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SI Text
Taxonomic Considerations. The recent discovery that Brachycephalus is allied with the eleutherodactylines (1, 2) has made uncertain the valid family name for Eleutherodactylus and its relatives. However, the decision to place all ~800 species of eleutherodactylines in the family Brachycephalidae (2, 3) was based on analysis of 16 eleutherodactyline species, only six of which were from South America (2). In our analyses here of many more species, Brachycephalus appears in a basal position in the trees and does not group significantly with any species or clade of eleutherodactylines (SI Figs. 5-13). The association of Brachycephalus with the eleutherodactyline lineage, in itself, does not require the use of the family name Brachycephalidae for the entire assemblage. If multiple families are recognized, the correct allocation of these names will depend on a more robust phylogeny, especially concerning the basal branches. For example, Brachycephalidae might be appropriate only for Brachycephalus, with one or more other family names applied to the eleutherodactylines. Also, the suggestion by Frost et al. (2) that two genera of eleutherodactylines might be close relatives of Brachycephalus was not supported in a recent morphological study (4). For these reasons, we use Brachycephalidae as a family name only for Brachycephalus and refer in this current study to the remaining assemblage of genera and species as "eleutherodactyline" frogs, pending further study.
Assignment of unsampled species to the major clades (genera) defined here (SI Table 2) was based largely on their species-group affiliation. Although our analyses show many changes in content for previously defined species groups, most species assigned to the same species group are placed here in the same genus. Exceptions include Phrynopus and species from southeast Brazil, which will require further systematic study (see below).
We restrict the generic name Eleutherodactylus Duméril and Bibron, 1841 (type species, Hylodes martinicensis) to the Caribbean Clade and continue recognition of the included subgenera. The subgenus Eleutherodactylus is now restricted to the West Indies, and Syrrhophus includes two species in Cuba (E. symingtoni and E. zeus), besides those in Middle America. Eleutherodactylus counouspeus will be assigned to a new subgenus elsewhere. We consider Eleutherodactylus planirostris rogersi as a species (Eleutherodactylus rogersi) distinct from E. planirostris because they were not found to be closest relatives.
Because sequence data are unavailable for the type species of Phrynopus Peters, 1873 (Phrynopus peruanus), it is not possible to allocate that generic name to the separate clades defined here as including species of "Phrynopus," and, therefore, we make no changes in their generic status. The molecular phylogeny (Fig. 2) places into question the validity of the morphological species groups, restricting us from making any assumptions about species believed to be closely related to P. peruanus based on morphology.
The Southeast Brazil Clade is a similar situation in that we lack sequence data for the type species (Eleutherodactylus verrucosus) of the earliest possible generic name for the group, Ischnocnema Reinhardt and Lütken, 1862. However, in this case, we have confidence that the generic name Eleutherodactylus (The Caribbean Clade) does not apply to any of the species, based on geography and morphology (5-7). Also, our analyses (Fig. 2) indicate that E. binotatus and, by implication, its close relative E. plicifer, belong to a separate genus that we will name elsewhere. These two species also have an unusually long first finger which appears to set them apart from almost all of the other species of Eleutherodactylus in southeast Brazil except E. paulodutrai and E. ramagii, which differ significantly in other characters (5, 6). Although the number of species in this study from the Southeast Brazil Clade is small (four), they represent several clusters of species, each sharing a suite of morphological traits indicating relatedness (5-9). By implication, this increases the size of the clade to a minimum of 12 species: E. epipedus, E. erythromerus, E. gualteri, E. guentheri, E. hoehnei, E. izecksohni, E. juipoca, E. nasutus, E. oeus, E. parvus, E. pusillus, and E. vinhai.
Until additional data are collected, we cannot say with confidence that E. verrucosus and the remaining species of Eleutherodactylus in southeastern Brazil belong to this Southeast Brazil Clade. Nonetheless, those remaining species form several clusters themselves (e.g., lacteus group and ramagii group) and some species, in the past, have been placed in the same morphological species group (6) along with species that we confidently place in the Southeast Brazil Clade. Until further study can clarify the relationships of these species, and considering the geographic isolation of this group, we assign all of the species of Eleutherodactylus in southeastern Brazil (excluding E. binotatus and E. plicifer) to the genus Ischnocnema.
For the large South American Clade, we resurrect the name Pristimantis Jiménez de la Espada, 1870 (type species, Pristimantis galdi). P. galdi has been placed in the unistrigatus group of Eleutherodactylus (10), a species group that contains species that appear in the South American Clade in our analyses. After this study was completed, we obtained sequence of this species and have confirmed that it is highly nested in the South American Clade. For the clade encompassing members of the anomalus, bufoniformis, and sulcatus groups, we apply the available name Limnophys Jiménez de la Espada, 1871 (type species, Limnophys cornutus). The clade including members of the dolops and nigrovittatus groups, as well as some species of Phrynopus, requires a new generic name, which will be described elsewhere. Further details on all of these taxonomic decisions will be treated elsewhere by us in a monograph on the classification of this complex of >800 species.
Geographic Distributions of Major Clades. Although the three major clades (the Caribbean Clade, the Middle American Clade, and the South American Clade) are strongly associated with geography, each includes species that occur in other areas. In the Caribbean Clade, 24 members of the subgenus Syrrhophus (all except Eleutherodactylus zeus and E. symingtoni) occur in North and Middle America. In the Middle American Clade, five species (Craugastor crassidigitus, C. fitzingeri, C. longirostris, C. opimus, and C. raniformis) occur in both Central and South America, and one (C. biporcatus) is endemic to South America. In the South American Clade, two species are endemic to the Lesser Antilles (Pristimantis euphronides and P. shrevei), nine are endemic to southern Central America (P. altae, P. cerasinus, P. diastema, P. hylaeformis, P. laticorpus, P. museosus, P. pardalis, P. pirrensis, and P. tigrillo), and nine are found in Central and South America (P. achatinus, P. caryophyllaceus, P. cruentus, P. gaigeae, P. moro, P. quidditus, P. ridens, P. taeniatus, and P. vocator). It can be assumed that those species of Craugastor and Pristimantis occurring in both Central and South America achieved their current distributions through Pliocene or later overland dispersal. However, the possibility of over-water dispersal for C. biporcatus or endemic Central American Pristimantis cannot be excluded. Our data set includes none of these endemics, so no determination based on times of divergence can be made. Thus, recent (<3 Mya) speciation or overland dispersal, followed by source-area extinction, or earlier over-water dispersal all remain possible explanations for the current distributions of these species.
Data Collection. In addition to broadly sampling the eleutherodactyline genera, our data set also spans a broad geographic range. Included were seven eleutherodactyline species from Southeast Brazil (plus one species of Brachycephalus), 116 from other parts of South America, 19 Middle American species, and 140 West Indian species. These regions respectively contain totals of 47, 477, 154, and 149 described species.
Tissue samples were hand-collected by using approved methods (Pennsylvania State University Institutional Animal Care and Use Committee approval 17632, for those collected by S.B.H.). They were frozen in liquid nitrogen or preserved in ethanol and kept cold during transport. Additional ethanol-preserved or frozen tissue samples were obtained from museum sources or other researchers. In the laboratory, samples were maintained at -80°C. In addition to sequences generated from these tissues, additional sequences were obtained from GenBank. SI Table 4 lists all individual specimens used in this study, including source, tissue collection number (if applicable), museum voucher number (if available), genes sampled, and corresponding GenBank accession numbers for each sequence. We chose the 12S and 16S genes because of their slower rate of evolution, as compared to other mitochondrial genes, in an attempt to avoid saturation problems (multiple nucleotide substitutions at the same site). The fragment of nuclear gene Tyr was chosen because it has proven informative in other anuran studies (2, 11). The fragment of nuclear gene Rag-1 is from the relatively faster-evolving first half of the gene.
Genomic DNA was extracted using the Qiagen DNeasy tissue extraction kit under the manufacturer's protocol. PCR amplification of samples was performed in 50-ml reactions using AmpliTaq DNA polymerase and ThermoPol buffer (New England Biolabs). For amplification of mitochondrial genes, each reaction contained ThermoPol buffer at 1×, dNTPs at 4 mM, forward and reverse primers at 1 mM, one unit of polymerase, and 1 ml of extracted DNA (more for low-quality tissue). For amplification of nuclear genes, dNTP was increased to 6.6 mM, polymerase to 2.5 units, and extracted DNA to 5 ml. Standard reaction conditions were an initial hold for 5 min at 94°C, followed by 40 cycles of 94°C for 30 s, 50°C for 30 s, and 72°C for 60 s. After 40 cycles, a final hold of 72°C for 7 min was performed before terminating the reaction at 4°C. For low- or nonyielding samples, annealing temperature was dropped from 50°C to 46°C. Primers used in PCR reactions were obtained from the literature or designed in the lab (see SI Table 5). Amplified PCR products were isolated by running on agarose gels and filtering with Millipore Ultrafree-DA gel filters or by vacuum filtration using Millipore Multiscreen filters.
Cycle sequencing was performed by using either the Amersham Pharmacia DYEnamic ET terminator cycle sequencing kit or ABI BigDye terminator cycle sequencing kit under manufacturers' guidelines. DNA sequencing was performed with an ABI 3100-Avant or 3730 genetic analyzer. Cycle sequencing and analysis were performed by the authors or by the Pennsylvania State University Nucleic Acid Facility. All fragments were sequenced in both forward and reverse directions.
Before analyses involving any sequence, all chromatograms were fully inspected, and all sequences were compared against their reverse complement to detect any call errors. Embedded primer sequences were deleted from all sequence fragments before assembly or alignment. Alignments of 12S and 16S sequences were created by using CLUSTAL X under default parameters. Resulting alignments were inspected for errors and compared against secondary structure models available from the European ribosomal RNA database. Regions of uncertain homology were excluded from analysis. Sequences for Rag-1 and Tyr were aligned by eye.
Phylogenetic Analyses. In certain cases, species deemed important to the study were missing large parts of data for one gene. This was due in part to the lack of available tissues or GenBank sequences and in part to the inability to amplify or sequence certain regions. In these instances, the available sequences were used, and unknown regions were coded as missing data. Additionally, 7 species in the 65-species data set were missing data for one of the two nuclear genes (SI Table 4). In the case of Limnophys anomalus, the sequence of the closely related species L. bufoniformis was substituted for the Tyr fragment because L. anomalus was the only species of Limnophys in the 65 species data set and Tyr is the least variable gene in our study.
Model choice for likelihood and minimum evolution analyses were based partly on the limitations of the software packages used. Our preferred model for all analyses was GTR + I + G. However, neither RAxML nor MEGA can employ this model. Therefore, substitute models available in each package close to the GTR + I + G model were chosen for likelihood and minimum evolution analyses.
Bayesian analyses were run for 20,000,000 (280-species), 10,000,000 (146-species), or 2,000,000 (65-species) generations with three heated and one cold chain. Chains were sampled every 1,000, 500, or 100 generations, respectively. The first 25% of samples were discarded as burn-in. To ensure that this was an adequate number of samples discarded for each analysis, plots of log likelihood vs. generation were produced for every Bayesian analysis. In all cases, the region of increasing log likelihood values was encompassed in the first 25% of samples. Convergence for each Bayesian analysis was assessed by using the program Tracer 1.3 (12) to obtain estimated sample sizes for each model parameter (six substitution frequency categories, four nucleotide frequency categories, g-parameter, proportion of invariant sites, tree length, and log likelihood). Estimated sample sizes of each parameter were >100 for both independent runs of nearly all analyses, except for a substitution frequency (ESS = 99) and a nucleotide frequency parameter (ESS = 90) for one of the 146-species runs, the tree length parameter (ESS = 98) for the other 146-species run, and the tree length parameter (ESS = 94) for one run of the 65-species nuclear + mitochondrial data set. All ESS values were >200 when the two independent runs of each analysis were combined.
For the 65-species dataset, ME, ML, and Bayesian analyses were run with the nuclear gene data set separately (data not shown). The same major clades appeared in all of these analyses.
Divergence Time Estimation. Geologic times and boundaries of periods used here are from a recent update (13). The five chosen calibrations were based on several independent lines of evidence. Jamaica did not become permanently emergent until 10 Mya (14, 15), setting a maximum time for basal divergences in the Jamaican clade (including representative species E. gossei and E. luteolus). The Hispaniolan South Island has a similar geologic history (16), setting a maximum time of 10 Mya for the basal divergence in the South Island clade (e.g., between representative species E. thorectes and E. caribe). An Eleutherodactylus fossil in amber from northern Hispaniola (17) is dated 15-20 Mya (18). The fossil is assumed to be in the subgenus Eleutherodactylus based on location (North Island of Hispaniola), age (older than uplift of South Island), and normal head width as compared with members of the subgenus Pelorius (which are wide-headed) (19). This establishes a minimum date for the origin of the lineage leading to the subgenus Eleutherodactylus on Hispaniola (divergence of the subgenera Pelorius and Eleutherodactylus). Australian hylids (pelodryadines) are most closely related to South American hylids (phyllomedusines), and probably arrived by overland dispersal via Antarctica (20, 21). The time window for this dispersal was 35-70 Mya (22-25), providing constraints for the divergence of representatives Litoria caerulea and Agalychnis callidryas.
Multidivtime requires prior estimates for rttm, rtsd, rtrate, rtratesd, brownmean, brownsd, and bigtime. The prior for the rttm (in-group root) parameter was set at 65 Mya based on recent data (26) that place hyloid family divergences near the K-T boundary. Rttmsd, the standard deviation of rttm, was set at 25 Mya. This is a conservative estimate, considering that the true date of the ingroup root is almost certainly less than one standard deviation from rttm in either direction. Higher values of rttm (80 Mya) and rttmsd (40 Mya) were also used to test their results on timing analyses. The prior for rtrate was set at 0.0075 for the mt analysis, 0.0015 for the nuclear analysis, and 0.0035 for the combined analysis. These values were obtained by first dividing the typical root-to-tip branch length for each gene (as determined by estbranches in the Multidivtime package) by rttm, and then taking the average of these values for the genes used in each respective analysis. As a conservative measure, the prior for rtratesd was set equal to rtrate in each analysis. The prior for brownmean was set at 0.0125, arrived at by dividing one by rttm. The prior for brownsd was set equal to brownmean because of the large measure of uncertainty in the prior for brownmean. Bigtime was set at 150 Mya. All other parameters (minab, newk, othk, thek) were maintained at default values. Analyses were run for 1,100,000 generations, with a sample frequency of 100 after a burn-in of 100,000 generations.
In addition to the times presented (SI Table 3), analyses were also performed with higher values for the rttm and rttmsd priors (see SI Text). Changing these priors had little effect on resulting dates, which differed by <3% when both rttm and rttmsd are increased, and by even less if only one of the priors was increased.
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Table 2. Taxonomic listing of all eleutherodactylines and
Brachycephalus (812 sp.)
|
"Eleutherodactylus" binotatus* |
Spix, 1824 |
|
"Eleutherodactylus" plicifer* |
Boulenger, 1888 |
|
"Eleutherodactylus" cruralis† |
Boulenger, 1902 |
|
"Eleutherodactylus" discoidalis† |
Peracca, 1895 |
|
"Eleutherodactylus" heterodactylus† |
Miranda-Ribeiro, 1937 |
|
"Eleutherodactylus" ibischi† |
Reichle, Lötters and de la Riva, 2001 |
|
"Eleutherodactylus" madidi† |
Padial, Gonzáles, and De la Riva, 2005 |
|
"Eleutherodactylus" pereger |
Lynch, 1975 |
|
"Eleutherodactylus" zongoensis† |
Reichle and Köhler, 1997 |
|
"Eleutherodactylus" adercus‡ |
Lynch, 2003 |
|
"Eleutherodactylus" araiodactylus‡ |
Duellman and Pramuk, 1999 |
|
"Eleutherodactylus" babax‡ |
Lynch, 1989 |
|
"Eleutherodactylus" dolops‡ |
Lynch and Duellman, 1980 |
|
"Eleutherodactylus" elassodiscus‡ |
Lynch, 1973 |
|
"Eleutherodactylus" fallaciosus‡ |
Duellman, 2000 |
|
"Eleutherodactylus" flavomaculatus‡ |
Parker, 1938 |
|
"Eleutherodactylus" latens‡ |
Lynch, 1989 |
|
"Eleutherodactylus" lucida‡ |
Cannatella, 1984 |
|
"Eleutherodactylus" lundbergi‡ |
Lehr, 2005 |
|
"Eleutherodactylus" mantipus‡ |
Boulenger, 1908 |
|
"Eleutherodactylus" nebulanastes‡ |
Cannatella, 1984 |
|
"Eleutherodactylus" nigrovittatus‡ |
Andersson, 1945 |
|
"Phrynopus" brunneus‡ |
Lynch, 1975 |
|
"Phrynopus" peraccai‡ |
Lynch, 1975 |
|
Adelophryne adiastola |
Hoogmoed and Lescure, 1984 |
|
Adelophryne baturitensis |
Hoogmoed, Borges, and Cascon, 1994 |
|
Adelophryne gutturosa |
Hoogmoed and Lescure, 1984 |
|
Adelophryne maranguapensis |
Hoogmoed, Borges, and Cascon, 1994 |
|
Adelophryne pachydactyla |
Hoogmoed, Borges, and Cascon, 1994 |
|
Atopophrynus syntomopus |
Lynch and Ruiz-Carranza, 1982 |
|
Barycholos pulcher |
Boulenger, 1898 |
|
Barycholos ternetzi |
Miranda-Ribeiro, 1937 |
|
Brachycephalus alipioi |
Pombal and Gasparini, 2006 |
|
Brachycephalus brunneus |
Ribero, Alves, Haddad, and Dos Reis, 2005 |
|
Brachycephalus didactylus |
Izecksohn, 1971 |
|
Brachycephalus ephippium |
Spix, 1824 |
|
Brachycephalus ferruginus |
Alves, Ribeiro, Haddad, and Dos Reis, 2006 |
|
Brachycephalus hermogenesi |
Giaretta and Sawaya, 1998 |
|
Brachycephalus izecksohni |
Ribero, Alves, Haddad, and Dos Reis, 2005 |
|
Brachycephalus nodoterga |
Miranda-Ribeiro, 1920 |
|
Brachycephalus pernix |
Pombal, Wistuba, and Bornschein, 1998 |
|
Brachycephalus pombali |
Alves, Ribeiro, Haddad, and Dos Reis, 2006 |
|
Brachycephalus vertebralis |
Pombal, 2001 |
|
Craugastor adamastus |
Campbell, 1994 |
|
Craugastor alfredi |
Boulenger, 1898 |
|
Craugastor amniscola |
Campbell and Savage, 2000 |
|
Craugastor anciano |
Savage, McCranie and Wilson, 1988 |
|
Craugastor andi |
Savage, 1974 |
|
Craugastor angelicus |
Savage, 1975 |
|
Craugastor aphanus |
Campbell, 1994 |
|
Craugastor augusti |
Dugès, 1879 |
|
Craugastor aurilegulus |
Savage, McCranie and Wilson, 1988 |
|
Craugastor azueroensis |
Savage, 1975 |
|
Craugastor batrachylus |
Taylor, 1940 |
|
Craugastor berkenbuschii |
Peters, 1870 |
|
Craugastor biporcatus |
Peters, 1863 |
|
Craugastor bocourti |
Brocchi, 1877 |
|
Craugastor bransfordii |
Cope, 1886 |
|
Craugastor brocchi |
Boulenger, 1882 |
|
Craugastor campbelli |
Smith, 2005 |
|
Craugastor catalinae |
Campbell and Savage, 2000 |
|
Craugastor chac |
Savage, 1987 |
|
Craugastor charadra |
Campbell and Savage, 2000 |
|
Craugastor chrysozetetes |
McCranie, Savage and Wilson, 1989 |
|
Craugastor coffeus |
McCranie and Köhler, 1999 |
|
Craugastor crassidigitus |
Taylor, 1952 |
|
Craugastor cruzi |
McCranie, Savage and Wilson, 1989 |
|
Craugastor cuaquero |
Savage, 1980 |
|
Craugastor cyanochthebius |
McCranie and Smith, 2006 |
|
Craugastor daryi |
Ford and Savage, 1984 |
|
Craugastor decoratus |
Taylor, 1942 |
|
Craugastor emcelae |
Lynch, 1985 |
|
Craugastor emleni |
Dunn and Emlen, 1932 |
|
Craugastor epochthidius |
McCranie and Wilson, 1997 |
|
Craugastor escoces |
Savage, 1975 |
|
Craugastor fecundus |
McCranie and Wilson, 1997 |
|
Craugastor fitzingeri |
Schmidt, 1857 |
|
Craugastor fleischmanni |
Boettger, 1892 |
|
Craugastor galacticorhinus |
Canseco-Márquez and Smith, 2004 |
|
Craugastor glaucus |
Lynch, 1967 |
|
Craugastor gollmeri |
Peters, 1863 |
|
Craugastor greggi |
Bumzahem, 1955 |
|
Craugastor guerreroensis |
Lynch, 1967 |
|
Craugastor gulosus |
Cope, 1875 |
|
Craugastor hobartsmithi |
Taylor, 1937 |
|
Craugastor inachus |
Campbell and Savage, 2000 |
|
Craugastor jota |
Lynch, 1980 |
|
Craugastor laevissimus |
Werner, 1896 |
|
Craugastor laticeps |
Duméril, 1853 |
|
Craugastor lauraster |
Savage, McCranie and Espinal, 1996 |
|
Craugastor lineatus |
Brocchi, 1879 |
|
Craugastor loki |
Shannon and Werler, 1955 |
|
Craugastor longirostris |
Boulenger, 1898 |
|
Craugastor matudai |
Taylor, 1941 |
|
Craugastor megacephalus |
Cope, 1875 |
|
Craugastor megalotympanum |
Shannon and Werler, 1955 |
|
Craugastor melanostictus |
Cope, 1875 |
|
Craugastor merendonensis |
Schmidt, 1933 |
|
Craugastor mexicanus |
Brocchi, 1877 |
|
Craugastor milesi |
Schmidt, 1933 |
|
Craugastor mimus |
Taylor, 1955 |
|
Craugastor monnichorum |
Dunn, 1940 |
|
Craugastor montanus |
Lynch, 1965 |
|
Craugastor myllomyllon |
Savage, 2000 |
|
Craugastor nefrens |
Smith, 2005 |
|
Craugastor noblei |
Barbour and Dunn, 1921 |
|
Craugastor obesus |
Barbour, 1928 |
|
Craugastor occidentalis |
Taylor, 1941 |
|
Craugastor olanchano |
McCranie and Wilson, 1999 |
|
Craugastor omiltemanus |
Günther, 1900 |
|
Craugastor omoaensis |
McCranie and Wilson, 1997 |
|
Craugastor opimus |
Savage and Myers, 2002 |
|
Craugastor palenque |
Campbell and Savage, 2000 |
|
Craugastor pechorum |
McCranie and Wilson, 1999 |
|
Craugastor pelorus |
Campbell and Savage, 2000 |
|
Craugastor persimilis |
Barbour, 1926 |
|
Craugastor phasma |
Lips and Savage, 1996 |
|
Craugastor podiciferus |
Cope, 1875 |
|
Craugastor polymniae |
Campbell, Lamar and Hillis, 1989 |
|
Craugastor polyptychus |
Cope, 1886 |
|
Craugastor pozo |
Johnson and Savage, 1995 |
|
Craugastor psephosypharus |
Campbell, Savage and Meyer, 1994 |
|
Craugastor punctariolus |
Peters, 1863 |
|
Craugastor pygmaeus |
Taylor, 1937 |
|
Craugastor raniformis |
Boulenger, 1896 |
|
Craugastor ranoides |
Cope, 1886 |
|
Craugastor rayo |
Savage and DeWeese, 1979 |
|
Craugastor rhodopis |
Cope, 1867 |
|
Craugastor rhyacobatrachus |
Campbell and Savage, 2000 |
|
Craugastor rivulus |
Campbell and Savage, 2000 |
|
Craugastor rostralis |
Werner, 1896 |
|
Craugastor rugosus |
Peters, 1873 |
|
Craugastor rugulosus |
Cope, 1870 |
|
Craugastor rupinius |
Campbell and Savage, 2000 |
|
Craugastor sabrinus |
Campbell and Savage, 2000 |
|
Craugastor saltuarius |
McCranie and Wilson, 1997 |
|
Craugastor sandersoni |
Schmidt, 1941 |
|
Craugastor silvicola |
Lynch, 1967 |
|
Craugastor spatulatus |
Smith, 1939 |
|
Craugastor stadelmani |
Schmidt, 1936 |
|
Craugastor stejnegerianus |
Cope, 1893 |
|
Craugastor stuarti |
Lynch, 1967 |
|
Craugastor tabasarae |
Savage, Hollingsworth, Lips, and Jaslow, 2004 |
|
Craugastor talamancae |
Dunn, 1931 |
|
Craugastor tarahumaraensis |
Taylor, 1940 |
|
Craugastor taurus |
Taylor, 1958 |
|
Craugastor taylori |
Lynch, 1966 |
|
Craugastor trachydermus |
Campbell, 1994 |
|
Craugastor underwoodi |
Boulenger, 1896 |
|
Craugastor uno |
Savage, 1984 |
|
Craugastor vocalis |
Taylor, 1940 |
|
Craugastor vulcani |
Shannon and Werler, 1955 |
|
Craugastor xucanebi |
Stuart, 1941 |
|
Craugastor yucatanensis |
Lynch, 1965 |
|
Dischidodactylus colonnelloi |
Ayarzagüena, 1985 |
|
Dischidodactylus duidensis |
Rivero, 1968 |
|
Eleutherodactylus (Eleutherodactylus) abbotti |
Cochran, 1923 |
|
Eleutherodactylus (Eleutherodactylus) amplinympha |
Kaiser, Green and Schmid, 1994 |
|
Eleutherodactylus (Eleutherodactylus) antillensis |
Reinhardt and Lütken, 1863 |
|
Eleutherodactylus (Eleutherodactylus) audanti |
Cochran, 1934 |
|
Eleutherodactylus (Eleutherodactylus) auriculatoides |
Noble, 1923 |
|
Eleutherodactylus (Eleutherodactylus) auriculatus |
Cope, 1862 |
|
Eleutherodactylus (Eleutherodactylus) barlagnei |
Lynch, 1965 |
|
Eleutherodactylus (Eleutherodactylus) bartonsmithi |
Schwartz, 1960 |
|
Eleutherodactylus (Eleutherodactylus) brittoni |
Schmidt, 1920 |
|
Eleutherodactylus (Eleutherodactylus) cochranae |
Grant, 1932 |
|
Eleutherodactylus (Eleutherodactylus) cooki |
Grant, 1932 |
|
Eleutherodactylus (Eleutherodactylus) coqui |
Thomas, 1966 |
|
Eleutherodactylus (Eleutherodactylus) eileenae |
Dunn, 1926 |
|
Eleutherodactylus (Eleutherodactylus) eneidae |
Rivero, 1959 |
|
Eleutherodactylus (Eleutherodactylus) flavescens |
Noble, 1923 |
|
Eleutherodactylus (Eleutherodactylus) fowleri |
Schwartz, 1973 |
|
Eleutherodactylus (Eleutherodactylus) glamyrus |
Estrada and Hedges, 1997 |
|
Eleutherodactylus (Eleutherodactylus) gryllus |
Schmidt, 1920 |
|
Eleutherodactylus (Eleutherodactylus) guantanamera |
Hedges, Estrada and Thomas, 1992 |
|
Eleutherodactylus (Eleutherodactylus) haitianus |
Barbour, 1942 |
|
Eleutherodactylus (Eleutherodactylus) hedricki |
Rivero, 1963 |
|
Eleutherodactylus (Eleutherodactylus) ionthus |
Schwartz, 1960 |
|
Eleutherodactylus (Eleutherodactylus) jasperi |
Drewry and Jones, 1976 |
|
Eleutherodactylus (Eleutherodactylus) johnstonei |
Barbour, 1914 |
|
Eleutherodactylus (Eleutherodactylus) karlschmidti |
Grant, 1931 |
|
Eleutherodactylus (Eleutherodactylus) lamprotes |
Schwartz, 1973 |
|
Eleutherodactylus (Eleutherodactylus) leberi |
Schwartz, 1965 |
|
Eleutherodactylus (Eleutherodactylus) locustus |
Schmidt, 1920 |
|
Eleutherodactylus (Eleutherodactylus) mariposa |
Hedges, Estrada and Thomas, 1992 |
|
Eleutherodactylus (Eleutherodactylus) martinicensis |
Tschudi, 1838 |
|
Eleutherodactylus (Eleutherodactylus) melacara |
Hedges, Estrada and Thomas, 1992 |
|
Eleutherodactylus (Eleutherodactylus) minutus |
Noble, 1923 |
|
Eleutherodactylus (Eleutherodactylus) montanus |
Schmidt, 1919 |
|
Eleutherodactylus (Eleutherodactylus) parabates |
Schwartz, 1964 |
|
Eleutherodactylus (Eleutherodactylus) patriciae |
Schwartz, 1965 |
|
Eleutherodactylus (Eleutherodactylus) pinchoni |
Schwartz, 1967 |
|
Eleutherodactylus (Eleutherodactylus) pituinus |
Schwartz, 1965 |
|
Eleutherodactylus (Eleutherodactylus) poolei |
Cochran, 1938 |
|
Eleutherodactylus (Eleutherodactylus) portoricensis |
Schmidt, 1927 |
|
Eleutherodactylus (Eleutherodactylus) principalis |
Estrada and Hedges, 1997 |
|
Eleutherodactylus (Eleutherodactylus) richmondi |
Stejneger, 1904 |
|
Eleutherodactylus (Eleutherodactylus) ronaldi |
Schwartz, 1960 |
|
Eleutherodactylus (Eleutherodactylus) schwartzi |
Thomas, 1966 |
|
Eleutherodactylus (Eleutherodactylus) unicolor |
Stejneger, 1904 |
|
Eleutherodactylus (Eleutherodactylus) varians |
Gundlach and Peters, 1864 |
|
Eleutherodactylus (Eleutherodactylus) wetmorei |
Cochran, 1932 |
|
Eleutherodactylus (Eleutherodactylus) wightmanae |
Schmidt, 1920 |
|
Eleutherodactylus (Euhyas) acmonis |
Schwartz, 1960 |
|
Eleutherodactylus (Euhyas) adelus |
Diaz, Cadiz and Hedges, 2003 |
|
Eleutherodactylus (Euhyas) albipes |
Barbour and Shreve, 1937 |
|
Eleutherodactylus (Euhyas) alcoae |
Schwartz, 1971 |
|
Eleutherodactylus (Euhyas) alticola |
Lynn, 1937 |
|
Eleutherodactylus (Euhyas) amadeus |
Hedges, Thomas and Franz, 1987 |
|
Eleutherodactylus (Euhyas) andrewsi |
Lynn, 1937 |
|
Eleutherodactylus (Euhyas) apostates |
Schwartz, 1973 |
|
Eleutherodactylus (Euhyas) armstrongi |
Noble and Hassler, 1933 |
|
Eleutherodactylus (Euhyas) atkinsi |
Dunn, 1925 |
|
Eleutherodactylus (Euhyas) bakeri |
Cochran, 1935 |
|
Eleutherodactylus (Euhyas) blairhedgesi |
Estrada, Diaz and Rodriguez, 1998 |
|
Eleutherodactylus (Euhyas) bresslerae |
Schwartz, 1960 |
|
Eleutherodactylus (Euhyas) brevirostris |
Shreve, 1936 |
|
Eleutherodactylus (Euhyas) caribe |
Hedges and Thomas, 1992 |
|
Eleutherodactylus (Euhyas) casparii |
Dunn, 1926 |
|
Eleutherodactylus (Euhyas) cavernicola |
Lynn, 1954 |
|
Eleutherodactylus (Euhyas) corona |
Hedges and Thomas, 1992 |
|
Eleutherodactylus (Euhyas) cubanus |
Barbour, 1942 |
|
Eleutherodactylus (Euhyas) cundalli |
Dunn, 1926 |
|
Eleutherodactylus (Euhyas) cuneatus |
Cope, 1862 |
|
Eleutherodactylus (Euhyas) darlingtoni |
Cochran, 1935 |
|
Eleutherodactylus (Euhyas) dimidiatus |
Cope, 1862 |
|
Eleutherodactylus (Euhyas) dolomedes |
Hedges and Thomas, 1992 |
|
Eleutherodactylus (Euhyas) emiliae |
Dunn, 1926 |
|
Eleutherodactylus (Euhyas) etheridgei |
Schwartz, 1958 |
|
Eleutherodactylus (Euhyas) eunaster |
Schwartz, 1973 |
|
Eleutherodactylus (Euhyas) furcyensis |
Shreve and Williams, 1963 |
|
Eleutherodactylus (Euhyas) fuscus |
Lynn and Dent, 1943 |
|
Eleutherodactylus (Euhyas) glandulifer |
Cochran, 1935 |
|
Eleutherodactylus (Euhyas) glanduliferoides |
Shreve, 1936 |
|
Eleutherodactylus (Euhyas) glaphycompus |
Schwartz, 1973 |
|
Eleutherodactylus (Euhyas) glaucoreius |
Schwartz and Fowler, 1973 |
|
Eleutherodactylus (Euhyas) goini |
Schwartz, 1960 |
|
Eleutherodactylus (Euhyas) gossei |
Dunn, 1926 |
|
Eleutherodactylus (Euhyas) grabhami |
Dunn, 1926 |
|
Eleutherodactylus (Euhyas) grahami |
Schwartz, 1979 |
|
Eleutherodactylus (Euhyas) greyi |
Dunn, 1926 |
|
Eleutherodactylus (Euhyas) griphus |
Crombie, 1986 |
|
Eleutherodactylus (Euhyas) guanahacabibes |
Estrada and Rodriguez, 1985 |
|
Eleutherodactylus (Euhyas) gundlachi |
Schmidt, 1920 |
|
Eleutherodactylus (Euhyas) heminota |
Shreve and Williams, 1963 |
|
Eleutherodactylus (Euhyas) iberia |
Estrada and Hedges, 1996 |
|
Eleutherodactylus (Euhyas) intermedius |
Barbour and Shreve, 1937 |
|
Eleutherodactylus (Euhyas) jamaicensis |
Barbour, 1910 |
|
Eleutherodactylus (Euhyas) jaumei |
Estrada and Alonso, 1997 |
|
Eleutherodactylus (Euhyas) jugans |
Cochran, 1937 |
|
Eleutherodactylus (Euhyas) junori |
Dunn, 1926 |
|
Eleutherodactylus (Euhyas) klinikowskii |
Schwartz, 1959 |
|
Eleutherodactylus (Euhyas) lentus |
Cope, 1862 |
|
Eleutherodactylus (Euhyas) leoncei |
Shreve and Williams, 1963 |
|
Eleutherodactylus (Euhyas) limbatus |
Cope, 1862 |
|
Eleutherodactylus (Euhyas) lucioi |
Schwartz, 1980 |
|
Eleutherodactylus (Euhyas) luteolus |
Gosse, 1851 |
|
Eleutherodactylus (Euhyas) maestrensis |
Díaz, Cádiz, and Navarro, 2005 |
|
Eleutherodactylus (Euhyas) michaelschmidi |
Díaz, Cádiz, and Navarro, 2007 |
|
Eleutherodactylus (Euhyas) monensis |
Meerwarth, 1901 |
|
Eleutherodactylus (Euhyas) nubicola |
Dunn, 1926 |
|
Eleutherodactylus (Euhyas) orcutti |
Dunn, 1928 |
|
Eleutherodactylus (Euhyas) orientalis |
Barbour and Shreve, 1973 |
|
Eleutherodactylus (Euhyas) oxyrhyncus |
Duméril and Bibron, 1841 |
|
Eleutherodactylus (Euhyas) pantoni |
Dunn, 1926 |
|
Eleutherodactylus (Euhyas) paulsoni |
Schwartz, 1964 |
|
Eleutherodactylus (Euhyas) pentasyringos |
Schwartz and Fowler, 1973 |
|
Eleutherodactylus (Euhyas) pezopetrus |
Schwartz, 1960 |
|
Eleutherodactylus (Euhyas) pictissimus |
Cochran, 1935 |
|
Eleutherodactylus (Euhyas) pinarensis |
Dunn, 1926 |
|
Eleutherodactylus (Euhyas) planirostris |
Cope, 1862 |
|
Eleutherodactylus (Euhyas) probolaeus |
Schwartz, 1965 |
|
Eleutherodactylus (Euhyas) rhodesi |
Schwartz, 1980 |
|
Eleutherodactylus (Euhyas) ricordii |
Duméril and Bibron, 1841 |
|
Eleutherodactylus (Euhyas) riparius |
Estrada and Hedges, 1998 |
|
Eleutherodactylus (Euhyas) rivularis |
Diaz, Estrada and Hedges, 2001 |
|
Eleutherodactylus (Euhyas) rogersi |
Goin, 1955 |
|
Eleutherodactylus (Euhyas) rufifemoralis |
Noble and Hassler, 1933 |
|
Eleutherodactylus (Euhyas) schmidti |
Noble, 1923 |
|
Eleutherodactylus (Euhyas) sciagraphus |
Schwartz, 1973 |
|
Eleutherodactylus (Euhyas) semipalmatus |
Shreve, 1936 |
|
Eleutherodactylus (Euhyas) simulans |
Díaz and Fong, 2001 |
|
Eleutherodactylus (Euhyas) sisyphodemus |
Crombie, 1977 |
|
Eleutherodactylus (Euhyas) tetajulia |
Estrada and Hedges, 1996 |
|
Eleutherodactylus (Euhyas) thomasi |
Schwartz, 1959 |
|
Eleutherodactylus (Euhyas) thorectes |
Hedges, 1988 |
|
Eleutherodactylus (Euhyas) toa |
Estrada and Hedges, 1991 |
|
Eleutherodactylus (Euhyas) tonyi |
Estrada and Hedges, 1997 |
|
Eleutherodactylus (Euhyas) turquinensis |
Barbour and Shreve, 1937 |
|
Eleutherodactylus (Euhyas) varleyi |
Dunn, 1925 |
|
Eleutherodactylus (Euhyas) ventrilineatus |
Shreve, 1936 |
|
Eleutherodactylus (Euhyas) warreni |
Schwartz, 1976 |
|
Eleutherodactylus (Euhyas) weinlandi |
Barbour, 1914 |
|
Eleutherodactylus (Euhyas) zugi |
Schwartz, 1958 |
|
Eleutherodactylus (Pelorius) chlorophenax |
Schwartz, 1976 |
|
Eleutherodactylus (Pelorius) hypostenor |
Schwartz, 1965 |
|
Eleutherodactylus (Pelorius) inoptatus |
Barbour, 1914 |
|
Eleutherodactylus (Pelorius) nortoni |
Schwartz, 1976 |
|
Eleutherodactylus (Pelorius) parapelates |
Hedges and Thomas, 1987 |
|
Eleutherodactylus (Pelorius) ruthae |
Noble, 1923 |
|
Eleutherodactylus (Syrrhophus) albolabris |
Lynch, 1991 |
|
Eleutherodactylus (Syrrhophus) angustidigitorum |
Taylor, 1940 |
|
Eleutherodactylus (Syrrhophus) cystignathoides |
Cope, 1817 |
|
Eleutherodactylus (Syrrhophus) dennisi |
Lynch, 1970 |
|
Eleutherodactylus (Syrrhophus) dilatus |
Davis and Dixon, 1955 |
|
Eleutherodactylus (Syrrhophus) grandis |
Dixon, 1957 |
|
Eleutherodactylus (Syrrhophus) guttilatus |
Cope, 1979 |
|
Eleutherodactylus (Syrrhophus) interorbitalis |
Langebartel and Shannon, 1956 |
|
Eleutherodactylus (Syrrhophus) leprus |
Cope, 1879 |
|
Eleutherodactylus (Syrrhophus) longipes |
Baird, 1859 |
|
Eleutherodactylus (Syrrhophus) marnockii |
Cope, 1878 |
|
Eleutherodactylus (Syrrhophus) maurus |
Hedges, 1989 |
|
Eleutherodactylus (Syrrhophus) modestus |
Taylor, 1942 |
|
Eleutherodactylus (Syrrhophus) nitidus |
Peters, 1870 |
|
Eleutherodactylus (Syrrhophus) nivicolimae |
Dixon and Webb, 1996 |
|
Eleutherodactylus (Syrrhophus) pallidus |
Duellman, 1958 |
|
Eleutherodactylus (Syrrhophus) pipilans |
Taylor, 1940 |
|
Eleutherodactylus (Syrrhophus) rubrimaculatus |
Taylor and Smith, 1945 |
|
Eleutherodactylus (Syrrhophus) rufescens |
Duellman and Dixon, 1959 |
|
Eleutherodactylus (Syrrhophus) saxatilis |
Webb, 1962 |
|
Eleutherodactylus (Syrrhophus) symingtoni |
Schwartz, 1957 |
|
Eleutherodactylus (Syrrhophus) syristes |
Hoyt, 1965 |
|
Eleutherodactylus (Syrrhophus) teretistes |
Duellman, 1958 |
|
Eleutherodactylus (Syrrhophus) verrucipes |
Cope, 1885 |
|
Eleutherodactylus (Syrrhophus) verruculatus |
Peters, 1870 |
|
Eleutherodactylus (Syrrhophus) zeus |
Schwartz, 1958 |
|
Eleutherodactylus counouspeus |
Schwartz, 1964 |
|
Euparkerella brasiliensis |
Parker, 1926 |
|
Euparkerella cochranae |
Izecksohn, 1988 |
|
Euparkerella robusta |
Izecksohn, 1988 |
|
Euparkerella tridactyla |
Izecksohn, 1988 |
|
Geobatrachus walkeri |
Ruthveni, 1915 |
|
Holoaden bradei |
Lutz, 1958 |
|
Holoaden luederwaldti |
Miranda-Ribero, 1920 |
|
Ischnocnema bilineata |
Bokermann, 1975 |
|
Ischnocnema bolbodactyla |
Lutz, 1925 |
|
Ischnocnema epipeda |
Heyer, 1984 |
|
Ischnocnema erythromera |
Heyer, 1984 |
|
Ischnocnema gehrti |
Miranda-Ribeiro, 1926 |
|
Ischnocnema gualteri |
Lutz, 1974 |
|
Ischnocnema guentheri |
Steindachner, 1864 |
|
Ischnocnema henselii |
Peters, 1870 |
|
Ischnocnema hoehnei |
Lutz, 1958 |
|
Ischnocnema holti |
Cochran, 1948 |
|
Ischnocnema izecksohni |
Caramaschi and Kisteumacher, 1989 |
|
Ischnocnema juipoca |
Sazima and Cardoso, 1978 |
|
Ischnocnema lactea |
Miranda-Ribeiro, 1923 |
|
Ischnocnema manezinho |
Garcia, 1996 |
|
Ischnocnema nasuta |
Lutz, 1925 |
|
Ischnocnema nigriventris |
Lutz, 1925 |
|
Ischnocnema octavioi |
Bokermann, 1965 |
|
Ischnocnema oeus |
Heyer, 1984 |
|
Ischnocnema paranaensis |
Langone and Segalla, 1996 |
|
Ischnocnema parva |
Girard, 1853 |
|
Ischnocnema paulodutrai |
Bokermann, 1975 |
|
Ischnocnema pusilla |
Bokermann, 1967 |
|
Ischnocnema ramagii |
Boulenger, 1888 |
|
Ischnocnema randorum |
Heyer, 1985 |
|
Ischnocnema sambaqui |
Castanho and Haddad, 2000 |
|
Ischnocnema spanios |
Heyer, 1985 |
|
Ischnocnema venancioi |
Lutz, 1958 |
|
Ischnocnema verrucosa |
Reinhardt and Lütken, 1862 |
|
Ischnocnema vinhai |
Bokermann, 1975 |
|
Limnophys anatipes |
Lynch and Myers, 1983 |
|
Limnophys anomalus |
Boulenger, 1898 |
|
Limnophys bufoniformis |
Boulenger, 1896 |
|
Limnophys cadenai |
Lynch, 1986 |
|
Limnophys cerastes |
Lynch, 1975 |
|
Limnophys cheiroplethus |
Lynch, 1990 |
|
Limnophys cornutus |
Jiménez de la Espada, 1871 |
|
Limnophys helonotus |
Lynch, 1975 |
|
Limnophys ingeri |
Cochran and Goin, 1961 |
|
Limnophys laticorpus |
Myers and Lynch, 1997 |
|
Limnophys necerus |
Lynch, 1975 |
|
Limnophys necopinus |
Lynch, 1997 |
|
Limnophys ruizi |
Lynch, 1981 |
|
Limnophys sulcatus |
Cope, 1874 |
|
Limnophys zygodactylus |
Lynch and Myers, 1983 |
|
Oreobates choristolemma |
Harvey and Sheehy, 2005 |
|
Oreobates quixensis |
Jiménez de la Espada, 1872 |
|
Oreobates sanctaecrucis |
Harvey and Keck, 1995 |
|
Oreobates sanderi |
Padial, Reichle, and De la Riva |
|
Oreobates saxatilis |
Duellman, 1990 |
|
Oreobates simmonsi |
Padial, Reichle, and De La Riva, 2005 |
|
Phrynopus adenobrachius |
Ardila-Robayo, Ruiz-Carranza, and Barrera-Rodriguez, 1996 |
|
Phrynopus adenopleurus |
Aguayo-Vedia and Harvey, 2001 |
|
Phrynopus bagrecitoi |
Lynch, 1986 |
|
Phrynopus barthlenae |
Lehr and Aguilar, 2002 |
|
Phrynopus boettgeri |
Lehr, 2006 |
|
Phrynopus bracki |
Hedges, 1990 |
|
Phrynopus bufoides |
Lehr, Lundberg, and Aguilar, 2005 |
|
Phrynopus columbianus |
Werner, 1899 |
|
Phrynopus cophites |
Lynch, 1975 |
|
Phrynopus dagmarae |
Lehr, Aguilar, and Köhler, 2002 |
|
Phrynopus heimorum |
Lehr, 2001 |
|
Phrynopus horstpauli |
Lehr, Köhler, and Ponce, 2000 |
|
Phrynopus iatamasi |
Aguayo-Vedia and Harvey, 2001 |
|
Phrynopus juninensis |
Shreve, 1938 |
|
Phrynopus kauneorum |
Lehr, Aguilar, and Köhler, 2002 |
|
Phrynopus kempffi |
De la Riva, 1992 |
|
Phrynopus montium |
Shreve, 1938 |
|
Phrynopus nanus |
Goin and Cochran, 1963 |
|
Phrynopus parkeri |
Lynch, 1975 |
|
Phrynopus paucari |
Lehr, Lundberg, and Aguilar,2005 |
|
Phrynopus peruanus |
Peters, 1873 |
|
Phrynopus peruvianus |
Noble, 1921 |
|
Phrynopus pesantesi |
Lehr, Lundberg, and Aguilar, 2005 |
|
Phrynopus pinguis |
Harvey and Ergueta, 1998 |
|
Phrynopus tautzorum |
Lehr and Aguilar, 2003 |
|
Phrynopus thompsoni |
Duellman, 2000 |
|
Phrynopus wettsteini |
Parker, 1932 |
|
Phyllonastes carrascoicola |
De la Riva and Köhler, 1998 |
|
Phyllonastes duellmani |
Lehr, Aguilar, and Lundberg, 2004 |
|
Phyllonastes heyeri |
Lynch, 1986 |
|
Phyllonastes lochites |
Lynch, 1976 |
|
Phyllonastes lynchi |
Duellman, 1991 |
|
Phyllonastes myrmecoides |
Lynch, 1976 |
|
Phyllonastes ritarasquinae |
Köhler, 2000 |
|
Phyzelaphryne miriamae |
Heyer, 1977 |
|
Pristimantis aaptus |
Lynch and Lescure, 1980 |
|
Pristimantis acatallelus |
Lynch and Ruíz-Carranza, 1983 |
|
Pristimantis acerus |
Lynch and Duellman, 1980 |
|
Pristimantis achatinus |
Boulenger, 1898 |
|
Pristimantis actinolaimus |
Lynch and Rueda-Almonacid, 1998 |
|
Pristimantis actites |
Lynch, 1979 |
|
Pristimantis acuminatus |
Shreve, 1935 |
|
Pristimantis acutirostris |
Lynch, 1984 |
|
Pristimantis aemulatus |
Ruíz-Carranza, Lynch and Ardila-Robayo, 1997 |
|
Pristimantis affinis |
Werner, 1899 |
|
Pristimantis alalocophus |
Roa-Trujillo and Ruíz-Carranza, 1991 |
|
Pristimantis albericoi |
Lynch and Ruíz-Carranza, 1996 |
|
Pristimantis altae |
Dunn, 1942 |
|
Pristimantis altamazonicus |
Barbour and Dunn, 1921 |
|
Pristimantis anemerus |
Duellman and Pramuk, 1999 |
|
Pristimantis angustilineata |
Lynch, 1998 |
|
Pristimantis aniptopalmatus |
Duellman and Hedges, 2005 |
|
Pristimantis anolirex |
Lynch, 1983 |
|
Pristimantis anotis |
Walker and Test, 1955 |
|
Pristimantis anthrax |
Lynch, 2001 |
|
Pristimantis apiculatus |
Lynch and Burrowes, 1990 |
|
Pristimantis appendiculatus |
Werner, 1894 |
|
Pristimantis aracamuni |
Barrio-Amores and Molina, 2006 |
|
Pristimantis ardalonychus |
Duellman and Pramuk, 1999 |
|
Pristimantis ashkapara |
Köhler, 2000 |
|
Pristimantis atrabracus |
Duellman and Pramuk, 1999 |
|
Pristimantis atratus |
Lynch, 1979 |
|
Pristimantis aurantiguttatus |
Ruíz-Carranza, Lynch and Ardila-Robayo, 1997 |
|
Pristimantis aureolineatus |
Guayasamin et al. 2006 |
|
Pristimantis avicuporum |
Duellman and Pramuk, 1999 |
|
Pristimantis avius |
Myers and Donnelly, 1997 |
|
Pristimantis bacchus |
Lynch, 1984 |
|
Pristimantis baiotis |
Lynch, 1998 |
|
Pristimantis balionotus |
Lynch, 1979 |
|
Pristimantis baryecuus |
Lynch, 1979 |
|
Pristimantis batrachites |
Lynch, 2003 |
|
Pristimantis bearsei |
Duellman, 1992 |
|
Pristimantis bellona |
Lynch, 1992 |
|
Pristimantis bernali |
Lynch, 1986 |
|
Pristimantis bicolor |
Rueda-Almonacid and Lynch, 1983 |
|
Pristimantis bicumulus |
Peters, 1863 |
|
Pristimantis bipunctatus |
Duellman and Hedges, 2005 |
|
Pristimantis bisignatus |
Werner, 1899 |
|
Pristimantis boconoensis |
Rivero and Mayorga, 1973 |
|
Pristimantis bogotensis |
Peters, 1863 |
|
Pristimantis boulengeri |
Lynch, 1981 |
|
Pristimantis brevifrons |
Lynch, 1981 |
|
Pristimantis briceni |
Boulenger, 1903 |
|
Pristimantis bromeliaceus |
Lynch, 1979 |
|
Pristimantis buccinator |
Rodriguez, 1994 |
|
Pristimantis buckleyi |
Boulenger, 1882 |
|
Pristimantis cabrerai |
Cochran and Goin, 1970 |
|
Pristimantis cacao |
Lynch, 1992 |
|
Pristimantis cajamarcensis |
Barbour and Noble, 1920 |
|
Pristimantis calcaratus |
Boulenger, 1908 |
|
Pristimantis calcarulatus |
Lynch, 1976 |
|
Pristimantis caliginosus |
Lynch, 1996 |
|
Pristimantis cantitans |
Myers and Donnelly, 1996 |
|
Pristimantis capitonis |
Lynch, 1998 |
|
Pristimantis caprifer |
Lynch, 1977 |
|
Pristimantis carlossanchezi |
Arroyo, 2007 |
|
Pristimantis carmelitae |
Ruthven, 1922 |
|
Pristimantis carranguerorum |
Lynch, 1994 |
|
Pristimantis carvalhoi |
Lutz, 1952 |
|
Pristimantis caryophyllaceus |
Barbour, 1928 |
|
Pristimantis cavernibardus |
Myers and Donnelly, 1997 |
|
Pristimantis celator |
Lynch, 1976 |
|
Pristimantis cerasinus |
Cope, 1875 |
|
Pristimantis ceuthospilus |
Duellman and Wild, 1993 |
|
Pristimantis chalceus |
Peters, 1873 |
|
Pristimantis charlottevillensis |
Kaiser, Dwyer, Feichtinger and Schmid, 1995 |
|
Pristimantis chiastonotus |
Lynch and Hoogmoed, 1977 |
|
Pristimantis chloronotus |
Lynch, 1969 |
|
Pristimantis chrysops |
Lynch and Ruíz-Carranza, 1996 |
|
Pristimantis citriogaster |
Duellman, 1992 |
|
Pristimantis colodactylus |
Lynch, 1979 |
|
Pristimantis colomai |
Lynch and Duellman, 1997 |
|
Pristimantis colostichos |
La Marca and Smith, 1982 |
|
Pristimantis condor |
Lynch and Duellman, 1980 |
|
Pristimantis conspicillatus |
Günther, 1858 |
|
Pristimantis cordovae |
Lehr and Duellman, 2007 |
|
Pristimantis corniger |
Lynch and Suárez-Mayorga, 2003 |
|
Pristimantis corrugatus |
Duellman, Lehr, and Venegas, 2006 |
|
Pristimantis cosnipatae |
Duellman, 1978 |
|
Pristimantis cremnobates |
Lynch and Duellman, 1980 |
|
Pristimantis crenunguis |
Lynch, 1976 |
|
Pristimantis crepitans |
Bokermann, 1965 |
|
Pristimantis cristinae |
Lynch and Ruíz-Carranza, 1985 |
|
Pristimantis croceoinguinis |
Lynch, 1968 |
|
Pristimantis crucifer |
Boulenger, 1899 |
|
Pristimantis cruciocularis |
Lehr, Lundberg, Aguilar, and von May, 2006 |
|
Pristimantis cruentus |
Peters, 1873 |
|
Pristimantis cryophilius |
Lynch, 1979 |
|
Pristimantis cryptomelas |
Lynch, 1979 |
|
Pristimantis cuentasi |
Lynch, 2003 |
|
Pristimantis cuneirostris |
Duellman and Pramuk, 1999 |
|
Pristimantis curtipes |
Boulenger, 1882 |
|
Pristimantis danae |
Duellman, 1978 |
|
Pristimantis degener |
Lynch and Duellman, 1997 |
|
Pristimantis deinops |
Lynch, 1996 |
|
Pristimantis delicatus |
Ruthven, 1917 |
|
Pristimantis delius |
Duellman and Mendelson, 1995 |
|
Pristimantis devillei |
Boulenger, 1880 |
|
Pristimantis diadematus |
Jiménez de la Espada, 1875 |
|
Pristimantis diaphonus |
Lynch, 1986 |
|
Pristimantis diastema |
Cope, 1875 |
|
Pristimantis diogenes |
Lynch and Ruíz-Carranza, 1996 |
|
Pristimantis dissimulatus |
Lynch and Duellman, 1997 |
|
Pristimantis dorsopictus |
Rivero and Serna, 1988 |
|
Pristimantis douglasi |
Lynch, 1996 |
|
Pristimantis duellmani |
Lynch, 1980 |
|
Pristimantis duende |
Lynch, 2001 |
|
Pristimantis dundeei |
Heyer and Muñoz, 1999 |
|
Pristimantis elegans |
Peters, 1863 |
|
Pristimantis epacrus |
Lynch and Suárez-Mayorga, 2000 |
|
Pristimantis eremitus |
Lynch, 1980 |
|
Pristimantis eriphus |
Lynch and Duellman, 1980 |
|
Pristimantis ernesti |
Flores, 1987 |
|
Pristimantis erythropleura |
Boulenger, 1896 |
|
Pristimantis esmeraldas |
Guayasamin, 2004 |
|
Pristimantis eugeniae |
Lynch and Duellman, 1997 |
|
Pristimantis euphronides |
Schwartz, 1967 |
|
Pristimantis eurydactylus |
Hedges and Schlüter, 1992 |
|
Pristimantis exoristus |
Duellman and Pramuk, 1999 |
|
Pristimantis factiosus |
Lynch and Rueda-Almonacid, 1998 |
|
Pristimantis fallax |
Lynch and Rueda-Almonacid, 1999 |
|
Pristimantis fenestratus |
Steindachner, 1864 |
|
Pristimantis festae |
Peracca, 1904 |
|
Pristimantis fetosus |
Lynch and Rueda-Almonacid, 1998 |
|
Pristimantis flavobracatus |
Lehr, Lundberg, Aguilar, and von May, 2006 |
|
Pristimantis floridus |
Lynch and Duellman, 1997 |
|
Pristimantis frater |
Werner, 1899 |
|
Pristimantis fraudator |
Lynch and McDiarmid, 1987 |
|
Pristimantis gaigeae |
Dunn, 1931 |
|
Pristimantis galdi |
Jiménez de la Espada, 1871 |
|
Pristimantis ganonotus |
Duellman and Lynch, 1988 |
|
Pristimantis gentryi |
Lynch and Duellman, 1997 |
|
Pristimantis ginesi |
Rivero, 1964 |
|
Pristimantis gladiator |
Lynch, 1976 |
|
Pristimantis glandulosus |
Boulenger, 1880 |
|
Pristimantis gracilis |
Lynch, 1986 |
|
Pristimantis grandiceps |
Lynch, 1984 |
|
Pristimantis gularis |
Boulenger, 1898 |
|
Pristimantis gutturalis |
Hoogmoed, Lynch and Lescure, 1977 |
|
Pristimantis hamiotae |
Flores, 1994 |
|
Pristimantis hectus |
Lynch and Burrowes, 1990 |
|
Pristimantis helvolus |
Lynch and Rueda-Almonacid, 1998 |
|
Pristimantis hernandezi |
Lynch and Ruíz-Carranza, 1983 |
|
Pristimantis huicundo |
Guayasamin, Almeida-Reinoso, and Nogales-Sornosa, 2004 |
|
Pristimantis hybotragus |
Lynch, 1992 |
|
Pristimantis hylaeformis |
Cope, 1875 |
|
Pristimantis ignicolor |
Lynch and Duellman, 1980 |
|
Pristimantis illotus |
Lynch and Duellman, 1997 |
|
Pristimantis imitatrix |
Duellman, 1978 |
|
Pristimantis incanus |
Lynch and Duellman, 1980 |
|
Pristimantis incertus |
Lutz, 1927 |
|
Pristimantis incomptus |
Lynch and Duellman, 1980 |
|
Pristimantis infraguttatus |
Duellman and Pramuk, 1999 |
|
Pristimantis inguinalis |
Parker, 1940 |
|
Pristimantis insignitus |
Ruthven, 1917 |
|
Pristimantis inusitatus |
Lynch and Duellman, 1980 |
|
Pristimantis ixalus |
Lynch, 2003 |
|
Pristimantis jaimei |
Lynch, 1992 |
|
Pristimantis johannesdei |
Rivero and Serna, 1988 |
|
Pristimantis jorgevelosai |
Lynch, 1994 |
|
Pristimantis juanchoi |
Lynch, 1996 |
|
Pristimantis jubatus |
Garcia and Lynch, 2006 |
|
Pristimantis kareliae |
La Marca, 2005 |
|
Pristimantis katoptroides |