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Contribution à l’étude des Amphibiens de Guyane française. VIII. Validation d’Atelopus spumarius Cope, 1871, et désignation d’un néotype. Description d ’Atelopus spumarius barbotini nov. ssp. Données étho-écologiques et biogéographiques sur les Atelopus du groupe flavescens (Anoures, Bufonidés)
... We observed more than a single color at a locality, with individuals exhibiting high variation in color and pattern, especially in Amapá. Lescure (1981) defined three morphs forA. hoogmoedi in the Guianas, with varying colors of dorsal pattern, but mostly based on the variation of the size of the sinuous dorsolateral longitudinal bands. ...
... We found the same variation in different populations of A. hoogmoedi in Brazil. Populations of Amapá exhibit thin dorsolateral bands (Figure 2a-f) similar to the morph "A" of Lescure (1981) from Attachi-Bacca, French Guiana. In Tapajós river, south Amazonas river, individuals exhibited color bands very variable in width with the predominance of color extending from the ventral surface the snout being more colored than black (Figure 2i), similar to the morph "B" of Lescure (1981) from Surinam. ...
... Populations of Amapá exhibit thin dorsolateral bands (Figure 2a-f) similar to the morph "A" of Lescure (1981) from Attachi-Bacca, French Guiana. In Tapajós river, south Amazonas river, individuals exhibited color bands very variable in width with the predominance of color extending from the ventral surface the snout being more colored than black (Figure 2i), similar to the morph "B" of Lescure (1981) from Surinam. Specimens from Oriximiná, Pará has bands on the back that are wider than the black bands (Figure 2h and 2J) corresponding to the morph "C" of Lescure (1981). ...
Aposematic coloration plays an important role for species, as it serves as a clear signal of danger to visually-oriented predators regarding the potential toxicity of individuals displaying this condition. However, considerable gaps remain on this subject, and, primarily, empirical data are lacking to support robust discussions on the topic. The harlequin frog Atelopus hoogmoedi exhibits a range of aposematic colors across different populations, yet we still do not know how this characteristic was selected throughout evolution nor what the impacts of this characteristic are on the species’ biology, ecology, and behavior. Considering that this topic deserves further in-depth studies, particularly due to being one of the most threatened groups of vertebrates and still lacking research, we present possible insights to guide future investigations on this subject.
... nov. can be differentiated from A. hoogmoedi SS of French Guiana by a maximum SVL of 28.8 mm for females (minimum SVL 31.2 mm for females; [9]), tibia 47% of SVL (tibia varying from 43% to 45% of SVL among populations; [9]); dorsum light brown to reddish brown with light yellow or light green reticulation network (dorsum dark brown to black with yellow, orange, or pink reticulation network; [52,53]); and advertisement call duration of 689-840 ms (1190-1200 ms; [7]) consisting of 15-26 pulses (40-42 pulses; [7]). ...
... nov. can be differentiated from A. hoogmoedi SS of French Guiana by a maximum SVL of 28.8 mm for females (minimum SVL 31.2 mm for females; [9]), tibia 47% of SVL (tibia varying from 43% to 45% of SVL among populations; [9]); dorsum light brown to reddish brown with light yellow or light green reticulation network (dorsum dark brown to black with yellow, orange, or pink reticulation network; [52,53]); and advertisement call duration of 689-840 ms (1190-1200 ms; [7]) consisting of 15-26 pulses (40-42 pulses; [7]). ...
... The new species is distinguished from Atelopus franciscus by the larger size of females, a minimum SVL of 27.9 mm (maximum SVL of 26.5 mm; [9]); venter all white or white with a cream-colored gular region and head (venter and thighs red; [9]); and advertisement call duration of 689-840 ms (1340-1640 ms; [7]) consisting of 15-26 pulses (31-39 pulses; [7]). ...
We used integrative taxonomy to describe a new species of Atelopus from the lowlands of Central Amazonia in the region of Manaus, Amazonas, Brazil. The new species is geographically isolated from the southernmost species of Atelopus of the Guiana Shield. Atelopus manauensis species nova (sp. nov.) is characterized by the combination of the following characteristics: male snout-vent length range (SVL = 19.1-26.4 mm; n = 11); dorsal and lateral skin smooth; ventral surface entirely white or white with cream-colored gular region; fingers and toes lacking subarticular tubercles and fringes. The advertisement call of the new species has a call duration of 689-840 ms, contains 15-26 pulses, is emitted at an average pulse rate of 25.5 pulses per second, and has a dominant frequency ranging 3088-3610 Hz. The genetic divergence between the new species and its morphologically most similar congeners (A. spumarius and A. pulcher) is greater than 4%. Atelopus manauensis sp. nov. is closely related to species of the A. hoogmoedi complex inhabiting the Guiana Shield. The new species has a small geographic distribution (approximately 4500 km 2) in a landscape that is strongly threatened by the growth of Manaus, the largest city in Brazilian Amazonia. The new species is considered critically endangered and in need of urgent conservation measures.
... Detailed information on vocalizations oï Atelopus has been reported for only four species (Jaslow, 1979;Lescure, 1981;Asquith and Altig, 1987), and laryngeal morphology has been examined in three species (Martin, 1972). From our fieldwork, recordings made by colleagues, and a review of the tape archives of the National Museum of Natural History (USNM), the American Museum of Natural History (AMNH), and the Museum of Natural History, University of Kansas (KU), we have analyzed recordings of the calls of eight species of Atelopus, including seven species for which calls have not been reported previously. ...
... Few field recordings of Atelopus are available, and therefore some of the recordings utilized in this and other studies (Asquith and Altig, 1987;Jaslow, 1979;Lescure, 1981) were made from animals in captivity. We tested the potential for distortion of calls from animals calling from plastic bags by analyzing recordings of a single male of Hyla koechlini Duellman and Trueb calling in and out of a bag (this species has a pulsed call somewhat similar in structure to calls of Atelopus); our analyses showed no discernible difference between the two recordings. ...
... comm. Rivero, 1963;Ruiz-Carranza etal., 1988Peters, 1973Lescure, 1973, 1981Savage, 1972Savage, 1972Dunn, 1933This study Myers, pers. comm. ...
Vocalizations of frogs of the genus Atelopus include three discrete types of signals: pulsed calls, pure tone calls, and short calls. Repertoire composition is conservative across species. Repertoires of most species whose calls have been recorded contain two or three of these identifiable call types. Within a call type, details of call structure are very similar across species. This apparent lack of divergence in calls may be related to the rarity of sympatry among species of Atelopus and to the relative importance of visual communication in their social interactions.
... n.). All Guianan harlequin toads have shorter call durations (2049 ± 220 ms vs. 1490 ± 140 ms in A. barbotini; 1530 ± 220 ms in A. flavescens; 1490 ±150 ms in A. franciscus; 1190 ± 10 ms in the A. hoogmoedi complex from French Guiana; 1160 ± 390 ms in the A. hoogmoedi complex from Amapá, Brazil; 1060-1240 ms in the A. hoogmoedi complex from Pará, Brazil; Lescure 1981, Cocroft et al. 1990, Costa-Campos & Carvalho 2018. Tadpoles of the new species differ from those of A. flavescens, A. franciscus and the A. hoogmoedi complex in having the lower beak about twice as long as the upper (vs. ...
... n. is morphologically similar to its congeners of the flavescens-spumarius clade sensu Lötters et al. (in press), as well as A. andinusRivero, 1968, A. loettersi De la Riva, Castroviejo-Fisher, Chaparro, Boistel & Padial, 2011, and the A. tricolor Boulenger, 1902 complex (including the junior synonyms A. rugulosusNoble, 1921 and A. willimani Donoso-Barros, 1969) from the eastern Andean versant of Peru and Bolivia, and to A. palmatus Anderson, 1945 and A. planispina Jiménez de la Espada, 1875 from the eastern Andean versant of Ecuador. With respect to these superficially similar species, the new species differs as follows: from the nominal A. barbotiniLescure, 1981, A. flavescens Duméril & Bibron, 1841 (including its junior synonym A. vermiculatus McDiarmid, 1973, A. franciscus ...
For nearly four decades, harlequin toads, genus Atelopus, have suffered unparalleled population declines. While this also results in limited understanding of alphataxonomic relationships, these toads face an urgent need for advances in systematics to inform conservation efforts. However, high intraspecific variation and cryptic diversity have hindered a comprehensive understanding of Atelopus diversity. This is particularly exemplified among Amazonian populations related to A. spumarius, where decades of taxonomic work have not been able yet to unravel relationships between the many forms, while the names coined so far have led to taxonomic confusion leaving numerous lineages unnamed. A recent comprehensive phylogenetic study has revealed new insights into the systematics of harlequin toads with an emphasis on Amazonian forms, identifying several unnamed lineages. We here describe one of these evolutionary lineages as a new species, restricted to the Ecuadorian Amazon basin, in an integrative taxonomic approach using molecular, morphological, bioacoustic and larval information. With this, we contribute to a better understanding of Atelopus diversity as the baseline of conservation action.
... Atelopus barbotini in life shows a coloration similar to that of A. hoogmoedi nassaui. Ouboter & Jairam (2012: 46) distinguished the two solely on the basis of pink versus reddish color and a somewhat distinct pattern as described-for A. barbotini-by Lescure (1981a). However, central French Guianan specimens referable to A. barbotini are more variable and can look similar to A. hoogmoedi nassaui (compare Heselhaus & Schmidt 1988: 35 and Noonan & Gaucher 2005: 3023 with Ouboter & Jairam 2012. ...
... Another name needs to be discussed, A. pulcher hoogmoedi from French Guiana. Recent evidence implies that either it is a polytypic taxon or a species complex (most certainly not a subspecies of A. pulcher sensu stricto, an endemic taxon from the upper Amazon basin; Lötters et al. 2002) from the central-eastern Guiana Shield and adjacent portions of the lower Amazon basin, i.e., Guyana, Suriname, French Guiana, Brazil (Lescure 1981a;Lötters et al. 2010Lötters et al. , 2011Jorge et al. 2020;Da Silva et al. 2020). Various recent workers referred to it as A. hoogmoedi (complex) (e.g., Rueda-Almonacid et al. 2005;Costa-Campos & De Carvalho 2018;Mebs et al. 2018). ...
... The pulsed call is the most commonly known vocalization in harlequin toads, being present in at least 17 species (Asquith and Altig 1989;Cocroft et al. 1990;Ibáñez et al. 1995;Jaslow 1979;Lescure 1981;Lötters et al. 1999Lötters et al. , 2002 this study). The pulsed call of A. laetissimus consists of a short series of pulses (7-33 pulses) emitted rapidly, which is remarkably different from the pulsed calls of A. barbotini Lescure, 1981 (41- corded in a male-male aggressive interaction in A. chiriquiensis (Jaslow 1979). ...
... In any case, the predictions of these models should be interpreted with caution and they should be considered as a first approximation to the real distribution of the species, A. flavescens Duméril and Bibron, 1841 (45-58 pulses per call, 29.76-34.78 pulses/s, 2,500-3,000 Hz; Lescure 1981), A. franciscus Lescure, 1974 (31-39 Asquith and Altig 1987;Lescure 1981) or A. reticulatus Lötters, Haas, Schick, and Böhme, 2002 (27-32 pulses per call, 75-76 pulses/s, 3,282 Hz;Lötters et al. 2002) by having a higher number of pulses replicated more quickly at a lower dominant frequency. Likewise, it differs from the pulsed call of A. zeteki Dunn, 1933 (42-52 pulses per call, 115-146 pulses/s, 1,381-1,510 Hz; Cocroft et al. 1990), by having fewer pulses replicated more quickly at a higher dominant frequency. ...
Atelopus laetissimus is a bufonid toad that inhabits the mountainous areas of the Sierra Nevada de Santa Marta (SNSM), Colombia. This species is endemic and threatened, for which information about their ecology and distribution are crucial for the conservation of this toad. Here, we described the abundance, habitat use, vocalization and potential distribution in A. laetissimus from the San Lorenzo creek of the Sierra Nevada de Santa Marta. To this end, 447 individuals of A. laetissimus were analyzed during several sampling performed from 2010 to 2012. Against expectations, population density was significantly higher in the stream than riparian forest. Overall, A. laetissimus used seven different diurnal habitats with a high preference by leaf litter substrates and rocks. Rate of recaptures was decreasing lineally along the survey nights. We recorded two types of vocalization in A. latissimus. A short call characterized by a series of pulses like a buzz and another also short, but lacking pulses. According to our analysis, the areas with higher habitat suitability for A. laetissimus were located principally in the northern and northwestern region of the SNSM, which was in agreement with the findings reported in literature. Moreover, the modeling of data indicated a significant increase in habitat loss from 2013 to 2017. Our information should be considered as a starting point for the conservation of this valuable amphibian in the tropical landscapes of America.
... Boulenger (1882), Lescure (1981), Lindquist & Hetherington (1996) Single subgular MO Pereyra (pers. comm.) , Crump (1988), Savage (2002) Single subgular , Savage (2002) Atelopus zeteki Lindquist & Hetherington (1996), Grant et al. (2006) Single subgular Grant et al. (2006) Hartmann et al. (2005) Single subgular Heyer (1990) A.J. Elias-Costa and J. Faivovich (2019) -Vocal sacs in torrent-dwelling frogs. ...
Cascades and fast-flowing streams impose severe restrictions on acoustic communication, with loud broadband background noise hampering signal detection and recognition. In this context, diverse behavioural features, such as ultrasound production and visual displays, have arisen in the evolutionary history of torrent-dwelling amphibians. The importance of the vocal sac in multimodal communication is being increasingly recognized, and recently a new vocal sac visual display has been discovered: unilateral inflation of paired vocal sacs. In the diurnal stream-breeding Hylodidae from the Atlantic forest, where it was first described, this behaviour is likely to be enabled by a unique anatomical configuration of the vocal sacs. To assess whether other taxa share this exceptional structure, we surveyed torrent-dwelling species with paired vocal sacs across the anuran tree of life and examined the vocal sac anatomy of exemplar species across 18 families. We found striking anatomical convergence among hylodids and species of the distantly related basal ranid genera Staurois, Huia, Meristogenys and Amolops. Ancestral character state reconstruction identified three new synapomorphies for Ranidae. Furthermore, we surveyed the vocal sac configuration of other anuran species that perform visual displays and report observations on what appears to be unilateral inflation of paired vocal sacs, in Staurois guttatus-an extremely rare behaviour in anurans.
... Moreover, the structure of the advertisement call appears poorly adapted to a noisy environment. Contrary to forest stream dwelling Asian frogs [8,9,10], A. franciscus produces multi-pulsed units separated by short silences [11], typical of species living in open environments (ponds and paddy fields, [8]). The emission of advertisement calls in an absorbent and noisy environment imposes limits on the ability to communicate as calls suffer attenuation and degradation. ...
Atelopus franciscus is a diurnal bufonid frog that lives in South-American tropical rain forests. As in many other frogs, males produce calls to defend their territories and attract females. However, this species is a so-called "earless" frog lacking an external tympanum and is thus anatomically deaf. Moreover, A. franciscus has no external vocal sac and lives in a sound constraining environment along river banks where it competes with other calling frogs. Despite these constraints, male A. franciscus reply acoustically to the calls of conspecifics in the field. To resolve this apparent paradox, we studied the vocal apparatus and middle-ear, analysed signal content of the calls, examined sound and signal content propagation in its natural habitat, and performed playback experiments. We show that A. franciscus males can produce only low intensity calls that propagate a short distance (<8 m) as a result of the lack of an external vocal sac. The species-specific coding of the signal is based on the pulse duration, providing a simple coding that is efficient as it allows discrimination from calls of sympatric frogs. Moreover, the signal is redundant and consequently adapted to noisy environments. As such a coding system can be efficient only at short-range, territory holders established themselves at short distances from each other. Finally, we show that the middle-ear of A. franciscus does not present any particular adaptations to compensate for the lack of an external tympanum, suggesting the existence of extra-tympanic pathways for sound propagation.
Here the author makes a personal approach of his life standing interest in the genus Atelopus.
Vocalizations are one of the most important communication modalities in amphibian biology, and advertisement call is the type of vocalization mostly emitted by anurans (frogs and toads). In the present study, we carried out a scientometric analysis of the advertisement call in species of anurans in Colombia to determine the state of knowledge of this science in the country. We recorded the number of call descriptions and its trend throughout more than 60 years; in addition, we identified how many species there are with the call described per taxonomic family, geographic units and threat status. According to our review, between 1958 and June 30 of 2021, at least 296 studies have been published that describe the advertisement call of 307 species (of 785 in the country); only 130 descriptions come from recordings to individuals in Colombian populations. Leptodactylidae and Hylidae are the families with the highest percentage of species whose call has been described (84.6% and 68.4%, respectively). On the contrary, Bufonidae (19.5%) and Craugastoridae (12.9%) exhibit a low percentage of described calls. The Central and Eastern cordilleras were the regions with the best knowledge of advertisement calls, while the least known were the Caribbean – interAndean valleys, Orinoquía, and Sierra Nevada de Santa Marta. In terms of the threat status, the species with the least concern (LC) were the ones that presented the highest number of species with the call described. Despite an increase in the most recent two decades, our findings still reveal notable gaps in knowledge of the advertisement calls in Colombian anurans, which constitutes an incentive to develop future research on this subject. Finally, based on this meta-analysis, we highlight some recommendations that we hope that we hope stimulate new studies in ecoacoustics, using anurans in Colombia as a study model.
Anurans show a wide variety of anti-predator mechanisms, and the species of the Neotropical clade Leiuperinae display several of them. Most species of Edalorhina, Physalaemus and Pleurodema show eyespots, hidden bright colours, macroglands in a inguinal/lumbar position, defensive behaviours and/or chemical defence. We conducted a histological analysis of dorsal and lumbar skin and revised the colour patterns, defensive behaviours and glandular secretions to study the diversity and evolution of anti-predator mechanisms associated with macroglands. We describe 17 characters and optimize these in a phylogenetic hypothesis of Leiuperinae. In the most recent common ancestor of Edalorhina + Engystomops + Physalaemus + Pleurodema, a particular type of serous gland (the main component of macroglands) evolved in the lumbar skin, along with the absence of the Eberth–Katschenko layer. A defensive behaviour observed in leiuperines with macroglands includes four displays (‘crouching down’ behaviour, rear elevation, body inflation and eye protection), all present in the same ancestor. The two elements associated with aposematism (hidden bright colours and eyespots) evolved independently in several species. Our results provide phylogenetic evidence for the startle-first hypothesis, which suggests that behavioural displays arise as sudden movements in camouflaged individuals to avoid predatory attacks, before the origin of bright coloration.
Atelopus is a species-rich group of Neotropical bufonids. Present knowledge on bioacoustics in this genus is relatively poor, as vocalisations have been described in only about one fifth of the ca. 100 species known. All studied members of the genus produce vocalisations although, with a few exceptions, most species lack a middle ear. Nonetheless, hearing has been demonstrated even in earless Atelopus making bioacoustics in these toads an inspiring research field. So far, three structural call types have been identified in the genus. As sympatry is uncommon in Atelopus , calls of the same type often vary little between species. Based on recordings from the 1980s, we describe vocalisations of three Venezuelan species ( A. carbonerensis , A. mucubajiensis , A. tamaense ) from the Cordillera de Mérida, commonly known as the Andes of Venezuela and the Tamá Massif, a Venezuelan spur of the Colombian Cordillera Oriental. Vocalisations correspond, in part, to the previously identified call types in Atelopus . Evaluation of the vocalisations of the three species presented in this study leads us to recognise a fourth structural call type for the genus. With this new addition, the Atelopus acoustic repertoire now includes (1) pulsed calls, (2) pure tone calls, (3) pulsed short calls and (4) pure tone short calls. The call descriptions provided here are valuable contributions to the bioacoustics of these Venezuelan Atelopus species, since all of them have experienced dramatic population declines that limit possibilities of further studies.
The amphibian type specimens held in the National Museum of Natural History in Leiden are listed. A total of 775 type specimens representing 143 taxon names were encountered. The list provides the original name, the original publication date, pagination and illustrations, current name, type locality and notes on the type status.
The disturbance vicariance hypothesis (DV) has been proposed to explain speciation in Amazonia, especially its edge regions, e.g. in eastern Guiana Shield harlequin frogs (Atelopus) which are suggested to have derived from a cool-adapted Andean ancestor. In concordance with DV predictions we studied that (i) these amphibians display a natural distribution gap in central Amazonia; (ii) east of this gap they constitute a monophyletic lineage which is nested within a pre-Andean/western clade; (iii) climate envelopes of Atelopus west and east of the distribution gap show some macroclimatic divergence due to a regional climate envelope shift; (iv) geographic distributions of climate envelopes of westernand eastern Atelopus range into central Amazonia but with limited spatial overlap. We tested if presence and apparent absence data points of Atelopus were homogenously distributed with Ripley’s K function. A molecular phylogeny (mitochondrial 16S rRNA gene) was reconstructed using Maximum Likelihood and Bayesian Inference to study if Guianan Atelopus constitute a clade nested within a larger genus phylogeny. We focused on climate envelope divergence and geographic distribution by computing climatic envelope models with MaxEnt based on macroscale bioclimatic parameters and testing them by using Schoener’s index and modified Hellinger distance. We corroborated existing DV predictions and, for the first time, formulated new DV predictions aiming on species’ climate envelope change. Our results suggest that cool-adapted Andean Atelopus ancestors had dispersed into the Amazon basin and further onto the eastern Guiana Shield where, under warm conditions, they were forced to change climate envelopes.
Neotropical harlequin frogs, Atelopus, are a species-rich bufonid group. Atelopus monophyly has been suggested but intergeneric, interspecific and intraspecific relationships are poorly understood. One reason is that morphological characters of harlequin frogs are often difficult to interpret, making species delimitations difficult. Molecular analyses (DNA barcoding, phylogeny) may be helpful but sampling is hampered as most of the more than 100 Atelopus species have undergone severe population declines and many are possibly extinct. We processed mitochondrial DNA (12S and 16S rRNA) of 28 available ingroup samples from a large portion of the genus’ geographic range (Bayesian Inference, Maximum Likelihood). Our samples constitute a monophyletic unit, which is sister to other bufonid genera studied including the Andean genus Osornophryne. In contrast to previous morphological studies, our results suggest that Osornophryne is neither sister to Atelopus nor nested within it. Within Atelopus, we note two major clades with well supported subclades, one Amazonian–Guianan Clade (Flavescens-spumarius Clade plus Tricolor Clade) and an Andean–Chocó–Central American Clade (Varius Clade plus all other Atelopus). The first mentioned includes all species that possess a middle ear (i.e. stapes) except for A. seminiferus lacking it (like all remaining Atelopus). Previously proposed species groups based on frog-like versus toad-like overall appearance (i.e. Longirostris and Ignescens Groups) or phalangeal reduction in the thumb (i.e. Flavescens Group) are not monophyletic in our phylogeny, thus characters used to define them are not considered synapomorphies. We show that genetic divergence can be high between species belonging to different clades, in spite of their phenetic similarity (e.g. A. pulcher, Atelopus sp. 2). On the other hand, within the same clade, colour can vary tremendously, while genetic divergence is low (e.g. A. flavescens and allies). These observations demonstrate that Atelopus taxonomy is complicated and that an integrative approach is required before ‘splitting’ or ‘lumping’ nominal species.
We investigated the genetic structure of populations of Guianan harlequin toads (genus Atelopus) and their evolutionary affinities to extra-Guianan congeners. Phylogenetic analysis of mitochondrial cytochrome b (cyt b) and NADH dehydrogenase subunit 2 (ND2) gene sequences produced well-supported clades largely corresponding to the four recognized taxa in the Guianas (Atelopus spumarius hoogmoedi, Atelopus spumarius barbotini, Atelopus franciscus, and Atelopus flavescens). Our findings suggest that the Guianan A. spumarius represent distinct evolutionary lineages that merit distinction from Amazonian conspecifics, and that the status of A. flavescens and A. franciscus is somewhat less clear. Approximately 69% of the observed genetic variation is accounted for by differences between these four recognized taxa. Coalescent-based estimates of gene flow between taxa suggest that these lineages are largely isolated from one another. Negligible rates of migration between populations and significant divergence within such close proximity suggests that although the region inhabited by these taxa is almost entirely undisturbed, significant habitat heterogeneity exists as to have produced a remarkable diversification of Atelopus within the eastern Guiana Shield. These results contradict the commonly held view of the Guiana Shield as a 'refuge' whose stability during late Tertiary and Quaternary climatic fluctuations served as a biotic reservoir. Instead, we provide evidence that climatic fluctuations during this time had a diversifying effect within the Guianan region.
Atelopus chiriquiensis were observed during May and June, in 1976 in Chiriqui Province, Panama. Three distinct male vocalizations were recorded and analyzed and their contexts are described. Calling males oriented toward calling neighbors and occasionally wrestled to defend calling sites. Wrestling included an apparent submissive posture and the flipping of opponents.
Atelopus pulcher Blgr., 1882, is synonymized with A. spumarius Cope, 1871, and a new race of this species is described from Biabo Valley, Peru (A. s. andinus). The possible identity of some uncertain species of Atelopus from Peru and Ecuador is discussed.
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