitudinal transect (Colombia and China). The JSI average found for anurans in the Andean Colombianmountains was 0.43, while the JSI average for frogs of the Hengduan mountains in China was 0.60 (Table 4).Additionally, we compared the JSI average of frogs along Hehgduan mountains in China, from 400 to 5,000m, with another transect of similar altitudinal gradient in the Cordillera Central of Colombia (Bernal et al.2005), from 440 m to 4,300 m (Table 5). The average of this JSI was 0.38, which is lower than the JSI averageobtained for the China transect (0.55). Thus, these results agree with that mentioned above and with Huey’swork (1978). Because similarity indices between altitudinal bands in China are higher than in Colombia, itmeans that altitudinal distribution of these species are broader than those of Andean Colombian frogs. In fig-ure 2A it can be seen that effectively many of the Colombian Andean frogs have a restricted altitudinal range,between 0 to 500 m, in comparison with frogs of the China mountains which exhibit a broader elevationrange, mainly between 1001-1500 m (Fig. 2D).

In conclusion, data shown here agree with Huey’s (1978) reports of a latitudinal pattern of between-alti-tude faunal similarity, and consequently with their interpretation about Janzen’s hypothesis (1967): tropical

Locality Average AS Data obtained from:

Cordillera Occidental 0.33 This work

Cordillera Central 0.38 This work

Cordillera Oriental 0.47 This work

Tilarán, Costa Rica 0.15 Huey (1978)

Alta Verapaz, Guatemala 0.28 Huey (1978)

Sinaloa, México 0.46 Huey (1978)

Gómez Farias, México 0.70 Huey (1978)

Michoacán, México 0.60 Huey (1978)

Yosemite, California 0.84 Huey (1978)

Lassen Park, California 0.95 Huey (1978)



countries have a narrower fluctuation of their environmental temperature, lower than do temperate countries,so their ectothermic species should exhibit a restricted thermal tolerance and consequently a narrow altitudi-nal distribution.

TABLE 4. Comparison of between-altitude anuran similarity for two different latitudinal mountains. JSI: Jaccard Simi-larity Index.

TABLE 5. Latitudinal comparison of between-altitude anuran similarity for two similar transects. JSI: Jaccard SimilarityIndex.

Acknowledgements

MHB is very grateful to Instituto Colombiano para el Desarrollo de la Ciencia y la Tecnología, COLCIEN-CIAS and The Universidad del Tolima for the financial help to his PhD studies. This work was partially sup-ported by Fondo de Investigaciones de la Universidad del Tolima (project number 310105).

References

Acosta-Galvis, A.R. (2000) Ranas, salamandras y caecilias (Tetrapoda: Amphibia) de Colombia. Biota Colombiana, 1,289–319.

Bernal, M.H., Luna, V.F., Gallego, O. & Quevedo, A. (2007) A new species of poison frog (Amphibia: Dendrobatidae)from the Andean mountains of Tolima, Colombia. Zootaxa, 1638, 59–68.

Bernal, M.H., Páez, C.A. & Vejarano, M.A. (2005) Composición y distribución de los anfibios de la Cuenca del RíoCoello (Tolima), Colombia. Actualidades Biológicas, 27 (82), 87–92.

Bruhl, C.A., Mohamed, V. & Linsenmair, K.E. (1999) Altitudinal distribution of leaf litter ants along a transect in pri-mary forests on Mount Kinabalu, Sabah, Malaysia. Journal of Tropical Ecology, 15, 265–277.

Duellman, W.E. (1988) Patterns of species diversity in anuran amphibians in the American tropics. Annals of the Mis-souri Botanical Garden, 75 (1), 79–104.

Duellman, W.E. (1999) Distribution patterns of amphibians in South America. In: W.E. Duellman (Ed). Patterns of Dis-

Altitudinal bands (m) JSI for the Hehgduan mountains (China) JSI for the Andean Mountains (Colombia)

(1001-1500) – (1501-2000) 0.72 0.46

(1501-2000) – (2001-2500) 0.69 0.47

(2001-2500) – (2501-3000) 0.53 0.39

(2501-3000) – (3001-3500) 0.55 0.34

(3001-3500) – (3501-4000) 0.50 0.45

AVERAGE 0.60 0.43

Altitudinal bands (m) JSI for the Hehgduan mountains in China JSI for the Coello river watershed in Colombia

(0-500) – (501-1000) 0.20 0.69

(500-1000) - (1001-1500) 0.67 0.54

(1001-1500) – (1501-2000) 0.72 0.17

(1501-2000) – (2001-2500) 0.69 0.16

(2001-2500) – (2501-3000) 0.53 0.33

(2501-3000) – (3001-3500) 0.55 0.16

(3001-3500) – (3501-4000) 0.50 0.60

AVERAGE 0.55 0.38



tribution of Amphibians. A global Perspective. The John Hopkins University Press, Baltimore, pp. 255–328.Frost D.R. (2007) Amphibian Species of the World: An Online Reference. Version 5.1. Available from: http://

research.amnh.org/herpetology/amphibia/index.html (accessed 30 April, 2008). American Museum of Natural His-tory, New York.

Fu, C., Hua, X., Li, J., Chang, Z., Pu, Z. & Chen, J. (2006) Elevation patterns of frog species richness and endemic rich-ness in the Hengduan Mountains, China: geometric constraints, area and climate effects. Ecogeography, 29, 919–927.

Ghalambor, C.K., Huey, R.B., Martin, P.R., Tewksbury, J.J. & Wang, G. (2005) Are mountain passes higher in the trop-ics?. Janzen´s hypothesis revisited. Integrative and Comparative Biology, 46, 5–17.

Hedges, B.S., Duellman, W.E. & Heinicke, M.P. (2008) A replacement name for Isodactylus Hedges, Duellman, andHeinicke. Zootaxa, 1975, 67–68.

Huey, R.B. (1978) Latitudinal pattern of between-altitudinal faunal similarity: mountains might be “higher” in the trop-ics. The American Naturalist, 112, 225–229.

Janzen, D.H. (1967) Why mountain passes are higher in the tropics. The American Naturalist, 101, 233–249.Krebs, C.J. (1989) Ecological methodology. Harper & Row Publishers, New York. Lynch, J.D. (1998) New species of Eleutherodactylus from the cordillera occidental of western Colombia with synopsis

of the distributions of species in western Colombia. Revista de la Academia Colombiana de Ciencias Exactas Físi-cas y Naturales, 22 (82), 117–148.

Lynch, J.D. (1999) Ranas pequeñas, la geometría de evolución, y la especiación en los Andes Colombianos. Revista de laAcademia Colombiana de Ciencias Exactas Físicas y Naturales, 23(86), 143–159.

Lynch, J.D. (2005) Discovery of the richest frog fauna in the world. An exploration of the forests to the north of Leticia.Revista de la Academia Colombiana de Ciencias Exactas Físicas y Naturales, 29 (113), 581–588.

Lynch, J.D. (2006) The tadpoles of frogs and toads found in the lowlands of northern Colombia. Revista de la AcademiaColombiana de Ciencias Exactas Físicas y Naturales, 30 (116), 443–458.

Lynch, J.D., Ruiz-Carranza, P.M. & Ardila-Robayo, M.C. (1997) Biogeographic patterns of Colombian frogs and toads.Revista de la Academia Colombiana de Ciencias Exactas Físicas y Naturales, 21 (80), 237–248.

Lynch, J.D. & Suárez-Mayorga, A. (2004) Anfibios en el chocó biogeográfico. In: Rangel, J.O (Ed). Diversidad BióticaIV: El chocó biogeográfico/Costa Pacifica. Unibiblios. Bogotá, pp. 633–667.

McCoy, E.D. (1990) The distribution of insects along elevational gradients. Oikos, 58, 313–322.Navas, C.A. (2003) Herpetological diversity along Andean elevational gradients: links with physiological ecology and

evolutionary physiology. Comparative Biochemistry and Physiology Part A, 133, 469–485.Navas, C.A. (2005) Patterns of distribution of anurans in high Andean tropical elevations: insights from integrating bio-

geography and evolutionary physiology. Integrative and Comparative Biology, 46, 82–91.Olson, D.M. (1994) The distribution of leaf litter invertebrates along a Neotropical altitudinal gradient. Journal of Tropi-

cal Ecology, 10, 129–150.Patterson, B.D., Pacheco, V. & Solari, S. (1996) Distribution of bats along an elevational gradient in the Andes of south-

eastern Perú. Journal of Zoology, 240, 637–658.Poynton, J.C. (2003) Altitudinal species turnover in southern Tanzania shown by anurans: some geographical consider-

ations. Systematics and Biodiversity, 1, 117–126.Rueda, J.V., Rada, M., Sánchez, S., Velásquez, A. & Quevedo, A. (2006) Two new and excepcional poison dart frog of

the genus Dendrobates (Anura: Dendrobatidae) from the Northeastern flank of the cordillera Central of Colombia.Zootaxa, 1259, 39–54.

Ruiz-C., P.M., Ardila, M.C. & Lynch, J.D. (1996) Lista actualizada de la fauna de amphibia de Colombia. Revista de laAcademia Colombiana de Ciencias Exactas Físicas y Naturales, 20 (77), 365–415.

Savage, J.M., Myers, C.W. (2002) Frogs of the Eleutherodactylus biporcatus group (Leptodactylidae) of Central Amer-ica and Northern South America, including rediscovered, resurrected, and new taxa. American Museum Novitates,3357, 1–47.

Southwood, T.R.E. (1966) Ecological methods. Methuen, London.Stevenson, R.D. (1985) The relative importance of behavioural and physiological adjustments controlling body tempera-

ture in terrestrial ectotherms. The American Naturalist, 126, 362–386.Wiens, J.J., Kuczynski, C.A., Duellman, W.E. & Reeder T.W. (2007) Loss and re-evolution of complex life cycles in

marsupial frogs: does ancestral trait reconstruction mislead? Evolution, 61–8, 1886–1899.



Appendix 1

Check list and altitudinal distribution of the anurans described in the three Andean cordilleras. * Lowland spe-cies found in the Andes. E= Endemic species. + Full elevational range in Colombia, not only in the AndeanCordilleras. Classification follows Frost et al. (2007) with modifications by Wiens et al. (2007) and Hedges etal. (2008).

Family/Species CordilleraOccidental

CordilleraCentral

CordilleraOriental

+ Eleva-tional range

Family Aromobatidae EAllobates picachos Ardila-R., Acosta-G. & Coloma, 2000 1500-1600 1500-1600

EAnomaloglossus atopoglossus Grant, Humphrey, & Myers, 1997 1800-2260 1800-2260

Reobates palmatus (Werner, 1899) 1350-1500 1000-2520 350-2520

Family Brachycephalidae

Atopophrynus syntomopus Lynch & Ruiz-C. 1982 2780 2780

*Craugastor fitzingeri (Schmidt, 1857) 1070 2-1070

*Craugastor longirostris (Boulenger, 1898) 1060-1180 1360 0-1360

*Craugastor opimus (Savage and Myers, 2002) 1060 0-1060

*Craugastor raniformis (Boulenger, 1896) 1000-1510 1150 0-1510EHypodactylus adercus Lynch, 2003 2280 2280

Hypodactylus babax Lynch, 1989 1200-2250 1600-1970 1200-2250

Hypodactylus dolops Lynch & Duellman, 1980 1950 1150-1950 940-1950

Hypodactylus elassodiscus Lynch, 1973 2300-2900 2300-2900EHypodactylus latens Lynch, 1989 2720-3350 2720-3350

Hypodactylus mantipus (Boulenger, 1908) 1000-2400 1840-2100 1000-2100EPhrynopus adenobrachius Ardila-R., Ruiz-C. & Barrera-R., 1996 3100-3400 3100-3400

EPhrynopus columbianus (Werner, 1899) 1000-1300 1000-1300

EPhrynopus nanus (Going & Cochran, 1963) 3000-3850 3000-3850

EPristimantis acatallelus Lynch & Ruiz-C., 1983 1000-2680 1000-2680

*Pristimantis achatinus (Boulenger, 1898) 1000-1880 1020-1400 10-1880EPristimantis actinolaimus Lynch & Rueda-A., 1998 1940-2000 1940-2000

EPristimantis acutirostris Lynch, 1984 1740-2300 1740-2300

EPristimantis aemulatus Ruiz-C, Lynch, & Ardila-R., 1997 1410-1430 1410-1430

EPristimantis affinis (Werner, 1899) 2600-3100 2600-3100

EPristimantis alalocophus Roa-T. & Ruiz-C., 1991 2650-3800 2650-3800

EPristimantis angustilineatus Lynch, 1998 1700-2300 1700-2300

Pristimantis anolirex Lynch, 1983 1800-3550 1800-3550

*Pristimantis anthrax Lynch, 2001 1200 700-1200

Pristimantis apiculatus Lynch & Burrowes, 1990 1700-2020 1700-2020

Pristimantis appendiculatus (Werner, 1894) 2020 700-2020EPristimantis aurantiguttatus Ruiz-C., Lynch, & Ardila-R., 1997 1000-1940 1000-1940

EPristimantis bacchus Lynch, 1984 1700-2400 1700-2400

EPristimantis baiotis Lynch, 1998 1780-1940 1780-1940

EPristimantis batrachites Lynch, 2003 2180-2250 2180-2250



EPristimantis bellona Lynch, 1992 1000-1960 1000-1960

EPristimantis bernali Lynch, 1986 2530-2560 2530-2560

EPristimantis bicolor Rueda-A. & Lynch, 1983 1700-2240 1700-2240

EPristimantis bogotensis (Peters, 1863) 2410-3520 2410-3520

Pristimantis boulengeri Lynch, 1981 2200-2800 1750-3200 1750-3200

Pristimantis brevifrons Lynch, 1981 1430-2610 2460-2700 1430-2700

Pristimantis buckleyi (Boulenger, 1882) 3050 2600-3500 2600-3500

Pristimantis cabrerai Cochran & Goin, 1970 1140-1940 2000-2450 1140-2450EPristimantis cacao Lynch, 1992 2190-2600 2190-2600

EPristimantis calcaratus (Boulenger, 1908) 1700-2600 1700-2600

EPristimantis capitonis Lynch, 1998 2500-2800 2500-2800

*Pristimantis caprifer Lynch, 1977 1230 100-1230EPristimantis carlossanchezi, Arroyo, 2007 2400-2550 2400-2550

EPristimantis carranguerorum Lynch, 1994 1080-2060 980-2060

*Pristimantis caryophyllaceus (Barbour, 1928) 1070 50-1070

Pristimantis celator Lynch, 1976 1780-2600 1780-2600

*Pristimantis chalceus (Peters, 1873) 1000-1900 1780 10-1900

Pristimantis chloronotus Lynch, 1969 1900-3220 1900-3220

Pristimantis chrysops Lynch & Ruiz-C, 1996 1900-2150 900-2150

Pristimantis colomai Lynch & Duellman, 1997 1055-1120 1055-1120EPristimantis colonensis Mueses, 2007 2200-2750 2200-2750

EPristimantis cuentasi Lynch, 2003 2800 2800

Pristimantis curtipes (Boulenger, 1882) 2750-4400 2750-4400

Pristimantis degener Lynch & Duellman, 1997 1020-1400 1020-1400EPristimantis deinops Lynch, 1996 1230-2600 1230-2600

EPristimantis diaphonus Lynch, 1986 1180-1300 1180-1300

EPristimantis diogenes Lynch & Ruiz-C., 1996 1470-2100 1470-2100

EPristimantis dorsopictus Rivero & Serna, 1988 2000-2780 2000-2780

EPristimantis douglasi Lynch, 1996 1630-2670 1630-2670

Pristimantis duellmani Lynch, 1980 1800-2190 1780 1780-2190EPristimantis duende Lynch, 2001 3300-3600 3300-3600

EPristimantis elegans (Peters, 1863) 2600-3650 2600-3650

EPristimantis epacrus Lynch & Suárez-M., 2000 1000-1660 940-1660

Pristimantis eremitus Lynch, 1980 1780 1780

Pristimantis eriphus Lynch & Duellman, 1980 2100-2750 2100-2750

Pristimantis erythropleura (Boulenger, 1896) 1140-2470 1850-2600 980-2600EPristimantis factiosus Lynch & Rueda-A., 1999 1840-2150 1840-2150

EPristimantis fallax Lynch & Rueda-A., 1999 1180-1950 780-1950

EPristimantis frater (Werner, 1899) 1000-3000 900-3000

EPristimantis fetosus (Lynch, Rueda, 1998) 1800-2650 1800-2650

*Pristimantis gaigeae (Dunn, 1931) 1040-1200 10-1200

Pristimantis gladiator Lynch, 1976 2400-2750 2400-2750



EPristimantis grandiceps Lynch, 1984 2200-2400 2200-2400

*Pristimantis gularis (Boulenger, 1898) 1060-1770 1055 0-1770

Pristimantis hectus Lynch & Burrowes, 1990 1990-2610 1780 1780-2610EPristimantis helvolus Lynch & Rueda- A., 1998 1800-2000 1800-2000

EPristimantis hernandezi Lynch & Ruiz-C., 1983 2600 2600

Pristimantis illotus Lynch & Duellman, 1997 1230-1580 1230-1580EPristimantis ixalus Lynch, 2003 1300-1700 1300-1700

EPristimantis jaimei Lynch, 1992 1200-1580 1200-1580

EPristimantis johannesdei Rivero & Serna, 1988 1410-1800 1410-1800

EPristimantis jorgevelosai Lynch, 1994 1900-2200 1900-2200

EPristimantis juanchoi Lynch, 1996 1580-1990 1580-1990

EPristimantis jubatus Garcia & Lynch 2006 2550-2750 2550-2750

EPristimantis kelephas Lynch, 1998 2140-2610 2140-2610

*Pristimantis labiosus Lynch, Ruiz- C. & Ardila-R., 1994 1055 30-1055EPristimantis lasalleorum Lynch, 1995 3780-3850 3780-3850

Pristimantis laticlavius Lynch & Burrowes, 1990 1700-2020 1700-2020

Pristimantis lemur Lynch and Rueda-A., 1988 1800-1950 1800-1950

Pristimantis lentiginosus Rivero, 1984 1390-2300 1390-2300

Pristimantis leoni Lynch, 1976 2060-3400 2060-3400EPristimantis leptolophus Lynch, 1980 2400-3300 2400-3300

Pristimantis leucopus Lynch, 1976 2300-2900 2300-2900EPristimantis lichenoides Lynch & Rueda-A., 1997 2000-2450 2000-2450

Pristimantis loustes Lynch, 1979 1200 1200EPristimantis lutitos Lynch, 1984 1750 1750

EPristimantis lynchi Duellman & Simmons, 1977 1600-3580 1600-3580

EPristimantis maculosus Lynch, 1991 2000-2900 2000-2900

EPristimantis mars Lynch & Ruiz-C., 1996 1760-1790 1760-1790

*Pristimantis medemi Lynch, 1994 1000-2400 450-2400EPristimantis merostictus Lynch, 1984 2400 2400

EPristimantis miyatai Lynch, 1984 1720-2400 1720-2400

EPristimantis mnionaetes Lynch, 1998 3060-3080 3060-3080

EPristimantis molybrignus Lynch, 1986 1100-2350 800-2350

EPristimantis myersi (Goin & Cochran, 1963) 2800-3500 2800-3500

EPristimantis myops Lynch, 1998 1500-2250 1500-2250

EPristimantis nervicus Lynch, 1994 3100-3870 3100-3870

EPristimantis nicefori Cochran & Goin, 1970 2770-4180 2770-4180

EPristimantis obmutescens Lynch, 1980 2800-3500 2800-3500

Pristimantis ocellatus Lynch & Burrowes, 1990 1050-1580 1780 1050-1780

Pristimantis orpacobates Lynch, Ruiz-C., & Ardila-R., 1994 1140-2000 700-2000

Pristimantis padrecarlosi Mueses-C., 2006 1500–1950 1500-1950EPristimantis paisa Lynch & Ardila-R., 1999 1800-3100 1800-3100

Pristimantis palmeri (Boulenger, 1912) 1050-2440 1950 -2050 980-2440



EPristimantis parectatus Lynch & Rueda-A., 1998 1800-3100 1800-3100

*Prystimantis parvillus Lynch, 1976 1360 650-1360EPristimantis penelopus Lynch & Rueda, 1999 1010-1500 1010-1500

EPristimantis peraticus Lynch, 1980 2600-3460 2600-3460

EPristimantis permixtus Lynch, Ruiz-C. & Ardila-R., 1994 1950-3700 1950-3700

Pristimantis petersi Lynch, 1991 1950-2750 1410 1410-2750EPristimantis phalarus Lynch, 1998 2160-2250 2160-2250

Pristimantis phragmipleuron Rivero & Serna, 1988 2380 1800 1800-2380EPristimantis piceus Lynch, Ruiz-C. & Ardila-R., 1996 2600-3200 2600-3200

EPristimantis platychilus Lynch, 1996 1580-2600 1580-2600

EPristimantis polemistes Lynch & Ardila-R., 2004 2300-2320 2300-2320

EPristimantis polychrus Ruiz-C., Lynch, & Ardila-R., 1997 1410-1540 1410-1540

Pristimantis prolixodiscus Lynch, 1978 1800-2550 1800-2550EPristimantis ptochus Lynch, 1998 2080-2250 2080-2250

Pristimantis pugnax Lynch, 1973 2100-3300 2040-2380 2040-3300EPristimantis quantus Lynch, 1998 2110-2250 2110-2250

Pristimantis quinquagesimus Lynch & Trueb, 1980 1780-2600 1780-2600EPristimantis racemus Lynch, 1980 3030-3570 3030-3570

EPristimantis reclusus Lynch, 2003 2800 2800

EPristimantis renjiforum Lynch, 2000 2000-2800 2000-2800

EPristimantis repens Lynch, 1984 3150-3720 3150-3720

EPristimantis restrepoi Lynch, 1996 1790-2400 1790-2400

EPristimantis ruedai Ruiz-C., Lynch, & Ardila-R., 1997 1000-1870 1000-1870

Pristimantis sanguineus Lynch, 1998 1030-2130 620-2130EPristimantis satagius Lynch, 1995 3300-3850 3300-3850

EPristimantis savagei Pyburn & Lynch, 1981 1000-3000 600-3000

EPristimantis scoloblepharus Lynch 1991 2420-2800 2420-2800

Pristimantis scolodiscus Lynch & Burrowes, 1990 1780 1780EPristimantis scopaeus Lynch, Ruiz- C. & Ardila-R., 1996 3580-3600 3580-3600

EPristimantis signifer Ruiz-C., Lynch, & Ardila-R., 1997 1850-1860 1850-1860

EPristimantis silverstonei Lynch & Ruiz-C., 1996 1600-2500 1600-2500

EPristimantis simoteriscus Lynch, Ruiz-C. & Ardila-R., 1997 3350-3800 3350-3800

EPristimantis simoterus Lynch, 1980 3200-4300 3200-4300

EPristimantis siopelus Lynch & Burrowes, 1990 1780 1780

EPristimantis spilogaster Lynch, 1984 2200-2400 2200-2400

EPristimantis suetus Lynch & Rueda-A., 1998 1850-2780 1850-2780

EPristimantis sulculus Lynch & Burrowes, 1990 1780 1780

EPristimantis supernatis Lynch, 1980 1850-3200 1850-3200

EPristimantis susaguae Rueda-A., Lynch & Galvis, 2003 2530-2900 2530-2900

EPristimantis taciturnus Lynch & Suárez-M., 2003 2400-2700 2400-2700

*Pristimantis taeniatus (Boulenger, 1912) 1070-1880 1160-2150 1050-1120 0-2150EPristimantis tamsitti Cochran & Goin, 1970 1230-2380 1230-2380



Pristimantis thectopternus Lynch, 1975 1700-2540 1780-2300 1700-2540

Pristimantis thymelensis Lynch, 1972 3220-4150 3220-4150

Pristimantis torrenticola Lynch & Rueda-A. 1998 1800-2450 1800-2450EPristimantis tribulosus Lynch & Rueda-A., 1997 1800-2450 1800-2450

Pristimantis turbernasus Rivero, 1984 1950-2300 1950-2300EPristimantis uisae Lynch, 2003 2400-2700 2400-2700

Pristimantis unistrigatus (Günther, 1859) 2000-3230 2000-3230EPristimantis uranobates Lynch, 1991 2000-3600 2000-3600

EPristimantis veletis Lynch & Rueda-A, 1997 1850-2150 1850-2150

EPristimantis vicarius Lynch & Ruiz-C., 1983 2400-3300 2400-3300

*Pristimantis viejas Lynch & Rueda, 1999 1000-1375 1100-1502 565-1502EPristimantis viridicans Lynch, 1977 2000-2680 2000-2680

EPristimantis viridis Ruiz-C., Lynch, & Ardila-R., 1997 1500-1960 1500-1960

Pristimantis w-nigrum (Boettger, 1892) 1050-3000 1840-2800 1720-2450 800-3000EPristimantis xeniolum Lynch, 2001 3300-3600 3300-3600

EPristimantis xestus Lynch, 1995 4050 4050

EPristimantis xylochobates Lynch & Ruiz-C., 1996 2100-2250 2100-2250

E Pristimantis zoilae, Mueses, 2007 2060-2550 2060-2550

EPristimantis zophus Lynch & Ardila-R., 1999 2030-2680 2030-2680

Strabomantis anatipes (Lynch & Myers, 1983) 1600 100-1600

*Strabomantis bufoniformis (Boulenger, 1896) 1030-1780 20-1780EStrabomantis cadenai Lynch, 1986 1900 1900

Strabomantis cerastes Lynch, 1975 1000-2250 850-2250

Strabomantis cheiroplethus (Lynch, 1990) 1380-1900 800-1900

Strabomantis cornutus (Jiménez de la Espada, 1871) 1150-1800 1150-1800EStrabomantis ingeri (Cochran & Goin, 1961) 1700-3320 1700-3320

EStrabomantis necopinus Lynch, 1997 1800-2150 1800-2150

Strabomantis ruizi Lynch, 1981 1500-2000 1900-2000 1500-2000

*Strabomantis zygodactylus Lynch & Myers, 1983 1030-1490 230-1490

Family Bufonidae EAndinophryne atelopoides (Lynch & Ruiz-C., 1981) 2190 2190

EAtelopus angelito Ardila-Robayo & Ruiz-C., 1998 2900 -3000 2900-3000

EAtelopus carauta Ruiz-C. & Hernández-C., 1978 1140-1960 1140-1960

EAtelopus chocoensis Lötters, 1992 1900-2250 1900-2250

Atelopus ebenoides Rivero, 1963 2500-2600 2500-3700 2800-3650 2500-3700

Atelopus eusebianus Rivero & Granados-D., 1993 1050-1180 2780-3250 1050-3250EAtelopus famelicus Rivero & Morales, 1995 1050-1580 1050-1580

EAtelopus farci Lynch, 1993 2090 2090

EAtelopus galactogaster Rivero & Serna, 1993 1300-1800 1300-1800

EAtelopus guitarraensis Osorno-M., Ardila-R. & Ruiz-C., 2001 3400 3400

EAtelopus ignescens (Cornalia, 1849) 2000-3220 2000-3220

Atelopus longirostris Cope, 1868 1000-1200 800-1200



EAtelopus lozanoi Osorno-M., Ardila-R. & Ruiz-C., 2001 2340-3540 2340-3540

Atelopus lynchi Cannatella, 1981 1000-1410 800-1410EAtelopus mandingues Osorno-Muñoz, Ardila-R. & Ruiz-C., 2001 2580-3050 2580-3050

EAtelopus minutulus Ruiz-C., Hernández-C. & Ardila-R., 1988 1375-1560 1375-1560

EAtelopus mittermeieri Acosta-G., Rueda-A., Velásquez-Á., Sánchez-P. & Peña P., 2006

2525 2525

EAtelopus monohernandezii Ardila-R., Osorno-M. & Ruiz-C., 2002

2000-2200 2000-2200

EAtelopus muisca Rueda-A. & Hoyos, 1992 2800-3350 2800-3350

EAtelopus nicefori Rivero, 1963 1800-2670 1800-2670

EAtelopus pedimarmoratus Rivero, 1963 2600-3100 2600-3100

EAtelopus pictiventris Kattan, 1986 2600 2600

EAtelopus quimbaya Ruiz-C. Osorno-M. 1994 1650-2940 1650-2940

EAtelopus sernai Ruiz-C.& Osorno-M. 1994 2800-3100 2800-3100

EAtelopus simulatus Ruiz-C.& Osorno-M. 1994 2630-3000 2630-3000

EAtelopus sonsonensis Vélez-R.& Ruiz-C. 1997 1500 1500

EAtelopus subornatus Werner, 1899 2000-3020 2000-3020

Osornophryne antisana (Hoogmoed, 1987) 3400-3450 3400-3450

Osornophryne bufoniformis (Peracca, 1904) 2800-4700 2800-4700

Osornophryne guacamayo Hooogmoed, 1987 2410-2500 2410-2500EOsornophryne percrassa Ruiz-C.& Hernández-C. 1976 2700-3700 2700-3700

Osornophryne talipes Cannatella, 1986 2880-3630 2880-3630

*Rhaebo glaberrimus Günther, 1869 1000-1300 300-1300

*Rhaebo haematiticus Cope 1862 1050 75-1050

*Rhaebo hypomelas Boulenger, 1913 1480-1600 60-1600ERhinella cristinae Velez & Ruiz-C. 2002 1350-1470 1350-1470

*Rhinella granulosa (Spix, 1824) 0-1500ERhinella lindae Rivero & Castaño, 1990 1600-1800 1600-1800

ERhinella macrorhina Trueb, 1971 1840-2940 1840-2940

*Rhinella marina (Linnaeus, 1758) 1000-1840 1000-1385 1000-2000 0-2000ERhinella nicefori (Cochran & Goin, 1970) 2640-2800 2640-2800

ERhinella rostrata (Noble, 1920) 1890-2420 1890-2420

ERhinella ruizi Grant, 2000 2380-3100 2380-3100

ERhinella truebae Lynch & Renjifo, 1990 1840-2420 1840-2420

*Rhinella sternosignata (Günter, 1858) 1200-1700 1050-1820 400-1820

Family Centrolenidae ECentrolene acanthidiocephalum (Ruiz-C. Lynch, (1989) 1700-2200 1700-2200

Centrolene andinum (Rivero, 1968) 1450-2240 1450-2240ECentrolene antioquiense (Noble, 1920) 1850-2450 1850-2450

Centrolene audax (Lynch & Duellman, 1973) 1900 1900

Centrolene bacatum Wild, 1994 1900 1900

Centrolene buckleyi (Boulenger, 1882) 1900-3450 2020-3170 1650-3550 1650-3550

Centrolene geckoideum Jiménez de la Espada, 1872 1940-2430 2000-2450 2000-3024 1940-3024



Centrolene grandisonae (Cochran & Goin, 1970) 1230-2230 1900-2080 1230-2230ECentrolene guanacarum Ruiz-C.& Lynch, 1995 1800-1900 1800-1900

Centrolene heloderma (Duellman, 1981) 1890-2450 1890-2450ECentrolene huilense Ruiz-Carranza & Lynch, 1995 2190 2190

ECentrolene hybrida Ruiz-C.& Lynch, 1991 1410-2000 1410-2000

*Centrolene ilex (Savage, 1967) 1030-1060 50-1060

Centrolene lynchi (Duellman, 1980) 1840 1840ECentrolene medemi (Cochran & Goin, 1970) 1150-1750 980-1750

ECentrolene notostictum Ruiz-C.& Lynch, 1991 1730-2700 1730-2700

ECentrolene paezorum Ruiz-C. Hernández-C.& Ardila-R. 1986 3030 3030

Centrolene peristictum (Lynch & Duellman, 1973) 1780-1820 1970-2080 1970-2080ECentrolene petrophilum Ruiz-C.& Lynch, 1991 1180-2020 1180-2020

*Centrolene prosoblepon (Boettger, 1892) 1030-1960 1240-1616 1760 80-1960

Centrolene quindianum Ruiz-C.& Lynch, 1995 1400-2060 1900-2050 1400-2060

Centrolene robledoi Ruiz-C.& Lynch, 1995 1940-2050 2000-2800 1940-2800ECentrolene sanchezi Ruiz-C. & Lynch, 1991 2190 2190

Centrolene scirtetes (Duellman & Burrowes, 1989) 1900 1900ECochranella adiazeta Ruiz-C. & Lynch, 1991 1120-2060 1120-2060

ECochranella daidalea Ruiz-C. & Lynch, 1991 1630-2060 1630-2060

*Cochranella euknemos (Savage & Starrett, 1967). 1030-1940 100-1940

*Cochranella flavopunctata (Lynch & Duellman, 1973) 1000-1650 70-1650ECochranella megistra (Rivero, 1985) 1700-2100 1700-2100

*Cochranella punctulata Ruiz-C. & Lynch, 1995 1100 620-1100

Cochranella savagei Ruiz-C. & Lynch, 1991 1230-1800 1400-2050 1230-2050ECochranella solitaria Ruiz-C. & Lynch, 1991 1410 1410

*Cochranella spinosa (Taylor, 1949) 1030-1060 100-1060ECochranella susatamai Ruiz.C. & Lynch, 1995 1100-1650 450-1650

Cochranella xanthocheridia Ruiz-C. & Lynch, 1995. 1700-2060 800-2060

*Hyalinobatrachium aureoguttatum (Barrera-R. & Ruiz-C., 1989) 1030-1780 45-1780

*Hyalinobatrachium colymbiphyllum (Taylor, 1949) 1380-1880 1350-1650 100-1880EHyalinobatrachium esmeralda Ruiz-C. & Lynch, 1998 1600-1750 1600-1750

*Hyalinobatrachium fleischmanni (Boettger, 1893) 1100 60-1100EHyalinobatrachium ibama Ruiz-C. & Lynch, 1998 1660-2050 1660-2050

ENymphargus armatus Lynch & Ruiz-C., 1996 2140-2160 2140-2160

Nymphargus chami Ruiz-C. & Lynch, 1995. 1030-1280 800-1280E Nymphargus cristinae Ruiz-C. & Lynch, 1995 1480-2490 1480-2490

ENymphargus garciae Ruiz-C. & Lynch, 1995 1940-2600 1940-2600

Nymphargus griffithsi Goin, 1961. 1000-2430 1900-2450 420-2450ENymphargus ignotus (Lynch, 1990) 1280-2000 1280-2000

ENymphargus luminosus Ruiz-C. & Lynch, 1995 1140-1430 1140-1430

ENymphargus luteopunctatus Ruiz-Carranza & Lynch, 1996 1200-1500 1200-1500

ENymphargus nephelophila Ruiz-Carranza & Lynch, 1991 1660-2190 1660-2190



ENymphargus oreonympha Ruiz-Carranza & Lynch, 1991 2040-2380 2040-2380

ENymphargus posadae Ruiz-C. & Lynch, 1995 1940-2800 1940-2800

Nymphargus prasinus (Duellman, 1981) 1180-1510 900-1510ENymphargus rosada Ruiz-C. & Lynch, 1997 1430-2450 1430-2450

Nymphargus ruizi Lynch, 1993 2190-2850 2190-2850ENymphargus spilotus Ruiz-C. & Lynch, 1997 1850-1950 1850-1950

Family Dendrobatidae EAmeerega andina Myers and Burrowes 1987 2020 700-2020

Colostethus agilis Lynch & Ruiz-C, 1985 1280-2600 1280-2600EColostethus alacris Rivero & Granados-D., 1990 1400 1400

Colostethus brachistriatus Rivero & Serna, 1986 1500 1500EColostethus dysprosium Rivero & Serna, 2000 1300-2300 1300-2300

Colostethus fraterdanieli Silverstone, 1971 1550-2750 1800-2600 650-2750EColostethus furviventris Rivero & Serna 1991 1500 1500

Colostethus mertensi (Cochra & Goin, 1964) 2500-2610 2100-2610 2100-2610

*Colostethus pratti (Boulenger, 1899) 1600 110-1600E“Colostethus” ramirezi Rivero & Serna, 1995 1300-1400 1300-1400

EColostethus thorntoni (Cochran & Goin, 1970) 1480-2500 1480-2500

EColostethus ucumari Grant, 2007 2100-2500 2100-2500

EColostethus yaguara Rivero & Serna, 1991 1575 1575

*Dendrobates truncatus (Cope, 1861) 1000-1200 1600 350-1600

Hyloxalus abditaurantius (Silverstone, 1975) 1280-2060 1850-2110 1280-2110EHyloxalus borjai Rivero & Serna, 1995 1000-1500 1000-1500

Hyloxalus breviquartus Rivero & Serna, 1986 1700-1800 1950-2265 1700-2265EHyloxalus edwardsi Lynch, 1982 3070-3250 3070-3250

EHyloxalus fascianigrus Grant & Castro-H., 1998 1470-1960 1470-1960

Hyloxalus lehmanni Silverstone, 1971 2060 2060EHyloxalus pinguis Rivero & Granados-Díaz 2995 2995

Hyloxalus pulchellus (Jimenez de la Espada, 1875) 2000 2000EHyloxalus ramosi (Silverstone, 1971) 1240 1240

EHyloxalus ruizi Lynch, 1982 2410-2640 2410-2640

EHyloxalus saltuarius Grant & Ardila-R., 2002 1200-1600 1200-1600

EHyloxalus subpunctatus (Cope, 1899) 1750-4020 1750-4020

Hyloxalus vergeli (Hellmich, 1940) 1200-2100 1200-2100

*Oophaga lehmanni Myers & Daly, 1976. 1200 680-1200

*Phyllobates bicolor Duméril & Bibron, 1841. 1040-1580 50-1580

Ranitomeya bombetes Myers & Daly, 1980 1580-2270 1900-2050 1580-2270ERanitomeya daleswansoni Rueda-A., Rada, Sánchez-P., Velásquez-A. & Quevedo, 2006.

1800-2000 1800-2000

ERanitomeya dorisswansonae Rueda-A., Rada, Sánchez-P., Velásquez-A. & Quevedo, 2006

1780 1780

Ranitomeya opisthomelas Boulenger, 1899 1000-2200 1500-2640 1000-2640ERanitomeya tolimense Bernal, Luna, Gallego & Quevedo, 2007 1810 1810



ERanitomeya virolinensis Ruiz-C., & Ramirez-P., 1992) 1100-2200 1100-2200

ESilverstoneia erasmios Rivero & Serna 2000 1500-2000 1500-2000

*Silverstoneia nubicola (Dunn, 1924) 1540-1950 80-1950

Family Hemiphractidae

Cryptobatrachus fuhrmanni (Peracca, 1914) 1020-1675 1100-1680 900-1680ECryptobatrachus nicefori Cochran & Goin, 1970 1500 520-1500

Flectonotus pygmaeus (Boettger, 1893) 1140-1500 735-1500

Gastrotheca andaquiensis Ruiz-C., & Hernández-C., 1976 2000 1350-2000 1350-2000EGastrotheca antomia Ruiz-Carranza, Ardila-R., Lynch, & Restrepo, 1997

1140-2250 1140-2250

EGastrotheca argenteovirens (Boettger, 1892) 1650-3050 1650-3050

EGastrotheca aureomaculata Cochran & Goin 1970 2000-2600 2000-2600

Gastrotheca bufona Cochran & Goin, 1970 1430-1990 1990-2200 1430-2200

Gastrotheca dendronastes Duellman, 1983 1050-2090 1050-2090EGastrotheca dunni Lutz, 1977 2100-2720 2100-2720

Gastrotheca espeletia Duellman & Hillis, 1987 2530-3450 2530-3450

Gastrotheca guentheri (Boulenger, 1882) 1200-2010 400-2010

Gastrotheca helenae Dunn, 1944 2300-3250 2300-3250

Gastrotheca nicefori Gaige, 1933 1410-2200 1950-2500 1500-2450 400-1500

Gastrotheca orophylax Duellman & Pyles, 1980 2600-2900 2600-2900EGastrotheca ruizi Duellman & Burrowes, 1986 2250 2250

EGastrotheca trachyceps Duellman, 1987 2170-2540 2170-2540

Gastrotheca weinlandii (Steindachner, 1892) 1100-2370 1100-2370

Hemiphractus bubalus (Jiménez de la Espada, 1871) 1370-1655 300-1655

Hemiphractus fasciatus Peters, 1862 1430-1600 1600 300-1600

Hemiphractus johnsoni (Noble, 1917) 2000 1350-1660 1350-1660

Family Hylidae

*Dendropsophus bokermanni Goin 1960 1370-1900 200-1900

*Dendropsophus columbianus (Boettger, 1892) 1100-2350 1950-2100 950-2350

*Dendropsophus ebraccatus Cope, 1874 1620-1720 500-1720EDendropsophus garagoensis Kaplan, 1991 2000-2570 2000-2570

EDendropsophus labialis Peters, 1863 1600-3650 1600-3650

*Dendropsophus microcephalus Cope, 1886 1200 0-1200EDendropsophus padreluna Kaplan & Ruiz-C., 1997 1900-2060 1900-2060

Dendropsophus pelidna Duellman, 1989 2200-3000 2200-3000EDendropsophus praestans Duellman & Trueb, 1983 1750-2250 1750-2250

EDendropsophus stingi Kaplan, 1994 1400-2020 1400-2020

*Dendropsophus subocularis Dunn 1934 1520 805-1520EDendropsophus virolinensis Kaplan & Ruiz-C., 1997 1600-2400 1600-2400

Hylomantis buckleyi Boulenger, 1882 1370-2000 500-2000EHylomantis danieli Ruiz-C., Hernández-C., & Rueda-A., 1988 1640-1900 1640-1900

Hyloscirtus alytolylax Duellman, 1972 1700-2050 1600-1980 500-2050

Hyloscirtus bogotensis (Peters, 1882) 1980-2810 1750-3600 1750-3600



EHyloscirstus callipeza Duellman, 1989 1050-3000 1050-3000

EHyloscirtus caucanus Ardila-R., Ruiz-C. & Roa, 1993 2400-2700 2400-2700

Hyloscirtus colymba Dunn, 1931 1400-2250 1400-2250EHyloscirtus denticulentus Duellman, 1972 1800-2400 1800-2400

Hyloscirtus larinopygion Duellman, 1973 2140-2600 1950-3100 1950-3100

Hyloscirtus lascinius Rivero, 1970 1730-1960 1730-1960

Hyloscirtus lindae Duellman & Altig, 1978 2180-2660 2190-2450 2180-2660EHyloscirtus lynchi Ruiz-C., & Ardila-R., 1991 2540-3850 2540-3850

*Hyloscirtus palmeri Boulenger, 1908. 1030-1520 1010-2080 100-2080

Hyloscirtus pantostictus Duellman and Berger, 1982 2700 2700

Hyloscirtus phyllognathus Melin, 1941 1440-2190 600-2190EHyloscirtus piceigularis Ruiz-C., & Lynch, 1982 1750-2800 1750-2800

Hyloscirtus platydactylus Boulenger, 1905 1040-2550 1040-2550

Hyloscirtus psarolaimus Duellman and Hills, 1990 1950-2640 1950-2640EHyloscirtus sarampiona Ruiz-C., & Lynch, 1982 2190 2190

EHyloscirtus simmonsi Duellman, 1989 1100-2000 1100-2000

Hyloscirtus torrenticola Duellman & Altig, 1978 1440 1230-1750 740-1750

*Hypsiboas crepitans Wied- Neuwied, 1824 1000-1940 1100-2100 0-2100

*Hypsiboas lanciformis Cope, 1871 1600-1650 100-1650

*Osteocephalus buckleyi (Boulenger, 1882) 1660 300-1660

Osteocephalus carri (Cochran & Goin, 1970) 1000-1600 700-1600

Osteocephalus verruciger (Werner, 1901) 1150-2000 1150-2000

Phyllomedusa perinesos Duellman, 1973 1390 1390

Phyllomedusa venusta (Duellman & Trueb, 1967) 1250 130-1250

*Scinax ruber (Laurenti, 1768) 1200 0-1200

*Scinax x-signatus (Spix, 1824) 1470-1780 40-1780

*Smilisca phaeota (Cope, 1862) 1000-1430 1010-1560 0-1560

Family Leptodactylidae

*Leptodactylus colombiensis Heyer, 1994 1800 1200-1870 1130-2800 180-2800

*Leptodactylus fragilis (Brocchi, 1877) 1200 0-1200

*Leptodactylus fuscus (Schneider, 1799) 1200 1000-1300 0-1300

*Leptodactylus savagei Heyer, 2005 1250 0-1250

*Leptodactylus ventrimaculatus Boulenger, 1902 1200 50-1200

Family Microhylidae

Nelsonophryne aterrima (Günther, 1901) 1060-1530 1140-1630 300-1630

Family Ranidae

*Lithobates vaillanti Brocchi, 1877 1260-1430 1240 1304-1650 0-1650



Appendix 2

ICN numbers and particular references for specimens referred. ICN: Instituto de Ciencias Naturales de la Uni-versidad Nacional de Colombia

Familia Aromobatidae. Allobates picachos: 42617-62; 42824-7. Anomaloglossus atopoglossus: Acosta (2000). Rheo-bates palmatus: 669-700; 922-5; 928-9; 1262, 82; 1284-8; 1347-52; 1453-8; 1460-8; 1470-88; 1908-12; 1945-8;2027; 3244-6; 3552-3, 3559; 4015; 4071-2; 4094; 4100; 42124,6; 4262-5; 4350,56; 4373; 4404; 4410-5; 4428, 30,4432-3; 4585-7; 4604-28; 4645; 5025-7; 5162-4; 5436-8, 5440; 5442-5; 6082-6111; 6165; 8533-44; 8629-30, 8683-4; 8688-90; 8719,25; 14368; 15073-84; 18258; 18362-3; 19549; 20693; 21906-15; 22458-9; 23311-3; 33132-6;33260; 33469-70; 34934-45; 35450; 35478-90; 35531; 36573; 40721; 42499-502; 42505-11; 42574-5; 42587-9;43072-86; 43216-9; 43945-6; 43958-64; 44474-85; 44496-540; 44587-9; 44605-6; 45082-3; 45580, 2; 45707.

Familia Brachycephalidae. Atopophrynus syntomopus: 8611-8613. Craugastor fitzingeri: 19240-51. Craugastor longi-rostris: 31752; 32063; 37012; 38986-39000. Craugastor opimus: Savage and Myers (2002). Craugastor raniformis:17725, 9; 30474-92; 30852; 32565-84: 39392. Hypodactylus adercus: 47772. Hypodactylus babax: 13592; 16591-7;20713; 25965; 30864-5; 41289. Hypodactylus dolops: 23649-53; 23810-2; 23814-7; 24452-3. Hypodactylus elasso-discus: 49666-7; Ruiz et al. (1996). Hypodactylus latens: 775, 785-6; 6346; 8610; 39761. Hypodactylus mantipus:4945-6; 28637-8; 30331-3; 31748; 33347-53; 9161-73; 15579-80; 23670; 24605-22; 28637-8; 29486-7; 34991;41479-86. Phrynopus adenobrachius: 650; 773; 776, 789-94; 780-1, 784, 788, 796, 1856; 1861-2; 1866-7; 3207,3209-10; 3494; 6215, 6216-7; 6220-1; 6223-4; 6237; 18605-8; 18610; 18612, 18614; 18621; 20806; 26540-1.Phrynopus columbianus: Ruiz et al. (1996). Phrynopus nanus: 5269; 12503-4; 15096-140; 21188-91; 34316;42725-6. Pristimantis acatallelus: 2254; 7815-27; 16571; 16742-9; 18815-8; 18999-19005; 19063-69; 20216-24;25696-757; 25984-26010; 28976-91; 30337-44; 30515-8; 30681; 30760-1; 31294-5; 31659; 31667; 31674-6;32050-1, 56-57; 42790-1. Pristimantis achatinus: 1352; 17726, 28; 19307; 27582; 28027-31; 30316; 30452; 30589;30682-5; 30853-4; 31788; 33122; 47117-8; 36839-78. Pristimantis actinolaimus: 32287; 32295; 39941-68. Pristi-mantis acutirostris: 4355; 4379-80; 4497, 98; 5169; 5490; 11281; 12373-74. Pristimantis aemulatus: 19375-81.Pristimantis affinis: 7105-08; 14055-093; 22348-50; 26214; 32333-6. Pristimantis alalocophus: 25441-531, 549;29673-705; 33768-879, 881-885, 906; 37939-44. Pristimantis angustilineatus: 14104; 29247-63; 29275, 77-86;30298-9; 30301-2; 30554-66; 31296-304; 39598-617. Pristimantis anolirex: 10448-53; 10484-9; 10491-2; 10495-6;10504-5; 11374-11383; 11385-6; 11387-8; 15229-42; 15434-61,3; 15469-511; 26212; 33505-20; 33522-4. Pristi-mantis anthrax: 41697. Pristimantis apiculatus: Lynch (1998); Ruiz et al. (1996). Pristimantis appendiculatus:Lynch (1998); Ruiz et al. (1996). Pristimantis aurantigutattus: 16651-3; 16737-40; 16783, 94; 18869-70; 19363-5;19366-74, 89; 39158-9. Pristimantis bacchus: 4378; 4403; 5166; 5406-7; 5450; 5523; 5541; 6174-83; 6185-6;6188-9; 12389-97; 22478; 33137-9; 33148-66; 34231-2. Pristimantis baiotis: 16781-2; 19173-4, 6. Pristimantisbatrachites: 47887-92. Pristimantis bellona: 14118; 16330-47; 16580-6; 19213-8. Pristimantis bernali: 4844-48;10013-14. Pristimantis bicolor: 1160; 4631; 5191; 5300; 5404; 5452; 6190-1; 6196-6210; 6213; 8622-6; 12375-87;14456-81; 22479; 26321-2; 26543; 33167; 42753. Pristimantis bogotensis: 255, 263; 802-11; 813-5; 819-23; 826-95; 1026; 1029-30; 1102-3; 1115-24; 1413; 1849; 1850,2; 1886; 2494-7; 2781-92; 4824-6; 4857-78; 4897-4909;4911-7; 8723; 10877-900; 10902-17; 10921-66; 11037-42; 18219; 18391-410; 22418-39; 22487; 23325-7; 24443-6;38917; 42059. Pristimantis boulengeri: 25932,35; 3266, 70, 71; 6429-37; 6439-44; 6452-65; 6467, 70-75; 6477-79;6481-83, 86-89, 91, 93, 95, 96; 6501, 04-07, 16-18, 20-22, 24, 26-28, 44-51, 54, 62, 98, 99; 6604-06, 08-13; 15-30,32-36, 38-49; 6764; 6789-6801; 6803-14, 16-21, 23-36, 38, 47, 51, 55, 57-73; 7060, 65; 7277; 7948-8030; 8794;10024-51; 11527-630; 22770-83,85; 23577-98, 669; 24940-70; 25793, 883, 893; 26006, 121, 122; 28629-31, 35-6;33907; 34030-82; 35317-21; 36066-76; 37999; 38954-55; 39382-85; 40178-9; 41663, 815, 854-5; 41966-83;42016-25. Pristimantis brevifrons: 5212-29; 8172-3; 12493-4; 16784-93; 16799-805; 16819-56; 18762-3, 8; 19156-69; 19170-83; 21575; 25856-7; 25915-20; 28517-27; 28531-533; 28091-6; 28517-27; 28531-3; 29264-74; 31291-3;32071; 33330; 35906-9; 39059-61; 42816. Pristimantis buckleyi: 1549-55; 33428-30; 1539; 2082-84, 86, 88, 90, 92-97; 2498-9; 2502, 4, 76, 99; 6511, 66-7, 71, 73-4, 76; 6650-53, 55-58, 60-76; 6678-6702; 6704-07; 6881-83, 94-6,98-9; 6909, 10-12; 7690-7741; 11394-406, 11823; 12321-2; 12334, 40, 43-4; 12348-50, 52; 12355; 21737-876,21877-91; 21996-7; 22000-10; 24341-4, 24371, 73-78, 80; 24433, 47-48; 24462-64; 33428-30; 36822-4; 41780-812; 41889-93; 7690-7741. Pristimantis cabrerai: 16555-7; 19236-38; 16735-6. Pristimantis cacao: 25795-8. Pris-timantis calcaratus: 25795-8; 29066-90; 29293-311; 33394; 33360-2; 35846; 38934-5. Pristimantis capitonis:8124-45; 25860; 25938-40; 26008. Pristimantis caprifer: 38866-71; 38938-39. Pristimantis carranguerorum: 5128-30; 5132-45; 5345-9; 5351-63; 5601-03; 7166-71; 9402-37; 9439-68, 9470-1; 44953-7; 44966; 44970; 44975, 77,81, 83, 44985-6, 44989-90; 44995; 45002,5; 45014-15, 45024. Pristimanis carlossanchezi: 33502. Pristimantis car-ranguerorum: 5128-30; 5132-45; 5345-9; 5351-63; 5601-03; 7166-71; 9402-37; 9439-68, 9470-1; 44953-7; 44966;44970; 44975,77,81,83, 44985-6, 44989-90; 44995; 45002,5; 45014-15, 45024. Pristimantis caryophyllaceus:Lynch (1998). Pristimantis celator: Lynch (1998); Ruiz et al. (1996). Pristimantis chalceus: 17027; 14294-5;



16604-29; 19184-91; 20714-7; 28334-6; 28073-82; 30493; 31773-8; 32339; 32781-3; 37281-3; 12411-7. Pristiman-tis chloronotus: 26291-2; Ruiz et al. (1996). Pristimantis chrysops: 38834; 36900. Pristimantis colomai: 36809-12.Pristimantis colonensis: 26112; 49792-49805; Pristimantis cuentasi: 46187-9. Pristimantis curtipes: 22075- 77, 79;22154, 180. Pristimantis degener: 36808. Pristimantis deinops: 29369-70; 36917-9; 38936-7. Pristimantis diapho-nus: 13352; 38886-94. Pristimantis diogenes: 29371-2; 32705-14. Pristimantis dorsopictus: 1175-6; 9249-57;18745-54; 36532-37. Pristimantis douglasi: 15261-5, 15404-33; 15462; 15464-8, 15512-56. Pristimantis duell-mani: 8204-5; 25872; Lynch (1998). Pristimantis duende: 43855-72; 43873-76; 43878-88; 43893-98. Pristimantiselegans: 1024-5; 1027-8; 1394; 1452; 2493; 2632; 4057; 4254; 4823; 4879; 4910; 4918; 4959; 5741-4; 8681-2;8686; 8722; 11870; 14796-7; 26542; 33226; 34283; 35562-3; 35566; 40269; 40766; 41227; 41465. Pristimantisepacrus: 23654-7; 22088-172; 24115; 24121,3; 23654-5; 24094-5; 24097-8; 24101-2, 5; 24110; 23656-7; 24122-32;2413 4-9; 24147, 9; 24152-4; 24162-4; 24166; 24169; 24427-30; 24465-6. Pristimantis eremitus: Lynch (1998);Ruiz et al. (1996). Pristimantis eriphus: 26112; 47817-10; 49668-73; 49698-9; 49680-97. Pristimantis eryth-ropleura: 1214, 20, 24, 26; 3240-42; 3496; 5241; 13351; 14112-7; 14305-10; 14789-95; 16407-14; 16416-9; 16421-4; 16426-33; 16497-545; 16547-8; 16550-3; 16916-64; 17085-124; 18287-89; 19252-97; 19859-86; 19840-58;19887-927; 19928-20004; 20384; 20718-26; 20735; 21567-8; 21584; 27492-500; 28135-28200; 28234-28282,4,6-8; 28290; 28292-324; 28326-9; 29091-246; 29275; 31560-614; 30267; 30271-84; 30286-92; 30297; 30456-7;30834-7; 31680; 31192-225; 31870-928; 33123; 33017-112; 35848-60; 38872; 38895; 39069-85; 41615-48;42806,14; 43669-70; 5241; 9124-55; 15585-7; 20665-6; 21590-606, 20614-25; 23671-8; 24721-854; 27492-6;27498-500; 28626-8; 28642-8; 29091-156; 291578-79; 29157-246, 275, 3, 33711-717. Pristimantis factiosus:40030-160; 41474-78. Pristimantis fallax: 40792-797; 42420-21; 49796-7. Pristimantis fetosus: 32296-8; 40003-40029. Pristimantis frater: 5040-44; 5050; 5075-79; 5115-7; 7172-3; 9875-80; 21231-4; 39479-82; 39488-9; 39547-8; 40937-8; 40974. Pristimantis gaigeae: 2608; 5015; 8294. Pristimantis gladiator: 26113; 49700-3. Pristimantisgrandiceps: 12481-9; 29924; 33446. Pristimantis gularis: 19303-6; 45161. Pristimantis hectus: 25890; Lynch(1998); Ruiz et al. (1996). Pristimantis helvolus: 41664-9. Pristimantis hernandezi: 7489; 7848-61. Pristimantisillotus: 38947. Pristimantis ixalus: 47886. Pristimantis jaimei: 20739-46; 32818; 32851-54. Pristimantis johannes-dei: 19209-12; 30927; 32343-4. Pristimantis jorgevelosai: 15283-15403. Pristimantis juanchoi: 30345; 35063-84.Pristimantis jubatus: 52478-82. Pristimantis kelephas: 25931; 25933-36; 39635-6; 39618-74; 42454. Pristimantislabiosus: 36832. Pristimantis lasalleorum: Lynch (1998). Pristimantis laticlavius: Lynch (1998); Ruiz et al. (1996).Pristimantis lemur: 40786-91. Pristimantis lentiginosus: 10490, 93-94; 10497-503; 10506; 10508; 15520; 15259-60. Pristimantis leoni: 23141-4; 43823; 49704-13. Pristimantis leptolophus: 6492; 6744-63, 65, 69; 6772-88; 6822,37, 41, 45; 7003-08, 10-13; 7016-22, 24-33; 7035, 56, 61; 7067-75; 7276; 7292; 7295-6, 99; 7797-9; 11485-91;25695; 25921, 23-926; 41816-40, 41857-63; 41865-7; 41897-98. Pristimantis leucopus: Acosta (2000); Ruiz et al.(1996). Pristimantis lichenoides: 36546-7; 37171-205. Pristimantis loustes: Lynch (1998); Ruiz et al. (1996). Pris-timantis lutitos: 1435; 5192-3; 6214; 33169. Pristimantis lynchi: 960-976; 1932; 1934-5, 7; 4349; 4381-2, 6; 5131;5369-78; 5487; 5584; 5604-6; 5622-5; 5630-5; 5638-52; 5657, 5697-5700; 5805; 5818; 5820, 8, 5839-40; 5875-6;5878-5913; 5919-26; 5937; 5939-40; 5953-8; 6261-74; 9438; 9473-80; 23110-32; 31732-3; 33233-4; 33269-71;33296-314; 33327-8; 35451-64. Pristimantis maculosus: 8591-96; 36541-4, 36566; 37996-98; 38000. Pristimantismars: 30335-6. Pristimantis medemi: 5037-39,48; 5114; 9897-8; 9906; 21217-20; 21401; 39483-4; 39529-32;39534-8; 39544; 39573; 398828-9; 40588-605; 40958-61,3; 44935-7; 449839-45. Pristimantis merostictus: 1434;12466-480; 34233-242. Pristimantis miyatai: 5165; 5448-9; 6184; 8527; 12399; 12490-2; 15266-82; 22476-7;38738; 39108-20. Pristimantis mnionaetes: 11004-14. Pristimantis molybrignus: 7895-47; 17133; 17138; 25635-9;25643-54; 25660-3; 25665-69; 25861-7, 71; 25884-9, 92; 25937; 26011-12; 28097-100; 31750; 31976; 30855, 69;30872; 32054-5; 32654-7. Pristimantis myersi: 2503; 6484; 6500; 6677; 6743; 8196-203; 11349-51; 12201-4;12311-314; 24337-40; 25908-10; 26101; 33200-04. Pristimantis myops: 29326; 29331-8; 29342; 39684-717. Pristi-mantis nervicus: 11869; 11871-2; 26948-9; 32313-32; 40767-71; 41228; 41448-64. Pristimantis nicefori: 2614-6;3579-3610; 4054; 5819, 21, 23; 5830-2; 5844-5; 5927-36; 5938; 5952; 5959-74; 6280-7; 9713; 10253-79; 11369,11389; 20807-11; 20821-76; 20878-82, 20884-909; 20911-3; 20915-29; 20931-21087; 21090-146; 22260-346;22383-5; 22409-16; 31698. Pristimantis obmutescens: 2085, 7, 9; 6570; 6654; 6880, 84-93; 6897; 6900-8; 6911, 13-4; 8161-71; 11308-21; 23145-51; 5891, 907; 26001; 41902-37. Pristimantis ocellatus: 32857-63; Lynch (1998);Ruiz et al. (1996). Pristimantis orpacobates: 16472-96; 16630-34, 36-9; 16641-50; 16814; 17144; 17174, 94;20244-82; 21582; 27501; 28083-90; 28339-85; 29028-41; 30838-46; 30857-60-8; 32062; 33354-9; 35705, 35905;38835; 38896-8. Pristimantis padrecarlosi: 50072-86. Pristimantis paisa: 1178; 10001-2; 9992-8; 38280; 39566-7;43814-5. Pristimantis palmeri: 25657-9, 64, 70; 25689-93; 25813, 25; 28018, 23; 28515-6; 28528, 30; 29347-53;30285; 30348-53; 30531; 31258; 31271; 31677; 31781-7; 31856-63; 31275-8; 31286; 32880-90; 34994-6; 35833;35866-77; 369724; 39062-6; 41611-4; 41684-5; 9178-9181; 21573-4; 21648-655; 24680-717; 28516-7; 28528, 30;28534-44; 34992-96. Pristimantis parectatus: 9247; 10001; 41687-695. Prystimantis parvillus: 36825-31; 37011,13; 43822. Pristimantis penelopus: 15765; 42396; 42422-5; 42456-461. Pristimantis peraticus: 22489, 490; 40772-85. Pristimantis permixtus: 635-640; 642-53; 760-5; 771, 74; 945, 8; 952; 1162, 64, 66; 2024, 26; 2581-92, 94-5;3187-96; 3256-60, 62-65, 67-69, 72-3; 3491-3; 4113; 4278-81; 4734; 4782-943, 97, 98; 6226-8, 30-35; 6242-48, 50-



52; 6353-57; 8793; 8798; 8803; 8812; 8843-4; 8847-8; 8860; 8863; 8866; 8876; 8878; 8880-85; 8887-94; 8896-944;17127-30; 21607-8; 21676-8; 22491-7; 22784; 22813-32; 23517-547; 25005-40; 25535-6; 28674-736; 28738-744;28746-60,64-69; 29706-813; 34085-154; 35324-30; 35775; 35910-85; 36120-132; 36548-554; 37982-92; 38281;38926-29; 38953; 39404-21; 40180-82; 41487-510; 41683. Pristimantis petersi: 24238-9; 8210; 26114-6; 49714-25. Pristimantis phalarus: 39675-83. Pristimantis phragmipleuron: 18811; Ruiz et al. (1996). Pristimantis piceus:4104; 4841; 6236; 8566-7; 18801-4; 18820; 18596; 20412; 22573-5; 25537-47; 25794; 33756-67; 41856; 42010-1.Pristimantis platychilus: 8206; 16596; 25875; 36901-7. Pristimantis polemistes: 18808-10. Pristimantis polychrus:16730-4; 19219-35. Pristimantis prolixodiscus: 10102-19; 15157-81; 50087. Pristimantis ptochus: 39772-39827.Pristimantis pugnax: 22963-23009; 23171-87; 23868-87; 24438-40; 23171-87; 23189-23200. Pristimantis quantus:39762-71. Pristimantis quinquagesimus: Lynch (1998); Ruiz et al. (1996). Pristimantis racemus: 9034-84. Pristi-mantis reclusus: 46179-86. Pristimantis renjiforum: 5005-10; 5014; 13757-8. Pristimantis repens: Ruiz et al.(1996). Pristimantis restrepoi: 16588-94; 17037-8; 18276-7; 18413-6; 2884681; 28283-952; 39048-51; 42792;42808-15. Pristimantis ruedai: 16463-71; 17157-61; 18962-98; 27514; 28108; 30930-7; 37162-68. Pristimantissanguineus: 16635; 16750-6; 16872-5; 16879-83; 17137, 9; 19312-32; 19351-9, 61-2; 31936; 31948-75; 31977-32020; 32022-45; 37842, 44-51; 37827-62; 39718-27. Pristimantis satagius: 13722-3. Pristimantis savagei: 2982-4; 5045-7, 9; 5051-73; 5095-6; 5113; 5118-23; 9881-96; 9899-905; 9907-18; 12900; 20707; 21360-400; 26970-2;39485-7; 39582-4; 40492-575; 40584; 40902-35; 40962-73; 40975-79; 44946-51; 44958-65; 44971; 449974,6;44978-80, 2; 44984-88; 44991-3; 44996-45001; 4503-4; 4506-13; 45016-22; 45027-36; 45131-2. Pristimantisscoloblepharus: 8583-90; 40165-68. Pristimantis scolodiscus: Lynch (1998); Ruiz et al. (1996). Pristimantis sco-paeus: 22789-90; 22792; 22834; 40184-5. Pristimantis signifer: 20047-78. Pristimantis silverstonei: 29042-63;29373; 29637-8; 36908-16; 42793-803. Pristimantis simoteriscus: 21978-90; 22791; 22833, 35-48; 29670-1; 35125.Pristimantis simoterus: 757, 759, 766-70, 772; 4960; 6249; 9672; 18821-5; 18840; 32421-2. Pristimantis siopelus:Lynch (1998); Ruiz et al. (1996). Pristimantis spilogaster: 5411,3; 5453-4; 8528-31; 12420-54. Pristimantis suetus:10002; 41698-749. Pristimantis sulculus: Lynch (1998); Ruiz et al. (1996). Pristimantis supernatis: 6427-8; 6451;6480, 85, 94; 6510, 12-3; 6607; 6839-40, 42-44; 6984; 8031-8109; 8381; 8799-802; 8804-11; 8813-42, 45, 49-59;8861-2, 64-5; 8867-75; 8877, 79, 86, 95; 11322-9; 12799-800; 23201-241; 23575-6; 32716-50; 33689-96; 36560-1;41894-96 ; 49729-32. Pristimantis susaguae: 47712-714. Pristimantis taciturnus: 6523; 8207-09; 11689-702;25914, 22. Pristimantis taeniatus: 30347; 30847-9; 31290; 32067; 43323-4; 3551; 4115; 38269-72; 42397-408.Pristimantis tamsitti: 22948-51; 23632-41; 23818-58; 24441-2. Pristimantis thectopternus: 22488; 35841-2; 36117-8; 28105-7; 30315; 41602-9; 28338; 38950; 3233; 5240, 5295-9; 9156-60; 14337; 15588-93; 15615-18; 21656-58;23684-702; 23704; 24498-531; 24603; 26720; 28489, 95; 28641; 28644-48; 29488-508; 35841-2; 36117-8. Pristi-mantis thymelensis: 26349; 49733; Ruiz et al. (1996). Pristimantis torrenticola: 36557-8; 39969-99; 40001-02.Pristimantis tribulosus: 36559; 37169-70. Pristimantis tubernasus: 10483. Pristimantis uisae: 47882-5. Pristiman-tis unistrigatus: 1325-29; 2500; 4805-11; 36813-821; 43824; 47817-81. Pristimantis uranobates: 641; 2593, 6;3261, 74; 6229; 6253-4; 8568-82; 14424-31; 22705-69; 22786-88; 22793-812; 23573-4; 28529; 28546; 28762-3;35128-44; 35323; 35986-36065; 36088-110; 36565-9, 35572; 37945-57; 37960-81; 38956; 39554-60; 47817-81.Pristimantis veletis: 37206-21. Pristimantis vicarius: 6438; 6514; 6600-3; 6703; 6716-27; 6729-38; 6770-1; 6802;6875; 7059; 7278-91, 93-4, 97-8, 7300-3; 7742-88, 7890-4; 11413-49; 12190-8; 25855; 26005; 41813-4; 41853;41864. Pristimantis viejas: 42980-2; 42987-90; 42992; 42998-9; 42412; 42426-46. Pristimantis viridicans: 4780;7800-13; 25800-8; 25894-5. Pristimantis viridis: 16554; 37158-61; 19419-24; 30938-52; 37958-9. Pristimantis w-nigrum: 14292-302; 16312-20, 23; 16326-8; 16449-60; 16434-62; 17134-6; 17140-1; 17667-73; 18885; 18887-937;20727-33; 27490-1; 27501-13; 27925-28014; 28991-29020; 30238-42; 30303-14; 30317-30; 30494-502; 30590-600; 30602-80; 30752-58; 30870-1; 30873; 30238-42; 3033-14; 30317-30; 30494-502; 31226-56; 32751-80; 41263-68, 4351-2, 4387; 6187; 12398, 22849-66; 23859-64; 24426; 33251-2; 1872; 2022; 2079-80; 2501; 2526-29, 2531-58; 2560-75; 3197-99; 3403-05; 3490; 4129; 4800, 4849; 5232-9; 5489; 6347-52; 6468; 6552-3; 6558, 61, 68, 72;6948-79; 7096-101; 8110-123; 8276; 8945-9033; 9680; 9693; 1352-64; 12797-8; 14296-7; 14301-2; 15581-4;15623-4; 17125-6; 17156; 17651-66; 17699-724; 21610; 21626-34; 22067-9; 22073-4, 22576-90; 23152-168;23727-46; 24532-56; 24559-602; 25911;28470-488; 28490-4, 28639; 29587-626; 29631-6; 34435-6; 35322; 36119;36545; 38273-4; 40172-75; 41757-79; 42014, 43644-46; 49737-809. Pristimantis xeniolum: 43877; 43981. Pristi-mantis xestus: 32337. Pristimantis xylochobates: 28963-75. Pristimantis zoilae: 49751-59; 49761-3; 49766-70;49772-75; 49778-80, 82-83; 49785-88 Pristimantis zophus: 1179; 1208-13; 1215-19; 18591-3; 20141-200; 41670-82. Strabomantis anatipes: 12113; 32668-90; 32701-4; 12113. Strabomantis bufoniformis: 17049-53; 30510; 31766-52. Strabomantis cadenai: Ruiz et al. (1996). Strabomantis cerastes: 14096; 14103; 16601-3; 19205-6; 19208;29064-5; 30513-4; 33016; 37821-6. Strabomantis cheiroplethus: 16300-11; 16599; 18012-37. Strabomantis cornu-tus: 24240-1. Strabomantis ingeri: 2507; 4662; 6167-73; 7380; 10507; 10509-10; 12419; 13570; 13753; 32399;33168. Strabomantis necopinus: 24604; 37573-614. Strabomantis ruizi: 4933; 4961-2; 13967-9; 21566; 35086-9.Strabomantis zygodactylus: 30521-30.

Familia Bufonidae. Andinophryne atelopoides: 06373. Atelopus angelito: 33407-8; 33410-5. Atelopus carauta: 03180;03184; 13099; 16262-16269. Atelopus chocoensis: 28818-19. Atelopus ebenoides: 299; 339-61; 1257; 1270-1;



1298; 1411-2; 1415-6; 1418-9; 1926-31; 5571-81; 5712; 18292-4; 33205; 33232; 33272-94; 33315-20; 33329;34215; 38919-20; 393-409; 1198; 1535; 1540-1; 1560; 2425; 2620-27; 4717; 20010; 31765. Atelopus eusebianus:12272-302; 20010; 32946-33012,15; 33128; 33208. Atelopus famelicus: 32897-32922; 34210. Atelopus farci:14482-533; 32506. Atelopus galactogaster: 47894-97. Atelopus guitarraensis: 23348-51; 32432-3; 32530. Atelopusignescens: 376-7; 451-8; 460-8; 470; 474-80; 482-95; 1129-30; 1251; 1376-9; 1590-4; 1703-15; 3524-9; 3611-32;3634-5; 3639-53; 4144-52; 12807-8; 26326-48; 31458. Atelopus lozanoi: 378-392; 1125-6; 1254-5; 1296-7; 1299-1301; 1387; 1409-10; 1414,20; 13096-7; 21147-53; 31470-4; 32477-95; 33376-84. Atelopus lynchi: Ruiz et al.(1996). Atelopus mandingues: 19569; 21155; 22351; 26102; 34285-7; 34716-23; 34959; 45149-50. Atelopusminutulus: 4851-53; 5028; 13709-721; 7085-93; 9750. Atelopus mittermeieri: 7370-71; 7374-77; 7385; 7387-88;7790, 96,98; 7400-02; 12747-62; 12765-69; 52993. Atelopus monohernandezii: 5365-6; 5386; 5388; 5420; 5422-6;5496-7; 5526-8; 5542-5; 5679-80; 5705-6; 5709; 5987; 5989-6004. Atelopus muisca: 8669; 19567; 21154; 21158-9;31791-9; 32505; 33211; 33594-5; 34295; 34958. Atelopus nicefori: 621-25; 1163-71; 1256; 1269; 1292-93; 1304-05; 1353-67; 1380-81; 3484-85. Atelopus pedimarmoratus: Acosta (2000). Atelopus pictiventris: Acosta (2000);Ruiz et al. (1996). Atelopus quimbaya: 4759; 23339-47; 25827-43; 27300-2; 28774; 29819; 32475-76; 32507-32528; 35772; 36141-36151; 38001. Atelopus sernai: 4120; 4162; 4166; 4168; 4222; 4235; 4242-3; 4269; 4276-77;6359; 9868-72; 23703. Atelopus simulatus: 6708-10; 11331-33; 11346-7; 38930; 41899. Atelopus sonsonensis:37516-33; 37938; 45730. Atelopus subornatus: 2846, 47, 49, 53-72; 3542; 4045,58; 4156-59; 10852, 54, 57; 12794-96, 823-27; 18354, 55; 23360, 61; 24434; 28467; 31359, 435, 452; 34275-79; 35422; 8296-99. Osornophryne anti-sana: 12264. Osornophryne bufoniformis: 362-375; 1597; 1640; 1643; 1647-8; 2091; 2414; 6631; 6637; 6659;6711-15; 7562-87; 11500-8; 12246-7; 24432; 25896-905; 32927-35; 33375; 33405-6; 41852; 41900-1. Osor-nophryne guacamayo: 47811; 49834-7. Osornophryne percrassa: 319-41; 1596; 1629; 1639; 1641-2; 1922-25;2618; 2624-25; 3178-9; 3186; 3206; 3208; 3211-12; 3507; 3515; 4738; 10015-23; 17642-45; 18755-61; 33754.Osornophryne talipes: 12252-3; 12255; 12262-3. Rhaebo glaberrimus: 21221; 40804-5. Rhaebo haematiticus:32924-6. Rhaebo hypomelas: 27419; 27421-27; 30025-30; 30365. Rhinella cristinae: 23360; 23662; 26225-30;26232-40. Rhinella granulosa: Acosta (2000). Rhinella macrorhina: 9831; 13944,46-48; 33720-4; 41558-85.Rhinella marina: 4491; 5774-5; 12418; 12498; 14370; 22482-3; 26260; 31377-8; 39839; 40806-7; 41206; 42884;43297-8; 45555; 45698. 269-70; 13250; 20228-30; 20229-230; 28330-1; 30366; 326425-5; 41662; 34465. Rhinellanicefori: 10063-65; 13949; 41541-57. Rhinella rostrata: Ruiz et al. (1996). Rhinella ruizi: 4114; 4119; 4172-75;4177; 4241; 4266-68; 4601,3; 8374; 9817-30; 9832-34; 33720-24. Rhinella sternosignata: 4048; 8310; 23316;26261; 31347; 31364-73; 39140-48; 40808-25; 41204; 42463-98. Rhinella truebae: 14780.

Familia Centrolenidae. Centrolene acanthidiocephalum: 4395-97; 4405-07; 4669; 5183, 87, 88; 5255, 60, 62, 72-90;5367; 5429,31; 5495; 5704; 6006-12; 7347-8; 8447-58; 9717-20; 11278; 19531-2. Centrolene andinum: 1059-60;4360; 4582-4; 4632; 4728-9; 4955-7; 5185-6; 5292-3; 5432, 93; 5528; 5535-7,9; 5710; 6013-23; 6275; 7349-50;8434-45; 10525; 11059; 17861-85; 19737-8; 29908-21; 40413-8. Centrolene antioquiense: 9773; 35194; 36527, 9;37500-3. Centrolene buckleyi: 18334; 18337-43; 18622-40, 50; 28775-81; 43889-90, 1826-9; 2525; 2752-6; 2761;3290; 4744-51; 4934-5; 5564-70; 5701-3; 5722-7; 5802-4; 5914-8; 6322; 10231-2; 14910; 17860; 18075; 18205-7;18337-43; 33495; 6445-50; 6530-42; 6587-97; 6879; 7067-8; 7472-7540; 121829; 18172-76; 26962; 41868-79;42012-3. Centrolene geckoideum: 16169-70; 28830-36; 28838-45; 31305-08; 41291-95, 4633; 5557-63; 98; 7161-65; 8693; 24187 8694-8; 12138-41; 18336;29463-4; 31305-8; 36514-16. Centrolene grandisonae: 15952-73;17269-76; 17280-9719613-25; 19839; 26045, 76; 27712; 27722; 27769-90; 28794-95; 30031-35 30762-88; 31351;9752-54; 24655; 29466-8; 29470-75. Centrolene guanacarum: 11685-7. Centrolene heloderma: 7443-46; 26060-62;31103-14; 31115-29; 31130-63. Centrolene huilense: 7454-7; 7461-3. Centrolene hybrida: 5172; 5243; 5257; 5265;5333-40; 5481; 5599; 5620, 21; 9608-37; 10201-5; 10208-13; 13732; 13735-6; 13733; 17886-907; 18179, 203;18344-9; 19752; 24197-9; 42846-8. Centrolene ilex: 19196-204. Centrolene lynchi: 30891-4. Centrolene medemi:17857-9; 20005; 20791; 23642-7; 23898-9; 23902-4. Centrolene notostictum: 4374; 4383-5; 4409; 4770; 5341-43;5390-95; 5492,94; 5504-21; 5133-4; 5711; 6024-32; 7351-68; 8382-99; 8401-5; 8407-23, 25-32; 12604; 12606-7;12609-12; 12617-22; 12624; 12626-8; 12631-3; 12639; 12640-44; 14884-5; 14890-9; 14903-4; 14907; 14989;15029; 17973-18006; 18188; 19572-606; 19753; 29905-7; 33447-66; 34243-4. Centrolene paezorum: 11866. Cen-trolene peristictum: 15991; 9773; 12114. Centrolene petrophilum: 9564-70; 10192, 6; 17924-35; 19570-1; 19749-51. Centrolene prosoblepon: 16002; 16156-68; 19650-68; 19671-81; 19735; 17227-31; 27727-29; 27761-62;30036-37; 30820-33; 30895-99; 32089, 96, 99; 32661-62; 38865; 38880-83; 41408; 34779-805; 36879-80; 43503-11;26300-8. Centrolene quindianum: 27759-60, 9786-9799; 9801-4; 24873-924; 29433-51; 35097-101. Centrolenerobledoi: 16147-55; 41345-62; 9937-68; 9974-85; 17936-7; 17939-41; 17943-61; 17963-66; 17971-2; 28667-68;35102-5; 38111-17; 38149. Centrolene sanchezi: 24293-5. Centrolene scirtetes: 12149-80. Cochranella adiazeta:2758-60; 3293; 3554; 3556; 3558; 4361; 4367; 4670, 4673-4; 4718-27; 4958; 5011-3; 17908-23; 43147-8, 43202-3.Cochranella daidalea: 4958; 5344; 5491; 6036-9; 8459-63; 14911-7; 18008; 33467-8; 33596; 37252. Cochranellaeuknemos: 15983-8, 90; 17483-84; 19638-40; 27719-20; 27739-49; 30039; 30369; 30789-96; 30797-812; 31344.Cochranella flavopunctata: 5030, 83; 9571-2; 9607; 21235-6; 44787-8.Cochranella megistra: 17242-44; 27763-8;27718; 28796. Cochranella punctulata: 15800; 34745-59; 38095-7; 43512-13. Cochranella savagei: 19837; 21493-



4; 33218-20; 33342-3; 33364; 9767-9; 20763; 24925-32; 25969-72; 34271-4; 34926-30. Cochranella solitaria:24298. Cochranella spinosa: 19430-49. Cochranella susatamai: 34809-14; 38092-4; 43514. Cochranella xan-thocheridia: 27757. Hyalinobatrachium aureoguttatum: 4754; 15974-82; 17245-6; 17249, 51, 58-59; 17263-5;17505-23; 17525-39; 27741-51; 19744-46; 27752; 27747-56; 30153-4; 32110, 16-21; 32123, 25-26, 28;3888540755-59. Hyalinobatrachium colymbiphyllum: 15992, 94-98; 16000-01; 19561-66; 19685, 87; 19689-711;30156-76; 30377-9; 30878-90; 32100-04; 32106-09; 32111-15; 34727-44. Hyalinobatrachium esmeralda: 5031-5;9592-9603; 9638-9; 19607-12; 20708; 21237; 44785. Hyalinobatrachium fleischmanni: Ruiz et al. (1996). Hyalino-batrachium ibama: 5173; 5202-6; 5256; 5330; 6033-5; 10215-8; 10219-23; 11081-2; 12601-2; 14886-9; 14927;40606-19. Nymphargus armatus: 25000; 28037-72; 42807. Nymphargus chami: 19733; 32077-78. Nympharguscristinae: 17912-15; 17920; 17922-25; 18643-4; 18646-49; 18651, 19712-15; 19720-25. Nymphargus garciae:1645-6; 4752-3; 6877-8; 7451-3; 7458-60; 9755-66; 11715-20; 11722-51; 11752-60. Nymphargus griffithsi: 7464-68; 15904-17; 16003; 1606-86; 16871; 17482; 17486-503; 18335; 19546-47; 19626-34-36; 19726-30; 19838;26013-30; 28784-95; 29427-8; 30096-107; 30108-52; 30370-76; 30814-18; 31313-19; 31320-43, 9920-23; 24933-39; 29476, 78, 80-1; 36526, 30; 38125-46. Nymphargus ignotus: 14749-77; 15859-67; 15871-8; 15880-7; 15889-96;15898-903; 18007; 19641-4; 21496-533; 30040-8; 30050-3; 30055-95; 30367-8; 30567-68; 30570-6; 32095; 33238-48; 33333; 36969. Nymphargus luminosus: 15918-29; 15931-36; 19731-32. Nymphargus luteopunctatus: 20747;Ruiz et al. (1996). Nymphargus nephelophila: 24296-7. Nymphargus oreonympha: 20765-90. Nymphargusposadae: 7447-50; 11307. Nymphargus prasinus: 13419; 15937-51; 19645-7; 32080; 38884. Nymphargus rosada:30091; 34761-78; 35217-9; 36524-25. Nymphargus ruizi: 7469-71; 18180-6; 26031-7; 26038-44; 31345-50. Nym-phargus spilotus: 35155-8; 38073.

Familia Dendrobatidae. Ameerega andina: Ruiz et al. (1996). Colostethus agilis: 6403-21; 6516; 7618-25. Colostethusalacris: 20009. Colostethus brachistriatus: Ruiz et al. (1996). Colostethus dysprosium: Acosta (2000). Colostethusfraterdanieli: 10610; 16285; 1180, 83-86; 2023, 5; 2559; 3200-5; 17600; 18420-1. Colostethus furviventris: Acosta(2000). Colostethus mertensi: 8211-23; 43684-764; 15636. Colostethus pratti: 556-67. “Colostethus” ramirezi:Acosta (2000). Colostethus thorntoni: 26965-69. Colostethus yaguara: Acosta (2000). Dendrobates truncatus: Ruizet al. (1996); Bernal et al. (2005). Hyloxalus abditaurantius: 41651; 16272-7; 19483-94; 28035-6; 29482; 30579;32060-1; 15576-8; 18432-9; 21673-5; 23705-10; 24623-54; 24718-9; 29482; 9838-58. Hyloxalus borjai: Acosta(2000). Hyloxalus breviquartus: 6143-4; 6148-9; 8545; 21999; 22499-501; 6466-9; 21667-8. Hyloxalus edwardsi:2770; 6376-91; 8560-4; 21936-44; 35812-3; 3515; 3544-5. Hyloxalus fascianigrus: 35520-9. Hyloxalus lehmanni:21945-77; 6915-47; 9859-66; 17601-6; 18425-6; 18428-9; 23557; 23564; 24859; 29643; 32819; 34952-3. Hyloxaluspinguis: 20006. Hyloxalus pulchellus: Ruiz et al. (1996). Hyloxalus ramosi: Ruiz et al. (1996). Hyloxalus ruizi:4838-9; 5415-9; 25981. Hyloxalus saltuarius: 42512-6; 42663-70; 43464-7. Hyloxalus subpunctatus: 816-18; 824;977-1007; 1031-46; 1302; 1437-51; 1716-1777; 1853-4; 1885; 1887-90; 1893-4; 1898; 1904-7; 1933, 6, 1938-42, 4;2065-8; 2372-91; 2395-7; 2468-9; 2523-4; 2530; 2793-6; 2829-31; 3285-6; 3398-3400; 3402; 3973-93; 4155; 4245-8; 4250-3; 4255,4282; 4315-37; 4339-44; 4346-7; 4431-71; 4473-90; 4592; 4294-5; 4892-3; 5088-92; 5103-6;5198-5200; 5379; 5410; 5587; 5626-9, 5636-7; 5653-6; 5696; 5731-40; 5857-61; 5863-9; 5877; 5975-7; 6313-8;7174-99; 7234-7; 7272-5; 8303-6; 9469; 9482-4; 11283-4; 11462-5; 11806-8; 18076; 18201-2; 18220; 18222;18252, 18311-3; 18364-9; 18412; 20877; 20883; 20910,4; 22356-82; 20386-402, 20404-6; 20484-6; 23309; 20611;24436-7; 26130-9; 20155-8; 20164; 20915-42; 20963; 20995; 27024; 31686-97; 31699-710; 31712-20; 31722-31;32434, 97; 33177-9; 33265-8; 33295; 33322-3; 33365-71; 33389-403; 33431-6; 33686; 34083; 35342-92; 35395-406; 35425; 35443-9; 35465-77; 35517-9; 35532-9, 25561; 35564-5; 35567-619; 35672-3; 35814; 37017-20;37076-9; 37081-2; 42517-37; 42572-3; 42577-86; 42590-612; 42705-24, 27; 42829; 43784-95; 43965-44003;45564-7. Hyloxalus vergeli: 4930-1; 12815-22; 35423; 3556, 6297-6306. Oophaga lehmanni: 8284-5; 22181-3;39001-2. Phyllobates bicolor: 13101; 1509-13; 27304; 31867-69. Ranitomeya bombetes: 8281-3; 19820-4; 24299-306; 24860-72; 29483-5. Ranitomeya daleswansoni: 42308-37; 53277-8. Ranitomeya dorisswansonae: 53279;Rueda et al. (2006). Ranitomeya ophistomelas: 34616-9; 39759-60; 41593-4. Ranitomeya tolimense: 53372-3; Ber-nal et al. (2007). Ranitomeya virolinensis: 4256; 4588-91, 3; 4596-7; 5107; 5368; 5408-9; 5414; 5446-7; 5482-6;5499-5503; 6150-60; 6276-9; 8549-54; 9329-32; 9334; 14825-30; 16087-146; 18317-8; 22480; 26538-9; 26984-7;28408-10, 2; 33254-9; 42926-7. Silverstoneia erasmios: Acosta (2000). Silverstoneia nubicola: 16286; 30177;30686-88.

Familia Hemiphractidae. Cryptobatrachus fuhrmanni: 3483; 8738-92; 34621-48; 34650-59; 34807; 34913; 40161;37254; 38692-730; 42923-4. Cryptobatrachus nicefori: Ruiz et al. (1996). Flectonotus pygmaeus: 13789; 15209-28;43903. Gastrotheca andaquiensis: 23664-6; 24177-86. Gastrotheca antomia: 18187; 183299-31; 28019-22; 28201-33; 29426, 29-32; 30739-40; 38075-78; 39041; 42782-3. Gastrotheca argenteovirens: 1392; 3407; 4743; 5808;6255; 6311-12; 6368; 6422-3; 10052-54; 12206; 12809-10; 7081-84; 7095; 7541-55. Gastrotheca aureomaculata:2515-16; 2662; 2894. Gastrotheca bufona: 38689. Gastrotheca bufona: 38082-88. Gastrotheca dendronastes:12100-1; 19468-74; 20079-83; 27468-9; 28386-407; 28425-6; 30741-51; 30183-86; 30380; 32893-6; 35835-40;38916. Gastrotheca dunni: 1153; 1595; 9700. Gastrotheca espeletia: 12207-16; 10055-59; 41587-8. Gastrothecaguentheri: 33120; 38688; 38732-3; 47901-3. Gastrotheca helenae: 10541-62. Gastrotheca nicefori: 271; 4735;



4771-2; 5230-1; 5556; 5707; 5713-15; 5718-21; 5755-6; 5797-8; 5801; 7094; 7340; 8678-9; 12595; 13919; 14044;19548; 20084; 21905; 22450-2; 26194-7; 34251; 41661; 42754. Gastrotheca orophylax: Ruiz et al. (1996); Acosta(2000). Gastrotheca ruizi: 1633-4; 1637; 1641; 2611; 2891-2; 4736-7. Gastrotheca trachyceps: Ruiz et al. 1997.Gastrotheca weinlandii: 23628; 24449-50; 31558; 47899. Hemiphractus bubalus: 23629-31. Hemiphractus fascia-tus: 9327; 16271. Hemiphractus johnsoni: 36517.

Familia Hylidae: Dendropsophus bokermanni: 24001-85. Dendropsophus columbianus: 535; 5458-80; 11457-9;13428-30; 20748; 28101-4; 30381-96; 30928-9; 31665; 32692-5; 33334-41; 38922-5; 41652-60; 45576; 18448-50;21635-42; 21659; 38922-25. Dendropsophus ebraccatus: 5525; 6060-9; 18300-9; 43149-53; 43215. Dendropsophusgaragoensis: 12949-50; 12953; 17781-17805; 18361. Dendropsophus labialis: 958-9; 1048-77; 1154-9; 1247-8;1469; 1896-7; 1899-903; 1956-61; 2069-78; 2600-02; 4423-4; 4925; 5100; 5547; 5586; 5588-90; 5658-67; 5686-93;5728-30; 5745-8; 5810; 5856; 5983; 6260; 6375; 8517-23; 8680; 9711; 10294-9; 10318-40; 11461; 1166-8; 11809;14095; 15559; 17779-80; 18071- 4; 18194-6; 20818; 22352-3; 23352-3; 29948; 32400-1; 32414-6; 32923; 33223-4;33261-4; 33321, 6; 33404; 33610-40; 34228; 34348; 36621; 37080; 37083-4; 37092-7; 41229-30; 41235-8; 43806-8; 43819; 45510-1; 45563; 45717-20; 46915; 46917-20. Dendropsophus microcephalus: Ruiz et al. (1996); Bernalet al. (2005). Dendropsophus padreluna: 8628; 8631-46; 9604-6; 15840-2; 15844-8; 22011-66; 23310-312. Den-dropsophus pelidna: Ruiz et al. (1996). Dendropsophus praestans: 7556-61. Dendropsophus stingi: 14375; 15835-9; 15844-8; 17818-30; 17831-40; 17843-6; 18204; 49916-31. Dendropsophus subocularis: 28134. Dendropsophusvirolinensis: 4577-81; 4629,30; 5160; 5170-1; 5433-5; 5549-54; 5670-5; 5683; 5984; 6070-2; 6112-134; 8465-87;9653; 9716; 12512; 12528-9; 12533-5; 12525; 12541; 12543; 12551-2; 12554; 12560; 14344; 29949-30001; 34226-7; 35782; 38734-37; 39003-25; 43136-46; 45704; 45712. Hylomantis buckleyi: 5161; 5522, 29; 12402; 23658-9;23778; 23958-72; 33171; 42845. Hylomantis danieli: 16005. Hyloscirtus alytolylax: 21209-10; 21534-37; 32847-50; 35878-83; 38820-46; 41303-6; 12129-33. Hyloscirtus bogotensis: 303-6; 315-317; 1561; 1582,83; 1848; 2520-22; 2605; 2771-79; 3291, 92; 4416, 17, 19-22; 4993; 5380; 5457; 5548, 91; 5600, 76; 5806-7, 11, 15; 5829; 5850-4,8; 5870; 6256-59; 6344, 67; 6369-72, 92; 7379; 7433; 10870-1; 13751; 14799; 18228,62; 18357-9; 21732-6; 23317-8; 24381; 28468-9; 29925-34; 31317; 31543-8; 32501; 33225; 34298-304; 24349-51; 45509; 49568-9; 46914;12111. Hyloscirstus callipeza: 4051; 10454-461; 14936-941; 14965-77; 14983-5; 14987-94; 15001-07; 15 017-28;33471-88; 41466-72; 42395; 43027. Hyloscirtus caucanus: 7002; 7238-53; 11682; 12248. Hyloscirtus colymba:13876; 14099; 19495-530; 27435-44; 28130-3; 28820-27; 30714-7; 30234-6; 30397-418; 30699-738; 31354-8.Hyloscirtus denticulentus: 4769; 10300-317; 14986, 995-15000; 15042-071; 22472, 473; 26309-318; 29935-947;32496, 498; 33140, 172, 437-444, 489-493; 40735, 736; 7155; 8488-95; 8525; 9707, 08, 10. Hyloscirtus larinopy-gion: 13595; 28828-9; 28926; 31191; 42779; 6345; 8375; 9380-401; 9670,75; 15626,27; 18597-603; 23682-83;34433; 34970-72; 36133-38; 36518-19; 39728-29; 41880. Hyloscirtus lascinius: 14918-26; 14980-2; 15030-41;15557. Hyloscirtus lindae: 20795-6; 23786; 23865-7. Hyloscirtus lynchi: 15193-208; 33505, 521; 42728. Hyloscir-tus palmeri: 19475-80; 17207-10; 28128; 30419-24, 30426-38; 31781-7; 32872-9; 9175-77; 15780-99; 15817-19;39328-36. Hyloscirtus pantostictus: Ruiz et al. (1996). Hyloscirtus phyllognathus: 5124-7, 5175, 77-8; 5263-4,71;5312-29; 6364; 7442; 9511-32; 9535-46; 9552-7; 9746, 20797-804; 21900-904; 22462-70; 23905-10; 26241-2;44823-9; 45098-9; 45503-4. Hyloscirtus piceigularis: 4390-2; 4394; 5180; 5194; 5254, 5258; 5310; 6056-9; 7411-8;8524; 19536, 38; 22460-1. Hyloscirtus platydactylus: 9709; 14930-5; 32404-6; 33588-9; 42378. Hyloscirtus psaro-laimus: Acosta (2000); Ruiz et al. (1996). Hyloscirtus sarampiona: 07440-1. Hyloscirtus simmonsi: Ruiz et al.(1996). Hyloscirtus torrenticola: 22952-6; 23614; 23916-7; 23922-7. Hypsiboas crepitans: 4499; 5108-9; 5959,81-2; 7382-3; 8311; 8516; 8718; 11276; 12400-1; 12403-8; 14343; 15008-9; 22444-6; 33144-8; 38759; 40763-4;42885-95; 43033-62; 43188-95; 43251-2; 43256,9; 45551-2; 45554; 9246; 9677-8. Hypsiboas lanciformis: 5148-49.Osteocephalus buckleyi: 23648. Osteocephalus carri: 49932-48 ; 49950-61; 49963-81; 49982-87; 49990-91; 39259-60; 21244-46. Osteocephalus verruciger: 2492; 22957; 23765; 23773,75; 23942-57. Phyllomedusa perinesos:23772; 23779; 24188-9. Phyllomedusa venusta: 9258-62. Scinax ruber: Ruiz et al. (1996); Bernal et al. (2005). Sci-nax x-signatus: 2607; 5532; 5681-2; 5941; 6081; 8496-8501; 35124; 37098; 37222; 45557. Smilisca phaeota: 2021-4; 17217; 20205-8; 27387; 30439-51; 31811-14; 32691; 8671; 10060; 15774-78; 36887; 39369-70; 40259-60.

Familia Leptodactylidae. Leptodactylus colombiensis: 3236; 3557-60; 4493-5; 5150-1; 6161-4,6; 8226; 9558-63;9722-9; 18077-9, 17081; 18257; 19556; 21991-4; 22453-7; 33141-143; 33325; 42903-8; 43899-901; 1252; 7405-10; 26319. Leptodactylus fragilis: Bernal et al. (2005). Leptodactylus fuscus: 39832; 43070-1; Bernal et al. (2005).Leptodactylus savagei: 9934-6.

Familia Microhylidae. Nelsonophryne aterrima: 19482; 30237; 3250; 42928-9. Familia Ranidae. Lithobates vaillantii: 20283-96; 36888; 37463-74.


review and analysis of altitudinal distribution of the andean anurans in colombia

Documents

broader thermal

wider altitudinal

jaccard similarity

narrow altitudinal

universidad

broader altitudinal

broader altitudinal

restricted