10 p. : ill., map ; 24 cm.Includes bibliographical references (p. 10)

Dunn, Emmett Reid, 1894-1956.

American Museum of Natural History Scientific Publications

AMERIICAN MUSEUM1NOVITATESPUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORYCITY OF NEW YORK MAY 7, 1949 NUMBER 1416NOTES ON THE SOUTH AMERICANFROG GENUS EDALORHINABY EMMETT REID DUNNThe bizarre little frogs of this group were first made known toscience as "E. perezsi, n.g., n. sp." by Don Marcos Jimenez de laEspada in 1871, after his journey from Quito to the Rio Napo inEcuador. This trip, taken in order to trace the route of Orellanain his discovery of the Amazon, revealed a rich frog fauna. Prof.Orton, of Williams College, working in the opposite direction, alsocollected frogs of this group, and they were reported as Buboniasplicifrons, new genus, new species, by Cope in 1874. His speci-mens were taken at Nauta, on the Marafion in Peru. By 1882material from various sources had accumulated in the BritishMuseum, and in Boulenger's "Catalogue of Batrachia Salientia"(in which he synonymized Cope's genus with that of Jimenez de laEspada, but recognized both their species and added one of hisown, E. buckleyi) the range of the group was extraordinarily wellblocked out on a basis of 10 specimens and was in no way alteredby the report of a specimen by Muller in 1884.Thirty years later Boulenger described a fourth species (E.nasuta), on a basis of three specimens from just south of the pre-viously known range of the group in Peru. Thirty more yearselapsed, and Myers announced the receipt of a virtual topotype ofthe first-named form, adding the statement "records of it areextremely rare."As the entire literature contains references to only 15 specimens,and as I have examined 36 additional specimens in the collectionof the American Museum of Natural History, and one in theUnited States National Museum, some report on these would bein order.AMERICAN MUSEUM NOVITATSN-O.11Six Ecuadorian specimens, from three localities, are due to theactivities of Carlos Olalla and son. Thirty Peruvian specimens,from 12 localities, are due to the activity and the generosityof Dr. Harvey Bassler. This additional material only slightlyincreases the range of the genus, which may be stated as follows:north to San Jose Abajo, Ecuador (probably = San Jose de Moti,on a small northern tributary of the Napo, and practically on theEquator); west to Bafios, Ecuador, altitude 1831 meters; east toIquitos, Peru, altitude 106 meters; south to Valle, Peru (latitude80 S., in the Huallaga basin), and to Huancabamba, Peru, alti-tude 1900 meters (latitude 100 S., in the Ucayali basin). It isvirtually certain that the range will be extended to the Putumayodrainage in Colombia, probably to the upper Yavari drainage inBrazil, and possibly to the upper Beni drainage in Bolivia.Two generic names have been based on species of this group:Edalorhina JIMENEZ DE LA ESPADA, 1871, Jour. Sci. Acad. Lisboa, vol. 3, p.58 (monotype E. perezi Jimenez de la Espada, loc. cit.).Bubonias CoPE, 1874, Proc. Acad. Nat. Sci. Philadelphia, vol. 26, p. 124(monotype B. plicifrons Cope, loc. cit.).The following specific names have been based on specimens ofthis group:Edalorhina perezi JIMENEZ DE LA ESPADA, 1871, loc. cit. Banks of the RioNapo, Ecuador; type or cotypes presumably in Spain. 1875, Vert. Viaje Pa-cifico, p. 160, p1. 1, fig. 5. BOULENGER, 1882, Catalogue of the Batrachia Sali-entia ... in the ... British Museum, ed. 2, p. 227, pl. 16, fig. 2. MULLER, 1884,Verh. Nat. Gesellsch. BaSel, vol. 7, p. 281. TREWAVAS, 1933, Phil. Trans. Roy.Soc. London, vol. B222, p. 424, figs. 15-16 (hyoid and larynx). MYERS,1942, Proc. Biol. Soc. Washington, vol. 55, p. 151.Bubonias plicifrons COPE, 1874, loc. cit. Nauta, Peru; two cotypes, not yetlocated in any museum. 1889, Bull. U. S. Natl. Mus., vol. 34, pl. 71, figs. 30-31(skull and shoulder girdle).Edalorhina buckleyi BOULENGER, 1882, op. cit., pl. 16, fig. 1. Canelos, Ecua-dor; type a male in the British Museum.Edalorhina nasuta BOULENGER, 1912, Ann. Mag. Nat. Hist., ser. 8, vol. 10,no. 8, p. 190. Huancabamba, Peru "above 3000 feet"; cotypes three specimensin the British Museum.As will be shown hereinafter, I regard plicifrons and buckleyias conspecific with perezi: it therefore follows that I regardBubonias as congeneric with Edalorhina.Edalorhina perezi Jim'enez de la EspadaThe type of perezi, as described and figured by Jimenez de laEspada, had a dermal ridge between the eyes, lacked longitudinal2 NO. 1416SOUTH AMERICAN EDALORHINAwarts on the dorsum, and had vomerine teeth. The types ofplicifrons, as described by Cope, lacked the dermal ridge between/ lQw7~~ 3YFIG. 1. Edalorhina perezi (typical variety). A.M.N.H. No. 52843. Bafios,Ecuador. Natural size.FIG. 2. Edalorhina perezi (plicifrons-buckleyi variety). A.M.N.H. No.52847. Canelos, Ecuador. A. Dorsal view. B. Lateral view. Natural size.FIG. 3. Edalorhina perezi (intermediate variety). A.M.N.H. No. 42586.Pampa Hermosa, Peru. Natural size.the eyes, had longitudinal warts on the dorsum, and lacked vo-merine teeth. The type of buckleyi, as described and figured byBoulenger, resembled the types of plicifrons, but had vomerine1949 3AM8RICAN MUSEUM NOVITATESteeth. The American Museum collection has specimens thatresemble the types of perezi but that lack vomerine teeth.They thus display a hitherto unreported fourth combination ofcharacters.The interocular dermal ridge and relatively smooth dorsum ofthe perezi variety are illustrated by fig. 1 (a specimen fromBafios); the dorsal dermal characters of the plicifrons-buckleyivariety are shown by figure 2 (a topotype of buckleyi from Canelos).The majority of the collection can easily be allocated to one or theother of these two varieties, but three are somewhat intermediate.One of these, from Pampa Hermosa, is illustrated by figure 3.The "perezi" type of warting is present in 86.6 per cent of theEcuadorian specimens, and in 62.5 per cent of the Peruvian ones(the three Peruvian intermediates have been more or less arbi-trarily assigned to the "plicifrons" type).In Edalorhina the vomerine teeth are never well developed; atmost there are a few teeth on the tips of the backwardly project-ing prevomers. They are frequently present on one of the twopaired bones and absent on the other. Complete absence of vo-merine teeth was noted in 6.6 per cent of Ecuadorian specimensand in 37.5 per cent of Peruvian specimens.Although Cope's Peruvian types of plicifrons were said to havethe toes free, the American Museum specimens from Peru agreewith the forms described from Ecuador in having a rudimentaryweb. This is, however, more obvious in some states of preserva-tion than in others.To some extent specimens from Ecuador differ from specimensfrom Peru in the degree of development of a wart on the tip of thesnout (compare figs. 1 and 2 with fig. 3). In the majority of speci-mens from both countries this structure exists but is poorly de-veloped. It seems to be absent only in some Peruvian specimens,and well developed only in some Ecuadorian specimens. A der-mal appendage on the snout is, however, a conspicuous feature of aform of Edalorhina at the southern end of the generic range inPeru.The accompanying table (table 1) of known specimens is ar-ranged in four columns to show the four combinations of the twosorts of dorsal warting with presence versus complete absence ofvomerine teeth. For the 34 American Museum specimens andthe single National Museum specimen, the museum number isgiven, and the sex (if adult). The 12 specimens from the litera-4 NO. 14161949 SOUTH AMERICAN EDALORHINA 5TABLE 1LIST OF KNOWN SPECIMENS OF Edalorhina perezi"perezi" Warts "plicifrons" WartsVomerine Vomerine Vomerine VomerineTeeth Teeth Teeth TeethPresent Absent Present AbsentECUADORSan JoseAbajo 22183 e_Avila, 500 m. "Myers""Napo" "Espada"Bafnos 52843 eCanelos 52844 e "Boul." cCanelos 52845 eCanelos 52846 9 52847 eCanelos "Boul." 9 -Canelos "Boul." 9 - _Canelos "Boul." 9Sarayacu "Boul." -"Ecuador" "Boul." 9 -"Ecuador" "Muller" - -PERURio MarafionIquitos 43623 _'Nauta - - "Cope"Nauta - "Cope"Rio HuallagaValle 42956 yg. - 42957 9 int. 42958 c' int.Valle 42959 9 - 42961 9Valle 42960 i -_Valle 42962 yg.Rio UcayaliPampa Hermosa 42101 9 42353 yg.Pampa Hermosa 42129 9 42586 9 int.Pampa Hermosa 42130 9Campo Santa Clara 127182a e -Cashiboya 43456 9 43390 9 43385 9 -Upper Rio Cushabatay - 43428 9 43432 c'Rio Pisqui 43272 9 - 43563 c'Punga (tributary of Rio Ta-piche) - 42927 9Rio Bombo (tributary of RioTapiche) 42902 9Peru-Brazil frontier (Rio Ta-piche) - 42909 9 -Peru-Brazil frontier (Rio Uto-quinia) _ 43140 c'Puerto Mairo (Rio Pachitea) "Boul." 9Monte Alegre (Rio Pachitea) 43039 9 43032 9 43028 9Monte Alegre (Rio Pachitea) - 43040 ca U.S.N.M. number.AMERICAN MUSEUM NOVITATESture are denoted by "Espada," etc., and sex is given if mentionedby the authors.The Ecuadorian localities are arranged from north to south(see accompanying map, fig. 5); those from Peru are arranged bythe main river valleys.The first column shows the provenance and sex of 27 (17 Ameri-can Museum) specimens that agree, in the characters considered,with the description of perezi. The second column does thesame for six American Museum specimens that agree with thedescription of perezi in dorsal warting but that completely lackvomerine teeth. The third column lists the type of buckleyi andsix American Museum specimens, one of which (denoted by theabbreviation "int.") does not quite agree in the dorsal warting.The fourth column lists the cotypes of plicifrons and five AmericanMuseum specimens, two of which are aberrant in the direction ofperezi (also marked "int.").The first column contains 27 specimens, the second six, thethird seven, the fourth seven (in terms of personally examinedAmerican Museum specimens, 17-6-6-5). There is a fair geo-graphical spread of each category, except for column two whichcontains only Rio Ucayali specimens. The "perezi" warting ver-sus the "plicifrons" warting comes out in a ratio of 33:14 (Ameri-can Museum specimens 23:11). Presence versus complete ab-sence of vomerine teeth comes out 34:13 (American Museumspecimens 23:11). These ratios are not precisely the typicalMendelian 9: 3:3:1 of a dihybrid cross nor the 3:1 of a monohy-brid cross. But only 47 specimens are known; they are widelyscattered samples from a range of over 100 of latitude, the collec-tion may not be random sampling, and intermediates might beshifted from one column to another. It is quite possible that thespecimens in the first column are double dominants, that those inthe last are double recessives, and that those in the two middlecolumns show one dominant and one recessive character.I therefore relegate to the synonomy of perezi Jimenez de laEspada the specific names plicifrons Cope and buckleyi Boulenger,and refrain from describing as a fourth species the specimens incolumn two.Edalorhina nasuta BoulengerThis species is represented in the Bassler collection by twomales, A.M.N.H. Nos. 43006, 43007, 33 and 32 mm. long, respec-NO. 14166SOUTH AMERICAN EDALORHINAtively, from Monte Alegre, Rio Pachitea (Ucayali basin), Peru.Both have vomerine teeth, as did Boulenger's three types fromHuancabamba, which is farther south and higher up in the basinof the Pachitea. Boulenger's largest specimen was 38 mm. long.The nature of the dorsal warting, in the form of an "X," andthe fleshy, sharply pointed, projection on the snout (shown in thedrawings of No. 43006, fig. 4) are the only known differences be-tween this form and perezi. The five specimens all have the inter-ocular dermal ridge of typical perezi. A glance at the bottom of4a4bFIG. 4. Edalorhina nasuta. A.M.N.H. No. 43006. Monte Alegre, Peru.A. Lateral view, B. Dorsal view. Natural size.the tabulation of perezi specimens will show that three of the fourvariants of perezi occur with nasuta at Monte Alegre.After having placed plicifrons in the synonymy of perezi, for thereasons given, I find it somewhat difficult to explain why I havenot done the same for nasuta. However, it is an extremelyperipheral form, and no intermediate specimens are known. Itherefore leave it in the status in which I found it, a distinct species.1949 7AMERICAN MUSEUM NOVITATESEDALOR HI NA AS A GENUSParker (1927, p. 457) suggests that "Edalorhina may well be anoffshoot from the leptodactylid stock, which gave rise, by aFIG. 5. Map showing localities where Edalorhina has been taken.reduction of the vomer and ultimate loss of the vomerine teeth, toPhysalaemus; certainly the two are very closely related." Ananalysis of the characters of Edalorhina confirms Parker's sugges-NO. 14168SOUTH AMERICAN EDALORHINAtion, but does not indicate that Edalorhina is ancestral to Physalae-mus.The peculiar dermal developments on the eyelids, etc., are suigeneris. Males have a horny plate on the thumb, and another,separate, one on the carpal tubercle. Males lack the externalpigmented vocal sac, so markedly developed in Physalaemus.The omosternum is cartilaginous with a circular anterior expan-sion. The sternal style is bony, and the posterior, cartilaginouspart of the sternum is bifurcate. The quadratojugal is in suturalcontact with the maxilla. The prevomer has a backwardly andmedially directed process, which usually bears teeth. The ter-minal phalanges are simple. The tympanic membrane is welldeveloped. A brilliantly colored, but not precisely delimitedinguinal gland is present. The toes have dermal fringes and areslightly webbed at the base. There are two metatarsal tuberclesbut neither is of the flattened, digging type. The hyoid andlaryngeal skeleton of E. perezi, as reported by Trewavas, resemblesthat of Physalaemus cuvieri more than it does that of any otherknown leptodactylid frog. The musculature of the throat alsoresembles that of P. cuvieri, but the origins of the right and leftsternohyoideus muscles are well separated (in contact in Physalae-mus), and the interhyoideus posterior has a superficial layer withfibers running obliquely forward and medially (unique in Lepto-dactylidae).These characters indicate (a) a common ancestry for Edalorhinaand Physalaemus (that might be Leptodactylus, and that mightalso have given rise to Pleurodema; cf. Parker, 1927), Edalorhinaretaining more of the Leptodactylus-like traits; (b) a certain diver-gence or specialization on the part of Edalorhina, in a directionaway from either Physalaemus or Pleurodema.Edalorhina pustulata Shreve (1941, p. 80), based on Museum ofComparative Zoology No. 7666, taken at Guayaquil, Ecuador, byC. T. Brues, in 1913, does not seem to me to be congeneric with thespecies considered above.Judging from the description of the external anatomy anddentition of the unique type, it does not have any close similarityto either perezi or to nasuta. The description does, however,indicate a high degree of similarity with one, and with only one,known species from northwestern South America, Engystomopspustulosus, and I suggest that its relationships lie in that directionrather than with the species here allocated to Edalorhina.1949 910 AMERICAN MUSEUM NOVITATES NO. 1416LITERATURE CITEDPARKER, H. W.1927. A revision of the frogs of the genera Pseudopaludicola, Physalaemus,and Pleurodema. Ann. Mag. Nat. Hist., ser. 3, vol. 20, pp. 450-478.SHREVE, B.1941. Notes on Ecuadorian and Peruvian reptiles and amphibians with de-scriptions of new forms. Proc. New England Zool. Club, vol. 18, pp.71-83.

South American Edalorhina

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