More on the Atelopus (Amphibia, Salientia) from Western South America

MORE ON THE ATELOPUS (AMPHIBIA, SALIENTIA) FROM WESTERN SOUTH AMERICA

JUAN A. RIVERO Department of Biology University of Puerto Rico Mayaguez, Puerto Rico

ABSTRACT: 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.

FTER the publication of my paper on a “ch”, ends with a “tas” and has a “y” A the Atelopus from Colombia and in between, there is a good possibility that Ecuador g had the opportunity to examine Chyavetas may only be a misspelling of an additional number of specimens from Chayahuitas. these countries and from Peru. From these Both Pebas and Chayahuitas are within studies, it appears now that some of the the Amazon drainage system and both species mentioned can be synonymized, occur within the uniform, humid, tropical while others, whose affinities could not forest of the Amazonian Basin. The desc- be ascertained at the time, can now be ription of Atelopus puleher fits that of related to other forms occuring in those A. spumarius, except for the fact that countries. Cope speaks of “a broad band from orbit Atelopus pulcher Blgr., 1882, for exam- to groin composed of numerous aggregated ple, appears to be a strict synonym of annuli of greenish yellow which has the A. spumarius Cope, 1871. A. pulcher ca- appearance of green foam”, and neither me from Chyavetas, E. Peru, and A. spu- A. pulcher Blgr. nor any of the other marius from Pebas, about 400 miles to Atelopus from east of the Andes known the northeast. Chyavetas, to be ceriain, to the author, meet these specifications. was never found in a Peruvian map and no In A. pulcher Blgr. there is a solid brown Peruvian native could give reference to lateral band, and above this, a yellow this locality. However, a place by the name (green?) band usually containing brown of Chayahuitas was found near the margin dots, but no annuli. In fact, greenish or Rio Puma (5° 5’ S, 76° 10’ W), and about annuli with brown centers can only he 40 miles north of Balsapuerto. Considering found in some A. cruciger, and the appea- that the name Chayahuitas begins with rance of green foam can perhaps be most properly applied to the lateral pattern of Carib. J. Sci. 8 (1-2). March-June 1968. that species. This may be one of the rea- 19 20 JUAN A. RIVERO

sons why A. spumarius was long synony- certainty of the data is not possible to mized with A. cruciger. The possibility determine if there is any variation among that Cope may have confused the label and the rest of the Ecuadorean specimens assign to Peru material actually coming deposited at the A.M.N.H. (Banos-Cane- from Venezuela was considered, but aban- los, Cutuctu, Rio Upano, Chancha, Riobam- doned upon finding that apart from the ba). All of these animals show some “green foam”, the description does not variation from the Peruvian material,* fit A. cruciger while it does, in many de- but this may be due to the state of pre- tails, characterize A. pulcher. servation of the latter, as specimens from Material coming from as near as pos- Rio Oglan, Puyo (Pastaza), Veracruz, sible to the type locality of A. spumarius Miazal (Morona-Santiago) and Limon (Za- are AMNH 43405, 42334-36, from Tac- mora) U.S.N.M. are similar to the Peru- sha, Huachiyacu, Rio Morona, Loreto, Pe- vian from. Yet, Miazal is at the eastern ru. In all details, these specimens follow base of Cutucu, and Canelos is only about Cope’s description, but the lateral band 20 miles southeast of Puyo. Material from is green (above a brown one) and the dots Macas at the USNM is similar in snout within the band are of a solid brown co- shape to the Upano-Cutucu-Banos-Cane- lor. This seems to be the most common os (and Riobamba) form, but looks diffe- Atetopus in the departments of Loreto, rent from the Limon (Gral. Plaza) speci- Amazonas and San Martin of Peru. Spe- mens, which are from approximately the cimens have been examined from Rio same elevation along the same river sys- Bombo (E. Loreto), Utiquima and Roaba- tem. Thus, no conclusions regarding dif- ya (S.E. Loreto, at Rio Ucayali), Shepahua ferentiation of this species in Ecuador (S. Loreto, Rio Urubamba), Upper Riabo, can be offered, although one should be Cachiyacu and Rio Huallaga Valley (S. on the lookout for any possible change San Martin), Cachipuerto and Balsapuer- along the area of Macas and Rio Upano. to (N. San Martin), Manseriche, mouth of Apart from some slight variation in the Rio Santiago, and Rio Tambo (N.W. Lo- amount of webbing, in the pointedness of reto) and Rios Cenipe and Ayambis (N. the snout, in the abundance and size of Amazonas). The southernmost specimens the lateral spots and in the presence or known to the author come from Shepahua, absence of ventral spots, the population at the Cuzco Valley, and the northermost of the Amazonian region of Peru seems to be quite uniform. In the mountains from Rio Oglan, in Ecuador. Other Ecua- of San Martin, however, the species va- dorean specimens available for examina- ries somewhat, and for these, a new ra- tion are from Miazal, at the eastern base of the Cutucti Cordillera, Veracruz, Puyo, cial name is proposed. Some variation Banos-Canelos, Cutucti, Rio Upano, Chan- is already noticed in animals from Rio cha (Chimborazo) and Riobamba. Dr. Ja- Pisqui, but the greatest differentiation is mes Peters tells me that these last two shown by material from Biabo Valley and localities are undoubtedly incorrect, and from Tocache, in upper Rio Cachiyacu. that is probably what they are, as it is These valleys are enclosed by fairly high most improbable for this East Andean ridges, and the isolating effect of the species to occur west of the Andes (Chim- borazo) or as high as Riobamba. Yet, * In Peruvian specimens the sides of the snout the few small specimens labeled as coming are externally concave, opening up at the nostrils, while in some Ecuadorean material at the A. M. N. from Riobamba have a differently shaped H. (Banos-Canelos, Cututu, Rio Upano, Chancha, snout in which the eye diameter is slightly Riobamba), the sides of the snout are straight, for- longer than the distance between eve and ming a triangle with a more or less truncate tip. nostril. It seems that, no matter where When looked from below, the snout overhangs the mouth more in the Peruvian specimens. The nos- they are coming from, they have under- trils are, in these specimens, in elevated protuberan- gone some differentiation from the East ces, which explains why the snout is concave on the Andean population. Because of the un- sides. MORE ON THE ATELOPUS FROM SAUTH AMERICA 21

surrounding mountains, plus the effect of occurs in Sarayacu (AMNH 33891,3893) higher elevation, have apparently induced and may also have a transandean repre- differentiation of the mountain form. sentative at Balzabamba, in Bolivar, Ecua- Atelopus flavescens, from British Guia- dor AMNH 18229). This form has the na, belongs to the same group as A. spu- long, pointed snout, a post-orbital ridge marius, but it is separated from that spe- which forms an angle with the slightly cies by many miles of forest from where protruding rear margin of the frontopa- neither of them nor their representatives, rietals and a porous, rather indistinct, dor- have been collected. The two forms are solateral ridge. The color is as described very similar, however, and severance of a by Andersson, except for the fact that the probably once continuous population may light dorsolateral band is narrow and there have occured at not too distant a future. are no reddish spots inside or ramifi- cations of the light color into the brick In my paper on the Atelopus from Co- red upper areas. lombia and Ecuador (Carib. J. Sci., 1963, 3 (2-3) : 110) the possibility that Atelopus The two specimens from Sarayacu are palmatus Andersson be a synonym of A. granular on the pectoral region but the spumarius was considered. To be certain, one from Balzabamba is smooth. This there are a few characters in the descrip- last specimen is also slightly less webbed tion of A. palmatus that do not fit A. spu- than the eastern form, the tip of the snout marius. For one thing, the snout of A. is not triangular, the post-orbital ridge spumarius is not “as long as, or little is not distinct and the porous dorsolate- longer than, the diameter of the eye”, ral ridge is not apparent. In spite of but definitely longer, and there are no these differences, the two animals appear swollen protruding edges of the fronto- to be related and may, perhaps, be cons- parietals or glandular ridges behind the pecific. There is a good possibility that orbit. The rest of the description fits so the specimens from Sarayacu may repre- well, that one is inclined to disregard sent Atelopus palmatus Andersson. some of these characters as resulting from There is in Macas, E. Ecuador a small dessication or from a different way of tubercular species of Atelopus (AMNH taking measurements. 16717-16724) which cannot be assigned to I am still confused with two of the cha- either A. festae Peracca or A. carinatus racters mentioned by Andersson. He had Andersson, both of which occur in that an animal with an acuminate, projecting general region. It does not have a trun- snout and a long head (“considerably lon- cate snout and although the eye diameter ger than broad”). Yet, the eye diameter is larger than the distance between the was as long as, or almost as long as the eye and nostril, it cannot be said to be as snout. This does not seem right unless Andersson’s measurement of the snout long as the snout (as in A. carinatzs). was taken between eye and nostril. The The animal is of a yellowish brown color, other doubtful character is the extension sometimes with darker markings, and in of the adpressed hind heel, which, accord- this respect it differs from both A. cari- ing to Andersson, reaches in A. palmatus natus (solid brown above) and A. festae to “a little before de eye”. He doesn’t (brownish black). From A. festae it also say whether it is before it reaches the eye differs in size, in the wartiness of its skin or just beyond, that is, ahead of the eve. and in having the eye diameter definitely These characters discarded, or rather in- larger than the distance between eye and terpreted as if Andersson’s “snout” is nostril. from eye to nostril, and “before the eye” means anterior to the eye. there is another This form is perhaps most closely rela- form from eastern Ecuador that fits the ted to A. bufoniformis Peracca, from Pun description quite well. This is a frog that (near the Colombia-Ecuadorean frontier) 22 JUAN A. RIVERO and to A. nicefori Rivero,* from Caicedo, could mean any point in the Andean Cor- Colombia. With both species it shares dillera within the Department of Antio- the small size (25-30 mm), the tubercu- quia) and the distance, both vertical and lar skin and the proportion between eye horizontal, from the type locality, I would and distance between eye and nostril. It not hesitate to consider this animal as differs from A. bufoniformis in color strictly nonspecific with A. longirostris. (A. bufoniformis is brownish black above; In Mera, in the Napo Plastaza region brownish black with yellowish spots be- (now separated into the departments of low) and from A. nicefori in lacking a Napo and Pastaza) there is a form that tarsal fold. An occasional specimen of appears to belong to the same general the Macas form shows a color pattern that group as the above. Dr. James Peters is quite similar to that of A. nicefori but has examined this lot (AMNH 49943-46, unless this species represents a form of 449948-50) and believes that the locality very wide distribution, it is unlikely that record is incorrect and that these speci- an animal from Caicedo, 1800 m., in the mens agree with animals he has collected western face of the Andes of Colombia, in western Ecuador, which again, are may be the same as one occuring in Pun strictly conspecific with Cope’s Atelopus (2600-2800 m.) or in Macas (around 1000 longirostris. Yet, Cope did not call atten- meters in the eastern face of the Oriental tion to a mid-dorsal fold or to lateral spi- Andes). As the author did not have the nulosity, characters which are, however, opportunity to compare material from these present in the Mera specimens. These three localities or with a specimen from animals are fully webbed in the toes (Co- Aguadita (CNHM 81875-Sibate-Aguadita) pe’s were half webbed), there is a distinct tentatively, but probably incorrectly, in- spotting of the dorsum and the snout is cluded with A. nicefori (Rivero op. cit. pointed and protuberant although not qui- 115), no further comments are offered te as much as in the specimen from An- regarding this form. It appears, howe- tioquia. ver, that A. bufoniformis. A. nicefori, the Another form occuring in Cisneros, Co- form from Aguadita, and the form here lombia (CNHM 43850-51) and discussed mentioned belong to a group of related by the author in a separate paper (Rivero, frogs, the affinites of which can only be 1963: 112) should also be studied in con- ascertained by examining more material nection to this group of frogs. In all the from a good number of localities. forms included in the group there seems A specimen from Antioquia, Andes, Co- to be a mid dorsal fold, a long, pointed lombia. at the American Museum of Na- snout in which the nostrils are on distinct tural History (14149) is the closest thing protuberances, and a dark brown dorsal I have found to Atelopus longirostris coloration with yellow (green ?) spots on Cope (Vane de Quito), The animal has a the scapular region. These characters very long and pointed snout, the nostrils seem to indicate relationship among these are on two distinct lateral protuberances, various frogs. the fingers are slightly webbed, the toes The Ecuadorean collection of the Ame- a little more than half webbed and there rican Museum includes two groups of is a yellow (green?) marking on the sca- frogs coming from Rio Pescado in Gua- pular region and three or four others in yas. One of them is apparently made up the sacral region. If it were nor for the of specimens of A. spurrelli. The other, ambiguity of the locality (It is not known comprising 15 specimens, includes a frog if it refers to the town or to the depart- that in color pattern reminds one of ment of Antioquia and, of course, Andes, A. longibrachius Rivero (El Tambo, Gui- sito, Colombia). However, in this species * Described as plain brown on the basis of dry the dorsal markings are round and solid specimens but with markings when wet, as shown in photograph of the species (Rivero, 1963, Carib. J. while in A. longibrachius they are usually Sci., 3 (2-3): 248. sinuous, elongated or annular. There may MORE ON THE ATELOPUS FROM SAUTH AMERICA 23 be other differences, but since the type lity that the animals from Oyacachi re- material of A. longibrachius, is not present the same species is very doubtful. available for comparison at this time, There may be however, a place by the these specimens are, for the time being, name of Buenaventura in the east. Under considered as nonspecific with that spe- the circumstances, identification, of the cies. This spotted form is also similar to above mentioned material should be con- A. spurrelli (which may also show an sidered provisional. occasional spotted individual), but differs Still unaccounted for are Atelopus se- from it in snout shape (longer and more miniferus Cope, 1874, from, Balsapuerto pointed in A. spurrelli), in color pattern and Moyabamba, Peru, well within the and in having a shorter hind limb. territory of A. spumarius but differing A. bicolor Noble, from Cordillera Cutu- from it in having a solid brown dorsum, cu, east of Macas, 1800-2000 m. is cer- a brownish orange venter and a shorter tainly synonym of A. boulengeri Peracca, snout, and A. boussingaulti, (Thominot), from Gualaquiza and San Jose, E. Ecua- 1889, from Guayaquil-Latacunga, S. of dor. Four specimens from Turula, E. of Quito (?) which Dunn (personal commu- Macas, 2800 ft. (AMNH 13133-35, 33893) nication) and James Peters, who has and four from Ecuador (AMNH 17427, examinated the type, consider a synonym 17428, 17431) easily show the characters of A. longirostris Cope. of that species, which however, seems to The description of the subspecies pro- be somewhat variab1e insofar as the posed follows here: amount of webbing is concerned. In some of the Ecuador specimens, the snout, in Atelopus spumarius andinus sp. n. front of the eyes, is yellow, and in no. Type: AMNH 4300, a from Upper 17430, there is a yellow, sickle-shaped Biabo Valley, Peru. marking on the right side of the posterior portion of the back. Diagnosis: Atelopus spumarius andinus In my recent paper on Atelopus ignes- differs from the typical form in having a cens (Carib J. Sci., 5 (3-4) : 137-140), I granular skin and a different color mentioned A. pachydermus (Schmidt) as pattern. coming from Buenaventura, on the west- Description: Morphologically, the sub- ern coast of Colombia. The type locality species is very similar to the type species is actually “near Buenaventura, 5000 ft.” but the skin is densely spinulous, espe- and it was I who, on the basis that I could cially on the eyelids, the sides of the not find any other Buenaventura In Co- dorsum and the posterior half. The spi- lombia, concluded, perhaps prematurely, nules are not as common on the limbs. that A. pachyderms should have come The dorsolateral band and dorsal spots from the mountains north of the city. are tan rather than greenish yellow or Examination of a number of specimens green and the dorsal pattern in general is from Oyacachi, Napo-Pastaza, Ecuador different from that occuring in A. spu- (AMMH 20492-20503) which I have ten- marius (See Pl. II). tatively indentified as A. pachydermus, makes one doubt if Schmidt’s Buenaven- Measurements: (mm.) Snout-vent 28; tura was the western locality I originally head breath 8; head lenght 8.1 (from thought. However, no town by the name angle of the mouth) ; femur 13.2; tibia of Buenaventura has been found east of 12.3; foot 10.1. the Andes of Colombia, and no Colombian Paratypes: Belonging to this same form consulted believes there is such a town, are AMNH 42657, a from the same although some agree that there could be locality, and AMNH 42194, 43296, 43297, a small village with that name. 42657 from Tocache, Upper Rio Chachi- If Schmidt’s Buenaventura is the sea yacu, Sn. Martin, Peru and AMNH 43545 level, western town at 4°15’N, the probabi- from Lower Rio Pisqui, Loreto, Peru. 24 JUAN A. RIVERO

AMNH 42914 is a with the following herpetology of Tropical America. Proc. Amer. measurements: (mm.) snout-vent 34.9; Philos. Sot., 11: 553-559. head breadth 15; head length; femur 16; — 1874. On some Batrachia and Neognathi brought from the Upper Amazon by Prof. Orton. Proc. tibia 17.2; foot 14.7. Acad. Nat. Sci. Phila: 120-137. All the specimens are spinulous above NoBLE, G. K., 1921. Five new species of Salientia and in all, the light dorsal color is tan from South America. Amer. Mus. Novit, 29:1-7, rather than yellow or green. In all of 3 figs. them the dorsal pattern is different from PERACCA, MARIO G IACINTO , 1904. Viaggio del Dr. Enrico Festae Nell’Ecuador e regione vicine: the dorsal pattern of the typical form (see Rettilied Amfibii. Boll. Mus. Zool. Anat. Torino, Pl. 1). In all except the type, there is 19 (465): 1-41. considerable overlapping of the knees PETERS JAMES, 1955. Herpetological type localities when thighs and tibiae are placed perpen- in Ecuador. Rev. Ecuador Ent. Par., 2 (3-4): dicular to the body. 335-352. — 1960. The Snakes of Ecuador. Bull. Mus. Comp. Zool. Harvard, 112 (9):491-541. REFERENCES RIVERO, JUAN A., 1963. Five new species of Atelo- pus from Colombia, with notes on other forms from Colombia and Ecuador. Carib. J. Sci. 3 ANDERSSON, LARS GABRIEL, 1945. Batrachians from (2-3): 103-124, 3 pls. East Ecuador collected 1937, 1938 by Wm. Clar- — 1965. Notes on the Andean Salientian (Amphi- ke-Macyntire and Rolf Blomberg. Arkiv., for bia) Atelopus ignescens (Cornalia). Carib. J. Sci. Zoology, 37 (2): 1-88, 26 figs. 5 (34): 137-139, 1 pl. B OULENGER, G. A., 1882. Catalogue of the Batra- THOMINOT , ALEXANDRE , 1889. Observations sur chia Salientia in the collection of the British quelques reptiles et batraciens de la collection du Museum, ed. 2, London, i-xvi+l-503, 30 pls. Museum d’Histoire Naturelle de Paris. Bull Sot. COPE, EDWARD D., 1871. Eigth contribution to the Philom. Paris, 1 (8): 21-30.

FIGS . 1-8. Atelopus spumarius.—F IG . 1. U.S.N.M. 65488, from Macas, Morona-Santiago, Ecuador.— FIG. 2. A.M.N.H, 33915 from Cutucu, Santiago-Zamora, Ecuador.—FIG. 3-4 J.A.P. (James A. Peters) 7519, 7517, from Limon (Gral. Plaza), Morona-Santiago, Ecuador.—FIG. 5. A.M.N.H. 42334, from Tacsha, Loreto, Peru.—FIG. 6. A.M.N.H. 42527 from Roaboya, Loreto, Peru—FIG. 7a. J.A.P. 7514 from Limon.—7b. A.M.N.H. 21492 from Macas, 1600m.—7c A.M.N.H. 33915 from Cutucu.—7d A.M. N.H. 65487 from Macas.—FIG. 7. A.M.N.H. 10532 from Riobamba, Chomborazo, Ecuador. Note the difference in sonout shape between specimens from Macas-Cutucu (Figs. 1.2, 7b, c, d.) and those from elsewhere. Specimen A.M.N.H. 10532.—FIG . 8. Perhaps erroneously labeled as coming from Rio- bamba, has a shorter snout than other examples from Ecuador and Peru.

26 JUAN A. RIVERO

FIG. 1. A.M.N.H. 43200 Atelopus spumarius andinus, type, from Upper Biabo Valley, Loreto, Peru.— FIG. 2. A.M.N.H. 42657, A. spumarius andinus ptp. from Biabo Valley, Loreto, Peru.—FIG. 3. A.M. N.H. 432966, A. spumarius andinus prtp. from Tocache, Upper Rio Cachiyacu, Sn. Martin, Peru.— FIG. 4. A.M.N.H. 42914, A. spumarius andinus from Tocache.—FIG. 5. A.M.N.H. 20496, ¿Atelopus pachydermus from Oyacachi, Pueblo Viejo, Napo-Pastaza, Peru.—FIG. 6. A.M.N.H. 20493, ¿Atelopus pachidermus, from Oyacachi.—FIG. 7. A.M.N.H. 13133, Atelopus boulengeri from E. of. Macas, San- tiago-Zamora, Ecuador.

28 JUAN A. RIVERO

FIG. 1. A.M.N.H. 22198 Atelopus planispinus from top of Volcan Sumaco, Napo-Pastaza, Ecuador.— FIG. 2. A.M.N.H. 49950, ¿Atelopus longirostris from Mera, Napo-Pastaza, Ecuador.—FIG. 3. A.M.N.H. 49946, ¿Atelopus longirostris from Mera.—FIG. 4. A.M.N.H. 33891, ¿Atelopus palmatus from Sarayacu, Napo-Pastaza, Ecoador.—FIG. 5. A.M.N.H. 17738, Atelopus sp. (nr. longibrachius) from Guayas, Rio Pescado, Ecuador.—FIG. 6. A.M.N.H. 16717, Atelopus sp (nr. bufoniformis) from Cordillera Cutucu, E. of Macas Santiago-Zamora, Ecuador.