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Ff, io j ‘ ‘ a y ' t | J . 4 é 7 XK va hy : Wh : J : j "hi 0 on } ii z A, 4 4. 7 Won ah ta i as 4 eo ao tf \ 2) yy ‘/ WV i] ® vi | ul yi ) ‘ " , ts 5 Nia ia 7 iy, ‘ 7 ¥, 4 oA a wt wt Ds A it ef, Ma 1 % Ms i iy A ’ hi ia ae ey ts f i i ig 3 ~ ‘ Pi “i a ' , nas a iat it ye ny 1 al ha mf Digitized by the Internet Archive in 2011 with funding from California Academy of Sciences Library http://www.archive.org/details/occasionalpapers1/7univ Ve C/ f ¢ Ef se ae LI AN 40./- ‘i oe ee OCCASIONAL PAPERS .. eis a vic ee O. taurinus SS — SS Se HYLID FROGS, GENUS OSTEOCEPHALUS 19 Inner edge of third finger webbed to base of antepenultimate phalange; dorsum plain or marked with dark blotches or trans- verse bars; skull in adults smooth or slightly exostosed, lacking So GLP ES ENG, oS a ae FO 2 2. Skin on flanks areolate; dorsum in males bearing a mixture of large and small non-spinous tubercles; lips distinctly barred Spammers sgt OE eel PE ee Be O. buckleyi Skin on flanks smooth; dorsum in males bearing tubercles of Mmeegimnesize: lips mot barred 2.2.02 Sen 3 3. Dorsal pattern consisting of narrow transverse dark bars; dorsum in males bearing numerous small spinous tubercles _ O. leprieurii Dorsal pattern not consisting of transverse bars; dorsal tubercles RmeemeeneN 1a Oer te ee 4 4, Dorsum uniformly dark brown; venter heavily mottled with black, especially in females; dorsum in males bearing large, Weeumized tUDeTCIES 2-45. se eee O. verrucigerus Dorsum tan with irregular dark brown blotches; venter cream with fine brown reticulations; dorsum in males bearing few, small MupMiee mines. tubercles. 2.2). 5. re a he O. pearsoni ACCOUNTS OF SPECIES Osteocephalus buckleyi (Boulenger) Hyla buckleyi Boulenger, 1882:362 [Syntypes—BMNH 1947.2.13.36-39 from Sarayacu, Provincia Pastaza, Ecuador; BMNH 1947.2.13.40-41, 1947.2.13. 43-45 from Canelos, Provincia Pastaza, Ecuador; BMNH 1947.2.13.46 from “Paitanga” (= Pallatanga), Provincia Chimborazo, Ecuador (in error); Mr. Buckley collector; BMNH 1947.2.13.44 here designated as lectotype]. Hyla festae Peracca, 1904:39 [Holotype.—MIZS 2950 from “Valle de Santiago” (= lower Rio Zamora), Provincia Morona-Santiago, Ecuador; Enrico Festa collector]. New synonymy. Osteocephalus buckleyi—Goin, 1961:13. Hyla carri Cochran and Goin, 1970:211 [Holotype —FMNH 69702 from Acevedo, Rio Suaza, Departamento Huila, Colombia; Philip Hershkovitz collector]. New synonymy. Hyla cabrerai Cochran and Goin, 1970:215 [Holotype—USNM 152759 from Cano Guacaya, tributary of lower Rio Apoporis, Comisaria Amazonas, Colombia; Isadore Cabrera collector]. New synonymy. Justification of Synonymy.—Boulenger (1882:362) listed 11 spec- imens in his description of Hyla buckleyi. We have examined all of these and conclude that one (BMNH 1947.2.13.42) is O. leprieurii. Cochran and Goin (1970:213) restricted the type locality to Canelos, Provicia Pastaza, Ecuador; we here select BMNH 1947.2. 13.44 from that locality as the lectotype. This specimen is a male having a snout-vent length of 37.9 mm; the diameter of the tym- 20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY panum is 3.5 mm, 81.4 percent of the diameter of the eye. The type series, exclusive of BMNH 1947.2.13.42 (= O. leprieurii) consists of six males having snout-vent lengths of 37.9-44.6 (mean 40.4) mm, and four females having snout-vent lengths of 50.0-53.9 (mean 51.5) mm. The dorsum in the males bears a mixture of large and small tubercles, whereas the dorsum in females is nearly smooth. The skin on the flanks, especially the axilla, is areolate. The coloration consists of a creamy tan ground color with irregular reddish brown markings on the back and broad transverse bars on the limbs. The dorsal markings are narrowly bordered by creamy white; those on the back consist of an interorbital bar and a pair of longitudinal marks beginning in the scapular region and usually diverging pos- teriorly in the sacral region or converging into a broad median blotch. One specimen has a middorsal creamy white stripe from the tip of the snout to the vent. In all of the types large dark brown spots are present on the flanks and posterior surfaces of the thighs. The ventral surfaces are pale creamy tan with or without diffuse brown spots on the throat and chest. The holotype of Hyla festae is a female having a snout-vent length of 75.0 mm; the diameter of the tympanum is 3.9 mm, 57.4 percent of the diameter of the eye. The skin is smooth on the dorsum and areolate on the anterior part of the flanks. The dorsum is pale brown with a large median longitudinal dark brown blotch on the back and broad transverse bars, narrowly outlined by cream, on the limbs. Dark brown spots are present on the flanks; the pos- terior surfaces of the thighs are dark brown. The throat and belly are grayish white with irregular dark brown spots. The holotype of Hyla carri is a female having a snout-vent length of 66.1 mm; the diameter of the tympanum is 4.7 mm, 81.0 percent of the diameter of the eye. The skin on the dorsum is smooth with scattered small tubercles and areolate on the anterior part of the flanks. The dorsum is tan with irregular dark brown blotches on the back and transverse bars on the limbs; all dorsal markings are narrowly outlined by creamy white. Dark brown spots are present on the flanks; the venter and posterior surfaces of the thighs are tan without dark spots. The holotype of Hyla cabrerai is a female having a snout-vent length of 52.7 mm; the diameter of the tympanum is 4.0 mm, 76.9 percent of the diameter of the eye. The skin on the dorsum is weakly tuberculate and that on the anterior part of the flanks is areolate. The dorsum is creamy tan with dark brown markings (interorbital bar, reticulations on occiput, three longitudinal streaks on back, | HYLID FROGS, GENUS OSTEOCEPHALUS 21 and broad transverse bars on limbs). Irregular dark brown spots are present on the flanks. The venter is pinkish tan with small reddish brown spots on the throat and darker brown spots on the chest and belly. In their description of Hyla cabrerai, Cochran and Goin (1970: 217) stated: “This species, together with buckleyi and pearsoni certainly make a closely knit group. ... Both buckleyi and cabrerai have long hind legs, with the extended heel reaching to the tip of the snout, while in pearsoni the extended heel reaches only to the eye. H. buckleyi has the belly dusky, while it is heavily spotted in cabrerai and is reticulated in pearsoni. H. cabrerai seems to have the heaviest hands with the most webbing between the fingers; the other two species have the webbing reduced between the fingers.” The description of Hyla cabrerai was based on three specimens. We have examined the holotype and one paratype (WCAB 13284 from Territorio do Amapa, Brasil). Another paratype in the private collection of C. J. Goin from Cano Tui, between Mitu and Raudal de Yurupari, Comisaria de Vaupés, Colombia, was not examined. Cochran and Goin (1970:211) based their description of Hyla carri on one gravid female and stated: “A large Hyla with the vomerine teeth in two AA shaped patches between the somewhat squarish choanae; reduced webs between the fingers; and a pattern of dorsal dark blotches bordered by light margins. The species is not similar to any other species known in Colombia. It is perhaps most closely related to Hyla claresignata of Brazil, from which it can be differentiated by its more heavily spotted dorsum, larger tympanum, and lack of dark anal spots.” Except for the inclusion of the name in checklists, Hyla festae has not been mentioned in the literature since the original descrip- tion. The wholesale synonymization of names, which, on the bases of their published diagnoses, seem to apply to distinctly different species, is possible with the application of uniform criteria to the types and series of other specimens. In measurements and propor- tions the type specimens of the nominal taxa all fall within the range of variation exhibited by a series of 18 males and 15 females from Provincia Pastaza, Ecuador, except the ratio of the diameter of the tympanum to that of the eye in the female holotype of Hyla festae. In that specimen the ratio is 0.574, whereas the ratio in the 15 females from Provincia Pastaza is 0.587-0.905 (mean 0.736). Ventral coloration is the most variable character among the types. The venter in the type of Hyla festae is boldly spotted; it is 22 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY distinctly spotted in cabrerai, uniform tan in carri, and tan, flecked, or spotted in the type series of buckleyi. The ventral coloration in series of specimens from Amazonian Ecuador encompasses that ob- served in all of the types, except that of festae, which has more ventral spotting than any other individual. The webbing on the hand usually excludes the penultimate phalanges of the fingers, but in some specimens from Amazonian Ecuador the webbing encompasses the proximal parts of the penul- timate phalanges of the fingers. In a few of these specimens, the holotype of festae, and one paratype of cabrerai the webbing ex- tends to the middle of the penultimate phalanges of the third and fourth fingers. In the holotype of cabrerai the webbing extends to the middle of the penultimate phalanges of the third and fourth fingers and to the base of the disc of the second finger. The types of the nominal taxa and series of specimens from Guyana and Amazonian Ecuador display noticeable variation in dorsal coloration. The variety of dorsal patterns of all of the types is included in the variation displayed by the other specimens. All specimens have some amount of dark spotting on the flanks; all have vertically barred lips, on which a pale subocular spot usually is evi- dent. Probably the most unifying physical characteristic of all of the specimens is the nature of the skin on the anterior part of the flank. The skin is elevated amidst an irregular network of depres- sions. This areolate dermal condition is present in all specimens and does not occur in other species of Osteocephalus. The degree of tubercularity of the skin on the dorsum is variable and sexually dimorphic. All males are tubercular, whereas small females are smooth or have only a few scattered tubercles. Large females usually have pronounced tubercles on the eyelids and supra- tympanic fold. In their description of Hyla carri, Cochran and Goin (1970:211) misrepresented the nature of the dentigerous processes of the pre- vomers, which are angular, not A-shaped. Their suggestion that the Colombian Hyla carri is related to Hyla claresignata in southeastern Brasil is unfounded. The latter species is smaller (40 mm), has a yellow dorsum and venter, dark brown spots dorsolaterally, oblique dentigerous processes of the prevomers, small tympanum, and smooth skin dorsally. The ventral coloration of the type of Hyla festae resembles that of Osteocephalus verrucigerus, but the type differs from verru- cigerus by having areolate skin on the flanks and distinct dark mark- ings on the dorsum. In verrucigerus the skin on the flanks is smooth, HYLID FROGS, GENUS OSTEOCEPHALUS 23 and the dorsum is uniform dark brown, except for a tan snout in females. Comparisons of the types of the nominal species with series of specimens from Guyana, Colombia, Ecuador, and Pert: suggest strongly that the types are representative of one taxon, the oldest name for which is Hyla buckleyi Boulenger, 1882. Consequently, we place Hyla festae Peracca, 1904, Hyla carri Cochran and Goin, 1970, and Hyla cabrerai Cochran and Goin, 1970, as junior synonyms of Hyla buckleyi Boulenger, 1882. Diagnosis.—1) Size moderate, sexual dimorphism extreme; max- imum observed snout-vent length in males 48.1 mm, in females 75.1 mm; 2) skin on dorsum in males bearing a mixture of large and small non-spinous tubercles; 3) skin on flanks, especially anteriorly, areolate; 4) web usually extending only to base of antepenultimate phalange on inner edge of third finger; 5) dorsum pale tan or green with irregular, longitudinal, dark brown blotches, usually narrowly outlined with cream; 6) venter cream or tan, suffused with brown or marked with brown spots in some specimens; 7) lips marked with vertical brown and cream bars; 8) flanks creamy tan with irregular brown spots and/or diagonal marks; 9) dermal roofing bones of skull lacking exostosis; 10) dermal sphenethmoid absent; 11) nasals widely separated medially; 12) anteromedial margin of frontoparie- tal at mid-level of orbit; 13) frontoparietal fontanelle partially ex- posed; 14) palatine serrate; 15) parasphenoid bearing odontoids; 16) zygomatic ramus of squamosal extending approximately one- half of distance to maxillary arch; 17) transverse processes of third presacral vertebra narrower than sacral diapophyses; transverse processes of presacral vertebrae 3-8 subequal in width and narrower in males than in females; 18) intermandibularis and submentalis muscles independent; 19) supramandibular portion of inter- hyoideus extensively developed; associated skin forming broad loose fold. Osteocephalus buckleyi can be distinguished readily from all other species in the genus by the presence of areolate skin anteriorly on the flanks and by the rather boldly contrasting dorsal pattern. Furthermore, females are distinctive in having tubercles on the eyelids and supratympanic folds. Distribution.—The periphery of the Amazon Basin, in the Gui- anas and Territorio do Amapa in northeastern Brasil; the upper Amazon Basin from southern Colombia to east-central Bolivia; one locality (Acevedo) in upper Rio Magdalena drainage in Colombia (Fig. 8). All localities are at elevations of less than 700 m. Records 24 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY SOO l000 kilometers 50° Fic. 8. Distribution of Osteocephalus buckleyi (circles) and O. pearsoni (triangles ). for Pallatanga and Santiago in Provincia Chimborazo, Ecuador (high on the Pacific slopes of the Andes), are considered to be erroneous. 78 specimens from 40 localities. Remarks.—In life the dorsum is green with dark markings. A male (KU 123171) from Santa Cecilia, Ecuador, was: “Dorsum green with dark brown blotches. Anterior and posterior surfaces of thighs dull blue. Venter brown, flecked with white. Iris greenish bronze with brown horizontal triangles and ventromedian brown line.” (W. E. Dueliman, field notes, 16 June 1968.) A female (KU 126646 ) from Lago Agrio, Ecuador, was: “Dorsum pale green with darker green blotches and creamy yellow middorsal stripe. Lateral blotches bronze-tan. Flanks tan with black blotches. Anterior sur- faces of thighs dark brown. Dorsal and posterior surfaces of thighs and shanks tan with dark brown blotches. Webbing brown. Sub- orbital spot green. Postorbital bar black. Belly grayish brown in appearance—tips of granules white; intergranular spaces brown. Iris golden bronze with black flecks peripherally and median, hor- HYLID FROGS, GENUS OSTEOCEPHALUS 25 izontal, reddish brown streak.” (W. E. Duellman, field notes, 12 May 1969. ) No ontogenetic change in coloration has been noted. Osteocephalus leprieurii (Duméril and Bibron) Hyla leprieuriti Duméril and Bibron, 1841:553 [Holotype—MNHN 4629 from “Cayenne”; Mons. Leprieur collector]. Hypsiboas leprieurii—Cope, 1867:200. Hyla leprieurii britti Melin, 1941:42 [Holotype—NHMG 489 from the Rio Uaupés, north of the Rio Japu, Territorio do Amazonas, Brasil; Douglas Melin collector]. New synonymy. Hyla leprieurii leprieurii—Melin, 1941:42. Osteocephalus britti—Goin, 1961:13. Osteocephalus leprieurii—Goin, 1961:13. Justification of Synonymy.—The holotype of Hyla leprieurii is a female having a snout-vent length of 46.6 mm. The diameter of the tympanum is 3.7 mm, 69.8 percent of the diameter of the eye. The dorsal roofing bones are smooth, and the skin on the dorsum is smooth. The penultimate phalanges of the fingers are not included in the webbing. When we examined the specimen on 2 July 1969, it was slightly soft and somewhat faded to a peculiar grayish green color with faint darker transverse bars on the limbs. Duméeril and Bibron (1841:554) described the coloration, as follows: “The loreal region in black. A stripe of the same color extends from the posterior border of the orbit to the corner of the mouth, passing through the tympanum. All of the dorsal parts are grayish white with large transverse brown bands, which are more expanded and less reg- ularly outlined on the back than on the limbs. There is one of these on the occiput that is in a triangular shape. All of the venter is white.” (Free translation from French. ) The holotype of Hyla leprieurii britti is a male having a snout- vent length of 48.1 mm. The diameter of the tympanum is 3.6 mm, 65.5 percent of the diameter of the eye. The skin on the dorsum is tubercular; the tubercles are small on head and on the dorsal sur- faces of the limbs and slightly larger on the back. The penultimate phalanges of the fingers are not included in the webbing. Melin (1941:43) stated: “Above blackish brown with a very indistinct band between the eyes; iris with mottle of metalic lustre; hinder parts of upper jaw whitish; sides of body mottled with blackish brown; hind limbs (especially tibiae and tarsi) with narrow, diffuse cross bars; beneath whitish with slight brown mottle along jaw.” We examined the type on 17 February 1969; at that time it was dull brown above with faint, narrow, dark brown, transverse bars on 26 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY the back and dorsal surfaces of the limbs. A cream subocular spot was evident, and the venter was creamy white. Melin (1941:42) stated that the holotype of Hyla leprieurii britti “,.. resembles a good deal H. leprieurii Dum. & Bibr. As, however, it differs from the latter species by its very concave loreal region, small tympanum, and almost uniformly brownish colour, it may at least form a subspecies of leprieurii . . .” The pattern of narrow transverse bars on the backs of the holotypes of H. leprieurii and H. britti is a condition shared only by these two nominal taxa that are placed in Osteocephalus. Melin noted that britti differed from leprieurii in the depth of the loreal concavity and in the size of the tympanum. Neither of these differences is noteworthy in compari- son with series of specimens. The depth of the loreal concavity is a highly subjective character, and we note no differences between the types. The ratio of the diameter of the tympanum to the diameter of the eye is relatively smaller in both holotypes (0.698 in leprieurii— ? ; 0.655 in britti— ¢ ) than in series of fresh specimens from Lago Agrio, Ecuador (0.652-0.884, mean 0.785 in 17 males; 0.700-0.909, mean 0.790 in 20 females ). The smaller proportions in the types may be due to geographic variation or to shrinkage as a result of many years in preservative (130+ years for leprieurii; 45 for britti). Comparisons of the holotypes with series of specimens from Ecuador, Guyana, and Surinam indicate that one morphological species occurs throughout the upper Amazon Basin and the Guianas and that both type specimens are representatives of one species. Consequently, we consider Hyla leprieurii Dumeéril and Bibron, 1841, to be a monotypic species with Hyla leprieurii britti Melin, 1941, as a junior synonym. In their account of Osteocephalus leprieurii, Cochran and Goin (1970:323) stated: “The specimen described and illustrated (MCZ 28042) has been directly compared with the types of leprieurii, planiceps, and vilarsi by the junior author and there seems to be no doubt that all are conspecific. Another specimen (CNHM 69716) has been directly compared with the types of planiceps and vilarsi and these, likewise, are considered conspecific.” With this justifica- tion Cochran and Goin (1970:322) included Osteocephalus plani- ceps Cope, 1874, and Hyla vilarsi Melin, 1941, in the synonymy of Osteocephalus leprieurii. We do not concur with Cochran and Goin’s synonymy and con- tend that planiceps and vilarsi are synonyms of Osteocephalus taurinus; we give our reasons in the account of that species. We have examined the specimens listed as O. leprieurii by Cochran and HYLID FROGS, GENUS OSTEOCEPHALUS 27 Goin; several of them, including CNHM (= FMNH) 69716, are taurinus. Thus, due to Cochran and Goin’s confusion of two taxa, their comparisons of certain specimens with types has little meaning. Cochran and Goin did not include Hyla leprieurii britti in their synonymy of Osteocephalus leprieurii but did discuss the name in their account of Osteocephalus orcesi (= O. verrucigerus), as fol- lows (1970:319): “When we first examined one of the specimens we felt sure that we had Melin’s Hyla britti at hand, but on direct comparison with the type of britti the two proved to be different. After studying the type of orcesi (SUNHM 13150) we have no doubt that the specimens at hand are orcesi and that britti is a dif- ferent, probably valid species.” Diagnosis.—_1) Size moderate, sexual dimorphism evident; max- imum observed snout-vent length in males 48.4 mm, in females, 61.5 mm; 2) skin on dorsum in males bearing numerous, minute, spinous tubercles; 3) skin on flanks smooth; 4) web extending to base of antepenultimate phalange on inner edge of third finger; 5) dorsum tan or olive-brown with transverse brown or olive bars; 6) venter creamy white or pale tan without markings; 7) lips marked with creamy tan labial stripe and suborbital spot; 8) flanks pale tan with no markings; 9) dermal roofing bones of skull lacking exostosis; 10) dermal sphenethmoid absent; 11) nasals juxtaposed medially; 12) anteromedial margin of frontoparietal between mid- and an- terior levels of orbit; 13) frontoparietal fontanelle partially ex- posed; 14) palatine not serrate; 15) parasphenoid lacking odontoids; 16) zygomatic ramus of squamosal extending about one-half of distance to maxillary arch; 17) transverse processes of presacral vertebrae 3-8 about equal in width to one another and to sacral diapophyses; 18) intermandibularis and submentalis muscles con- nected; 19) supramandibular portion of interhyoideus forming simple tubular posterolateral extension; associated skin unmodified. Osteocephalus leprieurii differs from all other members of the genus by having transverse dark bars on the back. Two other hylids (Hyla lanciformis and multifasciata) in the Amazon Basin have transverse dark marks on the dorsum. Both of these differ from leprieuriti by having pointed snouts, much longer hind limbs, and smooth skin dorsally. Distribution.—The periphery of the Amazon Basin, in the Gui- anas and the upper part of the basin in southern Colombia, Ecuador, Pert, and extreme western Brasil (Fig. 9). Most localities are at elevations of less than 500 m, but the species ascends the lower 28 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY \ Foy = 0 500 ‘(000 / ) ees et ee = | NS : kilometers 6 73) ao 16° I6 ° ° : ° ° 78 74 70° 66 62 58° 54° 50° Fic. 9. Distribution of Osteocephalus leprieurii (circles) and O. verrucigerus (triangles ). Andean slopes to elevations of 1100 m. 265 specimens from 31 localities. Remarks.—Most adults of leprieurii have distinct transverse markings on the back; these are variable in width, extent, and ar- rangement. In some specimens, such as USNM 166557, some of the transverse bars are fragmented into spots; in a few specimens the dorsal pattern consists solely of small dark spots arranged in transverse rows. Such specimens have a dorsal pattern resembling that of some taurinus. The transverse nature of the dorsal markings is further modified in some specimens, such as USNM 166555, in which the dark bars are fragmented and oblique. Extreme ontogenetic change in color pattern is exhibited by this species (Fig. 10). Juveniles having snout-vent lengths of less than 28 mm have an olive-brown dorsum with a pale cream stripe across the head and broad, cream, dorsolateral stripes; transverse dark bars are absent on the body and limbs. Individuals having snout-vent lengths of 30-35 mm have dark brown transverse bars on HYLID FROGS, GENUS OSTEOCEPHALUS 29 Fic. 10. Ontogenetic change in color pattern in Osteocephalus leprieurii: a. KU 126644; b. KU 126640; c. KU 126625. x2. the back and limbs but still retain the light dorsolateral stripes, whereas the stripes are lost in larger individuals. Coloration in life of specimens from Lago Agrio, Ecuador: “In males the dorsal ground color varies from dark brown to ochre-tan; dorsal markings uniformly dark brown. Most specimens have dark brown and cream anal stripes; labial area cream-colored. Flanks vary from tan to white. Ventral coloration varies from salmon to tan to white. The iris is bronze with a greenish cast and black reticulations. In females the dorsal coloration is the same as in males, except that dark marks tend to be outlined with cream; venter tannish salmon.” (W. E. Duellman, field notes, 12 May 1969). Osteocephalus pearsoni (Gaige) Hyla pearsoni Gaige, 1929:3 [Holotype—UMMZ 57548 from the upper Rio Beni, below mouth of Rio Mapiri, Departamento El Beni, Bolivia; N. E. Pearson collector]. Osteocephalus pearsoni—Goin, 1961:13. Justification of Synonymy.—Goin (1961:13) suggested that Hyla pearsoni Gaige was an Osteocephalus, but Cochran and Goin (1970: 30 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 217) considered pearsoni to be a Hyla. The presence of exostosed dermal roofing bones, angulate prevomerine dentigerous processes, and the structure of the vocal sacs are characters which place the species in Osteocephalus. Diagnosis.—1) Size moderate, sexual dimorphism evident; max- imum observed snout-vent length in males 46.2 mm, in females 54.7 mm; 2) skin on dorsum in males bearing a few, small, scattered non- spinous tubercles; 3) skin on flanks smooth; 4) web extending to base of antepenultimate phalange on inner edge of third finger; 5) dorsum tan with irregular brown blotches; 6) venter cream with fine brown reticulations; 7) lips dark with pale vertical bar below eye; 8) flanks pale tan with round, brown spots; 9) dermal roofing bones of skull slightly exostosed; 10) dermal sphenethmoid absent; 11) nasals narrowly separated medially; 12) anteromedial margin of frontoparietal between mid- and anterior levels of orbit; 13) frontoparietal fontanelle covered; 14) palatine not serrate; 15) parasphenoid lacking odontoids; 16) zygomatic ramus of squamosal extending about one-half distance to maxillary arch; 17) transverse processes of third presacral vertebra approximately equal in width to sacral diapophyses; transverse processes of presacral vertebrae 3-8 subequal in width; 18) intermandibularis and submentalis muscles connected; 19) supramandibular portion of interhyoideus extensively developed; associated skin forming broad loose fold. Osteocephalus pearsoni can be distinguished most readily from other members of the genus by the brown reticulate pattern on the venter, round brown spots on the flanks, and smooth skin on the flanks. Also, it is the least tuberculate species in the genus. Distribution.—_Upper Amazon Basin and Amazonian slopes of the Andes in central Peru (1620 m in Rio Ucayali drainage) and northern Bolivia (less than 500 m in Rio Beni drainage) (Fig. 8). 6 specimens from 3 localities. Remarks.—The specimen from Yaupi, Peru (KU 136312) is a subadult female having a snout-vent length of 39.8 mm. In life the coloration was: “Dorsum light pinkish brown with large rich chocolate brown blotch from eyes to anterior tips of ilia; numerous small chocolate blotches on flanks; dorsal surfaces of thighs and shanks, canthus, and supraorbital region to insertion of forearm chocolate brown; supralabial border and short bar from eye to lip bronze-white; venter bronze-white with numerous tiny chocolate brown flecks [tending to form reticulations on throat and chest]; anterior and posterior surfaces of thighs light olive-brown; iris largely black with gold flecks.” (Thomas H. Fritts, field notes, 23 HYLID FROGS, GENUS OSTEOCEPHALUS 31 March 1970.) On the basis of this one subadult, it seems likely that reticulations on the venter develop with age. Osteocephalus taurinus Steindachner Osteocephalus taurinus Steindachner, 1862:77 [Holotype -NHMW 16492 from Barra do Rio Negro, Manaus, Territorio do Aimazonas, Brasil; Johann Natterer collector]. Osteocephalus flavolineatus Steindachner, 1862:80 [Holotype ——-NHMW 16495 from Cucui, Territorio do Amazonas, Brasil; Johann Natterer collector]. Trachycephalus (Osteocephalus) taurinus Steindachner, 1867:64. Osteocephalus planiceps Cope. 1874:122 [Holotype—ANSP 11399 from Nauta, Departamento de Loreto, Peri; James Orton collector]. New synonymy. Hyla taurina—Boulenger, 1882:363 [synonymized Osteocephalus flavolineatus Steindachner, 1862, with O. taurinus Steindachner, 1862]. Hyla planiceps—Boulenger, 1882:364. Hyla (Trachycephalus) vilarsi Melin, 1941:40 [Holotype —NHMG 488 from Taracua, Rio Uaupés, Territorio do Amazonas, Brasil; Douglas Melin collector]. (fide Bokermann, 1966: 64. ) Hyla depressa Andersson, 1945:73 [Holotype—NHRM 1966 from the Rio Pastaza watershed (? Provincia Pastaza), Ecuador; William Clarke-Mac- Intyre collector]. New synonymy. Justification of Synonymy.—The holotype of Osteocephalus taurinus is a female having a snout-vent length of 103.9 mm. The diameter of the tympanum is 6.8 mm, 77.3 percent of the diameter of the eye. The skull is strongly exostosed, and the lateral edges of the frontoparietals are elevated so as to form distinct ridges. The skin on the dorsum is smooth. When we examined the type on 5 August 1969, the specimen was soft and badly faded to a pale creamy tan with pale brown transverse bars on the hind limbs and spots on the flanks. Steindachner (1862:79) described the coloration of the type: “In the preserved specimen the dorsum of the entire body, including fore and hind limbs, is a light yellow-brown color, which becomes lighter towards the venter. The belly is whitish, as are the undersides of the arms and legs. The throat is indistinctly marbled with brown. Roundish dark brown flecks are randomly distributed in a considerable number along the side of the body up to the eye; the tympanum is more or less fully surrounded by brown. A few discrete spots, always more or less drawn out in length, on the sides of the body, are also found on the posterior part of the back. The dorsal surfaces of the fore and hind feet are marked with somewhat obliquely arranged brown transverse bands, which are more intensively colored near the margin than in the middle of the band.” (free translation from German. ) The holotype of Osteocephalus flavolineatus is a female having 32 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY a snout-vent length of 81.8 mm. The diameter of the tympanum is 6.0 mm, 71.4 percent of the diameter of the eye. The skull is strongly exostosed, and the lateral edges of the frontoparietals are elevated so as to form a ridge on each side. The skin on the dorsum is very weakly tuberculate. We examined the type on 9 August 1969 and found it to be in excellent condition. The color pattern is unchanged from that described by Steindachner (1862:81). The dorsum is tan with irregular brown blotches on the back, spots on the flanks, and transverse bars on the limbs. A narrow creamy white, middorsal stripe extends from the snout to the vent. The subocular area is creamy tan, and the venter is tan. Boulenger (1882:363) questionably synonymized flavolineatus with taurinus. We have observed that a middorsal cream stripe occurs in about 10 percent of the specimens of taurinus and in some specimens of buckleyi. This is a common color morph in many species of Eleutherodactylus. In the absence of distinguishing morphological characteristics we can only conclude that the middorsal stripe is a pattern variant and that Boulenger was correct in synonymizing flavolineatus with taurinus. The holotype of Osteocephalus planiceps is a male having a snout-vent length of 58.5 mm. The diameter of the tympanum is 4.9 mm, 77.8 percent of the diameter of the eye. The skull is moderately exostosed, and the lateral edges of the frontoparietals are distinctly elevated. The skin on the dorsum is tuberculate. Cope (1874:122) described the coloration of the type as follows: “Color above uniform dark brown, concealed surfaces on the limbs similar and without any markings. Sides a little varied with the white of the belt. A light border to the upper lip, and lighter line from the orbit to the angle of the mouth; dermal scapular fold pale edged. Femur and tibia with dark crossbands on the exposed surfaces.” We examined the holotype on 25 September 1969, and found it to be soft and rubbed. The coloration remains much the same as described by Cope, who provided no means of distinguishing planiceps from taurinus. The coloration and morphometric and structural characters of the type of planiceps all fall within the range of variation displayed by series of O. taurinus from the upper Amazon Basin. The type of Hyla vilarsi is a gravid female having a snout-vent length of 62.7 mm. The diameter of the tympanum is 4.8 mm, 73.8 percent of the diameter of the eye. The dorsal roofing bones of the skull are moderately exostosed, and the lateral edges of the fronto- parietals are distinctly elevated. The skin on the dorsum is smooth. HYLID FROGS, GENUS OSTEOCEPHALUS 33 Melin (1941:42) described the coloration of the holotype as follows: “Above uniform reddish brown; upper eyelids and sides of head darkish brown; below the rostral edge a narrow dark band, con- tinuing as a broader light-edged one through the eye and tympanum towards the base of the forelimb and then farther on continuing along the sides as a line of black spots; sides of upper jaw whitish with traces of dark cross bars (one distinct under the eye); sides of body darkish with black spots and marble, often on a whitish ground; thighs, tibiae, and tarsi each with two broad light-edged, dark cross bars on a brownish ground (less distinct on thighs ); sides of thighs finely mottled with brown; beneath whitish with small, sparse spots along jaw, on the chest and sides.” We examined the type on 17 February 1969, at which time the specimen was some- what desiccated, especially the hands and feet. The coloration re- mains much the same as described by Melin, except that he failed to note the presence of four elongate spots on the back. The status of the names Osteocephalus planiceps Cope and Hyla vilarsi Melin was confused by Cochran and Goin (1970:322), who assigned these names to the synonymy of O. leprieurii. Bokermann (1966:64) placed Hyla vilarsi in the synonymy of Osteocephalus taurinus without justification. The type specimens of both planiceps and vilarsi have moderately exostosed dermal roofing bones and distinct cranial ridges. The type of planiceps has moderately large tubercles on the dorsum, and the type of vilarsi has spots on the throat, chest, and flanks and longitudinal markings on the back. All of these features are characteristic of tawrinus and not of leprieurii, which lacks exostosis and cranial ridges and has trans- verse markings on the back, no spots on the throat, chest, and flanks, and in males has small dorsal tubercles. The type of Hyla depressa is a male having a snout-vent length of 69.8 mm. The diameter of the tympanum is 5.2 mm, 77.6 percent of the diameter of the eye. The dorsal roofing bones of the skull are moderately exostosed, and the lateral edges of the frontoparie- tals are elevated. The skin on the dorsum is tuberculate. The dorsum is dull brown with a broad darker brown longitudinal mark having indistinct lateral edges from the snout to the post-sacral area. A narrow cream middorsal line extends from the snout to the vent. The side of the head is dark brown, palest posteroventral to the orbit. The posterior surfaces of the thighs are dull brown; the flanks are pale brown, and the ventral surfaces are pale creamy tan. Dark brown transverse bars are present on the limbs. When we ex- amined the type on 3 January 1969, it was in excellent condition. 34 OCCASIONAL PAPERS. MUSEUM OF NATURAL HISTORY Andersson (1945:75) contrasted the type of Hyla depressa with leprieurii and buckleyi, but he did not compare his specimen with taurinus, from which it exhibits no distinguishing features. Osteocephalus taurinus is a widespread and variable species, and it has received several specific names. We are convinced that Osteocephalus taurinus Steindachner, 1862, is the oldest available name for this large Amazonian species. The following names are junior synonyms: Osteocephalus flavolineatus Steindachner, 1862; Osteocephalus planiceps Cope, 1874; Hyla (Trachycephalus) vilarsi Melin, 1941; Hyla depressa Andersson, 1945. Diagnosis.—1) Size large; sexual dimorphism evident; maximum observed snout-vent length in males 84.6 mm, in females 104 mm; 2) skin on dorsum in males bearing many moderately large, spinous tubercles; 3) skin on flanks smooth; 4) web extending to middle of antepenultimate phalange on inner edge of third finger; 5) dorsum brown usually with a large medial dark brown blotch or, less fre- quently, several dark spots; narrow middorsal yellow line present in some; 6) venter cream or tan with or without small, irregular brown flecks; 7) lips brown with vertical cream bar below eye in some, expanded into pale labial stripe posteriorly in some females; 8) flanks tan or cream with or without small, irregular brown spots; 9) dermal roofing bones of skull exostosed, casqued, and co-ossified (in large adults); 10) dermal sphenethmoid present; 11) nasals juxtaposed medially; 12) anteromedial margin of frontoparietals at mid-level of orbit; 13) frontoparietal fontanelle covered; 14) pala- tine serrate; 15) parasphenoid bearing odontoids; 16) zygomatic ramus of squamosal usually articulating with maxillary arch; 17) transverse processes of third presacral vertebra approximately equal in width to sacral diapophyses; transverse processes of presacral vertebrae 3-8 subequal in width; 18) intermandibularis and sub- mentalis muscles connected; 19) supramandibular portion of inter- hyoideus extensively developed; associated skin forming everted pouch. The moderately rugose dorsum (in males), large size, extensive webbing on the hand, and frontoparietal flanges in adults serve to distinguish taurinus from other members of the genus. Distribution.—_The Amazon Basin, the upper Orinoco Basin, and the Guianas. Most localities are below 500 m, but the species as- cends the lower Amazonian slopes of the Andes to elevations of about 1000 m (Fig. 11). A record from Caracas, Venezuela, and those from Provincia Carchi and Provincia Esmeraldas, Ecuador, are considered to be erroneous. The latter specimens were included HYLID FROGS, GENUS OSTEOCEPHALUS 35 4 > “Oe, 6G. 62° 58° 54° 50 Z ae _ 2 ae ae aed fe f ~ le 8 a las l as ae o S))__.-» e ®Yeo Ce eo. e e ® ( : \ od ) t @® _ «© e a Se ee: 2 a ff @ e @ ~ BQ Ly e y Ge e @. SQ : S . hge— 0 " eee L Tr ! pe er ® eee e ig ) e® 2 : j 2 J * { ° af --e@ P Z ee 16 ‘ Be ©) i / fe e ° 4 e 8 y * erry (oO OD e e a -@ i ) “@ 12° igre , . \ Ae i 0 500 _ 1000 < Os : Se SS e foo kilometers Ie" ier 78° 74 ° a) ee 3 3 3 2.70 66° “862 58 54 50 Fic. 11. Distribution of Osteocephalus taurinus. in a collection sold to the University of Illinois; contained in the collection are many common Amazonian species unknown from the Pacific lowlands. 516 specimens from 151 localities. Remarks.—This widespread species is highly variable in size and coloration. Striking differences in snout-vent length are evident in series from various parts of the range. The smallest calling males (CAS-SU 12351-6 from Rio Tapirapé, Brasil) have snout-vent lengths of 46.5-60.3 (mean 53.3) mm, whereas the largest (FMNH 140254, KU 92243-6, WCAB 9997, 10001, 10003-4 from Igarapé Marmelo, Brasil) have snout-vent lengths of 71.5-84.6 (mean 77.6) mm. Mean values of snout-vent lengths of males from other lo- calities are: Rio Pastaza drainage, Ecuador 73.8 mm, Surinam 67.7 mm, Rio Ucayali drainage, Peru 57.6 mm, and Guyana 55.5 mm. Although the difference between the smallest and largest adults is highly significant, populations bridging the gap do exist. Further- more, the geographic arrangement of small versus large frogs is a confusing mosaic. We have entertained the thought that we have included more than one species in taurinus, but on the basis of pre- 36 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY served specimens we are unable to detect consistent differences distinguishing two or more taxa. The coloration and pattern of taurinus are so variable that no one series of statements can describe samples drawn from the entire range of species. We have been unable to determine geographic trends in color pattern; instead the variation within a given sample can encompass the variety known in most other samples. Two minor exceptions do exist. A narrow middorsal light stripe is present in some individuals from throughout the range, but striped specimens are most common in the upper Amazon Basin. The absence of dorsal markings is uncommon in the entire species, but it is most frequent in individuals from the Guianas. A few individuals, such as KU 105230, have scattered white spots on the dorsum. The coloration of four males in life from Lago Agrio, Ecuador (KU 126652-5) was: “Dorsal ground color tan to dark brown with darker brown markings. Flanks creamy tan to yellow with brown or black flecks or mottling. Venter uniform creamy yellow or yellow with brown spots or reticulations. Iris greenish yellow with radiat- ing black streaks and a median, horizontal reddish brown streak.” (W. E. Duellman, field notes, 12 May 1969.) A female from Santa Cecilia, Ecuador (KU 123173), was: “Dorsum mottled olive-green and tan. Flanks tan with brown spots. Belly and throat creamy white, becoming tan posteriorly. Edge of upper jaw olive-green.” (W. E. Duellman, field notes, 16 June 1968.) Another female from Santa Cecilia (KU 123175), was: “Brown dorsally with cream- colored mottling. Transverse bars on legs darker brown with cream-colored edges. Margin of upper lip creamy yellow. Anterior and posterior surfaces of thighs tan. Flanks white with brown spots. Venter creamy white. Iris greenish bronze with heavy radiating reticulations of black.” (W. E. Duellman, field notes, 22 July 1968. ) The tendency for females to have a labial stripe posteriorly and the absence of dorsal tubercles in females has resulted in the iden- tification of many such specimens as O. leprieurii. Ontogenetic change in coloration is slight in taurinus. Most juveniles (less than 40 mm in snout-vent length) can be identified readily. There is a tendency for the dorsal markings of juveniles to consist of several small spots. Apparently with growth the spots usually coalesce, forming a large median blotch, but some adults retain the juvenile pattern. Cochran and Goin (1970:251) erroneously identified several juveniles from Colombia as Hyla palpebrogranulata Andersson. HYLID FROGS, GENUS OSTEOCEPHALUS 37 Osteocephalus verrucigerus (Werner) Hyla verrucigera Werner, 1901:601 [Holotype—ZMB 16589 from “Ecuador”; Richard Haensch collector]. Hyla riopastazae Andersson, 1945:72 [Holotype—NHRM 1960 from Banos, Rio Pastaza, Provincia Tungurahua, Ecuador; William Clarke-MaclIntyre collector]. Hyla orcesi Funkhouser, 1956:78 [Holotype—CAS-SU 13150 from Rio Pacayacu, tributary of Rio Cotapino, Provincia Napo, Ecuador; collector unknown]. Osteocephalus orcesi—Cochran and Goin, 1970:317. Osteocephalus verrucigerus—Trueb and Duellman, 1970:601 [Synonymized Hyla riopastazae Andersson, 1945, and Hyla orcesi Funkhouser, 1956, with Hyla verrucigera Werner, 1901]. Justification of Synonymy.—Trueb and Duellman (1970:605) discussed the assignment of the names in the synonymy of O. verrucigerus; only a brief resumé is given here. The extant type of Hyla verrucigera is a juvenile male having a snout-vent length of 32.0 mm. The dorsum is smooth except for tubercles on the eyelids; the skin is loose, and the body is soft. The specimen is faded to a pale brown; indistinct dark spots are present on the back, and transverse bars are evident on the limbs. The holotype of Hyla riopastazae is a gravid female having a snout-vent length of 64.7 mm. The dorsum is smooth. The dorsal ground color is pale brown with indistinct brown transverse bars on the limbs. The throat, chest, and belly are cream with brown spots and mottling. The holotype of Hyla orcesi is an adult male having a snout-vent length of 52.6 mm. The dorsum is heavily tuberculate. The dorsum is dark brown with faint transverse bars on the forearms and feet; the ventral surfaces are creamy brown. Trueb and Duellman (1970) provided conclusive evidence that the types of H. verrucigera, riopastazae, and orcesi are a juvenile, adult female, and adult male, respectively, of one species, the earliest available name for which is Hyla verrucigera Werner, 1901. Diagnosis.—1) Size moderate, sexual dimorphism evident; max- imum observed snout-vent length in males 54.3 mm, in females 65.8 mm; 2) skin on dorsum in males bearing large, keratinized tuber- cles; 3) skin on flanks smooth; 4) web extending to base of ante- penultimate phalange on inner edge of third finger; 5) dorsum uni- formly dark brown or black, with tan snout in females; 6) venter creamy white, heavily mottled with black or dark brown, especially in females; 7) lips marked with pale tan labial stripe and suborbital bar; 8) flanks dull reddish brown; 9) dermal roofing bones of skull lacking exostosis; 10) dermal sphenethmoid absent; 11) nasals 38 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY widely separated medially; 12) anteromedial margin of fronto- parietals at anterior border of orbit; 13) frontoparietal fontanelle covered; 14) palatine serrate; 15) parasphenoid bearing odontoids; 16) zygomatic ramus of squamosal extending approximately one- half of distance to maxillary arch; 17) transverse processes of third presacral vertebra approximately equal in width to sacral dia- pophyses; transverse processes of presacral vertebrae 3-8 subequal in width; 18) intermandibularis and submentalis muscles connected; 19) supramandibular portion of interhyoideus forming simple, tubular, posterolateral extension; associated skin unmodified. Osteocephalus verrucigerus can be distinguished from other members of the genus by its uniformly dark dorsum, heavily mottled venter, and large, spinous tubercles on the dorsum in males. Distribution._Lower Amazonian slopes (500-1840 m) of the Andes and on the western fringe of the Amazon Basin in Ecuador and Peru; one locality (Acevedo) in upper Rio Magdalena drainage in Colombia ( Fig. 9). 40 specimens from 13 localities. Remarks.—In life the dorsum in males is dull olive-green; the groin, anterior and posterior surfaces of the thighs, inner surfaces of limbs, and upper arms are dark brown. The ventral surfaces of the limbs are pinkish tan; the other ventral surfaces are pale creamy tan with reddish brown spots. The suborbital spot is pale greenish tan, and the iris is deep reddish brown. In females the dorsum is dull olive-brown; the anterior part of the head is tan, and the sub- orbital spot is yellowish tan. The groin and hidden surfaces of the limbs are dark reddish brown. The ventral surfaces of the limbs are brown; the throat and chest are creamy white, and the belly is reddish tan, both with dark brown mottling. Considerable ontogenetic change occurs in coloration. Juveniles are pale above with a dark median dorsal blotch and dark transverse bars on the limbs. The venter is white. The change consists prin- cipally of an increase in dark pigment and subsequent obliteration of the juvenile pattern. Tadpoles of this species have moderately long tails with low fins, robust bodies, two rows of labial papillae with median part of the upper lip bare, and two upper and five lower rows of teeth. Trueb and Duellman (1970) described the eggs, tadpoles, mating call, and variation in the adults. GENERIC RELATIONSHIPS Among the 33 genera currently recognized in the family Hylidae, there are two basic types of vocal sac structure (Duellman, 1970b), HYLID FROGS, GENUS OSTEOCEPHALUS 39 namely the subgular type and the lateral type. Only four hylid genera, all Neotropical lowland groups, are known to possess paired lateral vocal sacs; these are Osteocephalus, Argenteohyla, Phry- nohyas, and Trachycephalus. The geographical distributions and morphological characteristics of these four genera suggest that they are more closely related to one another than with any other hylid genera. Of the four genera, Osteocephalus is the most generalized in morphology, and, like Phrynohyas, has no specialized habits. Os- teocephalus and Argenteohyla are similarly distinguished from Phrynohyas and Trachycephalus on the basis of vocal sac structure. The vocal sacs of Osteocephalus and Argenteohyla are posterior and protrude posterolateral to the angles of the jaws when they are inflated, whereas those of Phrynohyas and Trachycephalus are more lateral and protrude posterior to the angles of the jaws when inflated. Although Osteocephalus and Argenteohyla have similar vocal sac structure, they are obviously distinct. The monotypic Argenteohyla is a rather specialized, semifossorial frog (Trueb, 1970b), charac- terized by smooth skin, moderate-sized digital discs, and a large inner metatarsal tubercle. The general architecture of the skull is not unlike that of Osteocephalus; the skulls of both are well roofed, broader than long, and characterized by posterolaterally oriented parasphenoid alae. Argenteohyla bears small, slightly curved pre- vomerine dentigerous processes in contrast to the large, angular processes of Osteocephalus. The skull of Argenteohyla shows spe- cializations, apparently adaptations to its semifossorial mode of existence, which further distinguish the genus from Osteocephalus. In comparison with Osteocephalus, the cranium of Argenteohyla is slightly depressed anteriorly, the roofing bones extensively casqued, and the palatines robust. Osteologically, Osteocephalus more closely resembles Phry- nohyas than either of the other two genera, but Osteocephalus and Phrynohyas are clearly distinct on the basis of their respective vocal sac structure. Like Osteocephalus, skulls of the members of the genus Phrynohyas are broader than long, have extensive dermal roofing bones, and have posterolaterally oriented parasphenoid alae. In contrast to Osteocephalus, the dentigerous processes of the prevomers are curved, rather than angular in Phrynohyas. Further- more, the latter genus is singularly distinguished from Osteocepha- lus, Argenteohyla, and Trachycephalus by having extensively de- 40 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY veloped parotoid glands that produce a viscous, milky volatile secretion. Trachycephalus is the most readily identifiable of the four genera under discussion. Members of this genus are large frogs with heavily casqued and co-ossified skulls (Trueb, 1970a). The dermal roofing bones bear omate and characteristic patterns of sculpturing. The medial ramus of the pterygoid does not articulate with the otic capsule, and the parasphenoid alae are laterally, rather than pos- terolaterally, oriented. A dermal sphenethmoid is present, and the parasphenoid bears odontoids. The basic structure of the skull has many characters in common with both Osteocephalus and Phry- nohyas. The obvious modifications of dermal roofing bones and of palatal and suspensory elements seem to be specializations adapting members of the genus Trachycephalus to their peculiar phragmotic habits. The vocal sac structure of Trachycephalus is like that of Phrynohyas and therefore further distinguishes it from Osteo- cephalus. Morphologically, Osteocephalus seems to be sufficiently diverse and generalized so as to represent a modern derivative of an an- cestral type which might have given rise to Phrynohyas, Trachy- cephalus, and Argenteohyla. The specialized vocal sac structure in Phrynohyas and Trachycephalus suggests that these two genera may be rather closely allied and represent a single phyletic line from an ancestral stock similar to Osteocephalus. Argenteohyla is quite distinct from Phrynohyas and Trachycephalus and apparently represents a distinct phyletic line from the ancestral stock. OCCURRENCE OF OSTEOCEPHALUS IN AMAZONIAN ECUADOR All of our observations on members of this genus have been made at four localities: 1) Santa Cecilia at an elevation of 340 meters on the Rio Aguarico, a tributary of the Rio Napo, 2) Lago Agrio, 330 meters, about 14 kilometers east of Santa Cecilia, 3) Puerto Libre, 570 meters, on the Rio Aguarico just east of its forma- tion by the confluence of the Rio Cofanes and Rio Chingua, and 4) south slope of the Cordillera del Dué, above the Rio Coca, 1150 meters. Osteocephalus leprieurii was found at all four localities, and buckleyi was found at all but the last; taurinus was found at Santa Cecilia and Lago Agrio, and verrucigerus was found only in the Cordillera del Dué. Our data are based on collections of 113 frogs and three lots of tadpoles, as well as observations on calling sites and young. The observations are summarized by species, as follows: HYLID FROGS, GENUS OSTEOCEPHALUS A] Osteocephalus buckleyi._No breeding activity was observed. Males were found only at night in March, June, and July. One was perched on a Heliconia leaf in a swamp at Puerto Libre, and two were on bushes in the forest at Santa Cecilia. A gravid female was found on a recently felled tree at Lago Agrio on the night of 12 May 1969. Osteocephalus leprieurii._Males were heard calling sporadically at Puerto Libre in July 1968, and at Santa Cecilia in May 1969. A small chorus was found on the night of 12 May 1969 at Lago Agrio, where the frogs were perched on branches of fallen trees over a temporary pool. The call is a soft rattling chuckle. In late April and May many gravid females and males with well-developed nuptial excrescences were obtained from trees as they were felled at Lago Agrio. The reproductive condition of the frogs indicates that they probably breed in May. One individual called nearly every night from a large tree at Puerto Libre between 4-17 July 1968. The tree was felled on the latter date, but no frog was found. Two nights later apparently the same individual called from a bromeiad at a height of about 10 m on a large bamboo adjacent to the felled tree; the frog was collected when the bamboo was cut down. Throughout the rainy months that we have worked in Ecuador (April-August) we have found occasional individuals perched on bushes or low trees at night. Large numbers of adults were ob- served only during a clearing operation which resulted in the felling of many large trees. Thus, it seems likely that leprieurii is a tree-top inhabitant. A partially digested adult male was removed from the stomach of a Hemiphractus proboscideus. At Santa Cecilia many recently metamorphosed young and ju- veniles were found in June and July 1968. Most of these were on low bushes or herbs in swamp forest at night; some were found in unfolded Heliconia leaves by day, and one was observed on the forest floor by day. Snout-vent lengths of 18 specimens are 12.3-17.0 (mean 15.1) mm. The smaller frogs were recently metamorphosed as evidenced by the melanophore deposits above the vent. The coloration of the young is strikingly different from that of the adults (see account of O. leprieurii), so the association of the young and adults was not made until individuals with intermediate patterns were obtained at Lago Agrio in May 1969. Probably juveniles ob- tained in June and July are the offspring of an April or May breed- ing. We have been unable to associate tadpoles with this species. Osteocephalus taurinus.—A small chorus occurred at Lago Agrio 42 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY on 12 May 1969. Males were calling from the ground adjacent to a small pool amidst recently felled trees. The males were very wary and, when approached, jumped onto limbs and ran up branches; this behavior was noted by Bokermann (1964). The call consists of a series of low-pitched, short notes—like a slow trill—four to six notes per call group. Call groups are repeated two, three, or four times followed by a lapse of several minutes. Although no amplec- tant pairs were found, several gravid females were collected at Lago Agrio in May, so it can be safely assumed that the species breeds in May. From April through July occasional individuals were observed on bushes and trees at night. During clearing operations at Lago Agrio several individuals were obtained from the tops of trees as they were felled. Osteocephalus_ verrucigerus.—Observations were made in a broad, shallow ravine, in which there was a small stream. On 2-4 August 1968, males were observed calling from low bushes and rocks at the edge of a quiet pool in the stream. The call consists of a series of well-pulsed, low-pitched, guttural notes produced at the rate of 5-10 per minute. One amplectant pair was found at the base of a bush adjacent to the pool on 3 August. Another female was found on a branch of a tree 2 m above the ground and 10 m from the stream. Tadpoles of this species were found in the quiet silt- bottomed pool. SPECIMENS EXAMINED The localities for each of the specimens examined are given in the following paragraphs. The arrangement of the data is as fol- lows: alphabetically by country, state (department or province), and locality; alphabetically by the first letter in the abbreviations for the museums, and numerically after each museum abbreviation. Specimens lacking precise locality data are listed first in the most restricted political unit possible; localities which have not been found on maps or the positions of which are not known to us are given in quotation marks. Where more than one specimen is in- cluded under one museum number, the number of specimens is given in parentheses after the museum number. Unless noted other- wise, all specimens are alcoholics. Osteocephalus buckleyi BOLIVIA: El Beni: Ivén, BMNH 1967.2070-1. Santa Cruz: Buenavista, CM 4333, 4339, UMMZ 66563-5. BRASIL: Amapd: No specific locality, WCAB 13284. COLOMBIA: Amazonas: Rio Guacaya, USNM 152759. Huila: Acevedo, HYLID FROGS, GENUS OSTEOCEPHALUS 43 Rio Suaza, FMNH 69702. Narifio: Rumiyacu, FMNH 54756. Meta: Rio Guejar, Campamento La Macarena, USNM 152199. ECUADOR: No specific locality, NHMW 6209, WCAB 35499. Chimbo- razo: Pallatanga, BMNH 1947.2.13.46; Santiago, FMNH 42529. Morona- Santiago: “Rio Santiago” (= Rio Zamora), MIZS 2950. Napo: Lago Agrio, KU 126646; Puerto Libre, Rio Aguarico, KU 123172; Santa Cecilia, AUM 8138, KU: 105208-9, 109506, 123171. Pastaza: Alpayacu, BMNH 1912.11.1.64; Canelos, BMNH 1947.2.13.40-1, 1947.2.13.43-5; Colonia Mena, Rio Conambo, ZSM 33/1962; Don Tomas, USNM 166014; Guaché, Rio Pastaza, AMNH 79986; Rio Bobonaza, USNM 166005; Rio Capahuari, USNM 166554; Rio Conambo at Rio Shiona-yacu, USNM 166018; Rio Copataza, upper Rio Pastaza, USNM 166007-13; Rio Pastaza, NHRM 1946; Rio Pucyacu, USNM 165997 (skeleton), 165998-6001; Rio Rutuno, USNM 166006; Rio Villano, USNM 166002-4; Sarayacu, BMNH 1947.2.13.36-9, MCZ 26090, ZMB 10166. GUYANA: Mazaruni-Potaro: Kartabo, AMNH 70971; Membaru River, upper Mazaruni River, UMMZ 85168; Oko Mountains, FMNH 26722-3. North West: Amakura River, Haulover, UMMZ 83558-9. Rupununi: Marudi River, AMNH 46233; Shudi-kar-wau, AMNH 49252. West Demerara: Dunoon, UMMZ 52449, 52508. PERU: Junin: Chanchamayo, BMNH 1911.12.13.79-80. Loreto: Andoas, AMNH 79984-5; Cashiboya, AMNH 43454; San Antonio, Rio Itaya, AMNH 43218. Puno: Yahuaramayo, BMNH 1913.2.25.7. SURINAM: Suriname: Powakka, CM 44217. SOUTH AMERICA: No specific locality, NHMW 6208. Osteocephalus leprieurii BRASIL: Acre: Tarauaca, FMNH 83247. Amazonas: Rio Javari, Benja- min Constant, CAS-SU 12620; Rio Uaupés, north of Rio Japu, NHMG 489. COLOMBIA: Amazonas: Gino-goje, lower Rio Apoporis, MCZ 28038, 28040-2, 28044, USNM 152136-8. ECUADOR: No specific locality, WCAB 35452-3; “Napo-Pastaza,’ USNM 166571. Napo: Avila, UMMZ 92093; south slope Cordillera del Dué, KU 123170; Lago Agrio, KU 125961-2 (skeletons), 126611-44, UMMZ 129326 (2); Limén Cocha, Rio Napo, KU 99210-6, UIMNH 63087-9, 63098, 63106-9, 63118-9, 64802-4, 64858, 87998-9, 88001-30, 88437-8, 88580, 88604-5, 89852-97, 89999-90000; Loreto, CAS-SU 11439, WCAB 36526; Puerto Libre, Rio Aguarico, KU 123190-1; Puerto Napo, UIMNH 55818-20; Rio Cotapino, UMMZ 92094; Rio Napo, UMMZ 92078; Santa Cecilia, AUM 8099, 8102, 8113-5, 8127-9, 8131, 8137, 8139-46, 8148, KU 105210-20, 109509-11, 111971, 122964-87, 123169, 126645. Pastaza: Canelos, BMNH 1947.2.13.42, KU 120915; Rio Alpayacu, UMMZ 92079; Rio Arajuno, USNM 166560-2, WCAB 40176; Rio Oglan, USNM 16655203, 166558; Rio Rutuno, USNM 166559; Rio Shilcayacu, below Puyo, USNM 166557; Rio Villano, USNM 166551. FRENCH GUIANA: No specific locality, MNHN 4629. Inini: Lunier River, MNHN 98/217. GUYANA: Mazaruni-Potaro: Kartabo, AMNH 70967-8, 70972, 70976. Rupununi: Shudi-kar-wau, AMNH 49255. West Demerara: Demerara Falls, BMNH 72.10.16.23, 72.10.16.37-8. PERU: Loreto: Estirén, Rio Ampiyacu, MZUSP 31033-4; Pebas, CAS-SU 3158, 3160; Roaboya, AMNH 43064. 44 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY SURINAM: No specific locality, MCZ 2036, RMNH 11468. Marowijne: Camp 3, RMNH 13045-6; Wane Creek North, RMNH 11469-70. Saramacca: Right Coppename River, RMNH 11467. Osteocephalus pearsoni BOLIVIA: El Beni: upper Rio Beni, below mouth of Rio Mapiri, MCZ 15565, UMMZ 57548, 67464-5; Rurrenbaque, UMMZ 57533. PERU: Pasco: Yaupi, KU 136312. Osteocephalus taurinus BOLIVIA: El Beni: Ivén, BMNH 1967.2040; Reyes, UMMZ 57532. La Paz: San Ernesto, Mapiri District, BMNH 1901.8.2.54. Santa Cruz: Buena- vista, AMNH 33951-2, 33958, BMNH 1927.8.1.19, 1927.8.1.118, FMNH 27091, UMMZ 63319-21, 63959(2), 63961(2), 66566(2), 66567, 66568(2), 66569 (2), 66570, 66571(2), 66575-6, 68196; Rio Mamore, 2 km N Boca Chaparé, AMNH 79324; Sara, CM 3840-1; Surutu, CM 3814-5. BRASIL: No specific locality: “Interior,” BMNH 74.7.16.8-9. Acre: Placido de Castro, MZUSP 6518; Tarauaca, WCAB 2496. Amazonas: Cucui, NHMW 16495; Manacapurti, ZMB 28492, ZSM 278/1925; Manaus, MCZ 56281, NHMW 16492; Maués, AMNH 69623, 76177; Taracua, NHMG 488, WCAB 18463-4. Mato Grosso: Mabuca, MZUSP 4272; Posto Coluene, Rio Xingu, WCAB 812; “Puerto Cabello,” AMNH 3154; Tapirapé, AMNH 73647- 62, CAS-SU 12351-6, MNHN 46/324. Pard: No specific locality, MPEG 623-6; Belém, KU 129866; Cachimbo, FMNH 175876, UIMNH 42149, WCAB 813; Cameta, NHMW 15892; Gurupa, BMNH 96.6.29.13; Ilha de Marajé, BMNH 1923.11.9.20-4; Ilha Mexicana, ZSM 111/1911, 112/1912; “Ponto Dois Indios,” BMNH 1939.1.5.5; Santarém, BMNH 75.10.22.1-4, MCZ 354. Rondonia: Abunda, CAS 49773-4, FMNH 64239; Forte Principe da Beira, WCAB 10230; Igarapé Marmelo, FMNH 140254, KU 84725 (skeleton), 92243-6, 92247-8 (skeletons), WCAB 9997, 10001, 10003-4; Porto Velho, MZUSP 16343. COLOMBIA: Amazonas: Gino-goje, lower Rio Apoporis, USNM 152139; Leticia, USNM 152010-1; Raudal de la Playa, lower Rio Apoporis, MCZ 28050; Rio Apoporis, MCZ 28060. Boyacd: Sutatenza, USNM 152054-6. Cundinamarca: Medina, MCZ 16269-71, USNM 152089-90, 152092-7, 152757. Meta: El Mico, Rio Guejar, USNM 152203; Rio Duda, Sierra de Macarena, AMNH 79914; Rio Guapaya, Sierra de Macarena, FMNH 81332; Rio Guaviari, Casa de Piedra, UTA No number. Putumayo: Rio Mecaya, FMNH 69711-4, 69716. Vaupés: Gomogoje, lower Rio Apoporis, MCZ 28048. ECUADOR: No specific locality, WCAB 35451, 35785; “Oriente,” UMMZ 90418. Carchi: below Salinas, USNM 166059. Esmeraldas: Carondelet, UIMNH 53560-9; Lagartera, Rio Caoni, UIMNH 53441, 53458-79. Morona- Santiago: Macuma, UIMNH 63142-3, 63145, 63147, 63151, 63154, 63157, USNM 166060. Napo: Avila, UMMZ 92077; Cuyabeno, UIMNH 63158, 90111; Lago Agrio, KU 126647-55; Limon Cocha, Rio Napo, AUM 8132-4, KU 99207-8, 99421-3, 99424 (skeleton), 99425, UIMNH 64801, 87798, 87800, 88032-5, 88576, 90066, 90082, 90102, 90104, 90314, 90984; Loreto, WCAB 35352; Rio Cotapino, UMMZ 92080; Rio Napo, UMMZ 84120; San José Abajo, AMNH 1295, 1449, 22180, 79990; Santa Cecilia, AUM 8117, 8150, KU 105230-3; south slope Volcan Sumaco, USNM 166570. Pastaza: No specific locality, ZSM 31/1956; Arajuno, USNM 165995; Bufeo, lower Rio Bobonaza, USNM 166046-8; Canelos, BMNH 80.12.5.179, 1947.2.13.48, UMMZ 89066; HYLID FROGS, GENUS OSTEOCEPHALUS 45 Don Tomas, Rio Bobonaza, USNM_ 166049-50; Montalvo, CAS-SU 10320, USNM 165987-9, 166058, 166566; 2.5 km SE Puyo, USNM 166051; Rio Arajuno, USNM_ 166043-5; Rio Arajuno (headwaters), USNM 166053; Rio Bobonaza, WCAB 3613-4, 35504; Rio Capahuari, USNM 165990, 166555-6; Rio Capahuari (headwaters), USNM 166057; Rio Conambo, USNM 166569, ZSM 28/1962, 35/1962; Rio Conambo at Rio Ollaguanga, USNM 166568; Rio Conambo:at Rio Shiona-yacu, USNM 166019, 166563-5; Rio Corrientes, USNM 195994, 166020-38, WCAB 3841-2; Rio Huiyo-yacu, Pico de Conambo, USNM 166052; Rio Pastaza, MCZ 19697; Rio Pastaza (drainage), NHRM 1966, USNM 165996; Rio Pindo, USNM 166039-41; Rio Pindo at Rio Tigre (village), USNM 165992-3, 166042; Rio Pucayacu, USNM 166054, 166056; Rio Rutuno, USNM 166055; Rio Solis, upper Rio Bobonaza, WCAB 39914; Rio Villano, USNM 165991, 166567; Sarayacu, BMNH 80.12.5.213, 80.12.5.239-40, MZUSP 323; Shell Mera, KU 99420. Zamora-Chinchipe: “Yani-Inzari,’ AMNH 43259, 43394; Zamora, AMNH 78928. FRENCH GUIANA: Cayenne: Crique Grégoire, UP 40; Maripa, Oyapok River, UP 72; Oyapok River, UZM 1473. Inini: Crique Gabrielle, UP 118-20. GUYANA: No specific locality: RMNH 1873(3), ZMB 3102(2). East Demerara: Atkinson Field, ASU 11622. Mazaruni-Potaro: Chinapora River, upper Potaro River, BMNH 1905.11.1.20-1; Kamakusa, AMNH 21416, 21418-9, 21422; Kartabo, AMNH 11689, 11691, 11697-9, 11703, 11706-8, 23107, 39730, 70966, 70969-70, 70973-5, USNM 118057; Moraballi Creek, Essequibo River, BMNH 1930.10.10.47-51; Oko Mountains, FMNH 26692-705; upper Potaro River, Tung District, BMNH 1905.11.1.40; Rockstone, FMNH 26591. North West: Amakura River, Haulover, UMMZ 83735. Rupununi: north of Acaray River, west of New River, KU 69747-8; Kuyuwini Landing, AMNH 46283; Pakaraima Mountains, BMNH 1933.6.19.49; Shudi-kar-wau, AMNH_ 10665, 39637, 49256(2). West Demerara: Demerara, CAS 54773-4; Demerara Falls, BMNH 72.10.16.16-22, 72.10.16.25-32; Dunoon, MCZ 4834, UMMZ 46736, 52493-4, 52502, 52504-5, 57271; Vryheid, BMNH 78.12.13.18. PERU: Amazonas: Rio Cenepa, AMNH 43400. Huanuco: Monte Alegre, Rio Pachitea, AMNH 43014, 43019. Loreto: Achinamisa, Rio Huallaga, AMNH 42178, 42502; Andoas, Rio Pastaza, AMNH 79991; Cashiboya, AMNH 43388, 43453; Estiroén, Rio Ampiyacu, CAS 93264-74, 93276, 93278-9, 93281, 93283-6, 93289, 93311, 93327; Igarapé Champuia, upper Rio Curiuja, MZUSP 10339; Iquitos, AMNH 42204, 42442, 43468, NHMW 6118; Lago de Mirano, mouth of Rio Napo, AMNH 42712, 43186; Nauta, ANSP 11399; Ollanta, AMNH 42865; Pampa Hermosa, Rio Cushabatay, AMNH 43124, 43146; Pebas, CAS-SU 6375; Pucallpa, MJP 101(2), 140(3); Punga, Rio Tapiche, AMNH 43194; “Rancho de Indiana, Iquitos District,’ MVZ 16890; upper Rio Abujao, AMNH 42908; Rio Itaya, AMNH 42755; upper Rio Pisqui, AMNH 43536; Rio Tapiche at Rio Contaya, AMNH 42983; Rio Utoquinia at Brasilian frontier, AMNH 43137; Sobral, Rio Tamaya, AMNH 43242; Yurimaguas, BMNH 84.2.18.50. San Martin: Cainarachi, AMNH 42763; Moyobamba, ZSM 19/1914. SURINAM: No specific locality, BMNH 70.3.10.67, NHMW 18433.3. Brokopondo: Afobaka, RMNH 16536; Brownsweg, RMNH 16537; Railway km. 121, RMNH 16534. Marowijne: Djai Creek, RMNH 16513-4; Maroni River, ZMB 8240, 8531; Nassaugebergte, RMNH_ 16517-33; Paloemeu, USNM 159025; Swamp Camp, RMNH 16515. Nickerie: Sipaliwini, RMNH 16538. Saramacca: Left Coppename River, RMNH 16535; Tibiti, RMNH 16516. 46 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Suriname: Berlijn, RMNH 15064; Powakka, CM 44226; Zanderij, CM 50568. VENEZUELA: Amazonas: Cerro Duida, UPR-M 2875; Cerro Marahuaca, UPR-M 114-5; Esmeralda, AMNH 23174; Iniridi, SMF 2640; La Culebra, MCZ 28572, UPR-M 117; Laguna, between Tama Tama and Esmeralda, UPR-M 2760; Rio Pescado, AMNH 23177; Tapara, UPR-M 113. Distrito Federal: Caracas, BMNH 51.7.17.182. Osteocephalus verrucigerus COLOMBIA: Huila: Acevedo, Rio Suaza, FMNH 69709-10. ECUADOR: No specific locality, ZMB 16589. Napo: Avila, UMMZ 90413; south slope Cordillera del Dué, KU 123176-88, 123189 (skeleton), 124208 (eggs), 124209-11 (tadpoles); L’Alegria, USNM 167472-3; Rio Pacayacu, tributary of Rio Cotapino, CAS-SU 13150; southeast slope Volcan Sumaco, CAS-SU 11442. Pastaza: Abitagua, CAS-SU 5067, FMNH 25791, 27619, UMMZ 90414, 92092; Alpayaca, Rio Pastaza, BMNH 1912.11.1.64; Mera, UMMZ 90412(4). Tungurahua: Banos, NHRM 1960. PERU: Ayacucho: La Mar, Sivia, Rio Apurimac, FMNH 39853. Huanuco: Rio Pachitea, midway between Puerto Victoria and Puerto Inca, CAS-SU 17745. Junin: Satipo, MJP 38. LITERATURE CITED ANDERSSON, L. G. 1945. Batrachians from east Ecuador collected 1937, 1938 by Wm. Clarke- MacIntyre and Rolf Blomberg. Arkiv Zool., 37A(2):1-88. BOKERMANN, W. C. A. 1964. Field observations on the hylid frog Osteocephalus taurinus Fitz. Herpetologica, 20:252-255. 1966. Lista anotada das localidades tipo de anfibios Brasileiros. S40 Paulo, 183 pp. BOULENGER, G. A. 1882. Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British Museum, ed. 2, London, xvi+503 pp. Cocuran, D. M. and C. J. Goin 1970. Frogs of Colombia. Bull. U.S. Natl. Mus., 288:xii+-655 pp. CopE, E. D. 1867. On the families of the raniform Anura. Jour. Acad. Nat. Sci. Philadelphia, 2: 189-206. 1874. On some Batrachia and Nematognathi brought from the upper Amazon by Prof. Orton. Proc. Acad. Nat. Sci. Philadelphia, 25:120-137. DUELLMAN, W. E. 1970a. Identity of the South American hylid frog Garbeana garbei. Copeia, (3):534-538. 1970b. The hylid frogs of Middle America. Monog. Mus. Nat. Hist., Univ. Kansas, 1:xi++753 pp. DuMERIL, A. M. C. and G. Brsron 1841. Erpétologie générale ou histoire naturelle compléte des reptiles, vol. 8. Paris, 792 pp. FITZINGER, L. 1843. Systema reptilium. Vienna, ix+106 pp. HYLID FROGS, GENUS OSTEOCEPHALUS 47 FUNKHOUSER, J. 1956. New frogs from Ecuador and southwestern Colombia. Zoologica, 91:73-80. GaAIcE, H. T. 1929. Three new tree-frogs from Panama and Bolivia. Occas. Papers Mus. Zool. Univ. Michigan, 207:1-6. Gon, C. J. 1961. Synopsis of the genera of hylid frogs. Ann. Carnegie Mus., 36:5-18. MELIN, D. 1941. Contribution to the knowledge of Amphibia of South America. Goteborgs Kungl. Vetensk.-och Vitterh.-Sam. Handl., Ser. B, 1(4):1-71. Peracca, M. G. 1904. Viaggio del Dr. Enrico Festa nell’ Ecuador e regioni vicine. Reptile ed amfibii. Boll. Mus. Zool. Anat. Comp., Univ. Torino, 19:1-41. STEINDACHNER, F’. 1862. Uber zwei noch unbeschriebene Batrachier. Arch. Zool. Anat. Fisiol., 2:77-82. 1867. Amphibien. Novara Expedition. Zool. Theil, 1, Vienna, 70 pp. TRUEB, L. 1970a. The evolutionary relationships of casque-headed treefrogs with co-ossified skulls (family Hylidae). Univ. Kansas Publ. Mus. Nat. Hist., 18:547-716. 1970b. The generic status of Hyla siemersi Mertens. Herpetologica, 26:254-267. TRUEB, L. and W. E. DuELLMAN 1970. The systematic status and life history of Hyla verrucigera Wemer. Copeia (4):601-610. TyYLer, M. 1971. The phylogenetic significance of vocal sac structure in hylid frogs. Univ. Kansas Publ. Mus. Nat. Hist., 19:319-360. WERNER, F. 1901. Ueber Reptilien und Batrachier aus Ecuador und Neu-Guinea. Verh. Zool.-Bot. Gesell. Wien, 50:593-614. UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly pub- lished in the above series are now published as Occasional Papers, Museum of Natural History. The Miscellaneous Pub- lications, Museum of Natural History, began with number 1 in 1946. Longer research papers are published in that series. Monographs of the Museum of Natural History were initiated in 1970. Institutional libraries interested in exchanging publications may obtain the Occasional Papers and Miscellaneous Publica- tions by addressing the Exchange Librarian, University of Kan- sas Library, Lawrence, Kansas 66044. Individuals may pur- chase separate numbers of all series. Prices may be obtained upon request addressed to Publications Secretary, Museum of Natural History, University of Kansas, Lawrence, Kansas 66044. OCCASIONAL PAPERS of the tt OF SU MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas NUMBER 2 APRIL 29, 1971 NOTEWORTHY RECORDS OF BATS FROM NICARAGUA, WITH A CHECKLIST OF THE CHIROPTERAN FAUNA OF THE COUNTRY By J. Knox Jones, Jr.,° James Date SmITH,” RONALD W. TURNER’ Nicaragua occupies a strategic position in Central America with respect to mammalian distributional patterns, but relatively little has been published concerning the fauna of the country and its zoogeographic relationships. The present paper records informa- tion on distribution, variation, and natural history of 40 species of bats from Nicaragua, 14 of which are here recorded for the first time from the country. Appended is a checklist of the chiropteran fauna of Nicaragua in which only primary literature with actual reference to specimens from the republic is cited. The specimens upon which this report is based are, with few exceptions, in the collections of the Museum of Natural History of The University of Kansas. Some of our material was obtained in 1956 by J. R. and A. A. Alcorn, field representatives of the Museum and sponsored by the Kansas University Endowment Association; most of the specimens, however, were obtained by field parties of which we were members that worked in Nicaragua in 1964, 1966, 1967, and 1968 under the aegis of a contract (DA-49-193-MD-2215) * Curator, Division of Mammals, Museum of Natural History, University of Kansas. * Assistant Professor, Department of Biology, California State College, Fullerton, California. * Assistant Professor, Department of Biology, St. Benedicts College, Atchison, Kansas. 2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY O 20 60 Kilometers Es es Fic. 1—Map of Nicaragua showing location of place-names associated with specimens reported in this paper. Localities, identified by number, are as follows: 1, Potosi; 2, Cosigitina; 3, Hda. Bellavista, Volcan Casita; 4, Chinandega; 5, San Antonio; 6, Jalapa; 7, Condega; 8, Yali; 9, Santa Maria de Ostuma; 10, San Ramon; 11, Matagalpa; 12, Dario; 13, Esquipulas; 14, Santa Rosa; 15, Boaco; 16, Teustepe; 17, Tipitapa; 18, Sabana Grande; 19, Managua; 20, Cuapa; 21, Villa Somoza; 22, Hato Grande; 23, Diriamba; 24, Guanacaste; 25, Mecatepe; 26, Nandaime; 27, Alta Gracia, Isla de Ometepe; 28, Mérida, Isla de Ometepe; 29, Rivas; 30, San Juan del Sur; 31, Sapoa; 32, Bonanza; 33, El Recreo; 34, Cara de Mono. between the U.S. Army Medical Research and Development Com- mand and The University of Kansas. Place-names associated with localities mentioned in the text from which specimens at Kansas were collected are plotted on Fig. 1. In the accounts that follow, departments in Nicaragua are listed alphabetically, but localities within each department are arranged from north to south; elevations are given in meters or feet, depend- ing on which was used on specimen labels. All specimens are in BATS FROM NICARAGUA 3 the Museum of Natural History of The University of Kansas unless noted otherwise. We are indebted to Drs. Charles O. Handley, Jr., and Ronald Pine of the U.S. National Museum (USNM ) for lending us certain critical specimens. ACCOUNTS OF SPECIES Saccopteryx leptura (Schreber, 1774) Two specimens from El Paraiso, 1 km N Cosigiiina, 20 m, Chinandega, on the Cosigiiina Peninsula, provide the fourth locality of record for this white-lined bat in Nicaragua. Jones (1964a:506 ) and Davis et al. (1964:375) earlier reported a total of eight speci- mens from the departments of Managua and Zelaya. The species is known as far north in Middle America as Chiapas (Carter et al., 1966:489 ). Our two bats, both females, were shot on the evening of 1 March 1968 as they foraged around a yard light. One carried an embryo that measured 8 mm (crown-rump), whereas the other was re- productively inactive. Peropteryx macrotis macrotis (Wagner, 1843) Four females (one young and three adult) captured 5 km N and 9 km E Condega, 800 m, in Madriz, on 23 June 1964, provide the first record of this small sac-winged species from Nicaragua. The bats were shot from daytime roosts in small, well-lighted, cave- like spaces formed among immense blocks of granite in a small patch of tropical deciduous forest surrounded by extensive pine- oak woodland. None of the adult females was reproductively active. Glossophaga soricina, Diphylla ecaudata, and a large nursery colony of Desmodus rotundus were found in association with the Perop- teryx. Measurements of our specimens agree closely with those re- ported for material from El Salvador (Felten, 1955:284) and Costa Rica (Starrett and Casebeer, 1968:3-4). Noctilio labialis labialis (Kerr, 1792) Specimens.—Boaco: 4 km W Teustepe, 140 m, 9. Chontales: Hato Grande, 13 km S, 8 km W Juigalpa, 60 m, 49. Rivas: 4 km S, 1.5 km E Alta Gracia, 40 m, Isla de Ometepe, 1; Finca Amayo, 13 km S, 14 km E Rivas, 40 m, 4. Zelaya: S side Rio Mico, El] Recreo, 25 m, 1; Cara de Mono, 50 m, 2. This species has been reported previously from Nicaragua by several authors. All our specimens were netted over small streams or shot as they foraged; parts of scarabids and lepidopterans were found in the mouths of several individuals shot at Finca Amayo. Twenty-six of 31 autopsied females taken in April were pregnant, 4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY each containing a single embryo—average crown-rump length 16.7 (5-26) mm. Testes of 15 males collected in April had an average length of 4.6 (2-7) mm, those of four taken in June, 5.2 (4-6) mm. We follow Cabrera (1958:55), Husson (1962:63), and Handley (1966b:758 ) in use of the subspecific name labialis, the type locality of which is the “Mosquito shore” of Nicaragua, rather than Peru as suggested by Hershkovitz (1949:433-434). Noctilio leporinus mexicanus Goldman, 1915 Specimens.—Chinandega: Potosi, 5 m, 2. Chontales: Hato Grande, 13 km S, 8 km W Juigalpa, 60 m, 4. Rivas: 4 km S, 1.5 km E Alta Gracia, 40 m, Isla de Ometepe, 4; Mérida, 40 m, Isla de Ometepe, 2; Finca Amayo, 13 km S, 14 km E Rivas, 40 m, 1. This fish-eating species, first reported from Nicaragua by Davis et al. (1964:376), apparently occurs throughout Middle America, al- though known from the region by comparatively few records. We have 13 additional Nicaraguan specimens as listed above. The two individuals (both females, one pregnant with an em- bryo that measured 20 mm) from Potosi were caught on 6 March in a mist net set over a large pool in a shallow estuarine stream; the mouth of the stream opened into the Gulf of Fonseca approximately 200 yards below our netting site. Other individuals of this species were observed as they foraged over large pools formed at high tide near the mouth of the stream. Our other specimens were caught in mist nets set over fresh water streams near Lake Nicaragua or along the shores of the lake. A female from near Alta Gracia, caught on 27 March, carried a single embryo that measured 41 mm, whereas one from Finca Amayo was lactating on 25 June. Four males taken on Isla de Ometepe in late March and early April had a mean testicular length of 9.5 (8-10) mm. Pteronotus davyi fulvus (Thomas, 1892) Specimens.—Chontales: Cuapa, 4. Matagalpa: 3 mi E San Ramon, 126. This small naked-backed bat has not been reported previously from Nicaragua. Autopsy of seven females collected on 9 May near San Ramon revealed that four were pregnant, each with a single embryo—average crown-rump length 25.0 (21-29) mm. The distribution of P. davyi is poorly known in much of Central America. The species was reported only recently from Costa Rica (Starrett and Casebeer, 1968:8) and is unknown from Panama. We assign our specimens tentatively to the subspecies fulvus on geographic grounds. BATS FROM NICARAGUA 5 Pteronotus parnellii fuscus (J. A. Allen, 1911) Specimens.—Boaco: Santa Rosa, 17 km N, 15 km E Boaco, 300 m, 1; Los Cocos, 14 km S Boaco, 220 m, 1. Chinandega: 6.5 km N, 1 km E Cosigiiina, 10 m, 1. Zelaya: Bonanza, 850 ft, 1; 2 mi SW Bonanza, 600 ft, 1; S side Rio Mico, El Recreo, 25 m, 1; Cara de Mono, 50 m, 4. Although this species is widespread in Middle America, it has been known previously from Nicaragua only by a specimen from “Chontales” ( Miller, 1902:402). All of our specimens were captured in mist nets. Two adult females (one taken on 28 February and the other on 8 March) each carried a single embryo (13 and 18 mm in crown-rump length, respectively ); a female taken in April was lactating and had a flaccid uterus suggesting relatively recent par- turition. Adult females captured in the months of June, July, and August evidenced no sign of reproductive activity. Two of these were in dark, fresh pelage, but one captured on 24 June and another on 28 July were molting. In both individuals, active molt evidently had slowed or subsided and remnants of the old pelage (rich ochraceous orange) were confined to a narrow strip at the lateral edges of the body. We tentatively apply the subspecies name fuscus to our Nicara- guan bats on geographic grounds; one of us (Smith) currently is investigating geographic variation in the genus Pteronotus. Pteronotus suapurensis (J. A. Allen, 1904) Specimens.—Chontales: Cuapa, 1. Matagalpa: 3 mi E San Ramon, 24. Zelaya: Cara de Mono, 25 m, 4. This relatively poorly known monotypic species occurs from southern Veracruz to the Amazon Basin. It has been reported from several localities in Central America including one in Nicaragua (Goodwin, 1942a:88). Three pregnant females from near San Ramon (9 May) carried embryos that had crown-rump lengths of 27, 27, and 28 mm, and two netted on 24 April at Cara de Mono each had an embryo that measured 22 mm. All of our specimens are in bright pelage that is fulvous red in color. Micronycteris megalotis mexicana Miller, 1898 Specimens.—Carazo: 3 km N, 4 km W Diriamba, 600 m, 5. Granada: Hda. Mecatepe [2 km N, 11.5 km E Nandaime], 5. Managua: 5 mi NW Managua, 1. Matagalpa: Rio Viejo, 7 mi WNW Dario, 4; 11 mi SE Dario, 1. Zelaya: S end Isla del Maiz Grande, 9. The individuals from Isla de Maiz Grande were shot on 30 June as they flew from small caves and crevices in rocky outcrops on the south end of the island. Of the three adult females obtained at 6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY TaBLeE 1.—Selected measurements of adults of two subspecies of Micronycteris megalotis from Nicaragua. Superscript numbers indicate fewer specimens averaged than indicated in left-hand column. 5 3 oe Ss x) i) S Py > Number of specimens P| Bo Sc Sa ane averaged or tos coe Sts) Se toms = catalogue number, ae oes 80D O'S Bac and sex a BS) Ono. N35 ate) — ace Micronycteris megalotis mexicana, west-central Nicaragua* Average 8 (36, 5@) 35.4 19.5° 95 8.0° (Os Minimum SHG) 18.8 9.1 eu toe Maximum 38.1 20.3 9.8 8.2 te Isla del Maiz Grande, Nicaragua Average 6 (324, 39) 35.6 19.4 9.2 7.8 7.4 Minimum 34.5 19.1 9.0 G7) fe | Maximum 37.0 19.8 9.3 7.9 5 Micronycteris megalotis microtis, Greytown, Nicaragua (holotype) USNM 16366/23364, ¢ ol .5 18.2 8.8 1.6 6.9 Rio Coco, Nicaragua (AMNH) Average 6 (34, 39) 33.6 18.9° — Lo Fi Minimum Aa | 18.8 — 7.4 7.0 Maximum 34.2 19.0 a 16 Vz. Bonanza, Nicaragua KU 96251, ¢ 32.3 18.4 8.8 7.6 7.0 * Specimens labeled with reference to Dario, Diriamba, and Managua. that time, two were lactating; each was accompanied by a young, the forearms of which measured 19.7 (male) and 21.3 mm (female). Two adult males from Isla del Maiz Grande had testes 2 mm in length. Four of the five specimens taken northwest of Diriamba were shot from a daytime retreat in a culvert; the fifth was caught by hand as it attempted to fly out of a hollow, fallen tree. Two adult females captured on 31 March were pregnant, each carrying a single embryo (13 and 14 mm in crown-rump length), whereas two obtained on 14 August showed no sign of reproductive activity. An adult male, also taken on 14 August, had testes 2 mm in length. Of three additional adult females, one captured on 3 June was pregnant (embryo 21 mm in crown-rump length), whereas two obtained on 14 April evinced no gross reproductive activity. The subspecies mexicana has not been reported previously from Nicaragua, although Gardner et al. (1970:715) recently extended its known distribution southward from Honduras (Goodwin, 1942c: 124) and El Salvador (Felten, 1956:180) to west-central Costa Rica. In Nicaragua, as apparently in Costa Rica, mexicana occupies BATS FROM NICARAGUA ig the Pacific versant, the Caribbean lowlands being inhabited by the smaller race, Micronycteris megalotis microtis Miller, 1898 (type locality, Greytown, Nicaragua—reported also from “Rio Coco,” Nicaragua, by J. A. Allen, 1910:110). Nicaraguan examples of M. m. mexicana are, on the average, considerably larger in cranial dimen- sions and length of forearm than are specimens of M. m. microtis (see Table 1). The series of microtis from “Rio Coco,” probably from the headwaters of that stream in the vicinity of San Juan de Rio Coco and thus near the divide between Caribbean and Pacific drainages, is intermediate in size between the two subspecies and suggests intergradation between them. As noted also by Gardner et al. (1970:715), we find no differences in length of ear between microtis and mexicana (contrary to Miller’s claim, in the original description, that microtis had noticeably smaller ears), nor do we note any consistent differences in color between the two races on the basis of the specimens at hand. Our specimens from Isla del Maiz Grande inexplicably agree rather closely in size (average but slightly smaller) with specimens of M. m. mexicana from western Nicaragua and elsewhere within the range of the subspecies, and are tentatively, therefore, referred to mexicana. Four specimens earlier reported (G. M. Allen, 1929: 130) from the same island also are relatively large (forearm 39mm). Further commentary on this insular population must await a time when additional material is available from Central America. Tonatia nicaraguae Goodwin, 1942 Specimens.—Boaco: Santa Rosa, 17 km N, 15 km E Boaco, 300 m, 1. Chontales: Hato Grande, 13 km S, 8 km W Juigalpa, 60 m, 1. Only four specimens of this seemingly rare species have been recorded previously from Nicaragua—one (the holotype) from Kanawa Creek, near Cukra, north of Bluefields, 100 ft, Zelaya (Goodwin, 1942b:205), another from 12.5 mi. S and 13 mi. E Rivas, 125 ft, Rivas (Davis and Carter, 1962:67-68), and two from 6 km N Tuma, 500 m, Matagalpa (Valdez and LaVal, 1971:248). Our specimens were caught in mist nets placed over small, quiet streams that were fringed with tall, gallery forest. The surrounding areas were savanna-like with small stands of secondary forest. An adult male taken on 21 April had testes that measured 5 mm in length, whereas those of another (9 August) were 3 mm. Handley (1966b:761) synonymized T. nicaraguae with Tonatia minuta Goodwin, 1942, a conclusion with which we agree. How- ever, the name nicaraguae (Goodwin, 1942b:205) has page priority over minuta Goodwin (op. cit.:206) and is the valid name for the 8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY species rather than minuta as used by Handley (see also LaVal, 1969:820; Gardner e¢ al., 1970:716; Valdez and LaVal, 1971:248). Measurements of the two males (that from Hato Grande listed first) are as follows: total length, 63, 60 mm; length of tail, 5, 6 mm; length of hind foot, 10, 9 mm; length of ear, 23, 23 mm; length of forearm, 34.5, 35.6 mm; weight, 11.4, 8.8 gms; greatest length of skull, 20.2, 20.56 mm; zygomatic breadth, 9.5, 9.5 mm; breadth of braincase, 8.5, 8.2 mm; postorbital breadth, 3.1, 2.9 mm; length of maxillary toothrow, 7.0, 7.1 mm; breadth across upper molars, 6.3, 6.5 mm. We have not compared our specimens directly with others from Middle America. On the basis of available measurements, they resemble material reported from Honduras (LaVal, 1969:820), averaging larger than other specimens for which measurements have been published (see especially Goodwin, 1942b:206; Davis and Carter, 1962:68; Davis et al., 1964:379; Gardner et al., 1970: 716-717). Phyllostomus hastatus panamensis J. A. Allen, 1904 Specimens.—Carazo: 3 km N, 4 km W Diriamba, 600 m, 1. Madriz: Daraili, 5 km N, 14 km E Condega, 940 m, 4. Managua: 3 km SW Tipitapa, 40 m, 1; 3 km N Sabana Grande, 50 m, 2; 2 km N Sabana Grande, 40 m, 1; 5 mi S Managua, 2. Matagalpa: La Danta, 1 km N, 5 km E Esquipulas, 760 m, 1. Nueva Segovia: 4.5 km N, 2 km E Jalapa, 630 m, 2; 1.5 km N, 1 km E Jalapa, 660 m, 1. Zelaya: Bonanza, 850 ft, 2; El Recreo, 25 m, 8. This large spear-nosed species has been reported previously from Nicaragua only from Matagalpa (Goodwin, 1942c:126). Phyllostomus hastatus evidently occurs throughout the country and is relatively common in some places. Two females taken 5 mi S Managua on 13 March each carried a single embryo (crown-rump length 27 and 32 mm). Lactating females were captured in June, July, and August. Trachops cirrhosus coffini Goldman, 1925 A male fringe-lipped bat netted over a small stream at Cara de Mono, 50 m, Zelaya, in the Caribbean lowlands, on 24 April, 1968, constitutes the second known occurrence of this species in Nicara- gua. Carter et al. (1966:491) earlier reported two males from Rio Coco, 64 mi NNE Jinotega, 1000 ft, Jinotega. The testes of our specimen were 4 mm long. Davis and Carter (1962:69), Carter et al. (loc. cit.), and Starrett and Casebeer (1968:11) did not apply a trinomen to bats of this species and noted overlap of measurements between T. c. cirrhosus (Spix) and T. c. coffini Goldman. However, until detailed com- BATS FROM NICARAGUA 9 parisons can be made, we follow Jones (1966:452) in retaining the subspecific name coffini. Chrotopterus auritus auritus (Peters, 1856) An adult male captured in a mist net set in a forest clearing at Santa Maria de Ostuma, 1250 m, Matagalpa, represents the first record of this bat from Nicaragua. The testes of our individual, taken on 1 July 1966, were 6 mm in length. Externally and cranially our Nicaraguan example closely resembles specimens of C. auritus from Veracruz and the Yucatan Peninsula. Handley (1966b:762) and Starrett and Casebeer (1968:12) expressed doubt as to the validity of the currently used subspecific names in this species. Comparisons of cranial and external measurements of the material at hand with those given in various published accounts—Burt and Stirton (1961), Starrett and Casebeer (1968), and Thomas (1905), for example—reveal little variation. Until additional information is available (especially as concerns the South American races), how- ever, we tentatively apply the subspecific name C. a. auritus to Middle American populations. Anoura geoffroyi lasiopyga (Peters, 1868) Four specimens of this species, netted on 24 and 25 July 1967 in a banana grove 1.5 km N and 1 km E Jalapa, 660 m, Nueva Segovia, provide the first record of this glossophagine from Nica- ragua. Two males each had testes 6 mm in length; one female evidenced no reproduction activity, whereas another carried an embryo 4 mm in crown-rump length. In addition to our material, there is a specimen in the British Museum (BM §8.6.22.4) from Cafetal “Concordia,” 4000 ft, Jinotega. It may be noteworthy that the two localities at which this bat is known both are in the highlands of the northern part of Nicaragua, and that we failed to take additional specimens in many hundreds of hours of netting in other places in the country. Choeroniscus godmani (Thomas, 1903) Godman’s bat was reported from Nicaragua by Handley (1966a: 86), who used the locality designation “E] Realejo” for the three specimens available to him. Actually, the three are from the nearby Hda. San Isidro, 10 km S Chinandega. We have taken three addi- tional specimens as follows: Santa Rosa, 17 km N and 15 km E Boaco, 300 m, Boaco (an adult female taken on 21 March, which was pregnant with an embryo measuring 16 mm in crown-rump length); 2 km E Yali, 900 m, Jinotega (an adult male netted on 3 10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY August, testes 4 mm in length); and Santa Maria de Ostuma, 1250 m, Matagalpa (a nonpregnant, adult female captured on 11 April). Bats of this species have been taken as far north as the western Mexican state of Sinaloa (Jones, 1964b:510). Lichonycteris obscura Thomas, 1895 Managua is the type locality of this rare long-nosed species and a specimen was reported from 6 mi W Rama, 50 ft, Zelaya, by Davis et al. (1964:380). Our collection contains three males, one from Jalapa, 660 m, Nueva Segovia, captured on the night of 27 July 1967 as it flew around a lighted room in a house, and two from the south side of the Rio Mico, El Recreo, 25 m, Zelaya, taken on 26 and 27 July 1966 (one was netted and the other caught by hand in the lighted room of a building at night). Selected measurements of the three males are, respectively: total length, 55, 58, 57 mm; length of tail, 7, 10, 9 mm; length of hind foot, 7, 9, 9 mm; length of ear, 10, 11, 11 mm; length of fore- arm, 30.9, 31.0, 30.8 mm; weight, 6.8, 6.8, 6.2 gms; length of testes, 1, 2,2 mm; greatest length of skull, 18.5, 18.0, 17.9 mm; breadth of braincase, 8.1, 8.4, 8.3 mm; length of maxillary toothrow, 5.7, 5.5, o.7 mm. Carollia castanea H. Allen, 1890 Specimens.—Boaco: Santa Rosa, 17 km N, 15 km E Boaco, 300 m, 7. Chontales: 1 km N, 2.5 km W Villa Somoza, 330 m, 4. Matagalpa: 1 km NE Esquipulas, 420 m, 1. Nueva Segovia: 7 km N, 4 km E Jalapa, 660 m, 4. Zelaya: S side Rio Mico, El Recreo, 25 m, 6; Cara de Mono, 50 m, 15. Bats of this species apparently are common in the Caribbean lowlands of Nicaragua; the highest altitude at which we have taken specimens is 660 meters. The only previous report of C. castanea from Nicaragua evidently is that of Davis et al. (1964:379), who mentioned it in passing (from Cacao, Zelaya) in an account of Macrophyllum macrophyllum. Two adult females each carried an embryo having crown-rump lengths of 20 (21 March) and 10 mm (27 July). Adult females evincing no gross reproductive activity were taken in February (one), March (one), April (four), June (three), July (two), and August (two). Four adult males netted on 24 April had testes with an average length of 7.0 (5-9) mm, whereas those of four other adult males taken in late June had an average of 4.0 (2-6) mm. In our collections from Nicaragua, we find at least three kinds of Carollia and we follow Handley (1966b:764-765) in tentatively assigning these to C. castanea, C. subrufa, and C. perspicillata. The BATS FROM NICARAGUA Li systematics of this genus is currently under study by Ronald H. Pine. In Nicaragua, castanea differs from subrufa (with which it has been confused in the past ) in being smaller, both externally and cranially, and much darker in color as well as in the additional characters mentioned by Handley (loc. cit.). Sturnira ludovici ludovici Anthony, 1924 An adult male (testes 6 mm) yellow-shouldered bat from Daraili, 5 km N and 14 km E Condega, 940 m, in Madriz, provides the only specimen thus far reported from Nicaragua. Sturnira ludovici evidently is relatively rare in Nicaragua and may be con- fined to the highlands in the north, whereas the smaller S. Lilium is abundant throughout the country and is the only other species of Sturnira represented in our collections. We provisionally refer our specimen to S. |. ludovici pending Luis de la Torre’s forthcoming review of the genus. Selected measurements are: total length, 77 mm; length of hind foot, 15 mm; length of ear, 19 mm; length of forearm, 44.8 mm; weight, 26.8 gms; greatest length of skull, 24.2 mm; zygomatic breadth, 14.2 mm; postorbital breadth, 6.3 mm; breadth of braincase, 10.5 mm; length of maxillary toothrow, 6.8 mm. Uroderma magnirostrum Davis, 1968 In the original description of U. magnirostrum, Davis (1968: 680) reported one specimen from Nicaragua—a female in our col- lection from 3 km N and 4 km W Sapoa, 40 m, Rivas. We have two additional specimens, both from the relatively dry northwest- em Departamento de Chinandega. On 8 July 1966, a male (testes 2mm) was netted under trees along a small stream at San Antonio, 35 m, and a female (one embryo 28 mm in crown-rump length) was netted on 4 March 1968 over a warm spring at a place 4.5 km N Cosigiiina, 15 m. Judging from published records, this species is limited in Middle America to the Pacific versant. Selected measurements of the three Nicaraguan specimens are, respectively: total length, 65, 64, 67 mm; length of hind foot, 12, 11, 14 mm; length of ear, 16.5, 16, 18 mm; length of forearm, 42.2, 41.7, 45.2 mm; weight, 16.2, 13.8, 21.4 (pregnant) gms; greatest length of skull, 22.7, 23.4, 23.8 mm; zygomatic breadth, 12.4, 12.9, 13.1 mm; postorbital breadth, 5.8, 5.5, 5.9 mm; mastoid breadth, 10.9, 11.1, 11.1 mm; length of maxillary toothrow, 7.9, 8.1, 8.6 mm. Vampyrops helleri Peters, 1866 Specimens.—Boaco: Santa Rosa, 17 km N, 15 km E Boaco, 300 m, 3. Carazo: 3 km N, 4 km W Diriamba, 600 m, 2. Chinandega: Potosi, 5 m, 1; 12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 6.5 km N, 1 km E Cosigiiina, 10 m, 1; 4.5 km N Cosigiiina, 15 m, 3; Hda. Bellavista, 720 m, Volcan Casita, 13. Chontales: 1 km N, 2.5 km W Villa Somoza, 330 m, 4. Granada: Finca Santa Cecilia, 6.5 km SE Guanacaste, 660 m, 4. Matagalpa: Finca Tepeyac, 10.5 km N, 9 km E Matagalpa, 960 m, 1. Nueva Segovia: 4.5 km N, 2 km E Jalapa, 680 m, 2; 1.5 km N, 1 km E Jalapa, 660 m, 2. Rivas: 2 km N, 3 km E Mérida, 200 m, Isla de Ometepe, 4; 4 km N, 4 km W Sapoa, 40 m, 1. Zelaya: S side Rio Mico, El Recreo, 25 m, 3. Only two specimens of this small white-lined species have been reported previously from Nicaragua—one from 1 km NW La Gatiada, 1300 ft, Chontales (Davis et al., 1964:383), and the other from 3 mi NNW Diriamba, Carazo (Jones, 1964a:507). This bat was relatively rare in collections until the last decade or so. Handley (1966b:766) reported V. helleri as abundant in the low- land areas of Panama, and we found the same to be true in Nicara- gua. We suspect that future investigations in Middle America will reveal this species to be a common member of lowland tropical communities. It is known as far north as southern Veracruz (Carter et al., 1966:494). Most of our specimens were captured in mist nets set over small streams bordered by gallery forest, or in banana groves. The range of ecological conditions in which this species was taken is represented by the semiarid environment of the Cosigiiina Peninsula in northwestern Nicaragua and the humid tropical forest (secondary growth) in the vicinity of El Recreo in the Caribbean lowlands. Pregnant females were captured in March, April, June, July, and August, indicating that this species probably breeds throughout much of the year. Vampyrodes major G. M. Allen, 1908 Specimens.—Boaco: Santa Rosa, 17 km N, 15 km E Boaco, 300 m, 8. Chontales: 1 km N, 2.5 km W Villa Somoza, 330 m, 2. Zelaya: S side Rio Mico, 25 m, 6. This large white-lined stenodermine was known previously from Nicaragua by a single specimen from an unknown locality (J. A. Allen, 1910:112). All of our specimens were caught in mist nets, which were set over streams at Santa Rosa and near Villa Somoza, and in a banana grove at El Recreo. Two pregnant females, cap- tured on 11 and 13 July at Santa Rosa, each carried an embryo (4 and 27 mm in crown-rump length, respectively ); one of two other adult females captured there on 9 August also had an embryo (35 mm in length) but the other evidently was reproductively quiescent. Testes of adult males varied in length from 3 to 10 mm on the following dates (testicular lengths in parentheses): 25 February BATS FROM NICARAGUA 13 (10 mm); 21 March (8, 8 mm); 17 June (3, 4mm); 13 July (6 mm); 27-28 July (4, 4 mm); 3 August (4 mm); 5 August (3 mm); 9 August (4mm). We follow Starrett and Casebeer (1968:12) in the use of the specific name major, rather than caraccioli as suggested by Cabrera (1958 ), Goodwin and Greenhall (1961), and Handley (1966b). Vampyressa nymphaea Thomas, 1909 A pregnant female (crown-rump length of embryo 5 mm) was captured in a mist net set in a small banana grove on the south side of the Rio Mico, El] Recreo, 25 m, in the Caribbean lowlands, on 27 February 1968. This specimen provides the first record of the big yellow-eared bat from Nicaragua. The species was recently reported for the first time from Costa Rica (Gardner et al., 1970: 721); it was characterized as uncommon in Panama by Handley (1966b:767). The one Costa Rican locality of record also is in the Caribbean versant. Selected external and cranial measurements of our female are: total length, 58 mm; length of hind foot, 11 mm; length of ear, 16 mm; length of forearm, 36.2 mm; weight, 12.3 gms; greatest length of skull, 21.1 mm; condylobasal length, 18.4 mm; zygomatic breadth, 12.3 mm; mastoid breadth, 10.5 mm; breadth across canines, 4.6 mm; breadth of braincase, 9.4 mm; length of maxillary toothrow, 7.2 mm; length of mandibular c-m3, 7.8 mm. Vampyressa pusilla thyone Thomas, 1909 Specimens.—Boaco: Santa Rosa, 17 km N, 15 km E Boaco, 300 m, 5. Chontales: 1 km N, 2.5 km W Villa Somoza, 330 m, 1. Managua: Hda. San José, 2. Matagalpa: 2 km N, 6 km E Esquipulas, 960 m, 2. The only previous record of occurrence for the small yellow- eared bat from Nicaragua is based on an adult female from Hda. La Cumplida, 670 m, Matagalpa (Starrett and de la Torre, 1964:60). Two individuals taken near Esquipulas in mid-March, a preg- nant female (crown-rump length of embryo 16 mm) and a male (testes 4 mm), were captured in nets set across trails cut through secondary forest. The wind was quite strong in this area at the time of our visit and only a few other species of bats—Glossophaga soricina, Artibeus jamaicensis, A. toltecus, A. phaeotis, Uroderma bilobatum, Sturnira lilium, Centurio senex, and Diphylla ecaudata —were taken in the same nets. The specimens from Boaco and Chontales were captured over small streams bordered by gallery forest. Four females collected at Santa Rosa on 21 March were pregnant; each carried a single embryo that measured 5, 18, 21, and 14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 30 mm in crown-rump length; a male taken on the same date had testes that measured 3 mm. Selected external and cranial measurements of two males, fol- lowed by the average (extremes in parentheses ) of six females are: length of forearm, 31.1, 30.8, 30.8 (30.0-31.4) mm; greatest length of skull, 18.9, 18.9, 18.5 (18.1-18.8) mm; zygomatic breadth, 11.0, 11.0, 10.6 (10.4-10.9) mm; mastoid breadth, 9.5, 9.2, 9.2 (9.0-9.3) mm; length of maxillary toothrow, 6.1, 5.9, 5.9 (5.7-6.1) mm. Chiroderma villosum jesupi J. A. Allen, 1900 Specimens.—Chinandega: 6.5 km N, 1 km E Cosigiiina, 10 m, 2; 4.5 km N Cosigiiina, 15 m, 7; Hda. Bellavista, 720 m, Volcan Casita, 5; San Antonio, 35 m, 2. Rivas: 2 km N, 3 km E Mérida, 200 m, Isla de Ometepe, 1. This species has been reported in Middle America from as far north as southern México. It evidently is uncommon in Costa Rica (see Gardner et al., 1970:722) and Panama (Handley, 1966b:767 ). Our material, all collected from mist nets and consisting of 16 spec- imens from the northwestern department of Chinandega and one from Isla de Ometepe in Lago de Nicaragua, constitutes the first report of this bat from Nicaragua. Four of five females taken in early March were pregnant; em- bryos averaged 26.0 (25-29) mm in crown-rump length. Four females taken in July carried embryos 14, 20, 23, and 25 mm in length. Testes of five adult males captured in March and April had an average length of 4.4 (3-7) mm, whereas those of two taken in July were 3 mm in length. Artibeus toltecus hesperus Davis, 1969 When Davis (1969) named A. t. hesperus, he assigned specimens only from as far south as El Salvador to the new subspecies, re- ferring the three Nicaraguan examples of the species at his disposal to the nominal race. On the night of 6-7 April 1968, one of us (Smith) netted bats on the south part of Isla de Ometepe at a place 2 km N and 3 km E Merida, 200 meters in elevation. One net was set across, and another parallel to, a small, boulder-strewn stream; the surrounding area was planted to coffee and had a good canopy of tall deciduous trees. Among the bats captured at this location were 10 A. toltecus that are referable to the subspecies hesperus, judging by their small size (Table 2). Six of our specimens are females and each carried an embryo (range in crown-rump length 20-28 mm). Three adult males had testes 5, 6, and 7 mm in length. External measurements (extremes in parentheses) of our series are: total length, 55.9 (51-60) mm; BATS FROM NICARAGUA 15 TABLE 2.—Selected measurements of two subspecies of Artibeus toltecus from Nicaragua. S 5 "es ToS fo) > : a8 a2 as ws Sas Number of specimens bo & pelea Stas 33 tome averaged a 2 oes bo 5 a © Rao and sex . aS C5.ona N.S =o Hes Artibeus toltecus toltecus, Departamento de Matagalpa Average 6 (36,39) 40.3 20.3 12.8 10:7 6.6 Minimum 38.8 19.8 11.8 10.5 6.5 Maximum 41.5 20.5 125 10.9 6.8 Artibeus toltecus hesperus, Isla de Ometepe, Rivas Average 10 (46,692 ) 38.0 19.4 11.5 10.2 6.3 Minimum 37.0 18.8 LE 2 9.8 Get Maximum 39.7 19.8 11.8 10.5 G5 length of hind foot, 10.7 (10-12) mm; length of ear, 14.8 (14-16) mm; weight of four males, 9.9 (8.8-11.5) gms; weight of six preg- nant females, 14.9 (12.7-16.9) gms. Artibeus toltecus toltecus (Saussure, 1860) Specimens.—Matagalpa: Santa Maria de Ostuma, 1250 m, 5; 2 km N, 6 km E Esquipulas, 960 m, 1. This bat has been reported from Nicaragua previously by Ander- sen (1908:300) and Davis (1969:28), based on a total of four spec- imens. We netted this species at Santa Maria de Ostuma in patches of cloud forest at a cafetal. The specimen from near Esquipulas was taken in a net placed across a trail in second growth forest. Two females collected on 11 April and one taken on 30 June were preg- nant (embryos 21, 26, and 12 mm, respectively, in crown-rump length). Testes of a male netted on 14 March were 7 mm in length, whereas those of two obtained on 11 April measured 4 and 7 mm. Selected measurements of our six specimens are given in Table 2. Artibeus watsoni Thomas, 1901 Specimens.—Chontales: 1 km N, 1.5 km W Villa Somoza, 330 m, 3. Nueva Segovia: 7 km N, 4 km E Jalapa, 600 m, 1. Zelaya: Bonanza, 850 ft, 6; S side Rio Mico, El Recreo, 25 m, 6; Cara de Mono, 50 m, 1. Davis (1970a:393-394) recently reviewed the systematic status of this small fruit-eating bat and recorded specimens from south- eastern Nicaragua; the species was first reported from the country by Andersen (1908:290), based on a specimen from the Escondido River. Our additional material reveals that A. watsoni occurs throughout eastern Nicaragua, the specimen from Nueva Segovia extending the known range as mapped by Davis (loc. cit.). 16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY A female from Bonanza (23 February ) carried an embryo 14 mm in crown-rump length, whereas one from El Recreo (26 February ) was not reproductively active; one of two females netted near Villa Somoza in early August was pregnant (embryo 21 mm in crown- rump length). Seven adult males collected in late February and early March had an average testicular length of 5.9 (5-7) mm; testes of two adults taken in late June and one captured in early August all measured 5 mm. The testes of young males (grayish pelage, partially unfused phalangeal epiphyses ) were 2 or 3 mm in length. Centurio senex senex Gray, 1842 Specimens.—Chinandega: 4.5 km N Cosigitina, 15 m, 1; San Antonio, 35 m, 5. Matagalpa: 2 km N, 6 km E Esquipulas, 960 m, 3. Nueva Segovia: 7km N, 4 km E Jalapa, 660 m, 1. Zelaya: S side Rio Mico, E] Recreo, 25 m, 1. Paradiso (1967) reviewed geographic variation in this unique bat, the type locality of which was restricted to Realejo, Chinan- dega, Nicaragua, by Goodwin (1946:327). Because additional ma- terial had not been reported from Nicaragua, Paradiso (op. cit.:598 ) felt it was “premature to restrict the type locality to a specific area in that country” (the holotype was obtained on the voyage of the H.M.S. Sulphur, which called at Realejo), and preferred the more general designation “west coast of Mexico or Central America.” In view of the fact that we now have specimens from but a few miles distant from Realejo (at San Antonio), we see no reason to contest Goodwin's restriction of the type locality to that place. Specimens from San Antonio were collected along a small stream, bordered by a bilevel gallery forest, in an area otherwise planted mostly to cane. Many trees of the lower level were covered by an extremely thick network of vines, which were interwoven with branches and supported fallen leaves and debris from the upper level. This situation led to formation of small “rooms” or “cubicles” under some shorter trees; the bats were shot as they hung from small branches under one such tree, which was in fruit. All of our other specimens were captured in mist nets. Pregnant females were taken on the following dates (crown- rump length of embryo in parentheses): 25 February (12 mm), 2 March (17 mm), 15 March (14 mm); a nonpregnant female also was taken on 15 March. Five males captured at San Antonio on 9 and 10 March had an average testicular length of 5.6 (5-6) mm. A male taken in July had testes 4 mm in length, whereas those of one obtained on 14 March were 5 mm long. Selected measurements (average, with extremes in parentheses ) of 11 adults (seven males and four females) are as follows: length BATS FROM NICARAGUA Vi of forearm, 42.5 (41.5-43.7) mm; condylobasal length (10 specimens only), 14.8 (14.5-15.0) mm; zygomatic breadth, 14.8 (14.4-15.1) mm; interorbital breadth, 5.0 (4.7-5.2) mm; breadth across upper molars, 10.6 (10.5-11.0) mm; length of maxillary toothrow, 5.0 (4.8- 5.3) mm. These measurements generally agree with those given by Paradiso (1967:600) for 20 individuals from Panama. Females in our series average slightly larger than do males in external and cranial measurements. Six males weighed an average of 22.9 (20.7- 25.1) gms; one nonpregnant female weighed 17.1 gms. Diphylla ecaudata Spix, 1823 Specimens.—Boaco: Los Cocos, 14 km S Boaco, 220 m, 5. Madriz: 5 km N, 9 km E Condega, 800 m, 1. Matagalpa: 2 km N, 6 km E Esquipulas, 960 m, 1. Our specimens constitute the first report of this species from Nicaragua. We follow Burt and Stirton (1961:37) in regarding D. ecaudata as monotypic. Specimens from Los Cocos (three males and two females) were captured in a mist net stretched across a large, quiet pool in a small stream. The banks supported well-developed gallery forest, the understory of which had been cleared for human habitation; grass- land ( grazed) and small stands of secondary forest obtained beyond the riparian habitat. Domestic ducks, a possible source of food, were observed sleeping along the bank of this stream and on top of large boulders situated in the stream. Males from this locality taken on 20 February, 4 April, and 18 July had testicular lengths of 5, 6, and 6.5 mm, respectively. Two adult females collected there on 4 April were reproductively inactive. An adult male (testes 6 mm) from near Condega was captured on 23 June in a daytime roost in a small, cave-like crevice (see account of Peropteryx marcotis), and one from northeast of Esquipulas (testes 5 mm) was netted on 14 March along a forest trail (see account of Vampyressa pusilla ). Natalus stramineus saturatus Dalquest and Hall, 1949 Specimens.—Granada: 6 km S Nandaime, 5. Zelaya: S side Rio Mico, El Recreo, 25 m, 2. This funnel-eared species occupies an extensive geographic range (northern México to Brazil) but appears to be relatively rare in Middle America to the south of Guatemala. Our specimens rep- resent the first of this species to be reported from Nicaragua. Both specimens from El] Recreo, adult males, were caught by hand at night after they flew through an open door into a small 18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY room, possibly seeking insects that were swarming around a light bulb. Those from near Nandaime (three males, two females) were caught in a mist net set over the mouth of a well in which they were roosting; the well was approximately 2 m in diameter, and the water level was about 5 m below the rim. The females were not reproductively active (6 August). We have compared our Nicaraguan material with a number of Mexicans specimens, including the holotype of N. s. saturatus and topotypes of N. s. mexicanus. We concur with Goodwin (1959) that in México there are two rather distinct subspecies, between which a broad zone of intergradation obtains. Our Nicaraguan specimens agree most closely with N. s. saturatus, and, until addi- tional comparative material is available from Middle America, we tentatively refer them to that race. Handley (1966b:770) and Starrett and Casebeer (1968:15), however, regarded mexicanus as the appropriate name for specimens from Panama and Costa Rica. Selected measurements of two males from El Recreo are: length of forearm, 41.2, 39.0 mm; greatest length of skull, 17.1, 16.5 mm; zygomatic breadth, 8.4, 8.5 mm; mastoid breadth, 7.7, 7.6 mm; breadth of braincase, 8.2, 8.1 mm; interorbital constriction, 3.2, 3.2 mm; length of maxillary toothrow, 7.3, 7.1 mm. Myotis albescens (E. Geoffroy St.-Hilaire, 1806) This handsome Myotis has been reported previously from Nica- ragua only from the Caribbean lowlands—from the Escondido and Prinzapolka rivers (Miller and Allen, 1928:203). We netted two specimens, both males, at Santa Rosa, 17 km N and 15 km E Boaco, 300 m, Boaco, in central Nicaragua on 13 July and 9 August 1967, under the same conditions described in the account of Myotis elegans. Testes of our specimens were 7 and 6 mm, respectively, in length. External and cranial measurements are as follows: total length, 84, 83 mm; length of tail, 31, 33 mm; length of hind foot, 9, 9 mm; length of ear, 15, 15 mm; length of forearm, 32.9, 35.1 mm; weight, 6.4, 6.2 gms; greatest length of skull, 13.9, 14.4 mm; zygo- matic breadth, 8.9, 9.0 mm; postorbital breadth, 3.8, 3.9 mm; breadth of braincase, 7.2, 7.3 mm; mastoid breadth, 7.4, 7.6 mm; breadth across upper molars, 5.6, 5.5 mm; length of maxillary toothrow, 5.3, 5.3mm. Myotis elegans Hall, 1962 The first specimen on record of this rare Myotis from Central America, a nonpregnant female, was taken on 11 July 1967 at Santa Rosa, 17 km N and 15 km E Boaco, 300 m, Boaco. It was BATS FROM NICARAGUA 19 captured in a mist net as it foraged over a small stream that sup- ported relatively well-developed gallery forest along the bank. The surrounding area was grassland (grazed), with small patches of tropical forest located on the sides of hills. Twenty other species of bats were taken at this same locality including Myotis argentatus and Myotis nigricans nigricans. External and cranial measurements of our specimen, followed in parentheses by those of the female holotype from Veracruz, are: total length, 71 (79) mm; length of tail, 32 (34) mm; length of hind foot, 7 (7.5) mm; length of ear, 11 (12) mm; length of fore- arm, 32.9 (33.0) mm; greatest length of skull, 12.5 (12.4) mm; condylobasal length, 11.6 (11.9) mm; zygomatic breadth, 8.2 mm; breadth of braincase, 5.8 (6.1) mm; postorbital breadth, 3.2 (3.2) mm; length of maxillary toothrow, 4.7 (4.6) mm. Our female weighed 3.2 gms. Myotis nigricans nigricans (Schinz, 1821) . Specimens.—Boaco: Santa Rosa, 17 km N, 15 km E Boaco, 300 m, 2. Chinandega: 6.5 km N, 1 km E Cosigiiina, 10 m, 1; San Antonio, 35 m, 1. Chontales: 1 km N, 2.5 km W Villa Somoza, 330 m, 1. Madriz: Daraili, 5 km N, 14 km E Condega, 940 m, 1. Nueva Segovia: 4.5 km N, 2 km E Jalapa, 680 m, 1. Rivas: 1 km NW Sapoa, 40 m, 1. This small Neotropical Myotis has been reported from Nicaragua only from the Caribbean lowlands of Zelaya (Davis et al., 1964: 379). Our records indicate that it is widely distributed in the republic but evidently nowhere common. Females taken on 5 March and 6 August each carried a single embryo (7 and 13 mm in crown-rump length, respectively), whereas one obtained on 21 July evidenced no reproductive activity. Richard K. LaVal currently is studying the Myotis nigricans complex; pending his revision our specimens are tentatively assigned to M. n. nigricans. Myotis simus riparius Handley, 1960 A male (USNM 52800) from the Escondido River above Blue- fields, originally reported by Miller and Allen (1928:203) as one of two M. albescens from that locality, provides the first record of the species from Nicaragua and the northernmost from Middle America. The cranial dimensions of this specimen, which compare well with those listed by Handley (1960:467) for the Panamanian holotype and paratype, are: greatest length of skull, 13.8 mm; zygomatic breadth, 8.9 mm; postorbital constriction, 3.5 mm; breadth of braincase, 6.7 mm; mastoid breadth, 7.4 mm; breadth across upper molars, 5.5 mm; length of maxillary toothrow, 5.2 mm. 20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY From Nicaraguan specimens of albescens, the skull of the speci- men of simus examined (skin not seen) differs most conspicuously in having a less inflated braincase, narrower postorbital region, and a distinct sagittal crest. Fptesicus furinalis gaumeri (J. A. Allen, 1897) Specimens.—Carazo: 3 km N, 4 km W Diriamba, 600 m, 7. Chinandega: Potosi, 5 m, 1; 6.5 km N, 1 km E Cosigiiina, 10 m, 1; San Antonio, 35 m, I. Chontales: 1 km N, 2.5 km W Villa Somoza, 330 m, 1; Hato Grande, 13 km S, 8 km W Juigalpa, 60 m, 3. Davis (1965:234) reported two specimens of this species from Nicaragua, one from Carazo (listed above) and the other from 1 mi SE Yalagitina, 2600 ft, Madriz. The only other specimen on record from Nicaragua is from Greytown (Miller, 1897:100). Except for two individuals that were shot as they foraged in early evening, our specimens were captured in mist nets stretched over water or as described below. Some of the bats captured northwest of Diriamba were taken in a net placed across the center of a water-filled, concrete cistern that was located in a large, open space used for drying coffee beans. Open at the top, the cistern was approximately 12 m in diameter, and the water level was approximately 3 m below the rim. Several Artibeus jamaicensis, A. lituratus, and Molossus pretiosus were caught in the same net. Aside from bats removed from the net, a few Eptesicus and Molossus were retrieved from the water where they fell, apparently stunned, after colliding with the wall of the cistern. The fact that a number of decomposed bats of these two species were observed floating in the water indicated that such entrapment was not caused by the placement of our net. A female pregnant with two embryos (crown-rump length 15 mm) was captured on 22 April, whereas one taken on 5 July was lactating. Adult males taken in March (two) and April (two) had testes 5, 7, 9, and 10 mm long, respectively. Rhogeessa tumida tumida H. Allen, 1866 Specimens.—Boaco: Santa Rosa, 17 km N, 15 km E Boaco, 300 m, 2. Carazo: 3 km N, 4 km W Diriamba, 600 m, 2. Chinandega: 6 km N, 1 km E Cosigiiina, 10 m, 3; Hda. Bellavista, 720 m, Volcan Casita, 1. Matagalpa: 1 km NE Esquipulas, 420 m, 1. This small vespertilionid occurs throughout much of Central America, but the available material still is insufficient to permit an accurate assessment of geographic variation in that region. All but one of our specimens were trapped in mist nets. Those captured BATS FROM NICARAGUA 21 near Diriamba were taken in a net stretched across a trail bordered by high cut-banks, whereas those from other localities were netted over small streams. The one bat from Bellavista was shot as it foraged in the evening along trees bordering the hacienda yard. Two females taken on 5 March each had swollen uteri and may have been in an early stage of pregnancy. Others taken on 13 July, 9 August, and 15 August were reproductively inactive. Males had testes that measured 2.5 (March), 3 (July), and 2 (August) mm. We have compared our material with representative specimens from Mexico of R. parvula and R. tumida, as defined by Goodwin (1958); we are not wholly convinced that these are valid species (rather than variable geographic races of the same species). Cer- tainly there is need for additional investigation of the problem. Our Nicaraguan specimens most closely resemble R. t. tumida (see Goodwin, op. cit.:3), to which they are tentatively referred. Selected measurements of two females from Boaco and a male from Matagalpa, followed by the average (and extremes) of three males and three females from western Nicaragua (Carazo and Chinandega), are, respectively: length of forearm, 30.4, 28.5, 28.3, 28.2 (27.5-29.0) mm; greatest length of skull, 12.8, 12.7, 12.8, 12.1 (11.8-12.5) mm; zygomatic breadth, 8.1, 8.4, 8.6, 8.1 (7.8-8.6) mm; mastoid breadth, 7.1, 7.2, 7.2, 6.8 (6.6-7.1) mm; breadth of brain- case, 6.0, 5.7, 6.0, 5.6 (5.3-6.1) mm; postorbital constriction, 3.3, 3.0, 3.3, 3.0 (2.9-3.2) mm; length of maxillary toothrow, 4.7, 4.7, 4.5, 4.3 (4.2-4.6) mm; weight, 4.4, 4.3, 3.8, 3.5 (3.1-3.9) gms. Tadarida laticaudata yucatanica (Miller, 1902) One adult male (testes 5 mm) of this free-tailed bat from Potosi, 5 m, Chinandega, on the Cosigiiina Peninsula, provides the only record of the genus Tadarida from Nicaragua. This specimen was netted after dark over a small stream that flowed into the Bay of Fonseca approximately 200 yards below our nets (see account of Noctilio leporinus). Other species taken in the same net included Carollia subrufa, Sturnira lilium, Eptesicus furinalis, Molossus ater, and Molossus molossus. Selected measurements of the male are: total length, 92 mm; length of tail, 31 mm; length of hind foot, 10 mm; length of ear, 15 mm; length of forearm, 39.2 mm; weight, 10.3 gms; greatest length of skull, 17.2 mm; zygomatic breadth, 10.6 mm; postorbital constriction, 4.0 mm; breadth of braincase, 8.6 mm; length of maxillary toothrow, 6.5 mm. 22 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Eumops auripendulus (Shaw, 1800) A broken skull (USNM 339917) of a female of this species from Hda. Mecatepe [2 km N, 11.5 km E Nandaime, ca. 40 m], Granada, represents the only known specimen of the genus Eumops from Nicaragua. This bat was obtained by M. K. Clark on 25 August 1964, but the conditions under which it was captured are not known. Available cranial measurements are: zygomatic breadth, 12.6 mm; breadth of braincase, 10.7 mm; postorbital breadth, 4.3 mm; rostral breadth, 7.4 mm; length of maxillary tooth- row, 9.1 mm; length of mandibular c-m3, 10.0 mm. Molossus ater nigricans Miller, 1902 Specimens.—Chinandega: Potosi, 5 m, 21; 4.5 km N Cosigiiina, 15 m, 2; El Paraiso, 1 km N Cosigiiina, 20 m, 17; Hda. Bellavista, 720 m, Volcan Casita, 4; Hda. San Isidro, 10 km S Chinandega, 20 m, 2; San Antonio, 35 m, 2. Nueva Segovia: Corozo, 15 km NNE Jalapa, 660 m, 1; 3.5 km S, 2 km W Jalapa, 660 m, 4. Only one specimen of this large molossid (from Volcan de Chinandega—J. A. Allen, 1908:670) has been reported from Nicara- gua. All of our specimens are from the northern or northwestern part of the country. Many were shot in early evening as they followed a straight, direct flight pattern (presumably from day- roosts toward foraging or watering areas); the series from Potosi was netted over a small stream as described in the account of Noctilio leporinus. We confidently refer our material to the species currently known as ater, although assignment to the subspecies nigricans is tentative. Of 21 females autopsied in the first week of March on the Cosigitina Peninsula, seven were pregnant, each with a single embryo (7-22 mm in crown-rump length, average 16.7). Ten males taken at the same time had an average testicular length of 5.7 (2-7) mm, whereas the testes of a male obtained on 9 March at San Antonio measured 8 mm. None of five females obtained in late July in Nueva Segovia evidenced reproductive activity, nor did two of three females taken in July and one taken in August from Chinandega; the fourth Chinandegan female, taken on 17 July, carried an embryo that was 36 mm in length. Two males from southern Chinandega (mid-July) had testes 7 and 4 mm long. Representative measurements of 10 specimens of each sex from Nicaragua are given in table 4. : { f BATS FROM NICARAGUA 23 Molossus molossus aztecus Saussure, 1860 Specimens.—Chinandega: Potosi, 5 m, 1; Hda. San Isidro, 10 km S Chinandega, 20 m, 1 (USNM). Boaco: Santa Rosa, 17 km N, 15 km E Boaco, 300 m, 7. Managua: 3 mi SW Managua, 8. Rivas: Rivas, 60 m, 4. This small free-tailed species has been reported from Nicaragua by Felten (1957:14), who listed two females from Corinto. Our records indicate that it is widely distributed, but of localized oc- currence. Specimens from Potosi and Santa Rosa were captured in mist nets over streams (as described in the accounts of Noctilio leporinus and Myotis elegans, respectively). The specimen from Hda. San Isidro was shot in flight, whereas those from Rivas were captured in a daytime retreat in a deep crevice in a concrete school building. We have no precise knowledge of the conditions under which bats from 3 mi SW Managua were obtained but suspect they were taken from a building. Females in our series were reproductively active at all times for which we have information—early March through mid-July—as fol- lows: a female from Potosi (6 March) carried an embryo that mea- sured 5 mm (crown-rump length) as did one from Santa Rosa (21 March, 17 mm in length); two of three females taken 3 mi SW Managua on 28 March were pregnant (embryos 13 and 15 mm), whereas each of two collected there on 3 May were gravid (embryos 27 and 30 mm); one female from Rivas (25 June) carried an embryo (30 mm) and another was lactating and accompanied by a small (forearm, 19.7 mm) hairless juvenile; and, finally, one of six females netted at Santa Rosa on 13 July was pregnant (embryo 23 mm), but the others evinced no reproductive activity. An adult male, taken at Rivas with the females mentioned above, had testes that were 6 mm in length. Bats of the genus Molossus are in need of thorough systematic study. We think most, if not all, mainland populations of small Molossus with pale-based hairs pertain to the species Molossus molossus, originally described from the Lesser Antilles. Although there is some variation in size and overall color among our Nicara- guan samples, all seem to represent that species. Because speci- mens from Nicaragua closely resemble examples of M. m. aztecus from Jalisco, México (Table 3), in external and cranial proportions, we tentatively refer them to aztecus. In our field experience in both Middle and South America, colonies of M. molossus tend to be extremely localized, with in- dividual populations or demes frequently separated by many miles of territory in which the species seemingly does not regularly occur. 24 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY TABLE 3.—Selected measurements of adult Molossus molossus. 3 iS 7 5 ¢ Number of S cs . 8 2s specimens + ae cs 2 Siu Ss - os averaged or 3GlaeaeGre Santa Rosa, Boaco, Nicaragua Average 6 (9) 36.0 17.5. 147 108 106 G1 Se 5a Minimum 35.5 17.3. 144 106 104 9.0 “S:G03saeeeee Maximum 36.7 178 15.1 110 108 92 SOs Rivas, Nicaragua KU 106291, ¢ 38.2 188 161 115 109 G4 “OR SGGeee KU 106290, 9° 39.6 174 150 110 105 91 “SG aga KU 106293, 9 37.2 173 148 10.7 102 9.0" “Sia °s eee The localized and presumably highly inbred populations may have diverged morphologically, in some cases at least, to a degree that mensural differences can be demonstrated even between samples from the same general geographic area. Localized variation and relatively marked secondary sexual variation (unrecognized by some earlier workers ), superimposed on geographic variation, have resulted in application of a relatively large number of names to these small Molossus. Felten (1957:13-14), for example, apparently used different specific names for males and females from El Salvador, and Gardner (1966) employed three different specific names for North American specimens. Only when material is available for a detailed study of variation throughout the Neotropics can the perplexing mosaic of characters in these small Molossus be assessed adequately. BATS FROM NICARAGUA 25 We have seen no specimens from Nicaragua that are identifiable as Molossus bondae, another relatively small species that has been reported from Greytown ( Miller, 1913a:89) and from elsewhere in Central America by other authors (Goodwin, 1942c:145; Handley, 1966b:772; Gardner et al., 1970:727). Our examination of the female holotype of M. bondae reveals that it is larger than M. molossus, corresponding in size to females recently reported from Costa Rica by Gardner et al. (loc. cit.), and that bondae has dark- based hairs. The two males reported by Goodwin (loc. cit.) from Honduras as bondae, would seem to be too small for that species, based on the measurements listed; also, these specimens allegedly have white-based hairs and probably represent M. molossus as here defined. Molossus pretiosus pretiosus Miller, 1902 Specimens.—Boaco: Los Cocos, 14 km S Boaco, 220 m, 28; San Francisco, 19 km S, 2 km E Boaco, 200 m, 3. Carazo: 3 km N, 4 km W Diriamba, 600 m, 25. Managua: 6 mi WSW Managua, 3. This relatively large mastiff bat has not been reported previously from Nicaragua. Specimens from several localities in Boaco were captured in mist nets over streams; most of those from northwest of Diriamba were shot in the early evening as they foraged high around large trees in a coffee finca, but several were netted over a water-filled concrete tank or found in the water in the tank as detailed in the account of Eptesicus furinalis. At Los Cocos, bats that we netted seemed to be emerging from a hollow located high in a tree over the stream. Selected measurements of M. p. pretiosus from Nicaragua, which compare favorably with those of topotypes from Venezuela, are listed in table 4 along with measurements of M. ater and M. sinaloae. The taxonomic relationships of M. pretiosus and M. ater are less than clear, and some authors (Handley, 1966b:773, for instance) have suggested that the two may be conspecific. What- ever their ultimate relationships may prove to be, two distinctive taxa seem to be present in Nicaragua; the larger is assignable to ater and the smaller to pretiosus, as currently understood. Further- more, the presumed presence of two large Molossus with dark-based hairs elsewhere in Central America (Dilford C. Carter, personal communication) and in southeastern México (Goodwin, 1956:4; Goodwin and Greenhall, 1964:20) argues for specific recognition of pretiosus. The species ater and pretiosus differ mainly in size (Table 4), some measurements clearly separating the two when sexual dimor- 26 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY TABLE 4.—Selected measurements of adults of three species of Molossus from Nicaragua. es 2 cz wu GS Ba Number of = 5 S° -feae specimens oa x § A "Sues eae S & averaged or af eo 2 pee Ss «sk §0 catalogue to a5 oo Erg aoe we «6S SB number, Fe Os abe 80S Da aa o and sex a OS Os NS m5 +8" es Molossus ater nigricans, Departamento de Chinandega, Nicaragua Average 10 (6) 49.5 232 20.2 14.3 11.0 8.2 6.1 Minimum 48.2, 22.8 19.9 14.1 10.5 8.0 6.0 Maximum BOM 23.0 20:5 14.9 12 8.3 6.3 Average 10 (2?) 49.0 92.0 19.2 13.6 10% 7.8 5.8 Minimum 47.1 2165 18.8 oS 10.5 7.6 5.6 Maximum Dilez 2235 20.0 14.1 11.0 8.2 6.1 Molossus pretiosus pretiosus, Departamento de Boaco, Nicaragua Average 8 (2) A451 IT 18.8 13.4 10.6 7.6 5.9 Minimum 44.1 2170 18.3 Peed 10.2 74 5.6 Maximum 46.1 2.9; 19.1 13.7 10.9 7.9 6.1 Average 23 (9?) 43.6 20.2 77 IZ LOW Wis: 5.2 Minimum 41.0 19.5 V7.2 12,2 95 ries! 5.0 Maximum 44.8 20.7 18.2 13.1 10.4 7.6 5.6 6 mi SW Managua, Nicaragua KU 70135, 6 44.8 NRT 18.6 13.0 10.2 Te 5.8 KU-70137. 6 45.1 21.6 19.0 13.2 10.7 ies 5.8 KU 70136, @ 43.5 19.9 ilies: pate 10.3 Teo 5.D 3 km N, 4 km W Diriamba, Nicaragua Average 7 (2) 44.8 212 18.5 13.0 10.5 7A 5.5 Minimum 43.8 207) 18.3 PAT 10.2 12 5.4 Maximum ABT pia ls: 18.8 13:3 10.9 16 ao Average 9 (9?) 44.0 20.7 18.0 12:6 10.2 we) Bad Minimum 42.0 20.0 17.4 12.0 10.0 7.0 5.0 Maximum 45.5 21.2 18.6 13:0 10.6 18 oo Molossus sinaloae sinaloae, E] Recreo, Nicaragua Average 4 (2) 47.6 D189, 18.6 123 9.8 Td 5.4 Minimum 46.7 20.7 18.3 12.0 9.7 7.6 iS Maximum 48.6 215 18.9 12:6 10.0 7.9 5.5 Average 10 (@ ) 47.5 20.2 17.8 1-8 9.7 res. Bak Minimum 46.2 20.0 17.6 1b) a) 9.5 TA 5.0 Maximum 48.3 20.4 18.0 12.0 9.8 7.9 5.5 phism is considered. Also, the average weights of 18 nonpregnant females and four males of pretiosus (all adults) collected on 20 February 1968 at Los Cocos, were 20.9 (14.6-23.8) and 27.0 (24.6- 31.7) gms, respectively, significantly smaller than corresponding figures for 11 nonpregnant females and nine males of M. a. nigricans taken two weeks later on the Cosigiiina Peninsula—29.1 BATS FROM NICARAGUA 27 (26.1-33.0) and 32.9 (29.3-35.1) gms. It is of note that we have not collected these two large species at the same localities in Nic- aragua, and it is possible that one competitively excludes the other in local situations. Among females from Departamento de Boaco, none of 18 col- lected on 20 February was pregnant, whereas three of four obtained on 4 April were gravid (embryos 13, 15, and 17 mm in crown-rump length) as was one (embryo 21 mm) taken on 18 July. Among those from northwest of Diriamba, two of three were pregnant on 30-31 March (embryos 7 and 13 mm in length), whereas only three of eight carried embryos (29, 31, and 34 mm) in mid-August at a time when flying young also were collected. Additionally, a female captured 6 mi WSW Managua on 27 March had an embryo that measured 11 mm. Four adult males collected on 20 February had an average testicular length of 5.7 (4-7) mm, whereas those of seven obtained in late March averaged 5.0 (4-6) mm; testes of two adults taken on 18 July were 6 and 6.5 mm, and those of three taken in the period 12-15 August were 5 mm in length. Molossus sinaloae sinaloae J. A. Allen, 1906 Specimens.—Boaco: Santa Rosa, 17 km N, 15 km E Boaco, 300 m, 1. Managua: 1 km N Sabana Grande, 1; 3 mi SW Managua, 1; 8 km SW Managua, 400 m, 3. Rivas: San Juan del Sur, 20 m, 1. Zelaya: S side Rio Mico, El Recreo, 25 m, 73. This mastiff bat can be distinguished easily from the other two large species of Molossus in Nicaragua in that the hairs of the dor- sum are relatively long and are pale, frequently white, basally. The species has been reported twice previously from Nicaragua— from Greytown by Miller (1913a:89) and by Goodwin and Green- hall (1964:13), who listed four individuals from Rio Grande. Most of our specimens were taken from buildings. A large series obtained in a house at El Recreo in late June 1966 consisted only of females and young, many of which were nearly full grown; some females still were lactating and at least two were pregnant (on 23 June—crown-rump length of embryos 12 and 14 mm). A group taken from another house at El Recreo (from cracks in the stone wall and from between rafters and the corrugated metal roof) in late July 1966 consisted of adult males, lactating females, and young. Similarly, an adult male, a pregnant female (embryo 14 mm _in crown-rump length), and a hairless juvenile were shot, on 16 _ February 1964, from a small colony in an attic of a large house 28 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY southwest of Managua; Glossophaga soricina also was taken in this attic. The testes of an adult male captured in June were 6 mm in length; those of four taken in July were 5, 5, 6, and 7 mm, and one obtained in August had testes that measured 3 mm. Selected measurements of M. s. sinaloae from El Recreo are given in table 4. CHECKLIST OF NICARAGUAN BATS Following is a checklist of the 68 species of bats thus far re- ported from Nicaragua. Citations are given only to publications that are primary sources of information on Nicaraguan specimens. In some instances, specimens were reported in the literature under a name different than the one here employed; we have mentioned earlier-used names where it seemed appropriate to do so. Taxa preceded by an asterisk are included in the present paper. Family EMBALLONURIDAE Rhynchonycteris naso (Wied-Neuwied, 1820).—J. A. Allen (1908:669, 1910: 110), Sanborn (1937:327), Davis et al. (1964:375, 379). Saccopteryx bilineata (Temminck, 1838-1839 ).—J. A. Allen (1910:110), San- born (1937:331), Jones (1964a:506). *Saccopteryx leptura (Schreber, 1774).—Davis et al. (1964:374), Jones (1964a:506). Cormura brevirostris (Wagner, 1843).—J. A. Allen (1910:110—specimens listed as Peropteryx canina from Pena Blanca, see Sanborn, 1937:348), Miller (1924:38), Sanborn (1937:349), Goodwin (1942c:119). Peropteryx kappleri kappleri Peters, 1867—J. A. Allen (1908:669, 1910:110 —as P. canina), Sanborn (1937:344). * Peropteryx macrotis macrotis (Wagner, 1843). Balantiopteryx plicata plicata Peters, 1867.—Jones et al. (1962:152), Davis et al. (1964:375), Jones (1964a:506). Diclidurus virgo Thomas, 1903.—Alston (1879-82:30). Family NOCTILIONIDAE *Noctilio labialis labialis (Kerr, 1792).—Goodwin (1942c:121, 1946:297), Davis et al. (1964:376), Kohls et al. (1965:344), Wenzel et al. (1966: 578). *Noctilio leporinus mexicanus Goldman, 1915.—Davis et al. (1964:376). Family PHYLLOSTOMATIDAE *Pteronotus davyi fulvus (Thomas, 1892). *Pteronotus parnellii fuscus J. A. Allen, 1911.—Miller (1902:402), Rehn (1904:203). *Pteronotus suapurensis (J. A. Allen, 1904).—Goodwin (1942a:88, 1946:299, 1953:246). Micronycteris brachyotis (Dobson, 1879).—Goodwin (1946:302—as M. sylvestris), Sanborn (1949:224—as M. platyceps), Goodwin and Green- hall (1961:231). Micronycteris hirsuta (Peters, 1869 ).—Valdez and LaVal (1971:247). * Micronycteris megalotis mexicana Miller, 1898.—G. M. Allen (1929:130). BATS FROM NICARAGUA 29 * Micronycteris megalotis microtis Miller, 1898.—Miller (1898:328, 1900:154), Lyon and Osgood (1909:263), J. A. Allen (1910:110), Poole and Schantz (1942:133). Micronycteris minuta (Gervais, 1856 ).—Valdez and LaVal (1971:248). Micronycteris schmidtorum Sanborn, 1935.—Davis et al. (1964:378). Macrophyllum macrophyllum (Schinz, 1821)—Davis et al. (1964:378), Jones (1964a:506). *Tonatia nicaraguae Goodwin, 1942.—Goodwin (1942b:205, 1946:305, 1953: 246), Davis and Carter (1962:67), Valdez and LaVal (1971:248). Phyllostomus discolor verrucosus Elliot, 1905.—Jones (1964a:507), Wenzel et al. (1966:598). *Phyllostomus hastatus panamensis J. A. Allen, 1904.—Goodwin (1942c:126), Wenzel et al. (1966:597). *Trachops cirrhosus coffini Goldman, 1925.—Carter et al. (1966:491). *Chrotopterus auritus auritus (Peters, 1856). Vampyrum spectrum nelsoni (Goldman, 1917).—Gray (1844:18), Dobson (1878:471), Alston (1879-82:39), J. A. Allen (1910:111), Goldman (1917:116), Goodwin (1946:310). Glossophaga commissarisi Gardner, 1962.—Davis et al. (1964:380), Jones (1964a:507). Glossophaga soricina leachii (Gray, 1844).—Gray (1844:18), Dobson (1878: 501), Alston (1879-82:44), J. A. Allen (1910:111), Miller (1913b:419), Alvarez (1963:400), Davis et al. (1964:380), Starrett and de la Torre (1964:57). *Anoura geoffroyi lasiopyga (Peters, 1868). *Choeroniscus godmani (Thomas, 1903 ).—Handley (1966a:86). *Lichonycteris obscura Thomas, 1895.—Thomas (1895:57), Davis et al. (1964:380). *Carollia castanea H. Allen, 1890.—Davis et al. (1964:379). Carollia perspicillata azteca Saussure, 1860.—Hahn (1907:112), J. A. Allen (1908:669, 1910:111), Davis et al. (1964:379), Starrett and de la Torre (1964:58). Carollia subrufa (Hahn, 1905).—Davis and Carter (1962:71), Davis et al. (1964:379). Sturnira lilium parvidens Goldman, 1917.—Davis et al. (1964:379), Starrett and de la Torre (1964:59), Jones (1964a:507 ). *Sturnira ludovici ludovici Anthony, 1924. Uroderma bilobatum convexum Lyon, 1902.—Davis et al. (1964:382), Jones (1964a:507), Davis (1968:695). Uroderma bilobatum molaris Davis, 1968.—Davis et al. (1964:382), Davis (1968:697). *Uroderma magnirostrum Davis, 1968.—Davis (1968:680). *Vampyrops helleri Peters, 1866.—Davis et al. (1964:383), Jones (1964a:507 ), Valdez and LaVal (1971:248). *Vampyrodes major G. M. Allen, 1908.—J. A. Allen (1910:112), Goodwin (1946:320). *Vampyressa nymphaea Thomas, 1909. *Vampyressa pusilla thyone Thomas, 1909.—Starrett and de la Torre (1964: 60). *Chiroderma villosum jesupi J. A. Allen, 1900. Ectophylla alba H. Allen, 1892.—H. Allen (1892:441, 1898:267), Lyon and 30 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Osgood (1909:266), Poole and Schantz (1942:129), Goodwin (1942c: 134, 1946:323), Goodwin and Greenhall (1962:4). Type locality (Rio Segovia) placed in Comarca de El] Cabo, Nicaragua, by Miller and Kellogg (1955:77). Artibeus inopinatus Davis and Carter, 1964.—Davis and Carter (1964:120). Artibeus jamaicensis paulus Davis, 1970.—Andersen (1908:266), Davis (1970b:121). Artibeus jamaicensis richardsoni J. A. Allen, 1908.—Andersen (1908:267), J. A. Allen (1908:669), Elliot (1917:168), Goodwin (1953:249), Davis and Carter (1964:120), Davis et al. (1964:379), Starrett and de la Torre (1964:60), Davis (1970b:116). Artibeus lituratus palmarum J. A. Allen and Chapman, 1897.—Andersen (1908:279), Davis and Carter (1964:120), Davis et al. (1964:379), Starrett and de la Torre (1964:61). Artibeus phaeotis palatinus Davis, 1970.—Davis (1970a:401 ). Artibeus phaeotis phaeotis (Miller, 1902).—Davis et al. (1964:379), Davis (1970a:399). * Artibeus toltecus hesperus Davis, 1969. *Artibeus toltecus toltecus (Saussure, 1860).—Andersen (1908:300), Davis (1969:28). *Artibeus watsoni Thomas, 1901.—Andersen (1908:290), Davis and Carter (1962:71), Davis (1970a:393). *Centurio senex senex Gray, 1842.—Goodwin (1946:327, restricted type lo- cality to El Realejo, Nicaragua). Desmodus rotundus murinus Wagner, 1840.—J. A. Allen (1908:670, 1910: 113) * Diphylla ecaudata Spix, 1823. Family NATALIDAE *Natalus stramineus saturatus Dalquest and Hall, 1949. Family THYROPTERIDAE Thyroptera discifera discifera (Lichtenstein and Peters, 1854).—Miller 1896: 111, 1931:411), Goodwin (1946:330). Family VESPERTILIONIDAE *Myotis albescens (E. Geoffroy St.-Hilaire, 1806)—Miller and G. M. Allen (1928:203). * Myotis elegans Hall, 1962. *Myotis nigricans nigricans (Schinz, 1821).—Davis et al. (1964:379), Mc- Daniel and Coffman (1970:223). * Myotis simus riparius Handley, 1960. *Eptesicus furinalis gaumeri (J. A. Allen, 1897).—Miller (1897:100), Davis (1965:234). Lasiurus borealis frantzii (Peters, 1871 ).—Davis and Carter (1962:73). *Rhogeessa tumida tumida H. Allen, 1866.—J. A. Allen (1910:113), Goodwin (1958:3). Family MOLOSSIDAE * Tadarida laticaudata yucatanica ( Miller, 1902). *Eumops auripendulus (Shaw, 1800). BATS FROM NICARAGUA 31 * Molossus ater nigricans Miller, 1902.—J. A. Allen (1908:670). Molossus bondae J. A. Allen, 1904.—Miller (1913a:89). * Molossus molossus aztecus Saussure, 1860.—Felten (1957:14). * Molossus pretiosus pretiosus Miller, 1902. *Molossus sinaloae sinaloae J. A. Allen, 1906.—Miller (1913a:89), Goodwin and Greenhall (1964:13). LITERATURE CITED ALLEN, G. M. 1929. Mammals. Pp. 129-130, in Vertebrates from the Corn Islands. Bull. Mus. Comp. Zool., 69: 127-146. ALLEN, H. 1892. Description of a new genus of phyllostome bats. Proc. U.S. Nat. Mus., 15:441-442. 1898. The skull and teeth of Ectophylla alba. Trans. Amer. Philos. Soc., 19:267-272. ALLEN, J. A. 1908. Mammals from Nicaragua. Bull. Amer. Mus. Nat. Hist., 24:647-670. 1910. Additional mammals from Nicaragua. Bull. Amer. Mus. Nat. Hist., 28:87-115. ALSTON, E. R. 1879-1882. Biologia Centrali-Americana, Mammalia. London, xx+220 (introduction by P. L. Sclater). ALVAREZ, T. 1963. The Recent mammals of Tamaulipas, México. Univ. Kansas Publ., Mus. Nat. Hist., 14:363-473. ANDERSEN, K. 1908. A monograph of the chiropteran genera Uroderma, Enchisthenes, and Artibeus. Proc. Zool. Soc. London, 204-319. Burt, W. H., and R. A. StirTON 1961. The mammals of El Salvador. Misc. Publ. Mus. Zool., Univ. Michigan, 117:1-69. CaBrera, A. 1958. Catalogo de los mamiferos de America del Sur. Rev. Mus. Argentino Cienc. Nat., Buenos Aires, 4:iv-+307. Carter, D. C., R. H. Pine, and W. B. Davis 1966. Notes on Middle American bats. Southwestern Nat., 11:488-499. Davis, W. B. 1965. Review of the Eptesicus brasiliensis complex in Middle America with description of a new subspecies from Costa Rica. Jour. Mamm., 46:229-240. 1968. Review of the genus Uroderma (Chiroptera). Jour. Mamm., 49: 676-698. 1969. A review of the small fruit bats (genus Artibeus) of Middle Amer- ica. Southwestern Nat., 14:15-29. 1970a. A review of the small fruit bats (genus Artibeus) of Middle Amer- ica. Part II. Southwestern Nat., 14:389-402. 32 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 1970b. The large fruit bats (genus Artibeus) of Middle America, with a review of the Artibeus jamaicensis complex. Jour. Mamm., 51: 105-122. Davis, W. B., and D. C. Carter 1962. Notes on Central American bats with description of a new subspecies of Mormoops. Southwestern Nat., 7:64-74. 1964. A new species of fruit-eating bat (genus Artibeus) from Central America. Proc. Biol. Soc. Washington, 77:119-121. Davis, W. B., D. C. Carrer, and R. H. PINE 1964. Noteworthy records of Mexican and Central American bats. Jour. Mamm., 45:375-378. Dosson, G. E. 1878. Catalogue of the Chiroptera in the collection of the British Museum. British Museum, London, xlii+-567. EELIOT,D..G. 1917. A check-list of mammals of the North American continent. .. . Suppl., edited by J. A. Allen, Amer. Mus. Nat. Hist., iv-+-192. FELTEN, H. 1955. Fledermiuse (Mammalia, Chiroptera) aus El] Salvador. Tiel 1. Senckenbergiana Biol., 36:271-285. 1956. Fledermause (Mammalia, Chiroptera) aus El Salvador. Tiel 3. Senckenbergiana Biol., 37:179-212. 1957. Fledermause (Mammalia, Chiroptera) aus El] Salvador. Tiel 5. Senckenbergiana Biol., 38: 1-22. GARDNER, A. L. 1966. A new subspecies of the Aztec mastiff bat, Molossus aztecus Saussure, from southern Mexico. Contrib. Sci., Los Angeles Co. Mus, Pit s1-5. GARDNER, A. L., R. K. LAVAL, and D. E. Wi1Lson 1970. The distributional status of some Costa Rican bats. Jour. Mamm., 51:712-729. GoLpMAN, E. A. 1917. New mammals from North and Middle America. Proc. Biol. Soc. Washington, 30:107-116. Goopwin, G. G. 1942a. New Pteronotus from Nicaragua. Jour. Mamm., 23:88. 1942b. A summary of recognizable species of Tonatia, with descriptions of two new species. Jour. Mamm., 23:204-209. 1942c. Mammals of Honduras. Bull. Amer. Mus. Nat. Hist., 79: 107-195. 1946. Mammals of Costa Rica. Bull. Amer. Mus. Nat. Hist., 87:271-473. 1953. Catalogue of type specimens of Recent mammals in the American Museum of Natural History. Bull. Amer. Mus. Nat. Hist., 102: 211-411. 1956. A preliminary report on the mammals collected by Thomas Mac- Dougall in southeastern Oaxaca, Mexico. Amer. Mus. Novit., 1757: IES; 1958. Bats of the genus Rhogeessa. Amer. Mus. Novit., 1923:1-17. 1959. Bats of the subgenus Natalus. Amer. Mus. Novit., 1977:1-22. | i ‘| BATS FROM NICARAGUA 33 Goopw1n, G. G., and A. M. GREENHALL 1961. A review of the bats of Trinidad and Tobago. Bull. Amer. Mus. Nat. Hist., 122:189-301. 1962. Two new bats from Trinidad, with comments on the status of the genus Mesophylla. Amer. Mus. Novit., 2080:1-18. 1964. New records of bats from Trinidad and comments on the status of > Molossus trinitatus Goodwin. Amer. Mus. Novit., 2195:1-23. Gray, J. E. 1844. Zoology of the voyage of H.M.S. Sulphur. . . . Mammalia (parts 1,2). London [not seen]. Haun, W. L. 1907. A review of the bats of the genus Hemiderma. Proc. U.S. Nat. Mus., 32:103-118. HANDLEY, C. O., Jr. 1960. Descriptions of new bats from Panama. Proc. U.S. Nat. Mus., 112:459-479. 1966a. Description of new bats (Choeroniscus and Rhinophylla) from Colombia. Proc. Biol. Soc. Washington, 79:83-88. 1966b. Checklist of the mammals of Panama. Pp. 753-795, in Ectoparasites of Panama (R. L. Wenzel and V. J. Tipton, eds.), Field Mus. Nat. Hist., Chicago, xii+861. HERSHKOVITZ, P. 1949. Mammals of northern Colombia. Preliminary report no. 5: bats (Chiroptera). Proc. U.S. Nat. Mus., 99:429-454. Husson, A. M. 1962. The bats of Suriname. E. J. Brill, Leiden, 282. JOnEs, J. K., JR. 1964a. Bats new to the fauna of Nicaragua. Trans. Kansas Acad. Sci., 67:506-508. 1964b. Bats from western and southern Mexico. Trans. Kansas Acad. Sci., 67:509-516. 1966. Bats from Guatemala. Univ. Kansas Publ., Mus. Nat. Hist., 16: 439-472. Jones, J. K., Jn., T. ALVAREz, and M. R. LEE 1962. Noteworthy mammals from Sinaloa, Mexico. Univ. Kansas Publ., Mus. Nat. Hist., 14:145-159. _ Kouts, G. M., D. E. SONENSHINE, and C. M. CLIFFORD 1965. The systematics of the subfamily Ornithodorinae (Acarina: Argasi- dae). II. Identification of the larvae of the Western Hemisphere and descriptions of three new species. Ann. Ent. Soc. Amer., 58:331-364. LAVAL, R. K. 1969. Records of bats from Honduras and E] Salvador. Jour. Mamm., 50:819-822. Lyon, M. W., and W. H. Oscoop 1909. Catalogue of the type-specimens of mammals in the United States National Museum, including the Biological Survey collection. Bull. U.S. Nat. Mus., 62:ix+325. 34 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY McDanlic., B., and C. C. COFFMAN 1970. The labidocarpid bat-mites of the United States (Acarina: Lis- trophoridea). Proc. Helminthological Soc. Washington, 37:223-229. MILLER, G. S., JR. 1896. The Central American Thyroptera. Proc. Biol. Soc. Washington, 10:109-112. 1897. Revision of the North American bats of the family Vespertilionidae. N. Amer. Fauna, 13:1-135. 1898. Descriptions of five new phyllostome bats. Proc. Acad. Nat. Sci. Philadelphia, 50:326-337. 1900. Note on Micronycteris brachyotis (Dobson) and M. microtis Miller. Proc. Biol. Soc. Washington, 13:154-155. 1902. Twenty new American bats. Proc. Acad. Nat. Sci. Philadelphia, 54:389-412. 1913a. Notes on the bats of the genus Molossus. Proc. U.S. Nat. Mus., 46:85-92. 1913b. Revision of the bats of the genus Glossophaga. Proc. U.S. Nat. Mus., 46:413-429. 1924. List of North American Recent mammals 1923. Bull. U.S. Nat. Mus., 128:xvi+673. 1931. Two new South American bats. Jour. Mamm., 12:411-412. MILLer, G. S., Jr., and G. M. ALLEN 1928. The American bats of the genera Myotis and Pizonyx. Bull. U.S. Nat. Mus., 144:viii+218. MILLER, G. S. Jr., and R. KELLocG 1955. List of North American Recent mammals. Bull. U.S. Nat. Mus., 205 :xii+954. PARADISO, J. L. 1967. A review of the wrinkle-faced bats (Centurio senex Gray), with description of a new subspecies. Mammalia, 31:595-604. Poo.e, A. J., and V. S. SCHANTZ 1942. Catalog of the type specimens of mammals in the United States National Museum, including the Biological Surveys collection. Bull. U.S. Nat. Mus., 178:xiii+-705. REHN, J. A. G. 1904. A study of the mammalian genus Chilonycteris. Proc. Acad. Nat. Sci. Philadelphia, 56: 181-207. SANBORN, C. C. 1937. American bats of the subfamily Emballonurinae. Field Mus. Nat. Hist., Zool Ser., 20:321-354. 1949. Bats of the genus Micronycteris and its subgenera. Fieldiana-Zool., S12215-233. STARRETT, A., and R. S. CASEBEER 1968. Records of bats from Costa Rica. Contrib. Sci., Los Angeles Co. Mus., 148:1-21. STARRETT, A., and L. DE LA TORRE 1964. Notes on a collection of bats from Central America, with the third record of Cyttarops alecto Thomas. Zoologica, 49:53-63. BATS FROM NICARAGUA 35 Tuomas, O. 1895. On small mammals from Nicaragua and Bogota. Ann. Mag. Nat. Hist., ser. 6, 16:55-60. 1905. New Neotropical Chrotopterus, Sciurus, Neacomys, Coendou, Proechimys, and Marmosa. Ann. Mag. Nat. Hist., ser. 7, 16: 308-314. VALDEZ, R., and R. K. LaAVau 1971. Records of bats from Honduras and Nicaragua. Jour. Mamm., 52.:247-250. WENZEL, R. L., V. J. Tieton, and A. Krew icz 1966. The streblid batflies of Panama. Pp. 405-675, in Ectoparasites of Panama (R. L. Wenzel and V. J. Tipton, eds.), Field Mus. Nat. Hist., Chicago, xii+861. UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly pub- lished in the above series are now published as Occasional Papers, Museum of Natural History. The Miscellaneous Pub- lications, Museum of Natural History, began with number 1 in 1946. Longer research papers are published in that series. Monographs of the Museum of Natural History were initiated in 1970. Institutional libraries interested in exchanging publications may obtain the Occasional Papers and Miscellaneous Publica- tions by addressing the Exchange Librarian, University of Kan- sas Library, Lawrence, Kansas 66044. Individuals may pur- chase separate numbers of all series. Prices may be obtained upon request addressed to Publications Secretary, Museum of Natural History, University of Kansas, Lawrence, Kansas 66044. m3askMNeoe . AR Hf Ny JUN @ = 1971 ie AE ERS Ne pp RECEIVED oo of the = «MY OF STA MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas NUMBER 3, PAGES 1-62 MAY 26, 1971 QUANTITATIVE ANALYSIS OF THE ECOLOGICAL DISTRIBUTION OF A TROPICAL HERPETOFAUNA By MartTuHa L. Crump’ INTRODUCTION Possibly the rainforest environment is not so constant, equitable, and predictable as ecologists have assumed. Lloyd, Inger, and King (1968) suggested this possibility as a result of studies on amphibian and reptile diversity in tropical rainforests of Borneo. The ways in which species utilize environmental resources have long been of interest in ecology; recently some effort has been made to analyze the inherent properties of the rainforests as they relate to amphibians and reptiles. Schoener (1970) studied nonsynchro- nous spatial overlap of lizards, genus Anolis, in patchy habitats in the West Indies. Schoener and Gorman (1968) studied niche dif- ferences of three species of Anolis from the southern Lesser Antilles; Schoener (1968) also studied resource partitioning among anoles on South Bimini Island. Rand (1964) examined the ecological distribution of anoles in Puerto Rico. Rand and Humphrey (1968) studied ecological distribution and interspecific competition among lizards in the rainforest at Belém, Brasil. Duellman (1967) studied isolating mechanisms and resource partitioning in tree frogs in Costa Rica. Inger and Greenberg (1966) studied the relation be- tween niche overlap and interspecific competition for three species of frogs, genus Rana, in Sarawak. As indicated, the majority of studies have been carried out on specific genera; no extensive, quan- * Graduate Student, Museum of Natural History, University of Kansas. SS Ke 2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY titative ecological studies have been carried out in the New World tropics on an entire reptilian or amphibian community. This is a report on the ecological distribution of amphibians and reptiles undertaken at the Guama Ecological Research Area near Belém, Brasil. Field work was carried out from mid-January through July 1969, in April 1970, and in June and July 1970. Part of the resultant collection was given to the Museu Goeldi in Belém, and part is catalogued in the Museum of Natural History at the University of Kansas. The objectives of the present study are threefold: 1) to deter- mine the ecological distribution of 62 species of frogs, salamanders, and lizards within the rainforest environment of Belem; 2) to ana- lyze the environmental parameters affecting the distribution of species; and 3) to compare and contrast the major areas with re- gard to species composition. The ecological distribution of the herpetofauna presented here is based on data obtained in one small area in part of the year. A similar study carried out from August through January or in a different area probably would yield some- what different results. DESCRIPTION OF THE AREA Belém is located about one degree south of the equator, in the lower Amazon Basin, Estado do Para, Brasil; the elevation at the highest point is 12 m above sea level. The mean annual temperature is 26°C, and the average monthly temperature varies less than 2°C throughout the year. Seasonality is reflected through the temporal distribution of rainfall, yielding wet and dry seasons. The average annual rainfall (44 years) for the wet season, January through June, at Belém is 2028 mm, whereas that for the dry season, July through December, is 830 mm (Belem Virus Laboratory, 1967 An- nual Report). Belém is the headquarters for the Instituto de Pesquisas e Ex- perimentagao Agronomicas do Norte (IPEAN). An area of about 310 hectares of IPEAN property has been designated as the Guama Ecological Research Area (APEG). Most of my study was carried out in two of the APEG reserves. The Aura Reserve is part capoeira, part terra firme (Fig. 1), and part varzea forest (Fig. 2); some areas are transitional between terra firme and varzea forests. The Catu Reserve is a transect of igapé forest (Fig. 3) 1000x200 m. See figure 4 for spatial relationship of the reserves. The forest types are defined below. The reserves are divided into a network of 10X10 m quadrats, each marked with a numbered stake. For each observation or in- ANALYSIS OF ECOLOGICAL DISTRIBUTION 3a Pai ae Ses . " ). Well-drained forest on relatively a reserve Aur ra firme forest ( high ground. Photo by Roger Arle. nie: 1;. Ter 4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Fic. 2. Varzea forest (Aura reserve). Flooded daily by the back-up from the Rio Guama; predominance of Acai palm trees (Euterpe oleracea). Photo by Roger Arle. ANALYSIS OF ECOLOGICAL DISTRIBUTION 5 Fic. 3. Igapd forest (Catt reserve). Permanently flooded forest. The boardwalk provides easy access. 6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY BAIA de GUAJARA RIO GUAMA Fic. 4. Map of Mocambo and Aura Reserves in relation to Belém and the Rio Guama. The Mocambo Reserve consists of terra firme forest, surrounded by the Catt reserve of igapé forest (not indicated on map); the Aura Reserve consists of terra firme, capoeira, varzea, and transition forest areas. Some studies were carried out in the vicinity of the IPEAN headquarters and at the Agua Preta Reservoir ( Utinga Reserve ). dividual collected, the hectare and quadrat numbers were recorded, thereby assuring that all data were collected in the same spatial frame of reference. The distribution of water was determined and mapped for the capoeira, terra firme, and varzea study areas (Figs. 5-9); species distributions were superimposed on these maps to determine the associations of species with standing water. For the various quantitative analyses, 44 sampling plots, each 20X30 m, from four of the major forest areas were studied. The location of the 4 capoeira-terra firme transition, 19 terra firme-varzea transition, and 9 varzea plots relative to each other and to the distribution of water is shown in figures 10 and 11. The other 12 plots were in the igapé forest. For the purpose of an analysis of ecological distribu- tion, the rainforest at Belém was divided into seven major areas: Terra firme forest—Well-drained forest on relatively high ground that is never subject to flooding is called terra firme forest. It is a well-structured, complex, tropical rainforest. One 5.5 hectare area of terra firme forest (Mocambo Reserve) has been studied extensively by botanists. Cain et al (1956) found the area to be extremely complex, both in vegetation species richness and in vegeta- tion density; they estimated the density of trees exceeding 10 cm ANALYSIS OF ECOLOGICAL DISTRIBUTION 7 rl ale [4 | | @ Kd He 7 tial 1 EeGZe, BEAR AZE Ee TT | Oe @ [i mie | pve | Beeeseie! | | tt LOS 4 Zi) eS aS NT CNS WEN _U SESS ee meme TT | v4 ie Se eS ee 4+——+. . ANN WN e NS Z| : i DD ea RSE es opel Z EPA Sy eS PRD PA SES G BaSaB Ban Zeeaee BSS SET ao AN IEDR TBE EAP | ae | PCr D EY) ay x SD CON PNP OPN S RY PA | MS Pel neahed iG Ria BNt¢ RTI RY (RSD 07 Sas es aan : te / = = y TD) bs Semi Eta CSE Le Gee Bae ees ae eles ENS @) ae per pT Nee Tc NS Bae Pan rea } COLE er SSSRE Dey) 216S: | ort Ne bt Rae PAR PRED ebebetabke Mh | RT Fic. 5. Distribution of Bolitoglossa altamazonica in relation to distribution of water in terra firme, capoeira, and varzea transition forest. Each small square represents a quadrat, 10 x 10 m. Cross-hatched quadrats are those areas in which at least one frog, salamander, or lizard was observed by the author. Stippled areas represent terra firme-varzea transition depressions filled with standing water; non-stippled areas are better drained and usually are located on higher ground. Each dot represents the observation of at least one individual of Bolitoglossa altamazonica within the particular quadrat. ( in diameter to be 594 trees per hectare. Dr. Murca Pires, a botanist associated with IPEAN, identified 215 species of trees in this area. Hatheway (1967) estimated the canopy to be 80 percent closed, with an average canopy height of about 35 m. He distinguished three strata of vegetation. Beneath the nearly closed canopy is a 8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY SeRee Lzeae ih care Baec Bee BE-3)/SRRREERED eet ae=93 Ee py Has ie Soe geese elordegart a: teseet es oes ai SEEPS 708 2 CRRRREs Se ee BRN be Sie Sask BERS ee Bes pal | te ea tt ce Be et ta ee es Ve EN [LT BR DK oR SS SSSR Se posece Samael SERRE SP aed rt ES aeD SNe BRLSe Rhine S Sete Te [A le Sst ia Ora eeey AURORE Ea tart 14 iealhe] aH eee ci a ae wl@ cae shy PaRsascasenancnn tas ee (ra SAPRCL ONS PE . Ss ope seeeaeermooom 5s oa DE SS SERS BS SKS ZL BeReS tS aor mew Ee] et} is YI EL ERE SS Lt eT TY Sia ce st StF | |S tard asses WEE Haase EC AS Cemes) Ep a Seritisac acces SMS 8 SS a is BN ROD focal a | sf + POT Be saseeculee ra 1 Kop & ata Rea t= Ros SB Hite Re is “Bes w fe BIA “a! BS NY a an LN on ‘8 || I SS |_| a P~ = stad & Lae. | Lp MN EE Ore} cS D ald red As Ges a N las ree. MEWS Vie ees am ‘Ge | RESNNVENNY Pee ES BERSA2 SRS oO Sl TRS a Tee P | Faia ORD at | Oh a 4 ‘a aa etn | be. BEPss PN eget et te er stele aisles O Feo etc] | Bleed G2 RE SB RBI EK Bb. Be mat pall teh bg be Beth 41 OL pe Se HAS LACE A ARRIOLA é S ZEnnns ENCEUCN Ett are KY m ad a IN te NW eT TT IND NY NT LT Be Jena = 4 pepe ot SI SR et Lo SLSR REE RS Ds BS SS = hi a VERREERISS SS SAREPRS ae SNA URS aS SS Be Esper T TTT eh IN Et RS CEES? 2a Versace“ Ate Pa NSSSENNT NY SAIN TTT RINT Bie + oo 2s ae Bed aS Sie SB BINS RS CRAG PTNERY [TT ESE ESR (Be BSS FO Bs BS SEBS es SS Ef ARB BES aS? epsee Fic. 6. Distribution of Bolitoglossa altamazonica in relation to distribution of water in varzea forest. See figure 5 for explanation; in this figure stippled areas represent varzea depressions filled with standing water. deep layer of trees up to 20 m in height; the bottom, dense, scrubby layer extends to a height of about 1.5 m from the ground. Varzea forest—Swamp forest bordering the rivers is known locally as varzea. This forest is flooded daily by the back-up of the Rio Guama, due to tidal effect. The degree of flooding varies throughout the year and is correlated with rainfall. All aquatic environments in the immediate vicinity of Belém seem to be fresh- water (Humphrey, pers. com.). The “white water,” so called be- cause of the presence of sand, silt, and clay particles, yields a con- EE ANALYSIS OF ECOLOGICAL DISTRIBUTION 9 NNANY =] SRE N BREGANGNN No LN : an NOHO N fee : biel i : ee i SSG PRE a Ge hVaekas NN \ ‘ze | he N ali e aie Se ¢ GNA Cuma sss Digs ARS Rb 1 ges ieee iSite OF i Cd et 4 ( CRE LENABE micss eaaers ONO CEB: PSL tp | | te Cal 4 aK BN 4 ral = fx IC) : CHEE Buia 1: 1 OF Ee a Nt CEN JEG ee Ve SIT ES CTS Se SNE lay ew eS BEBSeom Teepe rear MASSee Fic. 7. Distribution of Leptodactylus marmoratus in relation to distribu- tion of water in terra firme, capoeira, and varzea transition forest. See figure 5 for explanation of symbols. tinual deposition of alluvium. The resultant alluvial varzea soil is rich, but has a low permeability. During the rainy season, parts of the varzea are flooded to a depth of 1 m or more. Depressions are present, resulting in differential drainage. Tall woody plants, palms, and giant aquatic herbs exist nearly side by side as a conse- quence of drainage patterns (Hatheway, 1967). There is a pre- dominance of palms in the varzea forest; the acai palm (Euterpe oleracea) is the most common tree. Lianas and epiphytes are com- mon, and moss as thick as 1 cm covers the trunks of trees up to 2 m 10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY SRBSRESERRESSEE See: GR aS SR SSSR RES. See Ae ZC a ge A0aa A 4 = aE ot C Con Fer se ED PA TS } Bees emeeeiennes SERS rd (a i Ba te nis ey ms HOSS N Wet 1 5 a4 Ds tod ON C a \ oa si i “nas Nee LAS : FY A Set ces SEEEE a ioe Be mM A} Wey sai =a a Perego S50NUENS SRS pe a a i an am. 4 am. 18 q = ' H ‘ee ro. =! t (oe Es pp ae eee it | a tI | ae Ss aS : Petal |b Bera | PE Pa MD 4) NAT, 7 core Bass RRR ep Pee i] ne iN Wet t ia Bel Fic. 8. Distribution of Gonatodes humeralis in relation to distribution of water in terra firme, capoeira, and varzea transition forest. See figure 5 for explanation of symbols. SSSR ea Oaths sw aaa iOT SS GAR Ieee ae: q Teer fn |e BARNABAS ( from the ground. Hatheway (1967) proposed that epiphytic mosses likely indicate high humidity in tropical forests. In the varzea, this high humidity probably results from constant evaporation from the moist ground and water. Hatheway estimated that the total density of trees over 10 cm in diameter is probably greater than 600 trees per hectare. The canopy is about 50 percent open, and the canopy trees are 30-35 m in height. Igap6 forest—This is forest that is permanently flooded with “black water,” so called because of organic residues. Hatheway | { > ———————— ANALYSIS OF ECOLOGICAL DISTRIBUTION ie mag PS fl A PIES [era ; De RIERA Re a RUT | RS ER G a ZEEE SVaRN Seas iss x Mog Awa BeNareet alee eee seit me Ae RRS ecu ae pS (alaed eo Cee eee ly K U | es | Sec esee Te ee 4) SESS oN be et | LPT sett LAroT tel rt a, a ‘ 3 rane i 2 eee ee ous aoe a}? eons Sores Ni Zee aes SO aos & cepeuee Ammen giimics aes oe (es MS Oe KP PN Siac BORE aos eh! my x. r aM DSTSSCSSRBIBE YS SEs ARN ax Si Pt RT od ao: Seats BSCS RSS RS SIRE BS Sat” Ed ae XQ | | ee N Nee TSN Rah Cae Ree fa] | Tey | lea BS STL bites Lea Lea ane PEEL SEER ERE h cp Sg MASE el ek [a poee2 CY NSESKNN Pett Bree eee Fic. 9. Distribution of Gonatodes humeralis in relation to distribution of water in varzea forest. See figure 6 for explanation of symbols. (1967) estimated the average depth of water to be 25 cm; beneath the water is another 25 cm of organic, water-logged muck, under- neath which is white clay. The area consists of many stagnant, foul-smelling, interconnected pools. Small islands of root masses project from the pools; much of the vegetation in the swamp forest is supported on these islands, although a few trees are rooted in the muck. There is no well-formed canopy, and other distinct vege- tational layers are difficult to distinguish. Most trees are small- crowned, slender dicots, rising above the thick mesh of tangled roots elevated to 3 m above the deep mud of the swamp. 12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY DOGO SBeSSaREeGelaeZg Beate Siete sieiai Ze 2 of oe Capoeira- 2 CA sGeREC ‘rt Terra Firme 22860 eehgeeeer <= sie : FSBURBEREGESETE Transition SSSREenEEs ATT Past talc cole alata SSSS585 858570 Rennes relseatZat a ONS SENENAUNEE IN [| a Terra Firme- _ Varzea dau Transition (Dry) | tea pi eee LKA EN LAN F= gS ae Aap Se a f Terra Firme- = 8 Varzea abba + (Wel) -- | to NG eS LAetot ee icy pera tk TEanCenaae . AS Ropes: x Pst Bere me .“s (RUEB TNITR BERP Ss NOPRss NC EY] iS EDpeSr rer THE SB Es CS EP | P| PZaR ea aise 2) SN A MN 258 5: BEE x sot ate a |g al_| ¢ SBBR i & COC EEE Pees a See Re LH 4 wee SSee hh iP . oe aa Eq B FCCECCCC REE BEPARRS Lp php 3 Fic. 10. Distribution of sampling plots in relation to distribution of water in terra firme, capoeira, and varzea transition forest. See figure 5 for explana- tion of symbols. The numbered plots, each 20 x 30 m, are indicated by heavy, straight lines; data from these plots were used in the contingency table analysis. Capoeira forest.—This is second growth forest on well-drained ground. The capoeira areas studied had relatively open canopies and fairly dense ground cover. Much of the area is composed of tall grasses and ferns; the forest floor in some sections is covered with brush and fallen logs. Capoeira-terra firme transition—Four plots were studied which are intermediate between capoeira and terra firme forest with re- ANALYSIS OF ECOLOGICAL DISTRIBUTION 13 SRRR NERA U7 2 eee He SRSRUE. CeI83 Be LU] Pees et Bap ta |_| tte 83/0 233 Bs Ant eT pes At et Cte | » le Sec ee coeaenl y | MELA a acl ree Nee | TT Cee ae EA = ea Ht es BRARE SSR = CRT NS DOSEN Eee Pt ty eraser TT epee Trae nS ih Ta pila § ee ete AT TTT Tea ee peti Tg Seenet gos Sueunce eeyserati cy aU a: PL IetD ty Nee Pe Canecmnn coh GSPSeS eet eS ie a Bale Pte teres Tacs Sra a SEP ace Bas ot Pi pe tt tt ett fe PN be A Sor = BAT a ee Big DEEe Cee LLL LLL eee ele Cd RS SBZpe as ee 3) GH aeee GEER Reed SACS el Sask PB SR See Bt OLR SR SnMmresckes tego) PPT TT oletcbolober okey | yy Se eee eee occas Sone ace MLE OA sk 7 a DB SR ZBo JUGMBGue Ese cues mam He RP NST Et =e. yes BT ea ed I ES en iz is Ia Bis] | aaa z oe fle yd 8 eis 2 Ei Saaaa tT er Pare aE Brs SEEBP Beer yes & feerec A CSS Poe ee at Sey Ae o7i ss eesduauaaeer ame PAS PEON te at nee PERCE EPP GT RE SENS Sms WS Aes sass Eta bit gle ! ar TA. . (V7 Naeger Res Cat pes Pa REP LT VAST ESE i T SAM RMS SENMPISE PEER IEPA Ped PEELE Dee te SSeS ee Fic. 11. Distribution of sampling plots in relation to distribution of water in varzea forest. See figures 6 and 10 for explanation. gard to characteristics of canopy and ground cover. The plots ex- hibit a greater vegetation density than typical capoeira, but less than typical terra firme forest. Terra firme-varzea transition—Plots in one area exhibit some characteristics of both terra firme and varzea forests, but differ noticeably in other ways. For instance, on well-drained ground there is a lower vegetation density than in typical terra firme forest. The flooded portion lacks the predominance of palms, characteristic of typical varzea forest. Corresponding to the varzea and terra firme forests respectively, some of the soil in the transition area is 14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY a poorly-drained, silty-clay alluvium, whereas other soil is a well- drained, heavy, yellow laterite. Open and edge.—All non-forest study sites are grouped in this category. Observations and collections were made in swampy areas in open fields, ponds along forest edges, and in second growth vege- tation bordering the IPEAN reserves. During the rainy season, the swamps and ponds contained water to a depth of about 1 m but were usually less than half as full during June and July. Dirt roads and roadside ditches on IPEAN property and sewage swamps within the city were examined weekly. COMPOSITION OF THE HERPETOFAUNA Three orders of amphibians: Gymnophiona (caecilians), Cau- data (salamanders ), and Anura (frogs) and four orders of reptiles: Amphisbaenia (amphisbaenids), Crocodilia (crocodilians), Squa- mata (lizards and snakes), and Testudines (turtles) are represented in the herpetofauna (116 species) of the Belém area; amphibians represent 35.4 percent of the herpetofauna, and reptiles 64.6 percent. The breakdown of species is as follows: caecilians—3, salamanders —l, frogs—37, amphisbaenids—3, crocodilians—l, lizards—24, snakes—44, and turtles—3. Further field work probably will reveal several additional species of snakes, caecilians, and turtles, as well as species of other groups. METHODS Most observations and collections in the forests were made along paths and boardwalks constructed several years previously. There- fore, the data are biased to whatever extent the different species are influenced by the narrow, open areas maintained by continuous human activity. Species distributions necessarily reflect my sam- pling activity (Figs. 5-9). An approximately equal amount of field work was done by day and by night. Every frog, salamander, and lizard observed was recorded by species, date, and locality (including hectare and quadrat numbers from the labeled study areas). The distribution of each of the 62 species was plotted on quadrat maps. Although snakes, turtles, and caecilians were collected, the few numbers of specimens of these groups precluded their inclusion in the analyses. Environmental gradients affecting the distribution of species within four major forest areas was inferred by use of a contingency table analysis. Resource partitioning was studied by means of field ob- servations and analyzed by niche breadth and niche overlap anal- ANALYSIS OF ECOLOGICAL DISTRIBUTION 15 yses. Following the analysis of species distributions, the species compositions of the major areas were compared and contrasted by means of the Shannon species diversity formula, an equitability in- dex, and coefficients of communities (see appropriate sections ). Definitions of terms, as I am using them, and a brief discussion of techniques of analysis are given below. The analytical techniques are treated in detail in appropriate subsequent sections of this paper. Major areas.—The region studied can be divided into several geographical sections referred to as major areas. I have delimited the artificial boundaries in such a way that each area possesses a certain subjective uniformity with regard to physical environmental parameters, such as vegetational physiognomy, light intensity, water, and soil type. The quantitative analyses were carried out on data obtained from four major forest areas: 1) capoeira-terra firme transition; 2) terra firme-varzea transition; 3) varzea; and 4) igapo. Resource partitioning observations were carried out in the following major areas: 1) open and edge; 2) capoeira; 3) terra firme; 4) varzea; and 5) igapo. Habitat.—This term refers to the structural aspect of a niche; it is that portion of the physical environment in which an organism carries out its life processes. The physical environment supports species in three major ways: 1) vertical zonation; 2) horizontal distribution; and 3) temporal spacing. Community—A community consists of interacting populations of animals. Each of the major areas included in this analysis has a herpetofaunal community different from every other area. The interaction and organization of each community is expressed in terms of resource partitioning with regard to differential utilization of the environment in space and time, species diversity including both species richness and equitability components, and species com- position and relative abundance. Resource partitioning —This term refers to the differential utilization of the physical environment in space and time by dif- ferent species. The result of resource partitioning is highly efficient utilization of environmental resources. Niche.—This is an abstract concept referring to the habitat and biotic relationships of an animal. A niche can be thought of as a hypervolume, consisting of numerous dimensions (Hutchinson, 1957); the dimensions are physical factors and biotic relationships required by a species for survival. The physical factors of the en- vironment making up the structural component (habitat) of the niche exist independent of the species, but the entire niche, inclusive of the position (biotic relationship) of the animal, does not exist 16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY independent of the species. Therefore, the niche is a function of the species. Formation of a particular niche is ultimately dependent on the structural adaptations, physiological requirements and capabilities, and correlated behavioral patterns of the species. No two species have identical physical and biotic requirements, coupled with identical structural, physiological, and behavioral attributes, and therefore no two species have the same niche. Niche breadth.—This term is used to describe the spectrum of any given dimension of the niche hypervolume. For instance, one can speak of the food preference niche breadth of species A and B. If species A eats 10 different kinds of insects and species B eats only 2 kinds of insects, species A is said to have a broad food preference niche breadth and species B a narrow food preference niche breadth relative to each other. Niche breadth as used in this paper refers to the habitat niche breadth. Niche overlap.—This term refers to the situation in which two or more species have similar requirements with respect to some dimension of the niche hypervolume. Niche overlap is a measure of the association of two or more species. The measurements in this study were obtained indirectly by the degree of coexistence of the species in the various plots sampled. Habitat-generalists, intermediates, and specialists —Habitat-gen- eralists are species that utilize a broad spectrum of the environment, as indicated by the contingency table indices; they are found in all four major forest areas and have high habitat niche breadth scores (16.0-32.0), as calculated from Levins’ index. Habitat-specialists are species apparently restricted in their distributions to one or two of the major forest areas; they seem to live in a narrow range of the environmental spectrum and have low niche breadth scores (1.0- 4.0). All other species are referred to as habitat-intermediates. In most instances, the habitat-specialists are the least common species, whereas the habitat-generalists are the most abundant. Species diversity—The concept of species diversity consists of two components, species richness and equitability. The former is the number of species, and the latter is the evenness with which the individuals are distributed among the species. A community having a large number of species in which the abundance decreases grad- ually from the most to the least abundant species is considered to have a high species diversity. According to Whittaker (1970), niche differentiation results in greater species richness through time, whereas a narrowing of habitat distributions tends to increase spe- cies equitability. Some investigators propose that species richness depends primarily on the structural diversity of the habitat, whereas ANALYSIS OF ECOLOGICAL DISTRIBUTION U7 equitability is more dependent on the stability of physical condi- tions. Apparently the more complex the vegetation is vertically, the greater is bird species diversity (MacArthur and MacArthur, 1961; MacArthur, MacArthur, and Preer, 1962; MacArthur, 1964, 1965; and MacArthur, Recher, and Cody, 1966). Pianka (1967) proposed that spatial heterogeneity is the most important single factor determin- ing the number of species of lizards in any given area. One of the most commonly accepted formulas to measure species diversity is the Shannon function (Shannon, 1948). Pielou (1966) discussed its use and disuse. The formula is used to describe an infinitely large population and results in the average diversity per species. Coefficient of community.—The coefficient of community (CC) is a mathematical measure of relative similarity of samples from two communities (Whittaker, 1970). ANALYSIS OF ECOLOGICAL DISTRIBUTION As discussed in the preceding section, each major area is a complex of intrinsic physical environmental parameters, different from those in other areas. Each species is adapted to a particular range of each environmental gradient; the totality of environmental gradients forms the structural niche, or habitat, of the species. One must assume that habitat adaptation is based on the genetic make-up of the individuals of the species in terms of morphology, physiology, behavior, and life cycle. Based on the preceding assumptions, the following hypothesis can be stated: The 62 species of frogs, sala- manders, and lizards in the Belém area are distributed in such a manner that environmental resources are partitioned; the conse- quence of habitat differentiation is highly efficient utilization of the environment. Several techniques were used to study the ecological distribution and to test the hypothesis; others were used to compare and con- trast the species composition within each major area. To avoid con- fusion, each analysis is presented separately. Included in each sec- tion is an explanation of purpose and a presentation and discussion of results; where applicable, advantages and limitations of the anal- yses are indicated. The distribution of frogs, salamanders, and lizards as taxonomic groups within five of the major habitats is presented in table 1. The varzea has the highest species richness, with 38 (61.3%) of the 62 species occurring there. Next in terms of species richness is terra firme forest, with 36 species (58.1%). The area with the lowest value is capoeira, with only 20 species (32.2%). The mature forest 18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY TABLE 1.—Distribution of Amphibians and Lizards by Major Taxonomic Groups in Five of the Major Areas. The top numbers are the number of spe- cies of a taxonomic group in a given area; percentages are the proportion of the taxonomic group in the area. Numbers in parentheses indicate the sum of the coded relative abundance indices (Table 2) for the taxonomic group in the area. Terra Open & Group Firme Varzea Igapd Capoeira Edge Frogs and toads _______ 20 22. 13 8 24 ov species’ 22.2 4. Ux 54.0% 59.4% 35.1% 21.6% 64.9% (36) (56) (23) (12) (80) Salamanders «ee IL i 1 1 0 ISDeECles ee 100% 100% 100% 100% (4) (4) (2) (3) ees EUAN ke eR ee 15 15 8 Lt 10 DA species =e 62.5% 62.5% 33.3% 45.8% 41.7% (27) (29) (16) (19) (23) Total No. Species _- 36 38 22, 20 34 % Total Species (62) _.. 58.1% 61.3% 35.5% 32.2% 54.8% Sum Abundance Indices 67 89 4] 34 103 Average Species Abundance Index ___. 1.86 2.34 1.86 1.70 3.03 areas likely are highest in species richness due to the greater vegeta- tional diversity, yielding environmental heterogeneity, as contrasted to second growth areas (capoeira) having less structural complexity. Open and edge areas are relatively rich with 34 species (54.8%); 24 species of frogs (62.3% of the total anuran fauna) breed in the numerous ponds in these areas. Abundance indices for each species in each area were coded as follows: O=apparently absent (none observed); 1l=not commonly seen (1-4 observations); 2=moder- ately common (5-15); 3=common (16-25); and 4=abundant (26 or more observations). The average abundance index (obtained by dividing the sum of the abundance indices for all the species in a given area by the total number of species in that area) is much higher in open and edge areas (3.03) than the next highest which is the varzea forest (2.34); this is partially due to the large congrega- tions of breeding frogs in open and edge areas. In addition, popula- tion densities of lizards are higher in open areas than in the forest, although this may be due to censusing methods; lizards are more easily seen in open and edge areas than in the dense forest. The ecological distribution and relative abundance of each species of frog, salamander, and lizard are shown in table 2. It is evident that certain species have a much broader range of ecological distribution than do others. Figures 5-9 indicate the distribution of three species relative to the distribution of water. The salamander, Bolitoglossa ANALYSIS OF ECOLOGICAL DISTRIBUTION 19 TABLE 2.—Ecological Distribution of Frogs, Salamanders, and Lizards. Num- bers indicate relative abundance of a species within an area, coded 0-4 as follows: O=Apparently absent (none observed), 1=Not commonly seen (1-4 observations), 2=Moderately common (5-15), 3=Common (16-25), and 4=Abundant (26 or more observations ). Terra Open & Species Firme Varzea Igapd Capoeira Edge Beppe 0 I 0 0 0 Eleutherodactylus lacrimosus — 1 0 1 0 0 Leptodactylus marmoratus __- 4 4 0 3 0 Leptodactylus mystaceus __... 1 0 0 0 0 Leptodactylus ocellatus __.. 0 0 0 0 4 Leptodactylus pentadactylus — 1 0 0 0 0 Leptodactylus rhodomystax _ 1 2 0 0 0 Leptodactylus wagneri _______- i 4 4 0 2, Physalaemus ephippifer 4 4 ] 1 4 Physalaemus petersi _______. 2 4 0 0 2 maypormarimnus 0 0 0 0 4 Pao wmpnonus 4 4 0 2 0 Dendrobates trivittatus 2 0 0 i) 0 Dendrobates ventrimaculatus _ 0 2 3 0 0 Hyla baumgardneri _ 0 0 i 0 4 iia Dogsemani 0 0 0 0 4 OT _ i! 3 0 0 0 OD CG _—_—__ ie 2 3 2 ib 4 Wiyla geographica ___.__.__.___. 0 4 0 0 3) i _—_ a, 1 2 1 0 4 lye eranosa 1 2 3 0 0 Hyla leucophyllata ________.___ 0 2 1 0 4 Hyla melanargyrea __... 1 0 0 0 4 SUNT __——_— 0 0 0 0 4 Hyla multifasciata 0 2 1 0 4 MGTNGHG fo 1 0 0 ) 4 flamaniceps 0 i 0 0 4 vila rondoniae .._____.____._ 0 1 0 0 0 MN) ies i! 3 2) 1 4 gla sp. (large rubra) ___ 1 1 0 0 4 fuia sp. (rubra-like) —____.... 4 4 il 2 i) Osteocephalus taurinus __.- 0 0 0 0 i Phrynohyas venulosa ih 0 1 i 3 Phyllomedusa bicolor __...... 0 2 0 0 2 Phyllomedusa hypochondrialis 0 0 0 0 4 Phyllomedusa vaillanti ___. 0 i 0 0 0 Sphaenorhynchus eurhostus __. 0 0 0 0 3 Bolitoglossa altamazonica _____- 4 4 2 3 0 Gonatodes humeralis _.-_- 4 4 5 3S 0 Hemidactylus mabouia _.__.-.- 0 0 0 0 4 Thecadactylus rapicaudus __- 1 0 0 0 0 Lepidoblepharus festae __._. 0 il il 0 0 Anolis fuscoauratus __._...-.-.---- 3 3 1 1 0 mois ortoni 9 1 0 0 0 0 manos punctatus if 1 0 1 0 fiana iguana 0 I 0 0 i Per umorg = 3 1 i 2 0 Polychrus marmoratus _____- 1 1 1 il 0 20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY TABLE 2.—(Concluded) Terra Open & Species Firme Varzea Igapd Capoeira Edge Tropidurus torquatus _______. 1 0 0 1 4 Uranoscodon superciliosa ____.... i! y} | 1 ] Mabuya mabouya ____-.-- 3 4 4 3 2 Alopoglossus carinicaudatus __. 0 i! 0 0 0 AMEICG GMeIOG —..2 ee if 0 0 3 4 Arthrosaura kochii __.-_-- ee! 0 0 0 0 Cnemidophorus lemniscatus _.. 0 0 0 0 3 Crocodilurus lacertina __.___. 0 1 0 0 0 Dracaena guianensis __._.___.__.. 0 0 0 0 ] Kentropyx calcaratus ______.. 4 4 4 2) 0 Leposoma percarinatum ____- 0 2 0 0 if Prionodactylus argulus ______.- 1 2 0 0 0 Tupinambus nigropunctatus _.. 1 ] 0 1 2 altamazonica, is abundant in capoeira, terra firme, and varzea areas, but few individuals are found in the terra firme-varzea transition area. Leptodactylus marmoratus occurs predominantly in well- drained areas, but Gonatodes humeralis tolerates wet and dry areas. RESOURCE PARTITIONING Extensive field observations were carried out in an effort to discern whether species do indeed partition environmental resources. Resources examined were those aspects of the habitat which are im- portant to the daily and seasonal activities of the species as follows: frogs—standing bodies of water, calling sites, and vegetation and ground area used for daily activities; salamanders—vegetation used for nocturnal activities; lizards—vegetation and ground area used for basking sites and other diurnal activities. For purposes of analysis, the environment can be divided into vertical and horizontal components such as arboreal (high and low), terrestrial, aquatic margin, and aquatic. The distribution of species in these subdivisions of each of the five major areas is shown in table 3. Most species studied are either low arboreal or terrestrial. TABLE 3.—Distribution of Species of Frogs and Lizards Within Subdivisions of Five of the Major Areas. Numbers preceding hyphens are frogs, and num- bers following hyphens are lizards. Terra Open & Subdivision Firme Varzea Igap6 Capoeira Edge Arboreal (high) 0-3 1-3 0-1 0-2 ]-1 Arboreal (low) ~..- 9-6 11-6 9-5 4-4 16-2 Miermestmal =) a. 9-6 7-5 3-2 4-5 4-7 Aquatic Margin ____. 1-0 2-0 1-0 0-0 3-0 PGT On hee ee ees 0-0 1-1 0-0 0-0 0-0 ANALYSIS OF ECOLOGICAL DISTRIBUTION 21 Spatial overlap among some species does exist. Salamanders and Hyla sp. (rubra-like) overlap greatly in their utilization of low vegetation in terra firme and varzea areas at night, presumably for obtaining food; also present, sharing the same vertical component, are numerous sleeping lizards (Gonatodes humeralis). Although these three species are the most abundant vertebrates using this aspect of the environment at night, the population densities appear to be so low that it is unlikely that significant interspecific competi- tion exists. There is evidence of breeding site partitioning in frogs, probably indicative of differing requirements of various species. Some tree frogs, such as Hyla baumgardneri, H. egleri, and H. goughi breed in diverse types of ponds and swamps, large or small, deep or shallow; apparently the frogs require only standing water and emergent vegetation. On the other hand, Hyla minuta, H. raniceps, and Phyllomedusa hypochondrialis are found in only some of the same areas as H. baumgardneri, egleri, and goughi. Hyla raniceps breeds only in larger bodies of water, at least 8 m by 15 m, usually at least 0.6 m in depth. Phyllomedusa hypochondrialis is restricted to ponds bordered by dense vegetation. The distribution of H. minuta is more difficult to interpret; the frogs occur in all types of areas, but without any regular pattern. For instance, numerous males call from one pond and not from a nearby pond having similar size, water depth, and emergent vegetation. The population density of this species appears to be lower than those of H. baum- gardneri, H. egleri, and H. goughi. Perhaps male H. minuta attract other males to an area for the purpose of forming breeding congrega- tions. This formation would be of greater importance to a less abundant species than to a more common one and would explain the fact that usually these frogs call in groups of at least 15 in- dividuals in contrast to H. baumgardneri, H. egleri, and H. goughi which often call in groups of 10 or less. Many species of frogs which breed sympatrically demonstrate calling site segregation (Tables 4 and 5). Most species characteris- tically call from a certain physiognomic type of vegetation, at a relatively uniform height from the water. The type of vegetation utilized is correlated with the body build and size of the animal. Large, heavy frogs generally call from the ground, sturdy vegetation near the ground, or from branches of trees; small frogs usually call from grass stems or leaves and small branches from emergent and edge vegetation. Some species have a broader range of calling sites than do others. For example, Hyla goughi commonly calls from both emergent and edge vegetation, 0.05-1.5 m above the water, OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 22 OOOO =a sg ae XXXXXXREKMKK =o a= SO ee SiS ici ae ie See DD hydoonay maT Kk == == BOOK KN em a a ee Sy ee HORE SRA NOL Ca Se bupu opi XXXXXXXXXKXKX XXKXXX XXXXXX XXXXXX XXXXXXXXXXKXK Bila ne Reg a a ae SSO XXXXXX XXXXXXXXXXXX XXXXKX XXXXXXXXXKXKK XKXKKKK ORE eat OL aa, ee pee sane ATE XXXXXX XXXXXX XXXXXX XXXXXXK XXXXXXXXXXXKXKXXKK Lee a ae Oe ee i uajsa DRT Shi el MXX XXX XXKKKK XXKKKK KAKA SO = DORGIOBOA MOT“ suputasaog DihTy BODO SOOO KOON KK Ooo MOL a he” Waupwounng vA i. ee ae XXXXXXXXXXXXXXXXXXXXKXXKK — HHH WOHVIOSIA MOT Y punois ———————————— Dy sofynu DAH XXXXXXXXXXXK —-—-—~ OOOO = Ge ee ae WOHIOZIA OT 3B PUNOIG (Diqns ose) “ds phy OL ®D2COL®SESOS4.©,9,6,0,8,6,9,8,©,6.0,0.6.8:9,6.8,9.0.9.6.6 >. 6 >.> Camm UCHUIOSOA NOPE PUONOIS, mn ay = es puqns DIA XXXXXXXXXXXK —---—- OOOO OC ON ee es ee ee gen Te punoig snjoyja90 smhjovpozdaT cas": =. = XXXXXXXXXXXXXXXKXK XXXXXXXXXKKK OO punts bafiddiyda snwavjoshyd XXXXXXXXXXXXXXXXXX XXKKXXK XXXXKK XXXKKK KKK eS eae EE OTL Ajn{ eun[ ABW ‘dy ‘Ie ‘qoyq ‘uel OIG SuTeD sotoedg ‘Aj[eoruoryoudg pure Ayjeoryeduisg Ayjenieg jsvo] ye Suy[eD 10/pue Surpse1g sso1yq Jo soloedg—F atavy, ANALYSIS OF ECOLOGICAL DISTRIBUTION 23 whereas Phyllomedusa hypochondrialis always calls from edge veg- etation usually 0.6-1.5 m above the ground or water. Complete seg- regation of calling sites does not exist for all species in all areas. Segregation is partially dependent on species composition at the site, relative abundance of the calling individuals, and on the size of the breeding site relative to the population densities. Generally, in large, mixed congregations segregation tends to break down, and the frogs call from whatever sites are available. Interspecific com- petition for calling sites is probably significant during times of much reproductive activity. Segregation is more pronounced in large areas with distinct physiognomic vegetational diversity than in smaller areas with less calling site diversity. A commonly accepted explanation for the evolution of partitioning of calling sites is the resultant tendency to reduce the chances of interspecific mating. However, because segregation breaks down in large, mixed congre- gations at the time it is most needed, I propose that calling site partitioning exists due to the structural and behavioral attributes of each species rather than as a necessary reproductive isolating mech- anism; advantages likely include improved mating efficiency and reduced energy expenditure. There is a definite replacement of several species of tree frogs at breeding sites because of calling site overlap. Hyla boesemani, H. multifasciata, H. raniceps, and H. rubra all call from thick clumps of emergent vegetation, usually within 20 cm of the water. Individuals of all four species call from the same swampy areas, but not all at the same time; the only two of these species ever found calling sympatrically and synchronically are H. boesemani and H. rubra, the two smaller species. Every congregation of Hyla sp. (large rubra) observed was found calling in association with H. rubra. Male Hyla sp. (large rubra) call from the ground or low, thick vegetation. They seem to be dominant over H. rubra as indicated by calling site displacement of H. rubra when the two species call sympatrically. Hyla rubra usually calls from low vegeta- tion, but when Hyla sp. (large rubra) is also calling from the area, the former calls from higher vegetation. Perhaps some syntopic species (species with similar habitats) coexist with minimum interspecific competition as a result of temporal partitioning of the environment, in terms of diel and sea- sonal activities. For example, the nocturnal gecko, Thecadactylus rapicaudus, is likely the temporal replacement for diurnal lizards feeding on similar species of insects and utilizing the same habitat. The two species of dendrobatid frogs use the same forest floor by day that several species of leptodactylids utilize at night. Frogs demon- OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 24 | | Kl KKK | punoig xX = XE = — xX ye = 4 — ye = xX — xX = X — xX = xX = b4 — xX = »« — 4 — MOT ys U0ne}IBIA AOdH ® NAdO ee = = my onl = SR. ae ae: cae anes snjysoyina snyouhysouavyds a +3 as ss X ae Bel ee ae sypiupuoysodhiy vsnpawophyg = a ae = e Sy a ae ee 40]091q Dsnpawopjhiyug fe L = <4) a nee I a iar a a (Diqns o81e[) “ds nj = X Rs A X eS ee ae Digns DIA ». e< = ob oa op. ae Se a a ae oe pupu vjhyT rt = ee = X Saba Se ee ee DIDVSD{yINWUW DIA me 4 a = is ae OC) Fe ae ee ee ee pynure DAY N a = =. = eee eras ee ee ee Daihpivupjau Dj ae _ = ae = an, Wik a re a pypyhydoona) vii ma X i: = X PA a eS ae Se psouvis DAH al = a= — X ey ee Veet ee ee ee ee yysnos DA = = oe X X a el, eee a ee paiydbvisoas Djh = X = a X = tae aS a se ee uajsa DIA - a ae ee XY 20) casas eer Oe ae a DIDIDIIDI DIA es as es ee =s =) S SAikae es oe Se tupulasa0q DIA am X Za as e a Wisco ae oe ee aupipswunDpg vii = = ad a Bs 2, a ee eee snuupu ofng es = = X = =a) © Bisse ee ee isiajad snuapjpshyd a fe a X = ae ae eee ee dafiddiyda snwavjpshyg X = = X = ee er Re kee ee wausvn snjhyovpojda'yT = 2° a — = See et ae tee a snjpjja90 snjhiyoppo.daT 13 s Ee: X = NO GN | Bae as ae aes SnyDLOULIDUL smfhyovopozdaT MOTT ystH MO'T YysIH PO | (0001) sotoedg UOQe}IBIA UONL}ISIA OdVDOI VAHZHUVA ‘S801, JO Satoadsg jo so}Ig Suy[eD jo suluonnieg—'G alav], 25 ANALYSIS OF ECOLOGICAL DISTRIBUTION OG VG G VG VG OG Ajn{ Burjeo Jo sousscid=y ‘poururexs sojeuroy y[Npe OU=Z ‘ATUO UOT}JVAIOSqO=[ ‘SI Bpod VY, ‘*ssO1y{ JO sapoA—) SuIpsoig UMOUY—'g ATAV] O”d VG O OG V VG OG oun G OG VG OG OV VG G OG Avy VG a. i G OW Vv VG VG OG G OG OG OVG a Od VG - a gq =e OV d dd V dvVG - OV d CVG VG OW d G VG Od O”d VG d a Mh d vv G a G G OG OG VG OVG OVG O" O” Od O” OV a d Od O” ‘Idy "IVI mc 1m AN H <= IANA QAnNa | ra Sr! ‘uel a snysoyina snyaufysouapyds BE Hubywa vsnpawohyd Of-00) = sypipuoysodiy psnpawoyhyg ie ee Lojon1g vsnpawonhyg We pg ste psojnuaa sphyouhsyg ee (ayy-pugns) ‘ds phy (ee === (Digns aB1e,) “ds opi ies oS eee piqns DAH (poe sdaouo. DIT OOL0ee eee puou Dj (Cpe es eee Dypvospfyynu vphTy NOL0G = eee pynumw Diy te) eo ares vaihsivupjau DAH Goa) == pyoyiydoona, vjAH Of. ee psounis DAH gj © =e rysnos vA 00 4 Sa poiydois0as DAH igg4, 9 waza Dj Al pyDLvIIDI DAH (cps Soa Se tupuasaog DIA 4001 = woupipsunog vj oe 2 ee SNYDINIDUALIUaA SayDgoLpuad ce 2 ae ae SIDA. ~SajDgoLpuad NEN) =e snwuoydhy ofng (et =e snuupw ofng Come «ee a wiajad snwappposhyd ASGaia ie Tata Sas lafiddiyda snwanjppshyd 40 eee wauspm smhyovpojidaT SU ariphie ee $njpj]290 smfiyovpojda I Nod Sr sngosounU snjfiyoppojdaT N soroeds ‘satoeds jo sayrusant 10/pue sojodpe} peumsaid jo souaseid=q ‘seroeds jo saprueAn{ 10/pue sajodprz uMOoUy JO souVseId=p ‘Sul[eo pue (seyoynjO 330 10/pue sojeutay plavis 10o/pue sired Surxejdure) surIpeerq fo soUeplAe=gq ‘so[eul 26 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY strate seasonal partitioning of the environment in terms of breeding activities (Table 6). Hyla multifasciata and H. raniceps both call from low vegetation in swampy, open areas; they are rather large tree frogs with similar mating calls. The former is a wet season breeder, whereas the latter is a dry season breeder. Most of the frogs are nocturnal (83.8%) and most of the lizards are diurnal (91.7%); the salamander is nocturnal. In the terra firme and varzea forests, 50 percent of the frogs, salamanders, and lizards considered as a group are diurnal; the distribution in the igap6o forest is similar, with 45.5 percent diurnal and 54.5 percent nocturnal. Most of the species in open and edge areas are nocturnal (73.5%); breeding tree frogs account for most of this distribution. On the other hand, most of the species in the capoeira area are diurnal (70%); over half of these species are lizards, many of which are heliotherms (Table 7). CONTINGENCY TABLE ANALYSIS The contingency table analysis technique, developed by Wil- liams (1952) as an extension of Fisher and Yates’ ideas for dealing with frequency counts in two-way tables, is employed here for two reasons: 1) to measure the degree of association between species and plots; and 2) to partition the species-plot association into in- dependent components representative of environmental gradients. The analysis was carried out on 20 species of frogs, salamanders, and lizards from 44 sampling plots, each 2030 m, from the capoeira- terra firme transition, terra firme-varzea transition, varzea, and igapo areas. All plots received approximately equal amounts of sampling time from mid-January to the end of July. The 20 species were the only species of frogs, salamanders, and lizards found within the boundaries of the particular plots analyzed. The total sample in- cludes 1218 individuals (Table 8). Most individuals were not re- moved from the habitat, so the relative abundance indices are pos- sibly inclusive of re-counted individuals; each observation was treated as a unit indicative of species-habitat association. TABLE 7.—Comparison of Activity Cycles of Amphibians and Lizards in Five of the Major Areas. Absolute number of species and percentage of species within each area are given. Period of Activity Terra Firme Varzea Igapo Capoeira Open & Edge Diurnal 18 19 10 14 9 50% 50% 45.5% 70% 26.5% Nocturnal ___... 18 19 ibs) 6 aD 50% 50% 54.5% 30% 73.5% 27 ANALYSIS OF ECOLOGICAL DISTRIBUTION (syo[d ZT ) odes] FI eI ILZ 9%g Si¢ Ge ieee mee r Or Z ts 9¢ OI 88 Sol II € Ga Me Rye I LI ae Me Ee ae ac aaa ILI 9% cs 5G. es A el oat ae 13 Z —) g ra Pit Ye ear ZOr (syoid 6) (s}ojd T) woRrsuesy, BVOZIV A, BOZIVA -OULITY VIII], IG (sjojd 7) UontIsuery, oUITY Bild T-eila0dey 060 sataedg ‘ON [¥}0 QIg] S[eNprAIpuy ‘ON [PIOL, Q 9 ee psoyvuiadns Uopodssouds (-) yy DIquUun DIY ee eee pinoqvw vhinqvyy ae wunjouiuvoiad puosodaT Cpl ees snyoip9j09 xhdo.uay eg sypiauny sapoyDUor) GG nse srypinvoasn{ syouy Of | =e ee DIUOZDULDYD DSsojsOJYOY so wiajad snuanppshyd 2 dafiddyyda snwapjppshyd SIl twauspn smfhyappojdaT iG == SNYDLOULIDUL smhyoopojda'T 09 0 (oyY-pugns) “ds ph Qe coccrccccccececccnncncncncncc cnet piqns DA pL cece psoubis DIA Gg cnn povydvisoas Djh [rr nneenenenennnerereen wajsa DINAH Gl waupiwsunDog DIA TT Clave a. ee SNADINIDULYUAA SayDgoLpuad QQ cme snuoydhy ofng fa satoeds ‘sisAjeuy a[qey, AoussunUOD IF pasn xLyeP v}eC IT X setoedg 9y} Jo AreuuINg—'g ATAV |, 28 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Terra Firme- Varzea Capoeira - Terra Firme 10 O lO 20 Index I Fic. 12. Scores on index I plotted against scores on index II for each of 44 plots. Each dot represents the position of a particular plot relative to the x and y axes. Index I is a moisture gradient from dry (negative) to wet (positive). Index II is a vegetation density gradient from dense ground cover (negative) to grassy ground cover (positive). The data were assembled into a species X plot table; the species frequency counts represent the number of individuals of each species which occurred in a particular plot. Williams (1952) showed that when actual environmental measurements were un- available, scores could be calculated from the data of the contin- gency table by simply using those sets of scores for which there is maximum correlation. The interpretation is feasible because the scores are adjusted to have a mean of zero and a variance of one. The computer print-out for the analysis consists of a series of indices, each representing an environmental gradient or a composite of such gradients. Each index maximizes the measure of association between the two sets of variables—species and plots. Index scores relative abundances are presented in table 9. The results are presented by Cartesian (x, y) scattergrams of two sets of scores (Figs. 12-17). In this way, two gradients (two indices) can be studied simultaneously and their interaction examined. Species or plots having similar index scores appear close together on the diagram. Thus, ecologically similar plots and species with similar distributions can be identified. No actual environmental measurements were taken; resource = ——= 29 ANALYSIS OF ECOLOGICAL DISTRIBUTION 00°¢ vHS 8e°o1 00°F 89°1G 10'CE ves 6o°S 879 8L'9 98°TT 9L OI 2 a Tes LOE O¢'P 9F'T 00°T 00'S Te'6 preg aypIN T62 96°0 99°6 26.6" vV'6 cy i- Te L 96'9- Te"9- 69°8e—- vO L- Ilv oy = yOUOO! = 2 OL See ea a ee ae 8 {20000 7OIO00 == = TD oes a tg GLE80'0 FELIO'O x25 & = Fo sD ee oS N Boo Ey Saety i s See eae See ce ele ~ oe Bs eae eG ¥ Bae a = = Le! =) 5 5 (a) ~ g 02 psoyio.adns uopossouns yp) PL OGL To Diquin DIN £S yp phinoqow vhinqoy p Uh acces wnyouiupvoiad puLosoda'y Ge ap snyoivaj09 xhdouuay ole ah SYyDIaUNY sapoyUos) an pie os smypinvoosn{ syouy 9¢ Cl DOUOZDULDYD DVSsoOjs0I0g 9] tac issajad snuapjpshyd oe wthleg sopiddwyde snuapppshyd cu ol uauspn smhyovpoyday O61 6 ~~ SNywLousLvUL snhyovpo}daT 09 aaa (oxY-puqns) “ds pA fe pigns mA] ot 9 —— psounis mh 6g ms een a paiydoisoas DA ll poo wajsa DA ZI @ waupiosunng DjA OT Z ~snyvjnovunyuaa sayqorpuaqd a smuoydhy ofng N zaqunyy satoedsg SOLOS 'g 9[qe], Ul poziiewUns ‘FF X QZ SUOIsUSUTIp ‘xIIyeU Beep Jord X satoeds ve 0} parjdde senbruyse} [voAyeue WOIF S}[NSeI 9Y} O1e SIOqUINNY ‘“Se10dg Y}peeIg 9YOIN' pue ‘saorIpuy sfqey, AoussuQuoD ‘esouepunqy eAQepeYy jo uonL[nqely— 6 ATAVv], 30 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 20 Capoeira- Terra Firme Transition Terra Firme- Varzea ea Transition 20 lO O lO 20 Index I Fic. 13. Scores on index I plotted against scores on index III for each of 44 plots. Each dot represents the position of a particular plot relative to the x and y axes. Index I is a moisture gradient from dry (negative) to wet (positive). Index III is the vertical distribution of species found within the plots from terrestrial (negative ) to low vegetation (positive ). requirements for each species were analyzed indirectly by assuming that a given sampling plot provides necessary resources for the species found therein. For this reason interpretation of the indices is inferential. Environmental gradients——The first four index scores from the contingency table analysis were analyzed in an attempt to: 1) de- termine the major limiting environmental parameters affecting the ANALYSIS OF ECOLOGICAL DISTRIBUTION 31 20 Capoeira - Terra Firme Transition e—4/4 10 Varzea = oO Ze} = lO 20 lO O Te) Index IL Fic. 14. Scores on index II plotted against scores on index III for each of 44 plots. See figures 12 and 13 for explanation of dots and indices. The three dots not included within forest boundaries all are igapo plots. distribution of species; 2) characterize the four major forest areas in terms of those limiting factors relevant to frogs, salamanders, and lizards; and 3) identify the habitat of each of the 20 species in terms of the environmental parameters represented by the indices. The first index indicates a moisture gradient from dry (low values) to wet (high values). Moisture probably is the most critical factor affecting the ecological distribution of amphibians and rep- tiles in the study area. Probably the next most critical limiting factor is the physiognomy of the vegetation. The second index is indicative of vegetation 32 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Index IL 10 O {@) 20 Index L Fic. 15. Scores on index I plotted against scores on index II for each of 20 species. See figure 12 for explanation of the indices. Each circle repre- sents the position of a particular species relative to the x and y axes. The species numbers are associated with species names in table 9, p. 29. ground cover. The spectrum is from dense ground cover (low values) to grassy areas (high values). The second index may also indicate light intensity, resulting from the structure and density of the vegetation. In general, areas with dense ground cover are darker habitats than are grassy open areas. The third index probably is a combination of factors affecting vertical distribution. Terrestrial species have low values, and spe- cies which inhabit low vegetation have high values. Lizards found on tree trunks and along the boardwalks have intermediate scores. The fourth index seems to be a composite of many factors. Some of the following may be involved, but no one of them is responsible for the separation of the plot or species scores: 1) temporal activity (diel and seasonal); 2) organism size; 3) phylo- genetic position of organisms; 4) heliophilous versus sciophilous or- ganisms; 5) niche breadth of organisms; 6) abundance of animals within plots; and 7) solitary organisms versus congregations. The fourth index segregates the following species pairs, which are similar on the basis of the first three indices: Hyla rubra and Hyla sp. (rubra-like), Hyla geographica and Uranoscodon superciliosa, and Hyla rubra and Bolitoglossa altamazonica. ANALYSIS OF ECOLOGICAL DISTRIBUTION 33 8) (13) 20 ()+281 H © ee © © © (2) © ©) 10 @ 20 C!) 10 O lO 20 Index L Fic. 16. Scores on index I plotted against scores on index III for each of 20 species. See figures 13 and 15 for explanation of indices and circles/species numbers respectively. When the scores of the first three indices for the 44 plots (Table 10) are plotted against each other, it is possible to characterize the four major areas in terms of the environmental gradients analyzed (Figs. 12-14). Likewise, when the species scores are plotted, it is possible to get some understanding of the habitat of each species in terms of the environmental gradients inferred from the indices (Figs. 15-17). Plot index scores—Four capoeira-terra firme transition plots 34 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY TaBLE 10.—Contingency table indices for each of 44 plots analyzed. Plot numbers are located on figures 10 and 11. Indices are plotted on figures 12-14. Plot No. Index I Index II Index III Nh niger. hen Te LA as 3.66 =6.77 10.62 7, Mise ap as ee eh =ola -7.83 20.81 PAL eh tate AL Sse teem —6.50 —2.36 8.60 1p cele ot eee ee ge es ee =—D.00 —4.80 14.94 1S pe eee net, ER et eee =714 0.88 —1.96 NIGH tress of ol oe De Ye BE =0:67 =)/ 80 8.52 Lif aR SD, Pee OD TENCE Ses 2.79 —2.38 Hi SShngs ee Nace oe ean =i G0 6.06 —1.55 (Ch Re RRs os 2k Ree 0 Veet -—10.78 4.77 —4,29 Opp ides She, oar th oe =19°62, 5.93 —2.88 AGA ie ia eee eae by Pas! —12.39 6.03 —3.31 sf ties otk ae la tate | ee ROR AN —10.17 3.26 —1.02 A ase ate ota RN tek =IOr35 ioe -3.19 Digit Met nace Thoma ES os OS =IG —0.88 —1 1.55 ge NY 1 Lat el 3.26 Seyi —13.31 Ogee eae cee ett dash oe, 200 lean —-9.13 Die tea ie Fe ery cet eg 1.87 -—6.29 25.82 OURS CE ee ee —5.26 —0.48 -10.31 SS () Pe Rem re ce 2 oe ASR oon OO 3.02 —1,95 -8.83 iL le oa se ee eS —0.67 =i45 —11.32 oie canbe rh aaa ies FSS S, 2.82 —2.87 -6.13 AD gine, Gem eecmeras, 0, elias ne —12.05 5.54 —2.63 ASS eG te ee ee —12.58 6.11 —2.95 AA psn iat Seity wn ty ha Me 6.64 -~9.18 5.09 is SG TaN co Ae a eA Se NG ae 0.55 —8.65 10.87 AG ees Pete, Bown eee 9.34 —11.48 92.55 |, leg ar eae Ue Saarinen aia al emy.) 5.89 —5.08 —4.86 BO). 22) One i 1.74 5.26 -—0.18 We fe aes Fal Re. . oie 8.56 =8.12 =13.45 SA ee eA ae ets Ae ee edb aT 6.33 —8.77 1153 LN HIN Rit atin APRNs ibe Haan 3.38 —4.85 1.81 5 EES dee ee he Ps ie, WU SRR ae 3.44 —5.34 —0.96 Gaal eda ace Oe Te DO095 41.36 13.27 6G) ext meee hte a a cn 14.31 —5.39 —14.28 Cillig ececnte ae das en, Oe ele niente 15.00 4.45 —5.85 GO se DERE) cat Lee 13.83 0.45 —5.74 GOs eit at aga be, Be 13.92 —6.85 —2.69 FO Bes SME cake OC, 5 Bile 15.77 —4.80 —1.69 F(A aah et UE Ne RTE A 12.38 4.32 —8.75 {2 pee Re RP Ie Bell 15.28 —5.04 -7.98 ike dk Mele Fy be gc 9.76 —4.10 —7.00 OA No 2th se Sens BN wees 0 adi BE pa veya se 8.98 4°30 —5.59 15 ne De Toned CNG A, RN ered 9.82 —4.66 -9.33 Ge, eee ee ee eet ae 12.64 —5.78 -—15.93 were studied. In general, these plots are characterized by low to mid-range values on the first index, fairly low on the second, and high on the third (Figs. 12-14). The area is towards the drier end of the moisture spectrum and near the denser end of the vegetation \ ‘ ie ANALYSIS OF ECOLOGICAL DISTRIBUTION 35 lO O 10) 20 Index IL Fic. 17. Scores on index II plotted against scores on index III for each of 20 species. See figures 12 and 13 for an explanation of indices and figure 15 for an explanation of circles. density spectrum. The value on the third index suggests that the herpetofauna of this area is predominantly found on low vegetation rather than on the ground. Nineteen plots are terra firme-varzea transition areas, and can be divided into two groups, dry transition and wet transition (Fig. 10). The two transition areas are clearly segregated when the index values are plotted against each other (Figs. 12-14). The entire transition zone is characterized by low to mid-range values on the 36 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY first index, high on the second, and low to middle on the third. The area represents an intermediate zone with regard to the physical environmental parameters, except on the second index, indicating that the ground cover is relatively grassy. One plot is extremely low on the third and fourth indices and segregates from the other transition plots. The low value on the third index is ex- plained by the many terrestrial leptodactylids found calling from temporary puddles. Nine varzea plots were analyzed (Fig. 11). There is a very small range of variation on the first and second indices, but a wide range on the third and fourth. In general, most plots have a fairly high value on the first index, low on the second, and from low to high on the third. The varzea is a wet environment with relatively dense ground cover; the organisms are neither predominantly ter- restrial nor inhabitants of low vegetation. Twelve igapo plots were studied. These plots have the highest values on the first index, indicating that the igapé is the wettest area. Most values on the second index range from low to middle and most on the third are low. The igapdé forest has a relatively dense to intermediate vegetation ground cover. Most of the lizards are ter- restrial or are found predominantly on the boardwalks. When the indices are plotted against each other, one plot is segregated from the other igapo plots by high values on the second and third indices (Figs. 12-14). The second index score is explained by the presence of large clumps of tall emergent grass in the plot. Two species of tree frogs not found elsewhere in the igapé forest utilize the grass for calling sites; this creates a higher third index score than those values for igap6é plots in which there are many terrestrial lizards. Species index scores—The Cartesian plots of index values (Figs. 18-21) and the bar diagrams (Figs. 22-24) illustrate that each spe- cies has requirements and tolerances with regard to the environ- mental gradients. Several trends represented by correlations be- tween species abundances and availability of a particular resource are evident (Figs. 22-24). For example, index I represents a mois- ture gradient; those species with the highest positive values are those found in association with wet areas. Each of the seven species with the highest scores (Leptodactylus wagneri, Kentropyx calcara- tus, Mabuya mabouya, Dendrobates ventrimaculatus, Hyla egleri, H. granosa, and H. baumgardneri) is most abundant in the igapo forest, less abundant in the varzea forest, still less common in the terra firme-varzea transition area, and rare in the capoeira-terra firme transition area, if found in the last two areas at all. The two species with extremely low negative scores on the first index (Lepto- ANALYSIS OF ECOLOGICAL DISTRIBUTION 37 dactylus marmoratus and Bufo typhonius) are more abundant in the terra firme-varzea transition area than in capoeira-terra firme transi- tion plots, contrary to what one might expect. The distribution is better understood when the second index scores are considered; both species have positive scores, but neither one is extreme. Ap- parently these terrestrial species inhabit relatively dry areas but avoid the open areas characteristic of capoeira forest in preference to the denser undergrowth of high ground, dry terra firme-varzea transition areas. In most instances, those species with scores closest to zero are the most abundant. This is probably because those species requiring neither extreme (considered generalized) are able to utilize more of the environment. If more of the environment is potentially available for exploitation by a species, it can be assumed that the potential carrying capacity of the environment for that species is greater than that for a specialized species restricted to a particular habitat. Gonatodes humeralis is the most abundant of the twenty species and has scores near zero on each of the four indices. The three next most abundant species, Leptodactylus marmoratus, Kentropyx calcaratus, and Bufo typhonius, have scores relatively close to zero on all indices except the first. Species with extremely high positive or low negative scores on index IV are relatively un- common. Most species of lizards do not have extreme values on any of the environmental gradients. Gonatodes humeralis, Kentropyx cal- caratus, and Mabuya mabouya are the only species found in all four major areas; none has extreme index values. The combined cumulative relative abundances (the three species from the four areas) represents 509 individuals, or 41.8 percent of the total sampled herpetofauna. Gonatodes humeralis is less abundant in the igapo forest than in the other three areas, whereas K. calcaratus and M. mabouya are most abundant in the igapé forest. The other four species of lizards are relatively uncommon in all of the areas. Anolis fuscoauratus, Leposoma percarinatum, and Plica umbra have no extreme index scores; the first two species are near zero on the moisture gradient, and P. umbra is near zero on the fourth index. Uranoscodon superciliosa is relatively generalized with respect to all the environmental gradients except vegetation density; the score on the second index is low, indicative of its occurrence in areas of dense vegetation. In general, the amphibians demonstrate more extreme environ- mental requirements than do the lizards. None of the thirteen OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 38 (osed }xou UO ponurzUuood ) oULIY vil9}-eIlo0deo Ul QOOT X souRpunqe saAr}e[e1 suede po}o[d soorpul se1Y} 9Y} JO YOR UO seIODS sofoedS ‘ET-9gT ‘sdIyY TT X9pu} IT X9pu| I X9pu} 02 Ol O O! Oc OG Ol O Ol O02 Ol O Ol b » ie 0 @| © © @ © © © @) 0 (Dy (894 I) To x9pu II Xepu| Ol 10) Ol ISUDJ] _DEZID/A -9WI4 Dia] e Oo oOo wa pajeodog /uol} OOOIX suUDpUNgY sAl{ojay UONISUDI] SUW4I4 Diva} - ANALYSIS OF ECOLOGICAL DISTRIBUTION 39 species is found in all four areas. The salamander, Bolitoglossa altamazonica, has a score near zero on the moisture gradient, but exhibits extreme scores on the second and third indices, indicative of the occurrence of individuals on low vegetation in relatively dense areas. Physalaemus petersi is generalized with regard to all of the environmental gradients. Bufo typhonius and Leptodactylus mar- moratus are specialized only with regard to the moisture gradient; they inhabit relatively dry areas. Hyla granosa is found in relatively open, very wet areas. This species is more abundant in the igapo forest than in the varzea forest; the fourth index score is almost zero. Leptodactylus wagneri is terrestrial, as indicated by the ex- tremely low third index score; the species is more common in the varzea and igapo forests than in the terra firme-varzea transition area, apparently due to the absence of permanent standing water in the transition area. The fourth index yields extreme values for several of the species of frogs. Physalaemus ephippifer, Hyla geographica, H. baumgardneri, Hyla sp. (rubra-like), and H. egleri all have low scores; Dendrobates ventrimaculatus and Hyla rubra have high values. Physalaemus ephippifer is terrestrial, found only in the terra firme-varzea transition area. Dendrobates ventrimacu- latus occurs in very wet areas of the varzea and igapo forests; the species is relatively uncommon in both areas. Hyla geographica and H. rubra are found in places of rather dense vegetation. The three most specialized species seem to be Hyla baumgardneri, H. egleri, and Hyla sp. (rubra-like). The first two species are found in very wet, open grassy areas, whereas Hyla sp. (rubra-like) is found in plots having intermediate values on the moisture gradient, with dense vegetation. All three species are found on low vegetation; all have extremely low values on the fourth index. Of the twelve species of frogs, the only abundant ones are Bufo typhonius, Leptodactylus marmoratus, and L. wagneri, all of which are terrestrial, and mainly forest inhabitants. Three species of tree frogs, Hyla baumgardneri, H. egleri, and H. rubra, are found principally in open, non-forested areas, where they congregate at ponds and swamps to breed, thus explaining their relative uncom- monness in the forest plots. transition (figure 18, upper) and terra firme-varzea transition (figure 19, lower). See figures 12 and 13 for an explanation of indices. Species numbers are associated with species names in table 9, p. 29. Numbers enclosed in squares indicate species that are habitat specialists; circles are habitat inter- mediates; diamonds are relatively uncommon generalists; triangles are moder- ately common generalists, and hexagons are abundant generalists. OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 40 ‘uoTeUR[dxe oF GT pue QT soinsy 0} IOJOY “JSOIOF VOZIVA OY} UL OYOT X VoURpUNqe oAT}e[AI ysUTeBse payo[d soorput so1Yy1 BY} JO YOR uO satoos soIDedg ‘OZ ‘DIq Ir x9 pul TI Xepu| T xapu] jS8404 DEZIDA Ul OOOIX BUDPUNGY sAI}D]EY 4] Z © = = = jam = 4 a) = << = So) eo) a e) Se) zs te eo) ce W > 4 < Z < IoJOy ‘uoTyeuRldxe 10J GT pue QT Sainsy 0} "yS910F Odes! oy} Ul QOOT X soURpuNqe saTRieI ysuTese poyo[d soorpur se14} 9Y} Jo YOR UO saiOOS so1roedg ‘TZ ‘oy If x9pu| O¢ O a+ O Ww O09 odpb| u! OOOIX e2uDpungy aniojay $sa104 42 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Relative Abundance XIOOO S10. 35 .26051I0 35 260410 35 260510 35 260 | Leptodactylus marmoratus Bufo typhonius Jee Saas Physalaemus petersi ee Plica umbra ; Gonatodes humeralis Physalaemus ephippiter Sos eee se Hyla sp. (rubra \ike) ae ate an Anolis fuscoauratus Bolitoglossa altamazonica IS Leposoma percarinatum ee ea ee ee Se ee ae geographica |. Se ae es es SS ranoscodon superciliosa Leptodactylus wagner! -—--—--—|—---—--- Hyla rubra } tKentropyx calcaratus alan ae ees Mabuya mabouya a Dendrobates ventrimaculatus —------------- Hyla egleri Sa Hyla granosa a a ES eS Hee aS Hyla baumgardneri Capoeira - Terra Firme- Terra Firme Varzea S Transition Transition Varzea Igapo 5 IS 25 35 Niche Breadth Fic. 22. Relationship of species scores on index I, relative abundance, and niche breadth scores for 20 species in each of the four major areas. Dots represent niche breadth scores. Horizontal bars indicate index scores and rela- tive abundance. ANALYSIS OF ECOLOGICAL DISTRIBUTION 43 Relative Abundance X|OOO fer, oo =6o02!0"°S5 =60310 35 =60510' 35 260 Hyla sp.(rubra-\ike) —e 2 SE a se ricer rae GBolitoglossa altamazonica - 2a ~Hyla geographica ae : Uranoscodon superciliosa Hyla rubra -—e-—-—-—-— Anolis fuscoauratus Leptodactylus wagner/. ee ee | ee SS Plica umbra_ SS eee percarinatum —-—@ — — — — ee - — — -— -|-— -— - - Physalaemus ephippiter Kentropyx calcaratus -—--—- Gonatodes humeralis eo — — Dendrobates ventrimaculatus O—-——— -e-— Mabuya mabouya Se SSeS Physolaemus petersi 5 == Bufo typhonius —@---—-— Leptodactylus marmoratus HH 10, © a cO £ @e———-—--—}- -—----- Hyla granosa Zs 5 eee Hyla egleri 215) 60 == — — | — — - — - - -/ -- -- Hyla baumgardner/ Cdpoeira- | Terra Firme- Terra Firme Varzea : Transition Transition Varzea Igapo 5 15 ES IS) Niche Breadth Fic. 23. Relationship of species scores on index II, relative abundance, and niche breadth scores for 20 species in each of the four major areas. See figure 22 for explanation. 44 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Relative Abundance XIOOO 410 35 260510 35 2605/10 $65. 250210 Gao ce —-@--—-—-—-— Physalaemus ephippifer Leptodactylus wagneri-4 ee ee Physalaemus peterst —e------ Leposoma percarinatum Kentropyx_ca/caratus 4 Sg SSS e — ——-—-—|Hyla geographica -—@-— —— Leptodactylus marmoratus oe eee Dendrobates ventrimaculatus Os cape Bufo_typhonius Or =8r 4} Gonatodes humeralis Mabuya mabouya SS Se ee es Se ees Hyla granosa H 5 x< ped) ae) £ —-@---——-- Uranoscodon superciliosa Oe — Plica umbra oo Se = SS Hyla rubra == Anolis fuscoauratus FS) ( } —e ome oe oe om ee oe 20 ———— — -]— — — — | - Hyla baumgardneri 25 ——— Bolitoglossa altamazonica e- - — — — —|-—— ---— — Hyla sp.(rubra-\ike) Capoeira- Terra Firme- Terra Firme Varzea Transition Transition Varzea Sie ee 25 35 Niche Breadth Fic. 24. Relationship of species scores on index III, relative abundance, and niche breadth scores for 20 species in each of the four major areas. See figure 22 for explanation. ANALYSIS OF ECOLOGICAL DISTRIBUTION 45 NIcHE BREADTH ANALYSIS Niche breadth is used in this paper to refer to habitat niche breadth and is presumed to be correlated with the range of en- vironmental tolerances. Niche breadth scores were calculated from the standard formula proposed by Levins (1967), where pj; is the proportion of occurrences of species j in plot i, niche breadth of species j (B;) equals: LB; = > pij* No actual environmental measurements were taken; resource requirements for each species were measured indirectly by assum- ing that a given sampling plot provides the necessary resources for the amphibians and lizards found therein. Although the niche dimension is referred to as being habitat, there may be certain latent biotic interactions influencing the distribution of species which are included in the niche breadth measurement. The limitation of using occurrence in sampling plots as an indirect method of measuring requirements of species is acknowledged. However, the analysis is the only one feasible due to the lack of direct physical environmen- tal measurements. The data are from the matrix (plot X species) used in the contingency table analysis. Niche breadth values are included in table 9. The niche breadth analysis used here was not meant to describe the entire niche of each species, but rather to de- limit the spectrum of the habitat dimension of the niche of each species. Three species of lizards (Gonatodes humeralis, Kentropyx cal- caratus, and Mabuya mabouya) have much higher niche breadth scores (16-32) than the next highest species, Leptodactylus wagneri (approximately 12). These three species of lizards are found in all four of the major areas, account for 41.8 percent of the entire sample of 1218 individuals, and do not have extreme scores on any of the contingency table indices representing environmental gra- dients. These lizards are considered to have wide niche breadths with regard to habitat requirements and tolerances and are referred to as habitat-generalists (Fig. 25). The relative abundances in each major area are plotted in figure 26. Five species can be considered habitat-specialists; all of them have niche breadth scores in the range of 1-4, indicating that they have very narrow tolerances and specialized requirements with regard to the environmental parameters measured indirectly by the analysis (Fig. 25). Each species is found in only one or two of the four major areas and is relatively uncommon. The cumulative 46 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Hyla baumgardner! (\O) ieee Hyla egleri (9) Uranoscodon superciliosa (2) Hyla granosa (\\) Leposoma percarinatum Hyla geographica (\9) INTERMEDIATES Hyla sp.(rubra-\ike) (49) Dendrobates ventrimacu/latus Hyla rubra (30) Flica umbra (\\) Bolitoglossa altamazonica (30) Physalaemus petersi (15) Physalaemus ephippifer (\7) Anolis fuscoauratus (\8) Bufo typhonius (\Ol) Leptodactylus marmoratus (\56) Leptodactylus wagneri (92) Mabuya mabouyo+; GENERALISTS (44) Kentropyx calcaratus (\I7) Gonatodes humeralis (257) 5 lO 15 20 25 210) yO) Niche Breadth Fic. 25. Niche breadth scores. The bars represent niche breadth scores. Numbers in parentheses indicate the accumulative relative abundance x 1000 in all of the four major areas. The dashed lines separate the species into habitat specialists, intermediates, and generalists. relative abundance of the five species in all of the areas is only 44 out of the total of 1218 individuals, or 3.6 percent. Two of the habitat-specialists are lizards (Leposoma percarinatum and Uranos- codon superciliosa), and three are frogs (Hyla baumgardneri, H. egleri, and H. granosa). Leposoma percarinatum, a secretive ter- restrial lizard found within the leaf litter by day, is probably more ANALYSIS OF ECOLOGICAL DISTRIBUTION 47 Gonatodes humeralis @ * Kentropyx calcaratus @——~—e Mabuya mabouya @—-—® 60 Capoeira- Terra Firme- Varzea Igapo Terra Firme Varzea Transition Transition Fic. 26. Relative abundance < 1000 of the three habitat generalists in each of the four major areas. widely distributed and more abundant than the data indicate. Uranoscodon superciliosa is found mainly near pools of standing water in the varzea forest. Hyla granosa is predominantly an igapo specialist, not found outside of the forest. The other two species of tree frogs, H. baumgardneri and H. egleri, are not primarily forest inhabitants, but are found abundantly in open areas; therefore, these two species are not specialized for the particular forest areas, but rather are dependent on standing water. For this reason the species have low niche breadth scores relative to the forest analysis. If a similar study were carried out in open areas, these species would probably have wide habitat niche breadth scores, for they are abundant and seem to have a wide range of environmental toler- ances in open areas. The remaining twelve species are considered to be habitat- intermediates (Fig. 25). In general, these species demonstrate intermediate niche breadth scores, corresponding to relatively few extreme values on the environmental indices from the contingency table analysis. They are generally more abundant and more widely distributed than the habitat-specialists, but less so than the habitat- 48 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY generalists. This category includes the one species of salamander, two lizards, and nine frogs. Several of the habitat-intermediates have niche breadth scores similar to those of the habitat-specialists. The artificial line separating the two groups is obviously based on more than niche breadth values; representation and _ relative abundance in the major areas were also considered. There seems to be a definite relationship between cumulative relative abundance and niche breadth scores (Fig. 27). In general, those species with wide habitat tolerances (high niche breadth values) are more abundant than those with narrow habitat toler- ances. The abundant generalist has the highest niche breadth value, the moderately common generalists have lower niche breadth values, and the five habitat-specialists have the lowest niche breadth values and are extremely uncommon. Another way of looking at the association is to plot index scores against niche breadth values (Fig. 28). All of the habitat-specialists have positive values on the first index, indicative of wet environ- ments. Three of the habitat-specialists are restricted to open, grassy areas. One of the specialists is terrestrial, and the other four are found predominantly on low vegetation. Three of the specialists have more extreme negative values on the fourth index than does the generalist having a negative value. The relationship of niche breadth values to both index scores and relative abundances within each major area is presented ( Figs. 22-24). When index scores are plotted against relative abundance values for each area, it is possible to characterize the areas with regard to species composition in terms of habitat-generalists, intermediates, and specialists (Figs. 18-21). The capoeira-terra firme transition area provides suitable habitat for the three habitat-generalists (one is moderately common and the other two are relatively uncommon), but the five habitat-specialists are absent. The terra firme-varzea transition area is composed of two habitat-specialists, the three generalists, and numerous habitat-intermediates. One of the gen- eralists is very abundant in this area, and the other two species are relatively uncommon; the two specialists are rare. Both of the specialists are found in one additional major area. The varzea area is represented by the three generalists (one abundant, one moder- ately common, and one relatively uncommon) and three habitat- specialists (all rare); one of the specialists is restricted to the varzea forest. Two of the generalists are moderately common in the igapo forest, and the third is abundant; three habitat-specialists inhabit the area, only one of which (Hyla baumgardneri) is restricted to the pee ae a ee 49 ANALYSIS OF ECOLOGICAL DISTRIBUTION ‘6 d “6 2[qe} UT souIeU SoToads YYA\ pozeloosse a1 slaquinu sotoedg ‘saroeds QZ ay} JO YORe 10F YYpRerq eyoru ysurese poyo[d QYOT X V:URpUNe oANK[eI VARINUMDDY “LZ ‘OI 002 OvI| OOOIX s2UDpUNgY d O02 {S!|DIDEdS 8} DIPOW 49} U| }S|[DJoUaH UOWWOD Aja}DJEpo|A| jSs|jOsouab juDpundy O| O¢e YypOoIg BUOIN OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 50 ‘uoleueldxe 10f GT pue QT seinsy 0} Jejay 6Z “d ‘6 2[qQe} UI soUreU soloeds YIM poyeloos “SB ale stoquinu satsedg ‘saroeds QZ 9Y} JO YOR 1oF YIpPRoiq Yoru Jsurese poyold [I] pure [J] seorpurl uO sa1ods saToedg “gz ‘OI TI xepu| TE xepu] O¢ Ol O O| Od O9 OG Ov O¢ Od Ol O Ol yypoesg sYdIN ANALYSIS OF ECOLOGICAL DISTRIBUTION 51 igapd. All three specialists have the highest positive scores of any of the igapé species on the first and second indices, indicating stand- ing water and areas of grassy, emergent vegetation; two species have the highest positive scores on the thtird index. The habitat- specialists in the igapé forest exhibit more extreme index scores and are more abundant than other specialists in other areas. NICHE OVERLAP ANALYSIS In a consideration of niche overlap, it is appropriate to ask: Proportionately, how often do species i and j occur together? Niche overlap can be crudely estimated by plot overlap if we assume that species requirements are intrinsic properties of the plots. The mea- sure does not indicate what the overlapping requirements of the species are, but merely that overlap exists. Niche overlap scores were obtained from a formula suggested by Horn (1966); pi; is the proportion of occurrences of species 7 in plot i. Overlap of species jand k ( «jk) is then estimated by the following: a fk = 2 Spy Dix! ( Spi + Upix") The index is from 0.0 (no overlap) to 1.0 (complete overlap). A high niche overlap value for two species indicates they are found together in the same plots. For example, Hyla baumgardneri and H. egleri have an overlap value of 0.971, the highest of any two species associations; these frogs breed in the same plots in the igapé forest. Other high correlations are Bufo typhonius and Leptodacty- lus marmoratus (0.928) and Hyla sp. (rubra-like) and Bolitoglossa altamazonica (0.913). Both species pairs usually occur sympatrically and therefore probably overlap greatly with regard to certain en- vironmental requirements. The following species pairs frequently occur together and have fairly high correlations, likely indicating similarities in environ- mental requirements: 1) Kentropyx calcaratus and Mabuya ma- bouya (0.718); 2) Hyla rubra and Anolis fuscoauratus (0.631); 3) Hyla egleri and H. granosa (0.589); 4) Leptodactylus marmoratus and Physalaemus petersi (0.586); 5) Leptodactylus marmoratus and Gonatodes humeralis (0.575); 6) Hyla baumgardneri and H. granosa (0.547); 7) Leptodactylus wagneri and Mabuya mabouya (0.539); 8) Hyla rubra and Uranoscodon superciliosa (0.533); 9) Bufo typhonius and Gonatodes humeralis (0.529); 10) Anolis fuscoauratus and Plica umbra (0.513); 11) Hyla geographica and Uranoscodon superciliosa (0.512); and 12) Bufo typhonius and a et ws - ne —=—eereee ——SS SSS SS SS SS Se 2868 “ee & or see See a elas es a s ca a T€9°0 FI 6890 P GIS 0 06 IL6°0 © TL60 ¥ 960 8ST 8660 6 = Zliks ee ces 0 0G LVS'O € 9€7'0 OT 689°0 9 LVS 0 9 8Le'0 9T 6¢9°0 ST ee 96¢°0 8I T9rv'0 8ST sOV'0 LI LOGO 8T SLTO 8I c9e0 G LOS’ 0 GI eS o2 o & cee'0 ET 6860 G VLE'O 8 cSl0-< LOTO &G 6860 9 T€¢0 61 = ere? SS a 08¢0 9T 8460 OT GIE0 G 960°0 9T 180°0 9T Isc 0 OL 8060 FI = sy a rs) 5 STG0 @ 90¢6°0 OT 8660 ST GLO0 GI 6700 OL OSc 0 0G €0¢0 €1 < SO eet 8810 ST vOTO S 81¢0 OT LS0°0 OL LvV00 L 6660 LI 8610 IT a < = B= O9TO 61 080°0 L Il¢0 €T 6S0°0 L eco Or Si STGO L Lohors 5 ieee, a, a vVTO S 9L0°0 ST 6060 FI 6900 6 000°0 0¢ 8810 8 c81T0 LI 8 8 =a Sel 0 8 VLO'O 0G GST 0 61 6700 LI 000°0 6I cst oO V GOTO 81 2 3 9 8 3 961 0 OL y900 LI VrTO L LvO'0 ST 0000 LT LOTO € €90°0 9T ea > a 5, S600 LI vs0'0 8 6010 IT se0'0 T 000°0 FI LEEOE ST cc0'0 ¥ Ss = Bes. ta 080°0 9 €20'0 €T VOTO 9 ST00 61 000°0 €I 160°0 ST €600 L = * es ps 6S00 ¥ STOO FI Sc0'0 GI 0100 FI 0000 GI LS0'0 VI 6c00 OL ae or S LV00 € 0000 6I 000°0 61 000°0 0G 000°0 IT 000°0 61 000°0 06 om SON YU €600 T 000°0 GI 000°0 6 000°0 €T 000°0 6 000°0 GI 000°0 9 =e po S68 6000 GI 000°0 IT 0000 ¥F 000°0 IT 000°0 8 0000 IT 0000 & a hina = a. 0000 IT 000°0 6 000°0 € 0000 8 000°0 & 000°0 6 000°0 € = © = 2 g 000°0 6 000°0 T 000°0 TI 0000 & 000°0 T 000°0 T 000°0 @ = = rs S S L soweds g sotoeds G satoadg p soweds ¢ sotoeds Z@ sotoeds I soroeds =~ = 2) a x = ‘depiaA0 JO sdIsap I9}e9IS B 9}BOIPUT Si1oq S = =| = 5 -uInU JOYysIy ‘UUIN[OO B UTY}IM poalopio oie Sal0dg ‘e[NULIOJ UIOFY oY} AQ poUTULIaJap sv ‘o10ds dep1aAo oyoru = 8 3 8 2 6 ay} AG peMoToF st (6G PGB ees) aAoqumnu soloods oY} UUIN[OD Yove uy ‘se10dg Ae[I9AGQ 9YOIN— TT AAV], _ Se eee a eee Ye} AS ca & 53 ANALYSIS OF ECOLOGICAL DISTRIBUTION T¢9'0 €TS'0 89F'0 SFr 0 ver 0 L8€'0 OLG'O 190 8060 G0G'0 9610 9910 esl 0 LS0°0 v0 0 C100 0100 000°0 000°0 PI setoeds L 61 06 eT cI 8 9T SI I a 6 GL LI G OL 9 V ET € €16'0 SFr 0 Lov 0 GEE 0 6660 16¢ 0 LIGO IT¢0 £060 961'0 v6r'0 8TTO LITO 601'0 €€0'0 610°0 000°0 000°0 000°0 ET sotoads 8 VI 61 L CT SI 0G q I 9T LI 6 G OL 9 at Et 7 € 9890 LOS'0 cIV 0 610 OLG 0 99T'0 LET O GL0°0 0L0°0 €V0'0 6600 S00 610°0 600°0 000°0 000°0 000°0 000'0 000'0 ZI Seeds 6 I LI ST Vl VI OI V 61 91 81 Ss el L 0G 8 9 € G panuyuoj— [] aATavV OIG 0 S610 6LT 0 VST O 9CT'O OGIO VITO 6010 €Or 0 G0 0 000°0 000°0 000°0 000°0 000°0 0000 000°0 000°0 000°0 IT soweds NOt Or 6ES 0 CSP 0 9EV 0 90€ 0 L9G 0 Isc 0 9060 LST O LET O Lol O 961 0 60T'0 €OT 0 LSO 0 670 0 vVVO O 6¢0'0 660 0 6100 0G Gl 8 ye el PD V c vi 6 I 61 QT setoads 6 setoeds cI60 €1 L8¢'0 FI PLEO & GLE 0 61 99¢°0 ST OVGO SI 6E6C0 0G Il¢0 LI 1610: 1 8810 G O9T'0 9T Sel 0 ZL L610 Ol 9ITO 6 ys0'0 9 000°0 GI 000°0 TI 000°0 F 000°0 € Q satoeds SPECIES DIVERSITY AND EQUITABILITY The Shannon index was used in the present analysis as a means of comparing the four major forest areas. The index is calculated as follows H’ ~ pe Pi log Pi = C/N (N logio N—Din; logio ni), v v OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 54 ces 0 L €1g'0 VI USO S L6v'0 €1 8970 FI LEO 8 9860 LI 0LG°0 ST 0S¢0 G Té¢o0 T 6E60 8 LLTO 6 LIGO €T v9TO IT OIG0 9I GIT 0 81 8810 81 O9TO L T8t 0.41 S210) 91 LET 0 OT 0L0°0 GI VL00 9 6100 OL 0000 6I SOO 0000 GI 000'0 0¢ 000°0 IT 000°0 LI 000°0 6 000°0 9 0000 7 0000 S 000°0 © 0000 € 0000 T 0000 G 0% Setoeds 61 Sseteds 8TL0 6ES'0 [9V'0 9670 96¢°0 OVE 0 160 19¢0 OVC 0 LOGO 88T'0 SLO c9TO 6ST'0 6710 cOT 0 0L0'0 6600 S600 QT setoeds 91 G L aT cl FI 8 V 0G c 61 q LI it 6 GI iLL GIV 0 GI 8IZ0 81 aaa 026 sOvV'0 & csV 0 OL 6¢9°0 I 9860 06 OSV 0 SI OSV 0 OT L9¢0 OT SLEO G ver 0 VI 8860 6 O8c0 L OVe'O 8I GCG 0 G OLG0 FI 6IE 0 GI IT¢0O 8 8G 0 9 90©°0 OT vOCO ST 8160 S 6660 €I v6.0 €1 OIG 0 06 OLGO 61 0610 OT 9610 €T 9960 8 G80 T 0610 LI 8660 S 6710 SI O9TO 8 vOGO LI S60 VI 8ST0 61 S810 L OGIO IT VALI) JUL ISTO 0G c60°0 2 960°0 F G20) tT 7900 9 [800 € 160°0 G 6700 F €90°0 I 9L0°0 9 000°0 61 €V0'0 GI LvV00 ¥V 000°0 € IV0'0 6 Sc0'0 © LT soetoeds QT setoads CT sotoads probability of en- papnjouojn— TT ATAV where H’ — average diversity per species, p; countering the i* species, C = the constant 3.321928 when using number of in- dividuals in the i species. The values are interpreted as follows. If in Area I there are 20 individuals of species A and 20 individuals of species B, then H’ ~ 2.00; there are two equally common species. If in Area II there are 40 individuals of species A and 10 individuals of species B, then H’ ~ 1.65; there are 1.65 equally common species. Area I is considered to have a higher species diversity than Area II. The data used were those in the original data matrix which also Base 2, N = total number of individuals, and n; served as the basis for the contingency table analysis and niche breadth and overlap analyses. ANALYSIS OF ECOLOGICAL DISTRIBUTION 55 The calculated species diversity values for the four major forest areas are as follows: capoeira-terra firme transition = 3.00; terra firme-varzea transition — 2.45; varzea = 3.02; and igapd = 2.72. Thus, in terms of the sampling plots analyzed, the varzea is the most diverse area with regard to species richness and evenness; the terra firme-varzea transition area is the least diverse. A deter- mination of statistical and biological significance of these values would require additional data, ideally with an equal number of plots sampled per major area. MacArthur (1957) proposed a “broken-stick model’ dealing with species equitability based on one million individuals of 200 species. The model is based on the equation = = Lc > Leas) 1=1 where 7, is the theoretical proportion of individuals in the r most abundant species (r = 1, 2,..., s), each theoretical proportion itself being obtained by summing over r terms (i = 1, 2,..., 1). By using this formula, it is possible to obtain an apportionment of the in- dividuals among the species in a sample in about as equitable a manner as ever occurs in nature. An advantage of MacArthur’s model is that there is no set of parameters into which data must conform; for each possible number of species (s), the equation generates a complete set of s proportions 7, (r = 1, 2,...,s). The model yields a curve whereby species abundances are graduated from the rarest to the most common. A maximum equitability curve, whereby for every sample size each species is equally abun- dant, can also be calculated. Species diversity values (H’) obtained from the Shannon index can then be compared to the broken-stick and maximum equitability curves. Any community falling between the two curves is considered to be extremely diverse. The species diversity values for the four areas were plotted in relation to the curve expected from the broken-stick model and the maximum equitability curve (Fig. 29). All areas fall to the left of the broken-stick distribution with the exception of the capoeira- terra firme transition area; the position of this area suggests that it is highly diverse in terms of species equitability. The validity of this model was questioned by Hairston (1969). He claimed that the broken-stick model lacks ecological meaning, because conformity to the model is largely a function of sample size. He demonstrated that large samples tend to make rare species even more rare and common species even more abundant; the reverse distortion is ap- parent in small samples. Because of the small sample sizes in the present analysis, the distribution of abundances may be distorted 56 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY |. Capoeira- Terra Firme Transition 2. Terra Firme- Varzea Transition I5- 3. Varzea g 4. |gapo < oO oO a 0 a) o ce) ES Broken = Zz | 2 3 4 Species Diversity (H') Fic. 29. Relationship of species diversity and numbers of species for each of the four major areas to MacArthur’s broken stick distribution and maximal diversity. such that rare species appear to be more common relative to abun- dant species than actually is the case. Lloyd and Ghelardi (1964) proposed an equitability equation for the measure of fit of observed relative abundances of species to those predicted by MacArthur’s broken-stick model, as follows: = sis; where s is the actual number of species and s’ is the theoretical number of species that should be present according to the broken- stick model at the actual diversity (H’), as calculated from the Shannon index. Maximum conformation to the model is 1.00. The following equitability indices were calculated from Lloyd and Ghelardi’s table: capoeira-terra firme transition = 1.10; terra firme- varzea transition — 0.58; varzea — 0.79; and igapo = 0.90. Because of inequitability in the distribution of individuals among the species, the sample from the capoeira-terra firme transition forest has a species diversity appropriate to a community with 10 percent more species than actually occur in the particular sample. On the other hand, the samples from the terra firme-varzea transition, varzea, and igapo areas have species diversities appropriate to communities with only 58, 79, and 90 percent as many species as actually occur in the respective areas. Therefore, only the capoeira-terra firme transition area is more diverse than would be expected by the ANALYSIS OF ECOLOGICAL DISTRIBUTION 57 TABLE 12.—Comparisons of Major Areas by Coefficients of Community. Num- bers in Roman are the number of shared species of amphibians and lizards between two major areas; numbers in bold face are the actual number of species in a given area; numbers in italics are the coefficient of community values. Major Terra Open & Areas - Firme Varzea Igapo Capoeira Edge era Firme ______ 36 24 16 20 16 Ga 0.480 38 19 16 18 oo 0.381 0.463 22 13 12 japocira 0.556 0.381 0.448 20 10 Open & Edge __. 0.296 0.333 0.273 OL227 34 broken-stick model (Fig. 29). Again, because of small sample sizes, the statistical and biological significance of this analysis is uncertain. COEFFICIENT OF COMMUNITY The coefficient of community (CC), used to measure the relative similarity of samples from two communities (major areas), is calculated, CG Ba) oe See eens where S,, is the number of species shared by samples A and B, S, is the total number of species present in sample A, and S, is the total number of species present in sample B. Coefficients were calculated for every two area combinations for five major areas: open and edge areas, capoeira, terra firme, varzea, and igapd. The distribution data used are found in table 2, consisting of 62 species of frogs, salamanders, and lizards. The coefficients are presented in table 12, in addition to the actual num- ber of species every two areas have in common. The varzea and terra firme forests have the most species in common (24), but the coefficient of community is the second highest (0.480). The capoeira and terra firme forests have 20 species in common and have the highest coefficient of community (0.556). The capoeira forest and open and edge areas have the fewest species in common (10) and have the lowest coefficient of community (0.227). Likewise, there is low similarity between the igapé forest and open and edge areas (12 shared species, with a coefficient of 0.273). SUMMARY AND CONCLUSIONS The ecological distribution of each of 62 species of frogs, sala- manders, and lizards was determined by means of continuous 58 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY sampling throughout the environment from mid-January to the end of July, 1969, two weeks in April 1970, and June-July, 1970. Each species exhibits a characteristic distribution, according to its genetic, morphological, and physiological make-up, its life cycle, its way of relating to the physical environment, and its interactions with other species. The contingency table analysis was used to obtain a measure of the association between species of amphibians and reptiles and their habitats (plots) and to partition this association into inde- pendent components (indices) which determine the distribution of species within four of the major forest areas. The components are interpreted as follows: the first is a moisture gradient; the sec- ond, a vegetation density gradient; the third, a vertical distribution gradient, and the fourth seems to be a composite of environmental parameters. Each species can be characterized in terms of the indices. Species with scores near zero are the most generalized with regard to the environmental parameters studied and are gen- erally the most abundant species; those species with extremely high positive or low negative scores are restricted to a particular range of the environmental spectrum and are relatively uncommon. The species of frogs exhibit more environmental extremes than do the lizards, indicating that the particular species of frogs studied have more narrow environmental tolerances than do the lizards included in the analysis. The environment likely produces greater restric- tions on frogs than on lizards in the carrying out of life processes due to basic physiological differences between the animals, resulting in more restricted distributions for frogs than for lizards. Niche breadth scores, as calculated from Levins’ formula, are presumed to be correlated with the range of environmental toler- ances. Three species of lizards have much higher habitat niche breadth scores than the other 17 species of amphibians and lizards; these three species are the only ones found in all of the major forest areas. There is a definite relationship between cumulative relative abundance and niche breadth values. In general, those species with wide environmental tolerances (high niche breadth scores) are more abundant than those with narrow tolerances (low niche breadth scores ). When niche breadth scores, abundance indices, and scores on the environmental gradients are analyzed together, three species are referred to as habitat-generalists, five species as habitat-spe- cialists, and the remaining twelve as habitat-intermediates. The generalists occur in all major areas, have high niche breadth scores, are abundant, and exhibit no extreme scores on the environmental ANALYSIS OF ECOLOGICAL DISTRIBUTION 59 gradients; the specialists are found in only one or two major areas, have low niche breadth scores, are relatively uncommon, and ex- hibit extreme requirements on one or more environmental gradient. Partitioning of resources is evident, in terms of both space and time. One of the most striking examples is the calling site segrega- tion among breeding male frogs in a given area. Habitat niche overlap can be estimated by plot overlap. Many species pairs have relatively high overlap values, thereby indicating that they fre- quently occur in association with each other. Assuming the plot is indicative of the requirements and tolerances of the species found therein, we can conclude that some niche overlap does exist. Four of the major forest areas were compared and contrasted by various analytical techniques. Each area was characterized by the contingency table indices. The capoeira-terra firme transition area is relatively dry and has rather dense vegetation; the herpeto- fauna predominantly inhabits low vegetation and tree trunks. The terra firme-varzea transition area can be divided into high ground transition and low, wet transition. The entire transition zone is an intermediate area with respect to the environmental gradients, ex- cept that in many areas the ground cover is less dense than that of the capoeira-terra firme transition area. The varzea plots are rela- tively wet and have fairly dense ground cover. The igap6 forest is the wettest area and has intermediate to relatively dense ground cover; most of the lizards are either terrestrial or are found on the boardwalks, and the frogs are found both on low vegetation and on the ground. Another way of comparing the areas is in terms of species rich- ness and equitability. Species richness values for five major areas are: varzea—38 species; terra firme—36 species; open and edge areas—34 species; igap6—22 species; and capoeira—20 species. Coefficients of community were calculated on these data to deter- mine relative similarity between every two areas. The highest similarity is between capoeira and terra firme forests (0.556), and the lowest is between capoeira and open-edge areas (0.227). Spe- cies diversity (Shannon index) scores were calculated from the contingency table data matrix; the results are: varzea—3.02; capoeira-terra firme transition—3.00; igapd—2.72; and terra firme- varzea transition—2.45. Equitability values were then calculated from the species diversity estimates (H’) and compared to Mac- Arthur’s broken-stick model. The capoeira-terra firme transition area has an equitability of 1.10, indicating that the area is more diverse than would be expected by MacArthur’s model. As dis- 60 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY cussed in the relevant section, the statistical and biological sig- nificance of this analysis is uncertain due to small sample size. The ecological requirements and tolerances characteristic of species in a community vary widely. Every species exploits the available resources in the most effective way possible for that par- ticular species. Some species accomplish this by specializing in part of the environment, whereas others are generalized and utilize a greater spectrum of environmental parameters. It is concluded that the niche segregation existent among the frogs, salamanders, and lizards living in various habitats within the rainforest at Belem permits many species to coexist with highly efficient utilization of environmental resources. ACKNOWLEDGMENTS I am extremely grateful to the many persons who made the field studies in Belém possible. Initial research was supported by A.R.O. Grant 7184-EN to the Smithsonian Institution and by the Museum of Natural History Director’s Fund, University of Kansas. Field work in 1970 was supported through a combination of monies from the W. G. Saul Fund, the Museum of Natural History Director's Fund, the Watkin’s Museum of Natural History Fund, and a grant from the National Science Foundation through the Committee on Systematic and Evolutionary Biology, all at the University of Kan- sas. I thank the officials of IPEAN (Instituto de Pesquisas e Ex- perimentacgao Agronomicas do Norte) for permission to carry out my studies in the APEG reserves (Area de Pesquisas Ecologicas do Guama). John P. Woodall, Director of the Belém Virus Laboratory, provided field and laboratory facilities. Many persons made my field work enjoyable and profitable; my thanks go to William E. Duellman, Philip S. Humphrey, Thomas E. Lovejoy, and Alan D. Crump for their guidance, help, and com- panionship in the field. I am also grateful to Carlos Cabega and the many other Brasilians who assisted with field collections; with- out their efforts my study would not have been possible. I am indebted to the numerous persons who helped with the synthesis of this paper. William E. Duellman aided with identifica- tion of specimens; William H. Hatheway ran my data through com- puter programs at the University of Washington. Philip $. Humph- rey, William E. Duellman, and Orley R. Taylor contributed nu- merous suggestions and ideas with regard to the writing of this paper. Also, I am grateful to Stephen R. Edwards, Paul B. Robert- son, and Janalee P. Caldwell for their comments and suggestions. ANALYSIS OF ECOLOGICAL DISTRIBUTION 6] Linda Trueb assisted with formats for illustrations and Thomas H. Swearingen executed most of the final drawings. Judy Macura typed the manuscript. Finally, I am grateful to all the many persons who provided endless encouragement throughout the course of study; special thanks go to Abraham Goldgewicht for his constant patience, under- standing, and confidence. LITERATURE CITED BELEM Virus LABORATORY, BELEM, PARA, BRAZIL 1967. Annual report. Unpublished. Cain, S. A., G. M. pe O. Castro, J. M. Pires, and N. T. pa Sitva 1956. Application of some phytosociological techniques to Brazilian rain forest. Amer. J. Botany, 43:911-941. DUELLMAN, W. E. 1967. Courtship isolating mechanisms in Costa Rican hylid frogs. Herpe- tologica, 23:169-183. Hairston, N. G., J. D. ALLEN, R. K. COLWELL, D. J. FuruyMa, J. HoweELt, M. D. Lupin, J. MATHIAS, and J. H. VANDERMEER 1968. The relationship between species diversity and stability: an experi- mental approach with Protozoa and bacteria. Ecology, 49(6):1091- ONS HatHeway, W. H. 1967. Physiognomic characterizations of three vegetational types at the Guama Ecological Research Area, Belém, Brazil. 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Nat., 98 :387-398. 1965. Patterns of species diversity. Biol. Rev., 40:510-533. 62 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY MacArtuur, R. H. and J. W. MacArruur 1961. On bird species diversity. Ecology, 42:594-598. MacArtuvr, R. H., J. W. MAcArTHUR, and J. PREER 1962. On bird species diversity: II. Prediction of bird census from habi- tat measurements. Amer. Nat., 96:167-174. MacArtuur, R. H., H. RECHER, and M. Copy 1966. On the relation between habitat selection and species diversity. Amer. Nat., 100:319-332. PIANKA, E. R. 1967. Lizard species diversity. Ecology, 48(3) :333-351. PreELou, FE. C. 1966. Shannon’s formula as a measure of specific diversity: its use and misuse. Amer. Nat., 100:463-465. RAnp, A. S. 1964. Ecological distribution in anoline lizards of Puerto Rico. Ecology, 45:745-752. Ranp, A. S. and S. S. HUMPHREY 1968. Interspecific competition in the tropical rain forest: ecological distribution among lizards at Belém, Para. Proc. U.S. Natl. Mus., 125 C3658) 17 SCHOENER, T. W. 1968. The Anolis lizards of Bimini: resource partitioning in a complex fauna. Ecology, 49:704-726. 1970. Nonsynchronous spatial overlap of lizards in patchy habitats. Ecol- ogy, 51(3):408-418. SCHOENER, T. W. and G. C. GoRMAN 1968. Some niche differences in three Lesser Antillean lizards of the genus Anolis. Ecology, 49(5):819-830. SHANNON, C. E. 1948. The mathematical theory of communication. In C. E. Shannon and W. Weaver, The mathematical theory of communication. Univ. Illinois Press, Urbana, 117 pp. WHiTTAKER, R. H. 1970. Communities and ecosystems. The Macmillan Company, Collier- Macmillan Limited, London, 162 pp. WitumaMs, E. J. 1952. Use of scores for the analysis of association in contingency tables. Biometrika, 39:274-289. UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly pub- lished in the above series are now published as Occasional Papers, Museum of Natural History. The Miscellaneous Pub- lications, Museum of Natural History, began with number 1 in 1946. Longer research papers are published in that series. Monographs of the Museum of Natural History were initiated in 1970. Institutional libraries interested in exchanging publications may obtain the Occasional Papers and Miscellaneous Publica- tions by addressing the Exchange Librarian, University of Kan- sas Library, Lawrence, Kansas 66044. Individuals may pur- chase separate numbers of all series. Prices may be obtained upon request addressed to Publications Secretary, Museum of Natural History, University of Kansas, Lawrence, Kansas 66044. ; o~ Ar © (13 34Lo Vln u BRA RAIN . ¥ | \ JUN 15 1971 \\ Pe mIAW/OoOCrs oC NOT Fe ECEIVED =< y, OCCASIONAL PAPERS EMy oF SWRA of the MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas NUMBER 4, PAGES 1-21 JUNE 3, 1971 A TAXONOMIC REVIEW OF SOUTH AMERICAN HYLID FROGS, GENUS PHRYNOHYAS By WILLIAM E.. DUELLMAN' Few large South American hylid frogs are so poorly known taxonomically as the widespread Phrynohyas venulosa (Laurenti, 1768) and its cogeners. Duellman (1956) included three South American species in the genus; four other species were recognized in Central America and México. Duellman (1966) and McDiarmid (1968) reduced all of the Middle American nominal species to synonyms of Phrynohyas venulosa. Duellman (1968) resurrected Hyla coriacea Peters from the synonymy of Phrynohyas venulosa and showed that coriacea was a valid species of Phrynohyas in the Guianas and Amazon Basin. Bokermann (1966) listed Hyla meso- phaea Hensel and Hyla imitatrix Miranda-Ribeiro in the genus Phrynohyas. The purposes of the present paper are to: 1) redefine the hylid genus Phrynohyas, 2) describe the geographic variation in the wide- spread Phrynohyas venulosa, 3) ascertain the assignments of several nominal species, and 4) present a synopsis of the species in the genus. Specimens examined are designated by the following abbrevi- ations of depositories: AMNH _ American Museum of Natural History ANSP Academy of Natural Sciences, Philadelphia BMNH _ British Museum ( Natural History ) CAS California Academy of Sciences CAS-SU | Stanford University collection, now in the California Academy of Sciences FMNH _ Field Museum of Natural History 1 Curator, Division of Herpetology, Museum of Natural History, University of Kansas. 2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY IRSNB Institut Royal des Sciences Naturelles Belgique KU University of Kansas Museum of Natural History LACM Los Angeles County Museum of Natural History MCZ Museum of Comparative Zoology, Harvard MLS Museo de La Salle, Bogota MJP Museo Javier Prado, Lima MNHN~ Museum National d’Histoire Naturelle, Paris MNRJ Museu Nacional, Rio de Janeiro MZUSP Museu de Zoologia, Universidade de Sao Paulo NHMG __ Naturhistoriska Museet, Goteborg NHMW _ Naturhistorisches Museum, Wien NHRM _ Naturhistoriska Riksmuseet, Stockholm RMNH _ Riksmuseet van Natuurlijke Histoire, Leiden SMF Senckenbergische Museum, Frankfurt UIMNH_ University of Illinois Museum of Natural History UMMZ University of Michigan Museum of Zoology USNM _ United States National Museum UZM Universitets Zoologiske Museum, Copenhagen WCAB Werner C. A. Bokermann, SAo Paulo, Brasil 7Z1US Zoologiske Institut, Universitet Stockholm ZMB Zoologisches Museum, Berlin ZSM Zoologisches Staatssammlung, Munich ACKNOWLEDGMENTS For permission to examine specimens in their care, I am grateful to James E. Bohlke, Werner C. A. Bokermann, F. W. Braestrup, Nelly Carrillo de Espinoza, Gaston F. deWitte, Josef Eiselt, Alice G. C. Grandison, Jean Guibé, Birgitta Hansson, Walter Hellmich, M. S. Hoogmoed, Robert F. Inger, Konrad Klemmer, Alan E. Leviton, Bertha Lutz, C. J. McCoy, Charles W. Myers, Hno. Nicéforo Maria, Gunther Peters, James A. Peters, Dorothy M. Smith, Paulo E. Vanzolini, Greta Vestergren, Charles F. Walker, Ernest E. Williams, John W. Wright, and Richard G. Zweifel. I am especially indebted to Linda Trueb for her astute osteo- logical observations and illustrations incorporated herein and for her aid in examining specimens in many museums. Study of specimens in European museums was made possible by a grant (No. 5063) from the Penrose Fund of the American Philosophical Society. PHRYNOHYAS FITZINGER, 1843 Phrynohyas Fitzinger, 1843:30 [type species, Hyla zonata Spix, 1824 (= Rana venulosa Laurenti, 1768) by monotypy and declaration of Intemational Commission on Zoological Nomenclature (Opinion No. 520, 1958) ]. Acrodytes Fitzinger, 1843:30 [type species, Hyla venulosa Daudin, 1803 (= Rana venulosa Laurenti, 1768) by monotypy; generic name suppressed by International Commission on Zoological Nomenclature (Opinion No. 520, 1958) for purposes of Law of Priority, but not for those of Law of Homonymy]. HYLID FROGS, GENUS PHRYNOHYAS 3 Scytopis Cope, 1862:354 [type species, Scytopis hebes Cope, 1862, by monotypy]. Diagnostic Definition—1) Skull slightly broader than long; 2) dermal roofing bones of skull well ossified, lacking exostosis and co-ossification; 3) nasals broad, overlapping anterior edge of sphen- ethmoid, not articulating with maxillaries: 4) frontoparietals hav- ing complete medial articulation, entirely covering frontoparietal fontanelle; posterolateral margin of frontoparietal lying just medial to epiotic eminence; 5) sphenethmoid well ossified with large portion exposed dorsally; 6) maxillary arch complete; pars facialis of maxil- lary moderately well developed but not articulating with nasal; 7) squamosal weakly articulating with prootic; zygomatic ramus extending about one-half of distance to maxillary; 8) pterygoid robust with complete articulations; 9) dentigerous processes of pre- vomers curved; 10) parasphenoid large, edentate, acuminate anteri- orly; alae oriented posterolaterally; 11) palatines moderately large, widely separated medially, bearing dentigerous ridge; 12) neural arches of vertebrae low, non-imbricate; 13) transverse processes of vertebrae subequal in width; width of processes on third presacral vertebra less than width of sacral diapophyses; 14) sacral diapophy- ses expanded; 15) ilial shaft long, cylindrical, with low dorsal pro- tuberance; 16) pubis cartilaginous; 17) clavicles expanded distally, strongly arched; 18) coracoids expanded distally and proximally, widely separated medially; 19) epicoracoids broadly overlapping; 20) suprascapula large, well ossified; cleithrum ossification incorpo- rated into suprascapula; 21) vocal sacs paired, lateral, behind angles of jaws; 22) submentalis muscle moderate in size, araphic; 23) inter- mandibularis muscle undifferentiated, lacking aponeurosis; 24) in- terhyoideus muscle forming large supramandibular lobes; no myointegumental contact between lobes and sacs; 25) postman- dibular septum short, attached to posterior end of intermandibular portion of interhyoideus; 26) length of vocal slits about one-fifth of length of mandible; 27) skin on dorsum thick, glandular, and smooth, granular, or tuberculate; that on venter heavily granular; 28) parotoid glands extensively developed in occipital and scapular regions; 29) diameter of tympanum 50-75 percent of diameter of eye; upper edge covered by supratympanic fold; 30) discs large, round; 31) fingers one-fourth to two-thirds webbed; toes about three-fourths webbed; 32) smooth, horny nuptial excrescences in breeding males; 33) pupil horizontal; 34) palpebrum clear; 35) snout-vent length of adults 50-114 mm; 36) known tadpoles having four upper and six lower rows of teeth. 4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 40 T BP coriocea @ 2 imitatrix A P. mesophaea @ PP venlosa ) @ ) %% e es ve at S ® eo.” OF \C ae eo @ eo Pe i —- De he ae 2 e 5 \ oo N ‘ ~ u \ J Nel X \ lOF ¢ & i = 20 16) 400 1000 KILOMETERS Fic. 1. Map showing locality records for the species of Phrynohyas. Content.—As defined here, the genus contains four species: P. coriacea (Peters), P. imitatrix (Miranda-Ribeiro), P. mesophaea (Hensel), and P. venulosa (Laurenti, 1768). Distribution —The lowland tropics from Sinaloa and Tamaulipas, México, southward to Chaco, Entre Rios, and Salta, Argentina, and Rio Grande do Sul, Brasil. The genus occurs on Trinidad and Tobago, but only two specimens are known from the Pacific low- lands of South America (Fig. 1). Most records of occurrence are at elevations of less than 500 m, but the frogs ascend the Amazonian slopes of the Andes to about 800 m and are known to occur at elevations of 1200 m in the Serra do Mar and Serra da Mantiqueira in southeastern Brasil. Comments.—As pointed out by Trueb and Duellman (1971:38), only four hylid genera have paired lateral vocal sacs; these are Argenteohyla, Osteocephalus, Phrynohyas, and Trachycephalus. Tyler (1971:349) provided evidence that the paired lateral vocal sacs were derived from single subgular sacs and stated: “Conceiv- ably the very highly specialized vocal sacs of Phrynohyas and Trachycephalus could have been derived from the simpler structures now found in some species of Osteocephalus.” Trueb and Duellman (1971:40) concluded that Phrynohyas and Trachycephalus probably HYLID FROGS, GENUS PHRYNOHYAS 5 were representatives of one phyletic line. Trachycephalus differs from Phrynohyas principally by having a heavily casqued, co-ossified skull and by lacking extensive parotoid glands. GEOGRAPHIC VARIATION IN PHRYNOHYAS VENULOSA In 1956 I noted the presence of variation in size and coloration in P. venulosa in the Amazon Basin; furthermore, I recognized, taxonomically two peripheral populations—P. hebes in the Gran Chaco and P. ingens in the Maracaibo Basin. Examination of many additional South American specimens reveals that the mosaic of variation is more complex than that described previously. Size——In South America very large individuals of P. venulosa occur in the Maracaibo Basin (females to 112.5 mm in snout-vent length) and in Amazonian Ecuador (females to 110.2 mm in snout- vent length). Equally large individuals are known from Panama and Costa Rica (Duellman, 1970:165). Specimens from other areas in South America are noticeably smaller (Table 1). Measurements and proportions of the limbs and head all are positively correlated with snout-vent length. Duellman (1956) suggested that there were interpopulational differences in the size of the tympanum, but data from larger series of specimens negate that suggestion (Table 1). Also, there seems to be no correlation between size and habitat, as suggested by McDiarmid (1968). Skin.—Considerable variation is apparent in the thickness and texture of the dorsal skin. Most specimens have scattered pustules dorsally; these are especially noticeable in young individuals. In a few specimens, such as KU 126067 from 2 km east of Leticia, Co- lombia, and especially MNRJ 4054 from Rio Branco, Brasil, the skin on the dorsum is highly pustulate. The parotoid glands result in thickened skin in occipital and scapular regions in most specimens from throughout the range. Many specimens from the upper Ama- zon Basin in Ecuador, Pert, and Bolivia have thick, leathery skin. McDiarmid (1968) suggested that extensive development of the parotoid glands is a seasonal phenomenon in frogs living in regions having wet and dry seasons, the glands being better developed in the dry season. Dates of collection are not available for many South American specimens of P. venulosa; the specimens having such data do not provide strong evidence in support of McDiarmid’s proposal. For example, well-developed glands are evident in specimens from the upper Amazon Basin where there is no prolonged dry season; SI ec © = 2 se a = ec = = = Zz ey © =. = es N 5 2 ip) fc fx] Ay < A 4 = Zz ° WY < S S) © 6 (9°¢) (67) 6 9-0'S GS-8 (SP) (L'P) 0'9-0'F 0'S-S'F (T'S) (T'S) g’c-0'S G'¢-0'S (9'8) (8'9) T'6-1'8 VL-9'S ($'F) : O'S-L'F (VP) (VT) LE-UP LY-6'S (9'9) (e'g) O'L-1'9 0'9-S'F (L'P) (SF) CP-0'F 0'9-0'F é 9 yysueT wunueduids 7, (OTS) 1'68-0°89 (O'eL) G’88-S'6S (Tek) CPVLO1L (9° LOT ) GOIT-8°c0I (9°99) 6'89-T'E9 (7 LOT) CCII-S' v6 (3°19) G'GL-0'S9 5 GE Sz) 0'98-3'F9 (0°69) 0'92-0'S9 (60) G'EL-S' 19 (L'L8) C'Z6-8'F8 (369) G'0L-6'€9 (€°99) 6'GL-S'8S (0'FS8) 0°88-0'S2 (6'FL) 0'°Z8-0'L9 2 Yqsue'T JUAA-ynNoUs N \ Rania > eee ae See eases a ree Aenseleg Ss a i i aig ae sh a TE Zar) ByUeS : eIATIOG ae so ee ae ee aes Pg eee tuog [HW ‘BIAllog SERRE REEEEREEEEEEEREEEEEEEEEEEEEEEEeeEEE ezeyseg-odeN, :1openoq ee. hay Oe Se ee ee > SS BIOTjO"T + BIQUIO[OT) EE 1950 of 2S Peres ho Ge ae ees a ee Woleq : [set Viele 2 ee Eo ae eae ulseq Oqlvorleyy -e[oNnZIUI A Goes eae ee ae ee pepruny 2 AyeooT ‘sasUeI PIAIOSGO MOTAq SasayzUered Ul UAAIS Ie SURIUT ‘sIOJOUIIT[IW UT 91e SyUOUTOINSeI|y ‘psojnuaa sphyoufisyg Ul syuoUIaINseay| Ule}9D Ul UOTVLIeA—'T ATAVL, HYLID FROGS, GENUS PHRYNOHYAS 7 Fic. 2. Dorsal color patterns in Phrynohyas venulosa. A. Normal, code 4, KU 92258. B. Divided, code 3, KU 126067. C. Spotted, code 2, NHMW 19013 (holotype of Hyla wettsteini). specimens from Napo and Pastaza provinces in Ecuador have more extensive glands than do individuals from the xeric Maracaibo Basin. Coloration.—All individuals of P. venulosa are brown or tan dorsally with or without darker markings. The venter is uniform grayish white or pale tan. The iris is golden with radiating black lines. There is considerable variation in dorsal color pattern, but most of the variants can be grouped into one of four categories; code numbers indicate a decreasing amount of dark pigment dorsally (Fig. 2): NORMAL: A large middorsal dark blotch extending from the occiput to the rump; blotch divided anteriorly by a longitu- dinal band of ground color in some specimens (coded 4). DIVIDED: As above, but blotch divided by a broad transverse band of ground color (coded 3). SPOTTED: Dorsal pattern consisting of irregular small dark spots (coded 2). PLAIN: Dorsum uniform brown (coded 1). The various color patterns are found throughout most of the range of the species in South America (Table 2). The only region in which all specimens have the normal pattern is Paraguay; the next highest percentage of specimens having the normal pattern is Trini- dad (93%). All four color pattern types were found in only four of 14 pooled samples; normal and plain patterns occurred in 12 sam- ples, and divided and spotted patterns occurred in eight samples. In the only large sample (151 specimens from Maracay, Venezuela) the following percentages occur: normal 14.6, divided 44.3, spotted 7.9, and plain 33.2. Each of the various patterns has been found in males, females, and juveniles. All four patterns, save the spotted one, are found in Central America. 8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY TABLE 2.—Variation in Dorsal Color Pattern in Phrynohyas venulosa. See text for explanation of patterns and scoring. Locality N Normal Divided Spotted Plain Score ‘Ramidae teins te eS Et 32 30 1 1 0 3.91 Venezuela: Maracaibo Basin... 8 0 0 7 { 1.89 Venezuela: Maracay ____- 151 22 67 12 50 2.38 Stuamanne 2 eee ne pa ats) 0 Z 4 3.28 BeasileBelenie = 2. oes” | ha 26 23 1 0 2 fs) Brasil: Mato Grosso! 2 15 DA 0 174 J 2.20 Brasil: Manaus: 22. 8 3 3 0 2 2.87 Colombia leucia®—.... 5 1 1 0 3 2.00 Ecuador: Napo-Pastaza 1653 0 1 0 12 1.25 Reruc Rebase tet 215. ena 10 4 0 4 2 2.60 Pera: Pocallpa 2. Sa 12 6 0 0 6 2.50 Bolivia: El Beni =. -- 2s 16 1 1 2 12 1.44 Bolivia: Santa: Criz, 22... 15 4 il 3 7 OAS Paraeuiay =... eee oes Oe 14 14 0 0 0 4.00 Remarks.—It is evident that the various characters considered to be of taxonomic value in this group vary independently of one another in P. venulosa. Refined quantitative analysis must await the acquisition of large series of specimens from throughout the range of the species. Currently, karyological, larval, and biosonic data are TABLE 3.—Alphabetical Synonymy of the Species of Phrynohyas. Trivial Name, Original Generic Name, Author, and Date Current Name adenodetma( Hyla) Lutz; 1968 ties: 2. oes eee ee P. venulosa bufona (Hyle) Spix, 1624 2.2 2. fae. 2 2 P. venulosa corasterias (Phrynohyas) Shannon and Humphrey, 1957 __- P. venulosa coriacea (Hyla) Peters, 1867... 2 ee ee P. coriacea hebes (Scytopis) ‘Cope; 1862, 2. Ooi) eis. eee P. venulosa imutatrix (Hyla) Miranda-Ribeiro, 1926 _... 2 ee ee P. imitatrix injlata (Acredutes) Laylor, BO44 2. 6p 2 P. venulosa ingens (Phrynohyas) Duellman, 1956 = eee P. venulosa latifasciata (Phrynohyas) Duellman, 19562... P. venulosa lichenosa (Hula) Ganther, 10509 2. eee P. venulosa macrotis (Hila) Anderxsson, 1945". 2325 Ei a ee P. venulosa mesophaca (Hyla) Hensel, 1867)... be. oom bss) sor) eee P. mesophaea modesta (Acrodytes) Taylor and Smith, 1945 P. venulosa migropunctata (gla) Boulenger, 1682 ee ee P. venulosa paemilata {(Hyla) Broechi, 1879 22. ae) Le ae P. venulosa palpebrogranulata (Hyla) Andersson, 1906 P. venulosa quadrangulum (Hyla) Boulenger, 1882... P. coriacea resinactnx ( Hyla) Coeldi, 1907 vz. ee ee P. venulosa spilomma (Hyla)’Gope: 1817 = See ee eee P. venulosa venulosa (Rana) Waaurenti, 1768 Lae. es ee P. venulosa vermiculata (Hyla) Duméril and Bibron, 1841 __...--- P. venulosa wettsteint (Hyla) AM, 1933 2.) ee ee P. venulosa gemigt (lglg) Ahi 4939 ie ae ee ee eee P. venulosa zonata (Hyla) Spix, 1824... 72. 2 eee ee P. venulosa HYLID FROGS, GENUS PHRYNOHYAS 9 B C Fic. 3. Anterior cranial roofing bones showing positions of nasals and proportion of dorsal exposure of the sphenethmoid. A. P. coriacea, KU 105249. B. P. imitatrix, KU 92249. C. P. mesophaea, KU 92257. D. P. venulosa, KU 138830. Abbreviations: fp = frontoparietal, n = nasal; sphenethmoid is stippled. lacking. Until such data are available for a more sophisticated analysis, we must be content to base conclusions on the present in- adequate series of preserved specimens, which have formed the basis for my broad definition of Phrynohyas venulosa. TAXONOMY OF PHRYNOHYAS Data taken from series of preserved specimens, some living frogs, skeletal preparations, and radiographs provide a basis for the recog- nition of four species in the genus Phrynohyas (Table 3). Diagnostic external morphological characters and features of coloration are used in the following key. Osteological characters that are important at the species level are discussed below. Nasals.—The size and position (relative to one another) of the nasals are distinctive (Fig. 3). In P. imitatrix and venulosa the nasals are juxtaposed; in mesophaea they are narrowly separated, and in coriacea they are widely separated. Sphenethmoid.—The sphenethmoid is well ossified in all species. The proportion of the element exposed dorsally shows interspecific differences (Fig. 3); the area of exposure is dependent on the rela- tive sizes and positions of adjacent dermal roofing bones—the nasals anteriorly and the frontoparietals posteriorly. The sphenethmoid is least exposed in P. imitatrix and moderately exposed in mesophaea and venulosa. A relatively large area of the sphenethmoid is exposed in coriacea, in which the nasals are widely separated. Prevomers.—The dentigerous processes are curved and separated medially in all species. The curvature is slight and the separation moderate in P. imitatrix and venulosa, slight and widely separated in coriacea, and strongly curved and narrowly separated in mesophaea. Sacral Diapophyses.—The diapophyses are dilated in all species; this dilation is symmetrical in P. mesophaea and venulosa and asym- metrical (greater dilation posteriorly) in imitatrix and coriacea. 10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY KEY TO THE SpEcIES OF Phrynohyas 1. Dorsal quadrangular mark outlined with white and terminating in sacral region; thighs lacking dark transverse bars; black post- tympanic spot present; red “flash-colors” in life P. coriacea Dorsal quadrangular mark, if present, usually not outlined with white; thighs usually having dark transverse bars (if blotches present on dorsum); black post-tympanic spot absent; no red “flash-colors’ ‘in. life. <2... bis oe 2 2. Dorsum and flanks dark brown with broad cream dorsolateral stripe from eyelid to groin; limbs unicolor or faintly barred CCE Lg Ne IES Po NET LE Ne P. mesophaea Dorsal pattern variable, but lacking dorsolateral cream strips; limbs usually distinctly barred, if blotch present on dorsum __ 3 3. Dorsal markings consisting of large brown blotch anteriorly and usually round spots or blotches posteriorly, darkest peripherally; limbs with distinct dark brown, light bordered bars; dorsal skin distinctly tuberculate 1.2 eee P. imitatrix Dorsal markings absent or consisting of large dark blotch (trans- versely divided in some) or small dark spots; limbs unicolor or barred, lacking light borders; dorsal skin variable —___ P. venulosa Phrynohyas coriacea (Peters) Hyla coriacea Peters, 1867:711 [syntypes, ZMB 5807 (3 specimens) from Surinam; H. Kappler collector]. Hyla quadrangulum Boulenger, 1882:367 [holotype, BMNH 1947.2.13.75 from “western Ecuador’; Mr. Fraser collector]. New synonymy. Phrynohyas coriacea—Duellman, 1968 :205. Distribution.—Surinam and the upper Amazon Basin in Ecuador and Pert at known elevations of less than 500 m (Fig. 1). The record for Machala on the Pacific coast of Ecuador probably is erroneous. 29 specimens from 11 localities. Remarks.—The three syntypes (ZMB 5807) were reported on by Duellman (1968). The holotype of Hyla quadrangulum (BMNH 1947.2.13.75) is an adult female having a snout-vent length of 57.5 mm and morphological characters which render it indistinguishable from P. coriacea. The color pattern consists of a dark brown qua- drangular dorsal blotch faintly outlined with cream on a tan dor- sum; the limbs are distinctly barred, and a dark post-tympanic spot is evident. HYLID FROGS, GENUS PHRYNOHYAS 1] The type locality of “western Ecuador” cannot be taken seriously; several species contained in the Fraser collections supposedly from western Ecuador have been found subsequently only in the Amazon Basin and eastern slopes of the Andes. Phrynohyas imitatrix (Miranda-Ribeiro) Hyla imitatrix Miranda-Ribeiro, 1926:77 [syntypes, MNRJ 154 (2 specimens) from Teresdpolis, Rio de Janeiro, Brasil; Alipio de Miranda-Ribeiro col- lector]. Phrynohyas mesophaea—Bokermann, 1966:55. Distribution —Mountains of southeastern Brasil—Serra do Mar, Serra da Bocaina, and Serra da Mantiqueira—in the states of Rio de Janeiro, Sao Paulo, and Santa Catarina (Fig. 1); the known localities are at elevations between 800 and 1200 m. Eight specimens from three localities. Phrynohyas mesophaea (Hensel) Hyla mesophaea Hensel, 1867:154 [holotype, ZMB 6810 from Porto Alegre, Rio Grande do Sul, Brasil; Reinhold Hensel collector]. Phrynohyas mesophaea—Bokerman, 1966:55. Distribution—Coastal lowlands of southeastern Brasil from Bahia to Rio Grande do Sul (Fig. 1). 259 specimens from 13 localities. Phrynohyas venulosa (Laurenti) Rana venulosa Laurenti, 1768:31 [based on a plate in Seba (1734, vol. 1, pl. 72, fig. 4)]. Hyla venulosa—Daudin, 1803:71. Hyla zonata Spix, 1824-41 [type formerly in ZSM, now lost, from Lago Tefé, Amazonas, Brasil; collector unknown]. Hyla bufonia Spix, 1824:42 [type formerly in ZSM, now lost, from Eca (= Tefé), Amazonas, Brasil; collector unknown]. Hyla vermiculata Duméril and Bibron, 1841:563 [holotype, MNHN 4797 from “Amerique Sept.”; presented by Mr. Harlan]. Phrynohyas zonata—Fitzinger, 1843:30. Acrodytes venulosa—Fitzinger, 1843:30. Hyla lichenosa Gunther, 1859 [lectotype, BMNH 1936.12.3.119 from Amazonas, Brasil; collector unknown]. Scytopis hebes Cope, 1862:354 [holotype, USNM 5837 from “Paraguay” (re- stricted to Asuncién by Duellman, 1956:16); T. J. Page collector]. New synonymy. Scytopis venulosus—Cope, 1866:85. Hyla spilomma Cope, 1877:86 [holotype, unknown, apparently lost, from Cosamaloapam, Veracruz, México; Francis Sumichrast collector]. Hyla paenulata Brocchi, 1879:21 [holotype, formerly in MNHN, now lost, from western (= southern ?) Guatemala; collector unknown]. Hyla nigropunctata Boulenger, 1882:366 [syntypes, BMNH _ 59.9.20.2 and 81.10.31.20 from Jalapa, Veracruz, México; Alphonse Hoege collector]. Hyla_ palpebrogranulata Andersson, 1906:14 [holotype, ZIUS A133 from Tatarenda, Santa Cruz, Bolivia; Erland Nordenskiold collector]. New synonymy. 12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Hyla resinifictrix Goeldi, 1907:135 [holotype, BMNH 1947.2.23.24 from San Antonio do Prata, Para, Brasil; Emil Goeldi collector]. Hyla zernyi Ahl, 1933:27 [holotype, NHMW 16503 from Taperinha, near Santarém, Para, Brasil; H. Zerny collector]. New synonymy. Hyla zeryni Ahl, 1933:27 [holotype, NHMW 16503 from Taperinha, near Santarém, Parad, Brasil; H. Zerny collector]. New Synonymy. Acrodytes inflata Taylor, 1944:63 [holotype, FMNH 100046 from La Venta, Guerrero, México; Edward H. Taylor collector]. Acrodytes spilomma—tTaylor, 1944:64. Acrodytes modesta Taylor and Smith, 1945:594 [holotype, USNM 115013 from Cruz de Piedra, near Acacoyaqua, Chiapas, México; Hobart M. Smith collector]. Hyla macrotis Andersson, 1945:70 [holotype, NHRM 1958 from the Rio Pastaza watershed, Ecuador; William Clarke-MaclIntyre collector]. New synonymy. Phrynohyas hebes—Duellman, 1956:16. Phrynohyas inflata—Duellman, 1956:19. Phrynohyas ingens Duellman, 1956:22 [holotype, UMMZ 55570 from La Fria, Pueblo Nuevo, Zulia, Venezuela; H. B. Baker collector]. New synonymy. Phrynohyas latifasciata Duellman, 1956:24 [holotype, BMNH 83.2.7.1 from Presidio, Sinaloa, México; Alfonso Forrer collector]. Phrynohyas modesta—Duellman, 1956:25. Phrynohyas spilomma—Duellman, 1956: 28. Phrynohyas corasterias Shannon and Humphrey, 1957:15 [holotype, UIMNH 67060 from 4.8 miles east of San Blas, Nayarit, México; Francis L. Humphrey and Frederick A. Shannon collectors]. Phrynohyas venulosa—Hemming, 1958:172. Hyla tibiatrix tibiatrix—Rivero, 1961:127. Hyla tibiatrix ingens—Rivero, 1961:131. Hyla venulosa venulosa—Rivero, 1964:493. Hyla adenoderma Lutz, 1968:3 [holotype, MNRJ 4054 from Rio Branco, a tributary of the Rio Madeira, Rondonia, Brasil; F. Casper collector]. New synonymy. Distribution—In South America—the Caribbean lowlands, in- cluding Trinidad and Tobago, the Amazon Basin, the Chaco, and northeastern Brasil (Fig. 1). Only one specimen is known from the Pacific lowlands; Duellman (1956:42) reported that specimen as being from Hacienda San Miguel, Milagro, Guayas Province, Ecuador, but James A. Peters, the collector, informed me that it actually came from 7 kilometers southeast of Buenavista, El] Oro Province, Ecuador. All specimens are from elevations of less than 800 m. 475 specimens from 107 localities. Remarks.—This widespread, variable species has received 19 names in 200 years. The earliest name, Rana venulosa Laurenti, 1768, was based on a plate in Seba’s “Thesaurus” (1734). Type specimens of Hyla zonata Spix, H. bufonia Spix, H. spilomma Cope, and H. paenulata Brocchi are no longer extant. I have examined the types of the other 14 nominal species. Duellman (1961) provided evidence that P. corasterias Shannon and Humphrey, 1957, was not specifically distinct from P. inflata (Taylor, 1944). Duellman (1966) showed that P. modesta (Taylor and Smith, 1945) was a color morph HYLID FROGS, GENUS PHRYNOHYAS 13 of P. spilomma (Cope, 1877). McDiarmid (1968) analyzed variation in Middle American Phrynohyas and concluded that P. spilomma (Cope, 1877), P. inflata (Taylor, 1944), and P. latifasciata Duellman, 1956, were synonymous with P. venulosa (Laurenti, 1768). My re- cent studies (Duellman, 1970) support McDiarmid’s conclusions; all Middle American populations seem to be correctly allocated to Phrynohyas venulosa. Of the names in the foregoing synonymy, Duellman (1956) listed Hyla vermiculata Dumeril and Bibron, H. lichenosa Giinther, and H. resinifictrix Goeldi in the synonymy of Phrynohyas zonata (Spix) [= P. venulosa (Laurenti) |]. Duellman (1956) included Hyla nigro- punctata Boulenger in the synonymy of Phrynohyas spilomma (Cope), now considered to be a synonym of P. venulosa (Laurenti). Duellman (1956) distinguished Phrynohyas hebes from venulosa by the wider dark limb bands in the former. Examination of addi- tional material from throughout the range of venulosa and especially from Argentina, Bolivia, Paraguay, and southern Brasil reveals that the supposed differences between venulosa and hebes are not con- stant. Thus, I now consider Scytopis hebes Cope, 1862, to be a junior synonym of Phrynohyas venulosa ( Laurenti, 1768). Duellman (1956) distinguished Phrynohyas ingens from venulosa chiefly on the basis of the larger size of the former and by its dorsal coloration consisting of small brown spots on a tan ground color, as opposed to a large dark blotch on the dorsum in venulosa. It is now apparent that the coloration considered to be diagnostic of ingens appears in populations throughout most of the range of venulosa (Table 2). Furthermore, the size of ingens is equaled or exceeded by some individuals from Ecuador and Costa Rica. Consequently, I now consider Phrynohyas ingens Duellman, 1956, to be a junior synonym of Phrynohyas venulosa (Laurenti, 1768). The holotype of Hyla vermiculata Duméril and Bibron (MNHN 4797) is an adult male having a snout-vent length of 65.2 mm. The specimen is soft and has been abraded. The dorsum is medium brown with distinct reddish brown marks; dark marbling is evident on the flanks and posterior surfaces of the thighs. The remnants of the color pattern are reminiscent of one of the color morphs of Phrynohyas venulosa common in northeastern South America. Hyla lichenosa Gimther was based on several juveniles; Smith and Taylor (1948:75) designated BMNH 1936.12.3.119 as the lecto- type. This specimen has the usual tuberculate skin of young Phrynohyas, and it has a color pattern resembling that of young P. venulosa from Belém, Para, Brasil. 14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY The holotype of Hyla resinifictrix Goeldi (BMNH 1947.2.23.24) is a male having a snout-vent length of 75.3 mm. The color pattern is well preserved. The ground color is golden tan; anteriorly, on the dorsum, there is a reddish brown blotch, which is narrowly sepa- rated from a large, posterior blotch. The limbs bear distinct cross bars; the skin on the dorsum is distinctly pustulate. In features of structure and coloration the holotype is like most individuals of P. venulosa from the lower Amazon Basin. The holotype of Hyla palpebrogranulata Andersson (ZIUS A133) is a juvenile having a snout-vent length of 32.3 mm. The skin on the dorsum is tubercular. The dorsum is yellowish tan with a large brown blotch beginning on the head and extending nearly to the vent. Broad brown cross bars are present on the limbs. A pair of pigment clusters above the anal opening suggest that the specimen is a recently metamorphosed young. The iris has the markings characteristic of P. venulosa; broad black streaks radiate from the pupil. This juvenile specimen certainly is a Phrynohyas venulosa. Cochran and Goin (1970) recognized a monotypic “Hyla palpebro- granulata group.” I have examined all of the Colombian specimens that they referred to H. palpebrogranulata and find that all are young Osteocephalus taurinus. The holotype of Hyla wettsteini Ahl (NHMW 19013) is a female having a snout-vent length of 61.3 mm and a moderately tuberculate dorsum. The dorsal surfaces of the head, body, and limbs are pale brown with scattered small brown spots (Fig. 2c). The pattern is identical to that of many individuals of P. venulosa from the middle and upper Amazon Basin and from Venezuela and Guyana. The holotype of Hyla zernyi Ahl (NHMW 16503) is a juvenile having a snout-vent length of 20.4 mm and only slightly tubercular skin on the dorsum. The dorsum is pale tan with a narrow, elongate brown blotch beginning on the eyelids and extending to the sacrum. The flanks are brown, and the limbs are marked with irregular dark brown cross bars. The upper lips are boldly marked by three vertical bars, the first two of which fuse with a broad, diffuse canthal stripe, whereas the third enters the orbit. A broad, postorbital brown stripe encompasses the tympanum and is confluent with the dark flanks. The edge of the lower jaw is dark brown interrupted by creamy white spots. In structure, this specimen is like juvenile P. venulosa; the color pattern is bold but falls within the range of variation of P. venulosa The holotype of Hyla macrotis Andersson (NHRM 1958) is an adult female having a snout-vent length of 110.2 mm. The skin on HYLID FROGS, GENUS PHRYNOHYAS 15 the dorsum is thick and glandular; the parotoid glands are extensive, diffuse, and form a moderately heavy supratympanic fold. The dorsum is uniform reddish brown; the loreal region is slightly paler brown, and the tympanum and groin are pale cream. The dentiger- ous processes of the prevomers are nearly transverse elevations be- tween the choanae and bear 10-10 teeth. Specimens resembling the type have been examined from several localities in Amazonian Ecuador; some of these have small dark spots on the dorsum and thus resemble the smaller spotted color morph known from Ama- zonian Brasil, Peru, and Bolivia. One of the large macrotis-like frogs was prepared as a skeleton; no osteological differences between that specimen and several of P. venulosa were apparent. Consequently, Hyla macrotis Andersson, 1945, is considered to be a junior synonym of Phrynohyas venulosa (Laurenti, 1768). The holotype of Hyla adenoderma Lutz, 1968 (MNRJ 4054) is a female having a snout-vent length of 66.0 mm. The skin on the dorsal surfaces of the head and body is greatly thickened and glandular, more so than in any other specimen of Phrynohyas that I have seen. The dorsal color pattern consists of a network of bold dark brown reticulations on a tan ground color (Lutz, 1968, fig. 1); dark brown cross bars are evident on the limbs. The following data supplement Lutz’s description: The discs on the hands are large and equal in diameter to the tympanum, the upper edge of which is covered by a heavy supratympanic fold. There are 7-9 prevomerine teeth on narrowly separated transverse elevations between the choanae. Although the iris is faded, there is a faint indication of radiating black streaks. This specimen seems to represent an extreme in glandular development of the skin and of a bold reticulated pat- tern. Otherwise, it is unquestionably a P. venulosa. Individuals having a fainter reticulated pattern are known from various localities in the Amazon Basin and Guianas. One specimen from Leticia, Colombia (KU 126067), has dorsal skin nearly as thick and glandular as that of the type of Hyla adenoderma, but the Colombian specimen has a large divided blotch on the dorsum. I consider Hyla adeno- derma Lutz, 1968, to be a junior synonym of Phrynohyas venulosa (Laurenti, 1768). SUMMARY Phrynohyas is one of four genera in the Hylidae having paired lateral vocal sacs behind the angles of the jaws; the other genera are Argenteohyla, Osteocephalus, and Trachycephalus. The four species of Phrynohyas exhibit differences in size, external structural 16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY features, coloration, and cranial osteology. Phrynohyas venulosa is widespread in the tropical lowlands from northern México to north- ern Argentina; the other three species have much more restricted distributions—P. coriacea in Surinam and the upper Amazon Basin, P. mesophaea on the coastal lowlands of southeastern Brasil, and P. imitatrix in the uplands of the coastal ranges of southeastern Brasil. Only P. coriacea and venulosa are known to occur sym- patrically. The variability in size, coloration, and texture of the skin on the dorsum in P. venulosa has resulted in 19 specific names being pro- posed for this species. Seven nominal species are herein relegated to the synonymy of Phrynohyas venulosa (Laurenti, 1768); these are: Scytopis hebes Cope, 1862; Hyla palpebrogranulata Andersson, 1906; Hyla wettsteini Ahl, 1933; Hyla zernyi Ahl, 1933; Hyla macrotis Andersson, 1945; Phrynohyas ingens Duellman, 1956; and Hyla adenoderma Lutz, 1968. Hyla quadrangulum Boulenger, 1882, is placed in the synonymy of Phrynohyas coriacea (Peters, 1867). RESUMEN Phrynohyas es uno de cuatro géneros en la familia Hylidae que poseén sacos vocales laterales apareados, localizados detras de los angulos de las articulaciones mandibulares; los otros géneros son: Argenteohyla, Osteocephalus, y Trachycephalus. Las cuatro especies de Phrynohyas exiben diferencias en tamafio, rasgos estructurales externos, coloracién, y osteologia cranial. Phrynohyas venulosa esta ampliamente distribuida en las tierras bajas tropicales, desde el norte de México al norte de Argentina; las otras tres especies tienen distribuciones mas restringidas—P. coriacea en Surinam y la parte alta de la cuenca amazénica, P. mesophaea en las tierras bajas de la costa del sureste del Brasil, y P. imitatrix en las tierras altas que costean el sureste brasilefio. Se sabe, que sdlo P. coriacea y venulosa ocurren simpatricamente. Diez y nueve nombres especificados han sido propuestos para P. venulosa, debido a la variabilidad del tamafo, coloracién, y textura de la piel dorsal de esta especie. Siete especies nominales son relegadas aqui a la sinonimia de Phrynohyas venulosa (Lau- renti, 1768); estas son: Scytopis hebes Cope, 1862; Hyla palpebro- granulata Andersson, 1906; Hyla wettsteini Ahl, 1933; Hyla zernyi Ahl, 1933; Hyla macrotis Andersson, 1945; Phrynohyas ingens Duell- man, 1956; y Hyla adenoderma Lutz, 1968. Hyla quadrangulum Boulenger, 1882, es puesta en la sinonimia de Phrynohyas coriacea (Peters, 1867). HYLID FROGS, GENUS PHRYNOHYAS iy SPECIMENS EXAMINED The localities for each of the specimens examined are given below. The arrangement of the data is as follows: alphabetically by country, state (department or province), and locality; alpha- betically by the first letter in the abbreviations for the museum, and numerically after each museum abbreviation. Specimens lacking precise locality data are listed first in the most restricted political unit possible; localities which have not been found on maps or which are too vague to be located precisely are given in quotation marks. Where more than one specimen is included under one museum number, the number of specimens is given in parentheses after the museum number. Unless noted otherwise, all specimens are pre- served in alcohol. Phrynohyas coriacea ECUADOR: “western Ecuador,’ BMNH 1947.2.13.75. El Oro: Machala, WCAB 40100. Napo: Lago Agrio, KU 126656; Limon Cocha, KU 107025, UIMNH 87799, 90315-6; Santa Cecilia, KU 105249, UMMZ 129323. Pastaza: “Rio Pastaza watershed,’ NHRM 1950 (7). PERU: Loreto: Pebas, CAS-SU 12622; Puerto Oriente, 5 km. above Contamana, Rio Ucayali, UMMZ 123915. SURINAM: No specific locality, BMNH 70.3.10.69, ZMB 5807 (3); “Spoor- ban,’ RMNH 9606. Marowijne: Albino, Rio Maroni, ZMB 7606 (3); Maroni, ZMB 7605 (2); Nassaugebergte, RMNH 7296. Phrynohyas imitatrix BRASIL: Rio de Janeiro: Teresépolis, MNHN 31/45-46, NHMW 18433.5, SMF 58269. Santa Catarina: Rio Vermelho, KU 92249 (skeleton), 92250-1. Sao Paulo: Sao José do Barreiro, Serra da Bocaina, SMF 62562. Phrynohyas mesophaea BRASIL: No specific locality, UZM 14223, ZMB 3122. Bahia: Santa Cruz, ZMB 30444. Guanabara: Rio de Janeiro, KU 92252-6, 92257 (skeleton), SMF 30024-9. Rio de Janeiro: Angra dos Reis, ZSM 60/1947; “Guapy,” SMF 39251- 3; 36 km from Rio Petrdpolis road, BMNH 1940.4.14.13; Teresépolis, SMF 2515, ZMB 26322. Rio Grande do Sul: Porto Alegre, ZMB 6256, 6810. Santa Catarina: No specific locality, UZM 14224; Blumenau, NHMW 6279-80; Hansa (= Corupa), BMNH 1928.11.5.89-106, SMF 2500-14, 30998, 31001-27; Jaragua, NHMW 6293 (4); Joinville, MNHN 6310 (5), NHMW 6282-3, 6287 (4), 6285 (21). 6289) (2); 6290 (6), 6291 (5), 6294 (5), 6295: (7), 6296 (6), 6297 (5), 6298 (6), 6299 (9), 6300 (6), 6301 (6), 6302 (7), 6303 (4), 6304 (3), 6305 (3), 6306 (7), 6307 (9), 6309 (3), SMF 2516-34, 5120, 5131; “Rio Humboldt,” BMNH 1910.7.26.3-6, 1923.6.1.84-91; “Rio Novo,” NHMW 6286 (5), 6295 (5). Phrynohyas venulosa AMERICA: No specific locality, MNHN 4797. ARGENTINA: No specific locality, USNM 73523. Chaco: Ciervo Petizo, KU 128948-50; Colonia Eliza, NHRM 1503. Entre Rios: Puerto Marquez, 2 km N La Paz, SMF 43634. Salta: Rio Pescado, KU 128946-7. BOLIVIA: No specific locality, NHMW 6177 (3). Cochabamba: Rio Chaparé, ZSM 9/1948. El Beni: Esperanza, BMNH 1920.11.29.15-16; Rer- 18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY renabaque, UMMZ 57528, 57531 (2). La Paz: Ixiamas, MCZ 10087, UMMZ 57529 (4), 57530 (3); Santa Ana de Movimas, BMNH 98.6.9.38. Santa Cruz: Buenavista, BMNH 1927.8.185-6, CM 2582, 3828, 3892-3, 3971, 3974, 3986, 4467-8, UMMZ 60526, 63960 (2), 66559-60; Rio de la Cal, ZMB 32107; Santa Cruz, BMNH 1904.10.29.177, 1907.10.31.72, 1927.8.1.87, CM 2514; Tatarenda, ZIUS Al3s. BRASIL: “lower Amazonia,’ USNM 28926-7. Amazonas: No specific ]Jo- cality, BMNH 1936.12.3.119; Aiapua, ZMB 30303; Fonte Boa, NHMW 6104; Manacapurt, Rio Solimdes, ZMB 37080, ZSM 23/1924; Manaus, AMNH 51755, IRSNB 2681 (14), KU 92258-61, UMMZ 66647; Rio Caicara, NHMW 6111 (2), 6119 (2), 13262 (2); Rio Juruéa, BMNH 1904.11.4.12; Rio Purts, ZMB 30307; Tefé, NHMW 6100. Bahia: No specific locality, BMNH 64.1.19.34, NHMW 6281. Sao Goncalo, 30 km SW Feira de Santana, KU 29438, MCZ 1531. Mato Grosso: Barro do Tapirapé, CAS-SU 12304-10, 12312, KU 40219; Chapada dos Guimaraes, BMNH 1903.3.26.54-6; Posto Jacari, Rio Xingu, KU 92262-3; Villa Murtinho, UMMZ 56770. Minas Gerais: La Filomena, NHMW 6102 (2), 6112 (2), 6114 (3), 6115; Manga, Rio Sao Francisco, UMMZ 108894; Piraporinha, USNM 98536-7; Sao Luis dos Caceres, CAS-SU 12298-9. Pard: Belém (= Para), BMNH 45.8.25.143, 96.1.7.2, CAS-SU 12390-3, NHMW 6098, ZMB 33189, ZSM 119/1911 (2); Cachoeira de Arari, Ilha de Marajé, ZMB 29999, ZSM 120/1911; Cameta, Rio Tocantins, NHMW 6099, 6107, 6113; “Ilha de Marajé,” BMNH 1923.11.9.26; IPEAN, 3 km E Belém, KU 127963-4, 128557-61, 130121-6; Livramento, CAS-SU 12383-7; “San Antonio do Prata,” BMNH 1947.2.23.24; Santarém, BMNH 56.3.25.14 (2); Taperinha, near San- tarém, NHMW 16503, 19013. Pernambuco: No specific locality, BMNH 58.11.28.48; Bonita, MCZ 4998. Piaui: Bosqus, BMNH 1926.5.26.4. Rondonia: Rio Branco, MNRJ 4054. Roraima: Serra Grande, below Boa Vista, Rio Branco, NHMW 15893. COLOMBIA: Amazonas: Leticia, KU 124897-900, 124955-6 (tadpoles); 2 km E Leticia, KU 126067. Caquetd: Morelia, ANSP 25313. Cundinamarca: Honda, MLS 1-2. Magdalena: Barranquilla, MLS 3; Fundacién, UMMZ 48263. Norte de Santander: Rio Zulia, N of Cucuta, MLS 4. ECUADOR: No specific locality, USNM 14054. El Oro: 7 km SE Buena- vista, USNM 192507. Napo: Avila, USNM 165979-80; Lago Agrio, KU 126657; Rio Cotapino, KU 138736, USNM _ 165977-8. Pastaza: Montalvo, CAS-SU 10318:9, USNM 165981 (skeleton), 165982-3; Rio Bobonaza, USNM 165984; upper Rio Oglan, USNM 165986; “Rio Pastaza watershed,’ NHRM 1958; Rio Villano, USNM 165988. FRENCH GUIANA: Cayenne: Cayenne, BMNH 53.2.89-90, NHMW 6101, 6103; Mt. Cabasson, LACM 44638. GUYANA: East Demerara: Georgetown, MCZ 2618, UMMZ 80495. Es- sequibo: Better Hope, BMNH 80.11.22.5-6. Mazaruni-Potaro: Kaieteur Falls, AMNH 23128. North West: Santa Rosa Island, Moruka River, UMMZ 55834. Rupununi: north of Acaray Mts., west of New River, KU 69778-82. PARAGUAY: No specific locality, AMNH 19916-7, 19919, USNM 5837; “Alto Paraguay,” ZMB 26181. Boquerén: Loma Plata, KU 73432. Central: Asuncion, BMNH 94.3.14.168; Patino, FMNH 10799. Concepcion: Apa-Berg- land, ZSM 137/1933. La Cordillera: San Berbardino, NHMW 6106 (2), ZMB 26054, 26115. Olimpo: Puerto 14 de Mayo, Bahia Negra, BMNH 98.6.3.19. PERU: Loreto: “Caseirio Libertad, Rio Amazonas,” MJP 514; Orellana, USNM 127168-9, 127180; Pebas, CAS-SU 3159, 6376, 12622; Pucallpa, MJP 84, 86 (3), 90, 92, 93 (2), 95, 97 (2), 99; Rio Ucayali, FMNH 3562, UMMZ 48210; Roaboya, AMNH 43316, 43534; “Transval, near Pebas,” CAS-SU 3162. SURINAM: No specific locality, BMNH 70.3.10.68-9, NHMW 6097, 6109 (2), USNM 14618. Brokopondo: Brokopondo, RMNH 16692 (2). Coronie: Coronieweg, RMNH 16693. Marowijne: Camp 3, RMNH 16694. Suriname: HYLID FROGS, GENUS PHRYNOHYAS 19 Berg en Dal, ZMB 7267; Coropina Creek, RMNH 16695; Kwattaweg, RMNH 16696; “Lelydorp,” RMNH 16697 (2); Paramaraibo, AMNH 4009, ANSP 2169- 70, BMNH 1946.4.2.24-25, MCZ 7662, RMNH 16698 (9), USNM 13820-1, ZMB 25966 (2), 26076. TRINIDAD AND TOBAGO: Tobago: Milford Bay, MCZ 4087; “Whim,” BMNH 1926.1.19.1. Trinidad: No specific locality, FMNH 41683, MCZ 3517, 5971; “Churchill-Roosevelt Hwy,” AMNH 55712-15 (16), 55717-19; “Cumpia,” SMF 2646-47; Fyzabad, BMNH 1929.11.25.6; “Imperial College of Agriculture,” BMNH 1932.9.16.41; “Mt. Harris,” FMNH 49701; “Mt. Thabor,” CM 5257; “Oarino River,’ BMNH 1929.11.25.4; “Piarco,” KU 84714; “St. Augustine,’ AMNH 55716. VENEZUELA: “Los Canales,” USNM 128792. Anzoatequi: Carapa, USNM 80612. Aragua: Maracay, ZSM 304/1929 (20), 305/1929 (46), 61/1930 (22), 120/1930 (4), 292/1933 (6), 43/1934 (10), 105/1935 (11), 196/1965 (5). Distrito Federal: Caracas, SMF 2645; La Guaira, USNM 22545, 27797. Mo- nagas: Caicara, AMNH_ 16904-6, 16911-2, USNM 36377; Caripito, USNM 117098-9. Trujillo: Savana de Mendoza, UMMZ 57397. Zulia: La Fria, Pueblo Nuevo, UMMZ 55567-70; Oropé, FMNH 2604. LITERATURE CITED AHL, E. 1933. Uber einige neue Frésche aus Brasilien. Zool. Anz., 104:25-32. ANDERSSON, L. G. 1906. On batrachians from Bolivia, Argentina, and Peru collected by Erland Nordenskiold and Nils Holmgren 1901-1902 and 1904-1905. Arkiv Zool., 3 (12):1-19. 1945. Batrachians from east Ecuador collected 1937, 1938 by Wm. Clarke- MacIntyre and Rolf Blomberg. Ibid, 37A (2):1-88. BOKERMANN, W. C. A. 1966. Lista anotada das localidades tipo de anfibios Brasileiros. Sao Paulo, 183 pp. BOULENGER, G. A. 1882. Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British Museum, ed. 2, London, xvi+-503 pp. Broccui, P. 1879. Sur divers batrachiens anoures de |”’Amerique Centrale. Bull. Soc. Philom. Paris, ser. 7, 3:19-24. Cocuran, D. M., and C. J. Goin 1970. Frogs of Colombia. Bull. U. S. Natl. Mus., 288:xii+655 pp. Coreg, E.. D. 1862. Catalogue of the reptiles obtained during the explorations of the Parana, Paraguay, Vermejo, and Uruguay rivers, by Capt. Thos. J. Page, U.S.N., and of those procured by Lieut. N. Michler, U.S. Top. Eng., Commander of the expedition conducting the survey of the Atrato River. Proc. Acad. Nat. Sci. Philadelphia, 14:346-359. 1866. On the structures and distribution of the genera of arciferous Anura. Journ. Acad. Nat. Sci. Philadelphia, ser. 2, 6:67-112. 1877. Tenth contribution to the herpetology of tropical America. Proc. Amer. Philos. Soc., 17:85-98. Daupin, F. M. 1803. Histoire naturelle et particuliere des reptiles. Paris, 8: 439 pp. DUELLMAN, W. E. 1956. The frogs of the hylid genus Phrynohyas Fitzinger, 1843. Misc. Publ. Mus. Zool. Univ. Michigan, 96: 1-47. 20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY DUELLMAN, W. E. 1961. The amphibians and reptiles of Michoacan, México. Univ. Kansas Publ. Mus. Nat. Hist., 15:1-148. 1966. Taxonomic notes on some Mexican and Central American hylid frogs. Ibid, 17:263-379. 1968. The taxonomic status of some American hylid frogs. Herpetologica, 24:194-207. 1970. The hylid frogs of Middle America. Monog. Mus. Nat. Hist., Univ. Kansas, 1: xi+-753 pp. DumMe_ni_, A. M. C., and G. Brsron 1841. Erpétologie générale ou histoire naturelle compléte des reptiles. Paris, S: (92 pp. FITZINGER, L. 1843. Systema reptilium. Vienna, ix+106 pp. GoELDI, E. 1907. Description of Hyla resinifictrix Goeldi, a new Amazonian tree frog peculiar for its breeding habits. Proc. Zool. Soc. London (1907):135- 140. GUNTHER, A. L. C. 1859. Catalogue of the Batrachia Salientia in the collection of the British Museum. London, xvi+160 pp. HEMMING, F-. (editor ) 1958. Opinion 520. Suppression under the plenary powers of the specific name tibiatrix Laurenti, 1768, as published in the combination Hyla tibiatrix, and of the generic name Acrodytes Fitzinger, 1843, and in- terpretation of Rana venulosa Laurenti, 1768 (Class Amphibia). Opin. Declar. Intl. Comm. Zool. Nomencl., 19: 169-200. HENSEL, R. F. 1867. Beitrage zur Kenntniss der Wirbelthiere Siidbrasiliens. Wiegmann’s Arch. Naturg., 33:120-162. LAURENTI, J. N. 1768. Specimen medicum, exhibens synopsin Reptilium emendatum cum experimentis circa venena et antidota reptilium Austriacorum. Vienna, 214 pp. Luiz, B. 1968. New Brazilian forms of Hyla. Pearce-Sellards Ser., 10:1-18. McDiarmip, R. W. 1968. Populational variation in the frog genus Phrynohyas in Middle Ameri- ca. Contr. Sci. Los Angeles Co. Mus., 134:1-25. MrirmANDA-RIBEIRO, A. DE 1926. Notas para servirem ao estudio dos Gymnobatrachios (Anura) bra- sileiros. Arch. Mus. Nac. Rio de Janeiro, 27: 227 pp. PETERS, W. 1867. Uber Flederthiere und Amphibien. Monats. Akad. Wissen. Berlin (1867) :703-712. Rivero, J. A. 1961. Salientia of Venezuela. Bull. Mus. Comp. Zool., 126:1-207. 1964. The distribution of Venezuelan frogs VI. The llanos and the delta region. Caribbean Jour. Sci., 4:491-495. 1734. Locupletissimi rerum naturalium thesauri accurata descriptio, et in- conibus artificissimis expressio, per universam physices historiam. Amsterdam, 1: xxxiv-+-178 pp, 111 pls. SHANNON, F. A., and F. L. HUMPHREY 1957. A new species of Phrynohyas from Nayarit. Herpetologica, 13:15-18. - seiner meree, i SOE HYLID FROGS, GENUS PHRYNOHYAS 21 SmiTH, H. M., and E. H. TayLor 1948. An annotated checklist and key to the Amphibia of Mexico. Bull. U.S. Natl. Mus., 194:iv+-118 pp. Spx, J. B. von 1824. Animalia nova, sive species novae testudinum et ranarum, quas in itinere per Brasiliam, annis 1817-1820. Monaco, 53 pp. Tay or, E. H. 1944. The hylid genus Acrodytes with comments on the Mexican forms. Univ. Kansas Sci. Bull., 30:63-68. TaAyLor, E. H,, and H. M. SMirH 1945. Summary of the collection of amphibians made in México under the Walter Rathbone Bacon Traveling Scholarship. Proc. U.S. Natl. Mus., 95 :521-613. TRuEB, L., and W. E. DUELLMAN 1971. 6 ~ aes) 6 S i “a ! 23 ary ey JS oy g Fic. 2. Hand (a) and foot (b) of Hyla mykter (KU 137553, mature male; holotype). Scale is 5 mm in length. The subarticular tubercles are round or oval; none is bifid. Distinct supernumerary tubercles are present on the proximal segments of the fingers but are indistinct on the palms; a few small pustules are present along the ventrolateral edge of the forearm but are not ~— raised into a ridge. There is a fold on the wrist. The length of the foot is 20.2 mm (Fig. 2b). The toes are moderately long, 1< 2< | 5 < 3 < 4, and about four-fifths webbed. The first, fifth, and PuateE 2. Hyla mykter: A. Guerrero, 40.1 mm snout-vent length (KU 137553, mature male; holotype). B. Guerrero, 50.9 mm (KU 137552, mature female; paratype). Hyla bogertae: C. Oaxaca, 43.3 mm (LACM 44403, mature female; paratype). D. Oaxaca, 49.0 mm (LACM 44402, mature female, para- ) type). = PLATE 2 NEW HYLA FROM GUERRERO, MEXICO Lt outer sides of the second and third toes are webbed to the base of the disc; the inner side of the second toe is webbed to the base of the penultimate phalanx, and the fourth toe and inner side of the third are webbed to distal end of antepenultimate phalanx. The inner metatarsal tubercle is distinct and elliptical; there is no outer metatarsal tubercle. The tarsal fold is short and faint. The subarticular tubercles are round or oval; none is bifid. Indistinct supernumerary tubercles are present on the soles and proximal seg- ments of the toes. The discs are moderately large, that of the fourth toe is 2.0 mm wide. The tibia length is 20.4 mm; the tibiotarsal articulation extends to the middle of the eye, and the heels overlap 5.7 mm when the legs are flexed and held at right angles to the body. The cloacal sheath is moderately long, grooved medially, directed posteroventrally, and opens just below the midlevel of the thighs; there is a transverse dermal fold above the sheath and large, distinct pustules on the thigh adjacent to its aperture. The skin is only slightly thickened; that of the dorsum and undersur- faces of arms, chest, and lower legs is smooth; that of the throat, abdomen, and undersurfaces of the thighs is granular. There is no enlarged axillary membrane. The dentigerous processes of the prevomers are each the same size as a choana, oval, and oriented transversely between the choanae, nearer one another than to choanae; there are 4-5 prevomerine teeth. The choanae are oval, each 1.0 mm long. Vocal slits are absent. The tongue is cordiform, slightly notched and one-sixth free posteriorly. As determined from radiographs, quadratojugals are absent. Color in life: The dorsum of the head and body is pale yellow- ish green, and the limbs are pale yellowish brown, all covered with black reticulations and spots; the flanks and rear of the legs and lower arms are mottled with black and pale yellow (Plate 2). The canthus and supratympanic fold are black-edged and the tympanum is brownish green. The iris is golden orange with black reticulations. The knee and ventrolateral edges of the forearms and legs are black-edged. The venter is mottled dusky brown, tend- ing toward deep purplish gray or chocolate brown around the perimeter, overlaid with some brassy pigment. The nuptial spines are dark brown. Color in alcohol: The body is dull grayish brown above with gray reticulations; gray mottling is present on the flanks. The gray or brown edging of venter and the yellow and green ground color of the dorsum are not evident. The ventral surfaces are pale yellow- ish cream with gray mottling. The iris is bluish, and the webbing is pale gray. The nuptial spines are brown. 12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Variation —The snout of the adult females (50.9 mm snout-vent length) as well as that of an immature male (30.8 mm) are pointed and possess a rostral keel. The female has only vestigial webbing between all fingers. The female has noticeably shorter legs and feet, proportionate to body length, than have either of the males. The female is also more robust than either male but has a slightly narrower head. The quadratojugal is lacking in all three individ- uals. Morphological variation in some mensural characters is given in table 1. Apparently only moderate color variation exists in this species (Plate 2). The adult female was pale gray-brown with a little dark flecking above, with a gray tympanum. There was a green wash along the upper rear edge of the thigh and in the groin, and some brassy overlay on the venter including throat; there were a few yellow spots or blotches along the flank and front edge of arm. The immature male was a deep olive-green dorsally mottled with black flecks and reticulations. Little metachrosis ‘was noticed, except that most individuals were dark greenish or dark yellowish tan if sluggish when first caught, and only later lightened to the colors described above. Habitat and Habits—These frogs were taken in a variety of locations, all in cool, moist, oak-pine or (at the lower elevations ) bamboo-tree fern cloud forests near Cerro Teotepec, at elevations between 1985 and 2520 meters. All specimens of Hyla mykter were taken in December. The female (which has large pigmented eggs) was found early in the morning in a bromeliad 2 m above ground in a pine tree, at the edge of a large clearing at Asoleadero (2520 meters). The vegetation and climate at this locality have been described by Musser (1964) and Adler (1965). The day was warm (21° C maximally) and dry, but temperatures fell to near, or below, freezing at night; there was considerable frost on the logs and boulders in the clearing at daybreak. The two males were taken along lush, cascading mountain streams at considerably lower elevations. The habitat at the type locality is dominated by tree ferns, bamboo, pine and oak, and dissected by deep ravines containing plant-shrouded streams (this is also the type locality for H. trux). The holotype was collected at 8:30 PM, fully exposed on vegetation directly over a stream; whereas, the smaller male was found at night in a bromeliad which had fallen along a wide, open stream. The climate at these localities was considerably warmer than at Asoleadero. None of the males was heard to call. Other frogs taken at the same time along these streams include Hyla trux, H. pentheter and Eleutherodactylus saltator. NEW HYLA FROM GUERRERO, MEXICO 13 Remarks.—The relationships of Hyla mykter appear to lie with presumably the most advanced members of the Hyla bistincta species group. These species (namely bogertae, crassa, pachy- derma, robertsorum and siopela) possess thick glandular skin, blunt heads, large webbed feet, and nuptial excrescences, and lack axil- lary membranes and vocal slits. We have not seen specimens of pachyderma or crassa, and have relied on Duellman (1970) for accounts of their characteristics. Hyla mykter differs from the latter two species and bogertae in having a weak tarsal fold, the outer fingers slightly webbed, a small spine-shaped quadratojugal, a rostral keel and a less robust body. It is distinguished from crassa by having feet four-fifths webbed and a well-defined tympanum. Hyla mykter differs from pachyderma in possessing smaller nuptial spines, a weak thoracic fold, and a well-defined tympanum in males (indistinct only in male pachyderma). The species is distinguished from bogertae*® by its rounded canthus and a gray-mottled belly. Hyla mykter differs from robertsorum in having a small quadrato- jugal, a more pointed snout with a better-developed rostral keel, more extensively webbed outer fingers, and, in males, a long cloacal sheath and less robust body shape. In two or three females of the robertsorum examined, the snout is slightly pointed, and in one there is a faint trace of a rostral keel; a few individuals of both sexes have a trace of webbing between the outer fingers, but in none of them is it as well developed as it is in siopela and mykter. Hyla mykter and siopela seem to be closely related because they share more characteristics than either does with other members of the subgroup. Both have a trace of webbing between the outer fingers and have a rostral keel (rarely absent in siopela), although the snout is more truncate (in both dorsal and side views) and the canthus more angular in siopela. When viewed from above, the nostrils of siopela are more anterior than are those of mykter, which are located dorsal to the margin of the upper lip. Nuptial excrescences are restricted to the first and second fingers in siopela (2 males examined) but at least some spines are found on all fingers in mykter (1 mature male); however, this difference may not hold in larger samples. The toes are slightly more fully webbed in mykter, although the difference is not as great as a comparison 3 The availability of specimens and colored.slides of the recently described Hyla bogertae has made it possible to illustrate certain features of this form not given in the type description. In an effort to maintain the uniformity of treatment used by Duellman (1970), we have included color illustrations of adults (Plate 2) and drawings of the hands and feet (Fig. 3) because of the usefulness of these features in hylid systematics. 14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY nas b Fic. 3. Hand (a) and foot (b) of Hyla bogertae (LACM 44401, mature male; paratype). Scale is 5 mm in length. of our drawing (Fig. 2) with that of siopela (Duellman, 1970, fig. 235-C) suggests; the latter figure shows an individual with less webbing than typifies most of the individuals we have seen of that species. The cloacal sheath of the single adult male mykter is longer than that of either of the two male siopela. Hyla mykter differs further in having a weak, instead of a well-developed, tarsal fold and in having proportionately longer legs. Hyla siopela and mykter are the only two members of this subgroup which typically have rostral keels and traces of webbing between the two outermost fingers. They also are the most thin-skinned and least robust of the six species. Of the remaining four species, robertsorum most closely NEW HYLA FROM GUERRERO, MEXICO 15 approaches siopela and mykter in these characteristics and _pre- sumably is closely related to them. The members of this subgroup of the Hyla bistincta species group typically are found in moist oak-pine forests at intermediate to high elevations in southern Mexico and can be subdivided further into two series. The species in the first are more robust, have more glandular skin and include bogertae (Oaxaca, 2650 m), crassa (Oaxaca, 2300 m), and pachy- derma (Veracruz, 1600 m); whereas in the second series the species are less robust, have thinner skin and include mykter (Guerrero, 1985-2520 m), robertsorum (Puebla and Hidalgo, 2250-3100 m) and siopela ( Veracruz, 2500-2650 m ). Several frogs referred to the Hyla bistincta group have been recorded from Guerrero. The types and only known specimens of Hyla chryses were collected between 2540 and 2600 meters near Asoleadero. (This species is readily distinguished from mykter by the well-developed axillary membrane, the more slender and thinner-skinned body, and the absence of nuptial spines in chryses. ) The three metamorphosing frogs from nearby Omiltemi (UIMNH 38023-25), referred to bistincta (by Duellman, 1964, and many subsequent authors including Adler, 1965), have been reexamined. All have tiny tailbuds (specimens collected in August, 1940), a mottled pattern on the dorsum, and long, virtually webless fingers. Newly metamorphosed individuals of chryses and mykter are not available; thus, these three specimens cannot be referred to any of the species known from Guerrero, although they seem to be allied to the Hyla bistincta group. They might even be referrable to an as-yet-unnamed form in this species group based on an examination of eight frogs (IPN CB 149-156) collected in April 1963 at Agua Fria, a logging camp about 10 km E of Cerro Teo- tepec; these are the same specimens referred to bistincta by Duell- man (1970, p. 698) as from “22 kilometers southwest of Yextla.” All are in a poor state of preservation and exact measurements cannot be made; the snout-vent lengths of the three males are about 38, 39 and 39 mm; those of females range from 34 to 39 mm. They lack the long cloacal sheath, vocal slits, and flank pattern charac- teristic of bistincta and pentheter, and also lack the axillary mem- branes found in chryses. Their fingers are long, and the males have nuptial excresences on the prepollex and smaller patches of spines on all other fingers; however, they have no webbing between the fingers, and lack the rostral keel and mottled venter characteristic of mykter. They were found by day in and along the edge of a pond at 2600 meters; their identity must await the acquisition of fresh material. 16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY One other frog from Guerrero (UMMZ 125376, an adult female from near Asoleadero ) also has been referred to bistincta (by Adler, 1965, and subsequent authors). A reexamination of this specimen shows this determination to be incorrect; the specimen lacks the bold markings on the flanks characteristic of bistincta, has a dark venter, is considerably smaller in size, and, according to radio- graphs, lacks a prepollical spine. It differs from mykter in the character of the prepollical spine, in having longer legs, and in several features of coloration; it is also dissimilar to the unnamed Agua Fria specimens in being larger, possessing reticulations on the flanks, and lacking a prepollical spine. Thus, apparently none of the published references of Hyla bis- tincta in Guerrero is, in fact, based on specimens clearly referrable to that species. Together with David H. Snyder, we collected four adult bistincta (KU 140420-23) in a moist cove along a dry, rocky canyon 4.5 km (by road) E of El Limén (about 6 km SW of Chilapa), Guerrero, at 1525 m elevation. The only other member of this species group known from Guerrero is H. pentheter, pre- viously known only from Oaxaca; we obtained three individuals (KU 140424-26) at an elevation of 2000 m in the vicinity of Cerro Teotepec. Etymology.—The specific name is from the Greek puxrjp, mean- ing nose, and is given in allusion to this frog’s distinctive snout. Distribution—Known only from localities in the vicinity of Cerro Teotepec in the Sierra Madre del Sur of Guerrero, at eleva- tions between 1985 and 2520 m. SUMMARY Two new frogs of the family Hylidae are described from the cloud forests of the Sierra Madre del Sur of Guerrero. They are members of two different species groups which have representatives primarily in the mountains of eastern and southern México. Both are illustrated (hands and feet; juveniles and adults in color). Hyla trux, a member of the H. taeniopus species group, is a large species (males to 81.0 mm snout-vent length) with a mottled brown dorsum, vocal slits and blue webbing in adult males, ab- sence of an axillary membrane, presence of a tarsal fold, nuptial spines on all fingers of mature males, and a hypertrophied web between toes one and two (in mature males). Specimens were taken at elevations between 1760 and 2120 m at several localities, all within 30 km of Cerro Teotepec. This species seems to be closely related to H. taeniopus Ginther of the states of Puebla, Veracruz, and Hidalgo. NEW HYLA FROM GUERRERO, MEXICO 17 Hyla mykter, a member of the H. bistincta species group, is a medium-sized species (males to 40.1 mm; females to 50.9 mm) with a yellowish olive or pale gray-brown dorsum, absence of vocal slits and axillary membranes, no quadratojugal, nuptial spines on all fingers of mature males, long fingers with vestigial webbing, and a weak rostral keel. Specimens were taken at elevations be- tween 1985 and 2520 m at several localities, all within 35 km of Cerro Teotepec. This species seems to be most closely related to H. siopela Duellman of the state of Veracruz. Hyla bogertae Straughan & Wright, a member of the H. bis- tincta species group recently described from Oaxaca, is illustrated (hand and foot; adults in color ). RESUMEN Dos nuevas ranas de la familia Hylidae, procedentes de las selvas nubladas de la Sierra Madre del Sur (México, Edo. Guer- rero), son descritas en el presente trabajo. Estas especies per- tenecen a dos grupos distintos de especies incluidas en el genero Hyla y con representantes el las Areas montafosas del sur y extremo este de México. Ambas especies son ilustradas (manos y pies; juveniles y adultos en color). Hyla trux es un miembro del grupo taeniopus; se trata de una especie grande, machos con 81 mm de longitud naso-ventral y dorso irregularmente manchado de marrén; fisuras vocales y membrana interdigital de color azul; membrana axilar ausente; pliegue tarsal presente. Espinas nupciales presentes en todos los dedos de machos sexualmente maduros. Membrana interdigital entre los dedos uno y dos, hipertrofiada. Los ejemplares fueron colectados en varias localidades ubicadas entre 1760 y 2120 m altura, todas ellas a unos 30 kms del Cerro Teotepec. Esta especie parece estar relacionada a H. taeniopus Giinther de los estados de Puebla, Veracruz e Hi- dalgo. Hyla mykter es un integrante del grupo bistincta. Es una es- pecie de talla media (longitud maxima de los machos, 40.1 mm; hembras 50.9). Coloracioén dorsal amarillo-olivaceo o marron- grisaceo claro. Fisuras vocales y membranas axilares ausentes; quadratojugales ausentes. Espinas nupciales presentes en todos los dedos de los machos maduros; dedos largos con membranas inter- digitales vestigiales. Cresta rostral débil. Los ejemplares fueron colectados en varias localidades ubicadas entre los 1985 y 2520 m altura, a unos 35 kms del Cerro Teotepec. Esta especie parece relacionada a H. siopela Duellman del Estado de Veracruz. 18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Hyla bogertae Straughan y Wright, miembro del grupo bis- tincta, recientemente descrita de Oaxaca, es ilustrada (mano y pié; adulto en color). ACKNOWLEDGMENTS AND METHODS We thank David H. Snyder (University of Notre Dame) for field assistance and for comments on the manuscript. William E. Duellman (University of Kansas) kindly provided relevant por- tions of his monograph of Middle American hylid frogs in advance of publication and loaned a large collection of comparative material (designated KU). Other museum abbreviations are: University of Michigan Museum of Zoology (UMMZ), University of Illinois Mu- seum of Natural History (UIMNH), Los Angeles County Museum of Natural History (LACM), Instituto Politechnico Nacional, Cién- cias Bioldgicas, México (IPN CB), United States National Museum of Natural History (USNM), Field Museum of Natural History (FMNH), American Museum of Natural History (AMNH), and British Museum (Natural History) (BMNH). Certain individuals loaned specimens from several of these institutions, namely Joseph T. Collins, Ticul Alvarez S., John W. Wright, Mrs. Dorothy M. Smith, and Charles F. Walker. James P. Collins and Arnold G. Kluge (both UMMZ) assisted in making x-rays. We thank Alfredo Sartorius Z. and his colleagues of Puerto del Gallo, Guerrero, for their hospitality, and Juan Luis Cifuentes L. (formerly of the Secre- taria de Industria y Comercio, México) who provided our collecting permit. Carlos Machado-Allison ( University of Notre Dame) trans- lated the summary into Spanish. Field work involved in this study was supported incidental to research on salamander reproductive biology assisted by a grant from the Society of the Sigma Xi. Methods of measurement of morphological features and of geo- graphic distances are as indicated in Adler (1965:16). SPECIMENS EXAMINED Hyla altipotens Duellman. Oaxaca: 33-37 km N San Gabriel Mixtepec, KU 101002-06, 101008; 104341-42 (skeletons ). Hyla bogertae Straughan & Wright. Oaxaca: 1.6 km S La Cofradia, LACM 44401-03. Hyla chaneque Duellman. Oaxaca: 4.2-16 km S Vista Hermosa, KU 86961-66, 86968, 86971; 84907-08 (skeletons); 8 km S Yetla, KU 86967. Hyla charadricola Duellman. Puebla: 14.4 km W Huachinango, KU 53813-15, 58415-22; 55624, 59813 (skeletons). Hyla chryses Adler. Guerrero: 38 km airline WNW of Chilpancingo, UMMZ 125372-73, 125375; KU 106306. Hyla mykter, new species. Guerrero: the type series, as listed above. NEW HYLA FROM GUERRERO, MEXICO 19 Hyla robertsorum Taylor. Hidalgo: El Chico Parque Nacional KU 71265- 95; 59824-25, 71757 (skeletons ). Hyla siopela Duellman. Veracruz: W slope Cofre de Perote, KU 100976- 80, 100982-85, 105628-29; 117428-29 (skeletons); Cofre de Perote, KU 117430 (skeleton ). Hyla taeniopus Ginther. Hidalgo: 3 km W Xochicoatlan, KU 53820-23, 53825-26; 55623, 59826 (skeletons); 2.5-4 km SW Tianguistengo, KU 53827- 30. Puebla: 3.7 km NNE Tezuitl4n, KU 53832-37, 57827; 55602-03 (skele- tons ). Hyla trux, new species. Guerrero: the type series, as listed above. Hyla, species uncertain. Guerrero: Omiltemi, UIMNH 38023-25; Agua Fria, 10 km E Cerro Teotepec, IPN CB 149-56; between Puerto Chico and Asoleadero, UMMZ 125376. LITERATURE CITED ADLER, K. 1965. Three new frogs of the genus Hyla from the Sierra Madre del Sur of México. Occas. Papers Mus. Zool. Univ. Michigan, 642:1-18, pl. 1. DuELLMAN, W. E. 1964. A review of the frogs of the Hyla bistincta group. Univ. Kansas Publ. Mus. Nat. Hist., 15:469-491. 1965. Frogs of the Hyla taeniopus group. Copeia 1965 (2):159-168. 1970. The hylid frogs of Middle America. Monogr. Mus. Nat. Hist. Univ. Kansas, 1:xii+-1-753, pl. 1-72. Musser, G. G. 1964. Notes on geographic distribution, habitat, and taxonomy of some Mexican mammals. Occas. Papers Mus. Zool. Univ. Michigan, 636: 1-22. UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly pub- lished in the above series are now published as Occasional Papers, Museum of Natural History. The Miscellaneous Pub- lications, Museum of Natural History, began with number 1 in 1946. Longer research papers are published in that series. Monographs of the Museum of Natural History were initiated in 1970. Institutional libraries interested in exchanging publications may obtain the Occasional Papers and Miscellaneous Publica- tions by addressing the Exchange Librarian, University of Kan- sas Library, Lawrence, Kansas 66044. Individuals may pur- chase separate numbers of all series. Prices may be obtained upon request addressed to Publications Secretary, Museum of Natural History, University of Kansas, Lawrence, Kansas 66044. Boe hilo CALIFORNIA .& ACADEMY OF SCIENCES JUL - 6 1972 LIBRARY OCCASIONAL PAPERS of the MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas NUMBER 8, PAGES 1-20 JUNE 28, 1972 VARIATION IN THE CENTRAL AMERICAN IGUANID LIZARD, ANOLIS CUPREUS, WITH THE DESCRIPTION OF A NEW SUBSPECIES By Henry S. Fitcu,' ANTHONY A. ECHELLE AND ALICE F. ECHELLE™ In the course of ecological studies of Costa Rican lizards con- ducted by the senior author from October 1967 to March 1970, Anolis cupreus Hallowell was found in larger numbers and at more localities than any other species of lizard. Approximately 3450 individuals of this small Pacific coast anole were processed and re- corded in the field in a mark-and-recapture program. Notable differences were observed between individuals, geographic popula- tions, and the sexes. The recent resurrection of Anolis hoffmanni Peters (a species long considered a synonym of A. cupreus) by Williams and Smith (1966) together with variation observed among our samples, suggested that more than one taxon was represented in our accumulated field records. In order to determine the num- ber of taxa involved, series of specimens were collected throughout the range of Anolis cupreus in Costa Rica and near the northern end of the range of the species in Guatemala. Additional preserved specimens from Costa Rica and western Nicaragua were examined in the University of Kansas Museum of Natural History (KU). Localities represented by our collections are shown in figure 1. Populations of A. cupreus at or near all Costa Rican localities shown in figure 1 were studied in all seasons, thereby providing informa- tion on population structure and other aspects of natural history. 2 University of Kansas Natural History Reservation, Lawrence, Kansas 66044 * University of Oklahoma Biological Station, Willis, Oklahoma 73462. 2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY The large series of specimens from those localities permitted study of individual variation and sexual differences. Knowledge of such factors enhances the validity of geographic comparisons. Fic. 1. Map of Central America showing probable range of Anolis cupreus (shaded) based on specimens collected by the authors and examined in life (dots), museum specimens, and published records and statements by Taylor (1956), Stuart (1955), and Peters and Donoso-Barros (1970). Inset below is a map of Costa Rica showing probable ranges of three subspecies of Anolis cupreus: A. A. c. cupreus; B. A. c. hoffmanni; C. A. c. spilomelas. The locali- ties from which series were examined are: 1. Playa del Coco, 2. Sardinal, 3. Tilaran, 4. Rio Higueron, 5. Rio Congo, La Irma, 6. Rio Naranjo, 7. Boca de Barranca, 8. Turricares, 9. San José, 10. Cartago, 11. Quepos. VARIATION IN ANOLIS CUPREUS 3 MATERIALS AND METHODS Our field data were gathered on six trips to Costa Rica (2 February to 28 March 1965, 13 October 1967 to 16 July 1968, 10 August to 8 September 1968, 18 January to 13 March and 8 August to 8 September 1969, 29 January to 14 March 1970) and one trip to Guatemala, 15 to 24 February 1971, by H. S. Fitch. A. F. Echelle participated in the 1967-68 field trip, and all three authors made the 1970 and 1971 trips together. Population samples, mostly based on live anoles, examined but not killed or removed, were obtained from Costa Rica as follows: on 13 days in eight different months at Boca de Barranca; on two days in one month at Cartago; on 26 days in nine months at La Irma; on 17 days in nine months at Quepos; on three days in three months at Rio Naranjo; on 75 days in 19 months at San José; on 15 days in nine months at Sardinal, and on three days in two months at Turrucares. The samples from Guatemala were obtained on 17, 18 and 19 February, 1971. The small samples that we used from Tilaran and Taboga are in the Museum of Natural History at the University of Kansas. Many variable characters were examined. Each character was found to have limitations, and each was used on relatively small but representative series of specimens. The conventional characters used in the classification of anoles are those that can be seen in preserved specimens. However, some characters that we found to be equally useful can be seen only in living or freshly killed anoles. Color and pattern, especially, were found to be useful for show- ing differences between geographic populations, between age groups, and between the sexes. Because these differences are ob- scured in preserved specimens, we recorded them only from live or freshly killed anoles. Snout-vent length was measured to the nearest millimeter, and most of the measurements were taken from live lizards or those that were freshly killed. Live weights of most of the anoles processed in 1970 and 1971 were recorded in the field with Oskar Ludi spring scales, to the nearest hundredth of a gram. Length of tail was also recorded for each anole, with regenerated parts noted separately. Relatively large amounts of data were therefore available for size and tail length, and both were found to show significant differences between populations and between the sexes. For comparison of size between populations, only adults were used. Females of 35 mm snout-vent length often contained uterine eggs or enlarged follicles while those that were smaller rarely contained either, hence females of 35 mm in length or larger were considered adults. Males of 40 mm or longer were considered 4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY adults; those that were smaller were found to be still growing rapidly. Other characters that were used were based upon preserved specimens. Number of scale rows around the body varied signifi- cantly between geographic populations and between the sexes, but recording them was tedious because of the small granular scales, and was inexact because of irregularities in the rows. Therefore the counts were made on relatively small series of museum speci- mens. VARIATION WITHIN AND BETWEEN POPULATIONS Size.—In A. cupreus, as in most iguanids, the male attains a larger size than the female (Figs. 2 and 3). In every series ex- amined, the males were somewhat larger in maximum size (Fig. 3). There is little or no weight difference between males and females of comparable size. Because of continued growth after attaining sexual maturity, the age structure and date of collection influence the comparisons. Most samples are similar in adult size (Fig. 3), but both sexes from Quepos and males from Escuintla are significantly larger. Respectively, these are the southern- and northernmost populations of A. cupreus sampled. Sexual difference in size is least in samples from San José and Cartago. e = Males ° =Females e e fe) 1°) e e e ° e WEIGHT (gm) Z2O5 +24 28 32 36 40 44 48 52 LENGTH (mm) Fic. 2. The relationship between weight and snout-vent length in Anolis cupreus collected at San José, San José Province, Costa Rica, February 1970. VARIATION IN ANOLIS CUPREUS 5 262 4 : 5| 202) . 8 7 I Quepos LENGTH (mm SVL) 39 Escuintia | a =e i an ,Cartago Sardinal Naranjo Boca La de Irma Barranca Fic. 3. Snout-vent lengths in adult Anolis cupreus, 35 mm or more in females (lower series of figures), 40 mm or more in males (upper series) from eight localities in Guatemala and Costa Rica. Dots indicate maximum lengths; horizontal lines represent means; bars show two standard errors on each side of the means; numbers are sample sizes. Tail length—Intact tails are proportionately longer in adults than in young. At Sardinal, Guanacaste Province, Costa Rica, the ratio of tail length to snout-vent length in 203 adults having snout- vent lengths of 40 mm or more is 1.60-2.00 (mean 1.77). Twenty hatchlings (probably in their first month of life) from the same locality have snout-vent lengths of 20 mm or less; the ratio of tail length to snout-vent length is 1.42-1.84 (mean 1.57). Adult males have relatively longer tails than females. Sexual dimorphism in tail length is less, if it exists at all, in hatchlings. In the adult series from Sardinal, the average ratio of tail length to snout-vent length is 1.80 in 112 males and 1.74 in 91 females. The series from the northern (Escuintla) and southern (Quepos) ends of the range have relatively long tails; whereas those from San José on the Meseta Central of Costa Rica are somewhat intermediate, and those from the lowlands of western Costa Rica (Sardinal, La Irma, Boca de Barranca) have relatively short tails (Table 1). The high ratio in females from Escuintla, slightly exceeding the male ratio, prob- ably is an artifact of the small sample size. Hind Limb.—No geographic variation could be demonstrated in our samples. In 77 per cent of 159 specimens, the fourth toe of the adpressed hind limb lies at a level between the anterior and OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 6 (Ze) (€@) (ZS) (€@) a (<3) (FG) (8) (8) be (IT) (03) uvoW ovjjoue’y [esi €€I-ScI SOI-r6 G6IT-VOL €ZI-L6 aes CcI-SIT OdI-66 O€T-8IT Col inl : OV I-86 osuey L LOI T cor 9OIT 9OIT it S'siI VCII Geel OSGI rae 0'0¢I 0'OGT URN (F) (8) (€T) (IT) = (9) (OT) (3) (3) ok (T) (OT) sopeule OSI-LGI SII-OOLT OGI-80I 844 mm, 2 >50 mm. Tibia Length (TL).—Measurement of length of tibia to nearest 0.1 mm; used as a ratio of snout-vent length. Head Width (HW).—Measurement of greatest width to nearest 0.1 mm; used as a ratio of snout-vent length. Webbing on Hand (WEB)*.—The extent of webbing on the inner edge of the fourth finger coded with respect to the point of termination of the webbing with respect to the distal subarticular tubercle: 1) proximal to tubercle, 2) at tubercle, 3) distal to tubercle. Subarticular Tubercles (TUB)*.—The structure of the distal subarticular tubercles on the third and fourth fingers was coded: 1) conical, single, 2) conical, bifid, 3) flat, bifid. Ulnar Fold (ULN)*.—A dermal fold on the ventrolateral edge of the forearm was coded: 1) absent, 2) present. Tarsal Fold (TAR)*.—A dermal fold on the ventrolateral sur- face of the foot was coded: 1) absent, 2) present. Calcar (CAL)*.—A triangular dermal appendage projecting posteriorly from the upper edge of the heel was coded: 1) absent, 2) small, 3) large. Snout Profile (SNO)*.—The shape of the snout in lateral view coded: 1) round, 2) truncate, 3) anteriorly inclined. Tympanum (TYM)*.—The upper edge of the tympanum is covered by a supratympanic fold, but the tympanic ring below the fold coded: 1) distinct, 2) covered with undifferentiated skin. Mental Gland (MEN).—A round glandular structure at the apex of the jaw was coded: 1) absent, 2) present. 4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Dorsolateral Stripe (DST)*.—A pale stripe extending from the edge of the eyelid to a point on the side of the body was coded: 1) absent, 2) present. Tarsal Stripe (TST )*.—Longitudinal stripes on the outer edge of the foot were coded: 1) absent, 2) white, 3) white above and brown below. Anal Stripe (AST) *.—Transverse stripes above the anus were coded: 1) absent, 2) white, 3) white above and brown below. Tooth Rows (TOO)*.—The number of upper/lower rows of teeth in tadpoles was coded: 1) 4/5, 2) 5/7, 3) 6/9, 4) 7-8/10. Labial Papillae (LAB)* —Labial papillae are continuous around the mouth in the tadpoles; the number of rows of papillae was coded: 1) one, 2) two. Serrations on Beak (SER)*.—The serrations on the cutting edge of the upper beak in tadpoles were coded: 1) none, 2) small, 3) medium, 4) large. Shape of Body (SHA)*.—The shape of the body of tadpoles in dorsal view was coded: 1) narrowly ovoid, 2) broadly ovoid, 3) narrow anteriorly and widening abruptly just anterior to eyes. Caudal Musculature (CAU)*.—The proportionate depth of the caudal musculature was coded: 1) <65%, 2) >65%. RESULTS For ease in discussion the nomenclature adopted in the follow- ing section on taxonomy is used throughout. The results of the discriminant analysis are discussed first, followed by the Phenogram Program and finally the mating calls. Multiple Discriminant Analysis Within-group Variation.—The amount of variation in snout-vent length is approximately the same in all species except H. bogotensis and H. platydactyla in which it is somewhat greater (Table 1). However, the amount of variation in the ratios of tibia length and head width to snout-vent length is no greater in these two species than in the others. Variation of all structural features, except presence or absence of ulnar and tarsal folds, occurs within groups; likewise, color pattern characters show within-group variation (Table 2). Tarsal and anal stripes are either present or absent within a group; if present, there may be one or two stripes (Hyla phyllognatha). The presence of a mental gland is the most variable structural feature, displaying variation in five of the 12 groups. The amount of webbing is the next most variable character; variation occurs in four groups. The variation within groups is not highly correlated with respect to the following characters—size, proportions, subarticular tubercles, snout shape, tympanum, and mental gland. Absolute positive cor- FROGS OF THE HYLA BOGOTENSIS GROUP 5 TaBLE 1.—Measurements and proportions of species in the Hyla bogotensis group. (Mean and one standard deviation given below observed range. ) Snout-vent Tibia Length/ Head Width/ Species Sex N Length SVL SVL H. alytolylax @ 13 32.1-37.0 0.443—0.525 0.298—0.331 34.85+1.51 0.474+0.024 0.316+0.010 qn 15 37.2—43.9 0.462—0.532 0.296—0.331 40.38 +2.23 0.496+0.021 0.310+0.010 H. bogotensis 3 7 29.4—43.0 0.491—0.514 0.329—0.365 35.20+5.58 0.502+0.010 0.345+0.012 °) 4 29.4—48.1 0.449—0.506 0.313—0.380 41.43 0.483 0.347 H. colymba 3 6 31.9-37.0 0.452—0.514 0:302—0.327 34.65+2.29 0.481+0.023 0.318+0.010 Q 3 31.4—39.1 0.486—0.493 0.310—0.340 36.23 0.490 0.325 H. denticulenta 64 il 44,2 0.493 0.303 Q 1 nya 0.513 0.308 H. phyllognatha 64 13 35 —34.0 0.443—0.524 0.301—0.348 32.96+0.69 0.488+0.028 0.319+0.013 g 3 30.0—39:3 0.488—0.506 0.326—0.331 36.87 0.495 0.329 H. platydactyla 3 on 26.5—39.4 0.464—0.540 0.323—0.383 34.36+3.05 0.488+0.016 0.347 +0.014 Q UI 30.6—42.3 0.457—0.506 03333—0356 38.41+3.64 0.491+0.016 0.342+0.009 relation exists between presence or absence of ulnar and tarsal folds, between tarsal folds and tarsal stripes, and between tarsal and anal stripes. Webbing is the most extensive and calcars are the largest in the species, H. denticulenta. Sexual Dimorphism.—No sexual dimorphism is evident in pro- portions, amount of webbing, nature of folds or calcars, snout shape, or nature of tarsal and anal stripes. Minor sexual dimorphism occurs in the nature of the subarticular tubercles in H. platydactyla, in the tympanum in H. bogotensis, and in the dorsolateral stripe in H. colymba (Table 2). Snout-vent length is greater in females than in males; snout-vent lengths of 77 males is 26.5-44.2 (x—34.43) mm, and of 37 females is 29.4-52.2 (x=39.63) mm. As noted above, the nature of the mental gland is highly variable. The gland is present in 44 of 77 (57.1%) males and in nine of 37 (24.3%) females. The structure and function of the mental gland are unknown; its development may be seasonal and associated with reproductive activity. Thus, its usefulness as a taxonomic character within the Hyla bogotensis group is questionable. Interpopulational Variation—The combination of the sexes in OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 6 co ee 1601 =! Goi -160T ~*~ —ae Swreo ys rang ae I I 1 1 1 Z I I I Z [i Bie* eG ee i, poem = toro. Evie ~~ soe Se ee BS weet Bick ZOU LeRT “Soert = ae ro hs ecole & ee I I 1 oT oT Z I i I SI I Le =~ mhgowphianjd “yy eg Terr 2 Dee HeOF So (Sa a to \e ee LLS'0 geae aeeeg a Geese met oeue a LOOK) beac ec an me tee L99'T eG eZ I oI I 1 1 Z g I Ciao f é eons 0s ate og ew es Let: Os) = = he as he LLBO ccm nes. a OT LO Ten COC 1 yy ee WP Ga LLO'T oS eZ I Z I SI 1 Z Z I ST 8=6&l—P Dywusoyhyd -H € € I I I g € Z Z € € I é € g I I I € € rd Z € € i 2 — pyuaynorquap “1 oj mete 0 Se, eet. Lee ee ee RO a ce nae TOs food. Payee ee ty a, Pee sop eee eS eS ts Z ra oI I I I Z Z I Z € é g Z Z Z I I I Z g I Z 9 ? pquihjoo *H I I I I Z Z I I I Z Z 4 é Se Oe or geo seso ~~ 7 mae = 5 ra nar a eee erut alba — ef) Se PS = I I i oI 1 Z I I I Z Z L 9 —-sysuajoS0q ‘Hy ST hee se aig go es ere emo: =. SSGei0 ss sf’, ar ce wen or MOOV aren pine. BS L98°S Z Z Z I I Z 1 Z rd I Ci. Sly a6 “ey 2 eee aie a, iad i 7 =e “= = 561g al ee oe oe aig Te Sr airs le! GEC Z Z Z Z I I I cd Z I oc = 81 ff ax oIio 7 IsV. ISL IS@ NAN WAL ONS TIVO 4@VL NIN @nl dam NS _ sopads (‘oBue1 pearesqo MOTE UeAIS UOT}eIAep piepuejs OUD pu UKOUT ‘BUIPOD puke SUOTJIADIGqe IOJ sIoJOvIeYD JO UOTUYecT 99¢ ) ‘dnois sisuajos0q vfyT 94} Ul UOTye1O[OO pue sloJORIeYO [eIN}oNI}]s UI UOTFeUeA—Z aATAV] FROGS OF THE HYLA BOGOTENSIS GROUP if the geographic samples provides an assessment of populational dif- ferences for taxonomic distinction. Each of the 14 characters, in combination with others, serves to distinguish one population from another, but, with the exception of snout-vent length in H. denticulenta, no one character distinguishes one population from all of the others. The stepwise discriminant analysis provided a multivariate analysis of 14 characters in the 12 groups (males and females were treated separately in each of the geographic samples ) and produced a two-dimensional plot of the first and second canonical variables (Fig. 1). The first canonical variable is weighted primarily on webbing, secondly on subarticular tubercles, and lastly on ulnar and tarsal folds, whereas the second canonical variable is weighted on snout-vent length, calcar, and dorsolateral stripe, in descending order. The plot clearly shows the clustering of sam- ples and the overlap (in most cases) of the sexes within the samples. Thus, on the basis of adult morphology, the samples can be treated as different taxa. Distance Phenograms In an attempt to determine the phenetic relationships of the six geographic samples, each of the six was treated as an OTU. Mean values for each of 11 characters (sexes pooled) were used. Body ratios and the nature of the mental gland were excluded from the analysis. The resultant distance phenogram (Fig. 2A) shows the separation of the phenetically similar H. bogotensis and H. platydactyla from the other taxa. Hyla denticulenta is separated from the remaining three species clumped in the middle of the phenogram. The arrangement of the taxa is compatible with the plot of the canonical variables (Fig. 1). Thus, the results of the Phenogram Program supports the results of the multiple discrim- inant analysis. The previous analyses were based on characters of the adult frogs. Tadpoles are available for all six geographic samples; these have been associated with adults on the basis of morphological char- acteristics of the metamorphosing young. The most obvious inter- populational differences are evident in the structures of the mouths (Fig. 3). The distribution of character states of five characters is shown in table 3. For purposes of analysis, the mean ratio of depth of caudal musculature to total caudal depth was used; the other characters are constant within populations and were coded. The phenogram of populations as OTUs differs only slightly from the arrangement based on adults (Fig. 2B). The major difference is that H. alytolylax is clustered with H. denticulenta, whereas in the phenogram based on adults H. alytolylax is grouped with H, colymba and H. phyllognatha., 8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Fic. 1.—Plot of first (horizontal axis) and second (vertical axis) canonical variables; solid dots are males and open circles are females. A. Hyla alytolylax. B. H. colymba. C. H. phyllognatha. D. H. denticulenta. E. H. bogotensis. F. H. platydactyla. A final analysis of both adult and larval characters produced a phenogram somewhat intermediate between the first two (Fig. 2C). Here it can be seen that H. bogotensis and H. platydactyla are divergent from the other taxa and that H. colymba and H. phyllognatha are close to H. alytolylax, whereas H. denticulenta is more distant. FROGS OF THE HYLA BOGOTENSIS GROUP 9 H. denticulenta H. alytolylax H. colymba H. phyllognatha H. platydactyla H. bogotensis . denticulenta . alytolylax . colymba . phyllognatha . Platydactyla ae ea Me Rae eee ac . bogotensis H. denticulenta H. alytolylax H. colymba H. phyllognatha H. platydactyla H. bogotensis 3.0 20 1.0 0.0 DISTANCE Fic. 2.—Distance phenogram of Hyla bogotensis group based on un- weighted analysis of 11 adult and five larval characters. A. Adult characters. B. Larval characters. C. Adult and larval characters. 10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY PO OI Sai MMe Che “yj... 3 Mimi “ops Uy) . Mj, w “Wy Mii Fic. 3.—Mouths of tadpoles of Hyla bogotensis group. A. H. platydactyla, KU 139521. B. H. alytolylax, KU 112358. C. H. phyllognatha, KU 143542. D. H. denticulenta, KU 139531. X8. The mouth of the tadpole of H. bogotensis is like that of H. platydactyla except that the former has much smaller serrations on the beaks; the mouth of the tadpole of H. colymba is like that of H. phyllognatha (see Duellman, 1970:330). Mating Calls The mating calls are known for all species, except H. bogotensis and H. denticulenta; the calls consist of a series of short, loud peeps. Analysis of tape recordings of calls of three species reveals notice- able differences in several parameters of the calls (Table 4, Fig. 4). Hyla colymba has the most distinctive call, differing from the others in notes per call group, duration of notes, pulse rate, and _ pitch; the second harmonic is dominant. The calls of H. phyllognatha and H. platydactyla are more nearly alike in having only one har- TABLE 3.—Characteristics of tadpoles of Hyla bogotensis group. Total Caudal Tooth Labial Serrations Length/ Musculature/ Species Rows _—‘ Papillae on Beak Body length Caudal Depth H. alytolylax 5/7 2 small 30.2% 63.1% H. bogotensis 4/5 1 medium 32.3% 66.7% H. colymba 6/9 2 small 35.6% 61.1% H. denticulenta 7-8/10 2 none 35.3% 57.2% H. phyllognatha 6/9 3) small 35.2% 60.0% H. platydactyla 4/5 il large 39.5% 80.0% FROGS OF THE HYLA BOGOTENSIS GROUP 11 monic and fewer, but longer, notes per call group. However, their calls differ in note repetition rate, pulse rate, and pitch. The ab- sence of analyzable recordings of the other three species precludes a complete biosonic analysis of the group. Nevertheless, the dif- ferences in the parameters of the calls of the three sepcies are of the magnitude separating both sympatric and allopatric species in other Neotropical hylids [see Duellman (1963), Duellman and Trueb (1966), Duellman and Fouquette (1968), and Duellman (1972) for examples]. TaBLE 4.—Comparison of mating calls of species in the Hyla bogotensis group. Character H. colymba H. phyllognatha H. platydactyla IN 3/9 4/13 1/16 Notes per 12-104 2-12 7-10 Call Group (40.3) (5:5) (8.8) Note Repetition 123-236 120-360 540 Rate (min) (179) Glo” Wr a Duration 0.05 0.11-0.13 0.09-0.17 C50) | ane (0.12) (0.14) Pulses per 120-160 260-280 170-180 Second (138) (279) (177) Harmonics 4 1 1 Fundamental 1760-1820 2550-2620 2700-2860 Frequency ( Htz) (1796) (2588) (2790) Dominant 3520-3640 2550-2620 2700-2860 Frequency ( Htz) (3592) (2588) (2790) “ N=individuals/notes. DISCUSSION The grouping of specimens into geographic samples provided an a priori assumption that the groups represented different taxa. This assumption was tested by discriminant analysis of 14 charac- ters in adults. The results were re-tested by subsequent independent and combined analyses of 11 characters of adults and five of tad- poles by means of the distance phenogram program. The conclu- sions are supported by data on mating calls. Although it is possible that some of the taxa (H. bogotensis and H. platydactyla; H. colymba and H. alytolylax) might be geographical variants (sub- species), there exists no evidence for gene flow between popula- tions. Consequently, the six recognized taxa are treated as species. The distributions of the species are broadly allopatric (Fig. 5). Two species, H. bogotensis and H. platydactyla, occur in the Andes at elevations of 2500-2900 m and 1600-2700 m, respectively. Hyla colymba inhabits both Caribbean and Pacific slopes at elevations of 560-1410 m in Costa Rica and Panama. The other species occur on Andean slopes: H. alytolylax at 800-1460 m on Pacific slopes, 12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Kilohertz Time in Seconds Fic. 4—Audiospectrograms of mating calls. A. Hyla platydactyla, KU Tape 1029; 15-20 km NW Mérida, Mérida, Venezuela; temperature unknown B. H. phyllognatha, KU Tape 1166; Rio Azuela, Napo, Ecuador; 18° C. C. H. colymba, KU Tape 599; ridge between Rio Jaqué and Rio Imamadé, Darién, Panama; 19° C. Band width 20 Htz. H. phyllognatha at 610-1740 m on Amazonian slopes, and H. denticulenta at 1400-2400 m on Caribbean slopes. The principal habitat requirement seems to be cool, rocky streams for development of the tadpoles. Due to lack of substantive information on direction of evolu- tionary change in many of the characters utilized, no quantitative phyletic analysis (Kluge and Farris, 1969) was attempted. The FROGS OF THE HYLA BOGOTENSIS GROUP 13 Kilometers Fic. 5.—Distributions of the species in the Hyla bogotensis group. A. H. alytolylax. B. H. colymba. C. H. phyllognatha. D. H. denticulenta. E. H. bogotensis. F. H. platydactyla. Lines enclose approximate known ranges, within which specific localities are indicated by dots; the range of H. phyllognatha extends to southern Pert. presence of dermal folds and calcars in adults and proliferation of tooth rows in tadpoles are certainly derived states (Duellman, 1970). The occurrence of extensive webbing and bifid subarticular 14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY tubercles are probably derived character states. Furthermore, it seems that the presence of two tarsal stripes may have evolved from a condition in which one stripe was present. Other characters of structure and coloration remain evolutionarily enigmatic. All evi- dence points to H. bogotensis and H. platydactyla being the most generalized species, that is, having fewest derived character states. Hyla denticulenta possesses an array of derived states in both larvae and adults and seems to be the most advanced species in the group. The remaining three species seem to be closely related and phylogenetically intermediate between the primitive H. bogotensis and H. platydactyla and the advanced H. denticulenta. Evidence from a variety of sources points to considerable Pleisto- cene climatic fluctuation and concomitant shifts in vegetation in the northern Andes (see Vuilleumier, 1971, for summary). This climatic fluctuation, together with orogenic changes in the Andes during the Pleistocene and volcanic activity to the Present, provided ample opportunity for isolation of populations on different slopes of the Andes (H. alytolylax, H. denticulenta, and H. phyllognatha), in disjunct interandean basins (H. bogotensis and H. platydactyla), and dispersal across presently uninhabited lowlands (H. colymba). TAXONOMY The inclusion of six species in the Hyla bogotensis group neces- sitates a modification of the definition of the group as given by Duellman (1970:327). The group can now be defined: 1) moder- ate-sized, stream-breeding frogs with males attaining snout-vent lengths of 45 mm and females 53 mm; 2) dorsum pale green or brown with or without dark flecks; 3) digits bearing small discs; 4) toes about three-fourths webbed; 5) axillary membrane absent; 6) quadratojugal articulating with maxillary; 7) sphenethmoid broad, not ossified anteriorly; 8) nasals small, widely separated medially; 9) frontoparietal fontanelle large; 10) tadpoles having long muscular tails, low fins, ventral mouths completely bordered by papillae, and 4/5—8/10 tooth rows; 11) mating calls consisting of series of short, loud peeps. Distribution—The combined distributions of the six species include the Andes and interandean valleys of western Venezuela and central and southern Colombia, Pacific and Amazonian slopes of Andes in Ecuador, Amazonian slopes in Peru, Caribbean slopes in northern Colombia, and Caribbean and Pacific slopes of high- lands in Costa Rica and Panama, with an elevational range of 560 to 2900 m (Fig. 5). Remarks.—Members of the Hyla bogotensis group superficially resemble another group of species inhabiting lower Andean slopes in Ecuador, Colombia, and Venezuela (H. palmeri Boulenger, H. albopunctata Boulenger, and H. lascinia Rivero). These frogs lack mental glands; the mating calls and tadpoles are unknown. FROGS OF THE HYLA BOGOTENSIS GROUP 15 KEY TO THE SPECIES IN THE HYLA BOGOTENSIS GROUP 1. Tarsal and ulnar folds and tarsal and anal stripes present; distal subarticular tubercle on fourth finger single, conical Sema, at mot Conical )> 2 Tarsal and ulnar folds and tarsal and anal stripes absent; distal subarticular tubercle on fourth finger bifid, conical ___. 5 2. Distal subarticular tubercle on fourth finger single, conical; calcar, if present, small; snout round or truncate; tarsal and gialestipes single-or doubles... 3 Distal subarticular tubercle on fourth finger bifid, flat; calcar large; snout anteriorly inclined; tarsal and anal stripes double, elatvabove, dark below H. denticulenta 3. Dorsolateral stripe absent; webbing in males not extending to distal subarticular tubercle on fourth finger, reaching Raigerele in females 2. oes. H. phyllognatha Dorsolateral stripe usually present; webbing in both sexes extending to or beyond subarticular tubercle on fourth ote! 6 ai aes ae Seen tie oi Sees 4 4. Dark pigment fine, scattered on dorsum; webbing usually extending beyond distal subarticular tubercle on fourth OS a eg eT 2 ad H. alytolylax Dark pigment coarse, dense on dorsum; webbing extending to distal subarticular tubercle on fourth finger H. colymba 5. Webbing extending to distal subarticular tubercle on fourth finger; dorsolateral stripe absent; dark pigment usually fine and scattered on dorsum __... H. bogotensis Webbing not extending to distal subarticular tubercle on fourth finger; dorsolateral stripe present or absent; dark pigment on dorsum forming discrete spots in adults Mt heidi Le sere Si evid triacetin si oe level) ol edosts H. platydactyla Hyla alytolylax new species Figure 6A Holotype—KU 111903 from Tandapi, Provincia Pichincha, Ecuador, 1460 m, obtained on 22 July 1967, by John D. Lynch. Paratopotypes.—KU 111892-6, 111898-902, 111906, John D. and Marsha Lynch, 15-24 July 1967; KU 117982-4, John D. Lynch, 1-3 March 1968; KU 120851-60, John D. Lynch and Gerald R. Smith, 27-28 July 1968; KU 132425-6, John D. Lynch, 6 August 1970. Diagnosis.—1) Webbing extending to or beyond distal subar- ticular tubercle on fourth finger; 2) distal subarticular tubercles on fingers single, conical; 3) ulnar and tarsal folds present; 4) calcar absent in males, absent or small in females; 5) snout in profile round in males, truncate in females; 6) tympanum distinct; 7) 16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Fic. 6.—Frogs of the Hyla bogotensis group. A. H. alytolylax, KU 111900, 9, 39.6 mm SVL. B. H. denticulenta, KU 133451, ¢, 44.2 mm SVL. C. H. phyllognatha, KU 143197, 6, 33.3 mm SVL. D. H. platydactyla, KU 133433, 6, 36.5 mm SVL. mental gland present in males, absent in females; 8) dorsolateral light stripe present; 9) tarsal and anal stripes white; 10) tadpoles having 5/7 tooth rows, two rows of labial papillae, and small serrations on beaks. Hyla alytolylax most closely resembles H. colymba, from which it differs by having slightly more webbing and less dense flecking on the dorsum; furthermore, the tadpoles of H. alytolylax have fewer tooth rows and a narrower snout. Hyla alytolylax also re- sembles H. phyllognatha, from which it differs by having more webbing, no or smaller calcars, and a dorsolateral light stripe; furthermore, the tadpoles of H. phyllognatha have more tooth rows and a broader snout. Description—N=13 4 6, 152 ¢; pertinent measurements and proportions given in table 1. Head slightly narrower than body; snout moderately short, round in dorsal aspect, round in profile in FROGS OF THE HYLA BOGOTENSIS GROUP 17 males, truncate in females; nostrils four-fifths distance from eye to tip of snout, not protuberant; internarial area slightly depressed; canthus round; loreal region concave; lips thin, round; top of head flat; eyes not greatly protuberant; supratympanic fold weak, curved downward toward angle of jaw, obscuring upper edge of tympanum; tympanic ring evident ventrally; tympanum slightly less than one- half diameter of eye (0.382-0.484, x=0.434, N=13 4 ¢; 0.425-0.500, x—0.457, N=15¢@ 2), separated from eye by distance slightly greater than diameter of tympanum. Axillary membrane absent; forearm robust, bearing ulnar fold; prepollical tubercle large, elliptical; palmar tubercle large, bifid; distal subarticular tubercle on fourth finger conical, single; super- numerary tubercles small, indistinct; fingers short, bearing small discs (that on third finger slightly larger than tympanum); length of fingers from shortest to longest 1-2-4-3; webbing vestigial be- tween first and second fingers, extending from base of penultimate phalange of second finger to middle of antepenultimate phalange of third, from base of penultimate phalange of third to base or middle of penultimate phalange of fourth finger. Hind limb moder- ately robust; dermal fold on knee; calcar usually absent; inner tarsal fold absent; outer tarsal fold extending entire length of tarsus; inner metatarsal tubercle elliptical, barely visible from above; outer tarsal tubercle absent; subarticular tubercles small, round; super- numerary tubercles minute, present proximally; toes long; length of toes from shortest to longest 1-2-3-5-4; discs slightly smaller than those of fingers; webbing extending to base of penultimate phalange of fourth toe, to discs on other toes. Anal opening directed posteriorly at upper level of thighs in both sexes; skin on belly and proximal posteroventral surfaces of thighs weakly granular; skin on other surfaces smooth; mental gland present in males. Tongue cordiform, shallowly notched posteriorly, barely free behind; dentigerous processes of prevomers antero- medially inclined posterior to small ovoid choanae; total number of prevomerine teeth 15-22 (x=18.8, N=13¢ ¢ ), 20-24 (x=21.8, N=15@ 2 ); vocal slit extending from midlateral base of tongue to angle of jaw; vocal sac bilobate, subgular. Coloration.—In life, adults have a greenish dorsum varying from pale green to greenish yellow, greenish gray, or greenish brown. In some individuals the anterior part of the dorsum is darker than the posterior part, and the loreal region usually is dark green. A cream line extends along the canthus, edge of eyelid, and supratympanic fold; a narrow white tarsal stripe and white anal stripe are present. The dorsal surfaces of the body and limbs have small scattered brown or black flecks and, in some individuals, minute white flecks. In some specimens faint cream reticulations are present on the dorsal surfaces of the body and thighs. The groin and concealed surfaces of the limbs are not pigmented. The 18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY venter is white, and in males the vocal sac is bluish green. The iris is creamy white. In preservative, the dorsum is creamy tan with black flecks, varying from few to many, scattered over the dorsum. Dorsolateral, tarsal, and anal white stripes are evident. The groin, hidden sur- faces of thighs, and all ventral surfaces are cream. Pale reticula- tions are evident posteriorly in some specimens. Tadpoles.—A typical tadpole in developmental stage 27 from Tandapi, Ecuador, has a body length of 15.2 mm and a total length of 42.6 mm. Body ovoid, widening abruptly just anterior to eyes, three-fourths as deep as wide; nostrils about midway between eyes and tip of snout; spiracle sinistral; spiracular opening directed posteriorly at a point just below midline about two-thirds length of body; cloacal tube dextral; caudal fin not extending onto body; caudal musculature robust, tapering gradually, extending nearly to tip of pointed tail; depth of caudal musculature at one-third length of tail 63 percent of total depth of tail. Mouth moderately large; lips indented posterolaterally, completely bordered by two rows of small labial papillae; five upper and seven lower rows of teeth; fifth upper row narrowly interrupted medially; beaks slender; lateral processes not expanded distally; upper beak bearing small peglike serrations; lower beak having small pointed serrations (Fig. 3B). In preservative, the body is dark brown above and dull gray below; the caudal musculature is cream with brown flecks or reticu- lations laterally. The dorsal musculature is cream with transverse brown bars. The fins are transparent, except for a few brown flecks on the outer edge of the dorsal fin. In life, the body is pale brown above and dark brown below; the caudal musculature is pale cream with brown mottling laterally and blotches dorsally. The iris is silvery cream. Etymology.—the specific name is derived from the Greek alytos meaning continuous, and the Greek lylax meaning babbler; the name is applied in reference to the long continuous calls of the species. Distribution.—Hyla alytolylax is definitely known from eleva- tions of 800 to 1460 m on the Pacific slopes of the Cordillera Occi- dental in Ecuador and southeastern Colombia. The northern lo- calities in Ecuador are along the Rio Pilatén, a tributary of the Rio Toachi, and the southern localities are in the Rio Playas drainage. Remarks.—At Tandapi, Ecuador, males were calling from vegetation over, and boulders in, cascading mountain streams in cloud forest in July 1967 and in March and July 1968. Gravid females were found on bushes along streams. Tadpoles were ob- tained from gravel-bottomed pools in fast-flowing streams. Meta- morphosing young were found on low vegetation along streams in July 1967, and two metamorphosed on 14 September from tadpoles obtained on 17 July 1967. Five young have snout-vent lengths of FROGS OF THE HYLA BOGOTENSIS GROUP 19 17.6-20.4 (x=18.4) mm. In life, the dorsal surfaces were pale green with a middorsal brown streak, expanded anteriorly into an interorbital bar in one individual. The mating call consists of a long series of short, loud whistles, reminiscent of the call of Hyla colymba. Analyzable recordings are not available. Two specimens (LACM 73000-1) from Camino de Yupe, Chocé, Colombia (420 m), possibly are referable to Hyla alytolylax, but because of certain differences, they have not been included in the analysis nor in the description. Both are males having snout-vent lengths of 39.0 mm and large mental glands. The dorsolateral stripe is absent, but tarsal and anal stripes are present; on either side of the anus is a triangular dermal flap, larger than the calcar. Perhaps these specimens represent an undescribed species that is intermedi- ate in certain characters between H. alytolylax and H. denticulenta. Duellman (1970:331) referred specimens of H. alytolylax to H. colymba. Hyla bogotensis (Peters) Hylonomus bogotensis Peters, 1882a:107 [Holotype—ZMB 10209 from Bogota, Departamento Cundinamarca, Colombia; Moschler collector]. Hyloscirtus bogotensis Peters, 1882b:127. Hyla bogotensis—Duellman, 1970:327. Diagnosis.—1) Webbing extending to distal subarticular tuber- cle on fourth finger; 2) distal subarticular tubercles on fingers bifid, conical; 3) ulnar and tarsal folds absent; 4) calcar absent; 5) snout in profile truncate; 6) tympanum distinct or covered in males, covered in females; 7) mental gland present or absent in males, absent in females; 8) dorsolateral light stripe absent; 9) tarsal and anal stripes absent; 10) tadpoles having 4/5 tooth rows, one row of labial papillae, and medium-sized serrations on beaks. Hyla bogotensis is like H. platydactyla in having bifid, conical subarticular tubercles, no ulnar and tarsal folds, no tarsal and anal stripes, and tadpoles with 4/5 tooth rows and one row of labial papillae; these characters distinguish these two species from other members of the group. Hyla bogotensis can be distinguished from H. platydactyla by the presence of more webbing and finer dorsal flecking in the former; moreover, the tadpoles of H. bogotensis have smaller serrations on the beaks than do those of H. platydactyla. Coloration.—I have not seen living individuals of this species, so I must rely upon descriptions given by others. Cochran and Goin (1970:315) noted that a specimen from Bogota, Colombia, was “...a yellowish brown with a definite pattern of more or less uniformly distributed pencil-like black spots on all the dorsal sur- faces. All ventral surfaces were a dirty gray without pattern, except for faint pepper-like flecks under the throat and a yellowish tinge to the under-margin of the jaws.” Philip A. Silverstone (field notes, 20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 6 January 1966) described a specimen (LACM 50562) from Paramo de Puracé, Colombia, as “Green all over; yellowish tinge on flanks and ventral surfaces of thighs. Blue on ankle and sole; web orange.” Juan A. Rivero (pers. com.) stated that a specimen (UPR-M 1800) from Paéramo de Palacio was solid brown above in life, but that the freshly preserved specimen was chartreuse above with dark brown dots; there was some yellow on the throat and ventral surfaces of the thighs. In preservative, most specimens are creamy tan with or without scattered dark flecks on the dorsum. Two individuals (FMNH 81916 and UPR-M 1800) have many larger black dots on the dorsum. Tadpoles.—A single tadpole (CAS-SU 22971) in developmental stage 37 has a body length of 21.0 mm and a total length of 68.0 mm. Body ovoid, two thirds as deep as wide; nostrils slightly closer to eye than to tip of snout; spiracle sinistral; spiracular opening directed posterodorsally on midline at about midlength of body; cloacal tube dextral; caudal fin not extending onto body; caudal musculature robust, tapering gradually; extending nearly to tip of pointed tail; depth of caudal musculature at one-third length of tail 66.7 percent of total depth of tail. Mouth moderately large; lips indented posterolaterally, completely bordered by single row of labial papillae; four upper and five lower rows of teeth; fourth upper row narrowly interrupted medially; beaks massive; lateral processes expanded distally; beaks bearing medium-sized serrations. In preservative the body is pale brown above and cream below. The caudal musculature is pale brown with a dorsolateral dark brown stripe on the proximal half of the tail. Distribution.—This species is known from subparamo regions in the Bogota Valley in the Cordillera Oriental and in the highlands east of Popoyan in the Cordillera Central in Colombia. All re- corded elevations are between 2500 and 2900 m. Remarks.—For many years this species was regarded as being generically distinct from Hyla on the basis of having rounded, in- stead of dilated, sacral diapophyses. Duellman (1970:328) noted that the type (ZMB 10209) actually had expanded sacral diapophyses. Subsequent examination of additional specimens supports the con- tention that the sacral diapophyses are not round in this species. The mating call is unknown. Dunn (1944:79) noted that a specimen was found in a bromeliad at Boquerén, Colombia. Philip A. Silverstone (field notes, 6 January 1966) found one individual on a leaf near a stream in subparamo by day. Notes by Anne Funkhouser accompanying a tadpole and partially metamorphosed young (CAS-SU 22970-1) state: “Fast mountain stream on road to Choachi from Bogota. Fairly common in area collected. Adults were never found, but young kept for several months were bright grass green above with random dark spots; turquoise shading to FROGS OF THE HYLA BOGOTENSIS GROUP 21 yellow ventrally; iris gold.”. The metamorphosing young (CAS-SU 22970) has a snout-vent length of 22.0 mm and a tail stub of 7.2 mm; in preservative, it is creamy tan with scattered brown flecks on the dorsum. Hyla colymba Dunn Hyla colymba Dunn, 1931:400 [Holotype—MCZ 10234 from La Loma, Provincia Bocas del Toro, Panama; Chester Duryea and Emmett R. Dunn collectors]. Hyla alvaradoi Taylor, 1952:882 [Holotype—KU 30886 from Moravia, Provincia Cartago, Costa Rica; Edward H. Taylor collector]. Synonymy fide Duellman (1966:267 ). Diagnosis.—1) Webbing extending to distal subarticular tuber- cle on fourth finger; 2) distal subarticular tubercles on fingers single, conical; 3) ulnar and tarsal folds present; 4) calcar absent; 5) snout in profile round; 6) tympanum distinct; 7) mental gland present in males, present or absent in females; 8) dorsolateral light stripe present in males, present or absent in females; 9) tarsal and anal stripes white; 10) tadpoles having 6/9 tooth rows, two rows of labial papillae, and small serrations on beaks. Hyla colymba most closely resembles H. alytolylax, from which it differs by having slightly less webbing and more dense flecking on the dorsum; furthermore, the tadpoles of H. colymba have more tooth rows and a broader snout. Hyla colymba differs from H. phyllognatha by having more webbing and usually a dorsolateral light stripe; the structure of the tadpoles of the two species is identical, but dark transverse blotches are present on the dorsum of the tail in H. phyllognatha, whereas the tail in H. colymba is nearly uniform brown. Coloration.—In life, most individuals are pale green with faint yellow flecks or scattered brown flecks on the dorsum. A faint creamy yellow stripe extends from the canthus, along the edge of the eyelid and on the supratympanic fold to a point above the arm. The anal, ulnar, and tarsal stripes are pale creamy yellow. The ventral surfaces of the limbs are pale green, and the throat is pale bluish green. The belly and chest are white, and the mental gland is white or creamy yellow. Some individuals are pale tan with brown flecks. The iris is pale brown. In preservative, the dorsum is pale creamy white, creamy tan, or pale brown. Minute dark flecks are present dorsally; usually these are most numerous on the head and middorsally on the body. The stripes are white, and the venter is creamy white. Tadpoles.—A typical individual in developmental stage 25 from Darién, Panama, has a body length of 15.1 mm and a total length of 37.3 mm. Body ovoid, four-fifths as deep as wide; nostrils mid- way between eyes and tip of snout; spiracle sinistral; spiracular opening directed posteriorly just below midline at midlength of 22 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY body; cloacal tube dextral; caudal fin not extending onto body; caudal musculature robust, tapering gradually, extending nearly to tip of pointed tail; depth of caudal musculature at one-third length of tail 61 percent of total depth of tail. Mouth large, ventrals; lips deeply indented posterolaterally, completely bordered by two rows of labial papillae; six upper and nine lower rows of teeth; sixth upper and first lower rows narrowly interrupted medially; outermost three lower rows shorter than others; upper beak broadly arched, bearing long slender lateral processes; lower beak broadly V-shaped; both beaks bearing small, blunt serrations. In preservative, the body is brown with faint white flecks. The caudal musculature is creamy tan; brown flecks are present on the musculature and dorsal fin. In life, the dorsum of the body is bronze-tan with golden lichenous flecks; the sides of the body are brown with gold flecks. The caudal musculature is tan; dark brown spots are present on the musculature and dorsal fin. This iris is dull bronze. Distribution—This species occurs in cloud forest and lower humid montane forest at elevations of 610-1200 m on the Caribbean slopes of the Cordillera Talamanca in Costa Rica and Western Panama and 560-1410 m on the Pacific slopes of the highlands in central and eastern Panama. Remarks.—Duellman (1970:328) gave a detailed account of H. colymba and discussed the nomenclatural history of the species; his reference to the occurrence of the species in Ecuador was based on material herein assigned to H. alytolylax. The mating call consists of a series of short high-pitched, cricket- like chirps. Calls have as many as 104 notes produced at a rate of 123-236 notes per minute. The second harmonic at about 3600 Hertz is dominant (Fig. 4C). Adults have been found on vegetation along streams, and males were found calling beneath boulders in streams. Tadpoles have been found in quiet pools in small rocky streams. Hyla denticulenta new species Figure 6B Holotype —KU 133451 from Charta, Departamento Santander, Colombia, 2400 m, obtained on 15 July 1970, by Stephen R. Edwards. Paratype —KU 133452 from Valdivia, Departamento Antioquia, Colombia, obtained on 3 August 1970, by Stephen R. Edwards. Diagnosis.—1) Webbing extending beyond distal subarticular tubercle on fourth finger; 2) distal subarticular tubercles on fingers bifid, flat; 3) ulnar and tarsal folds present; 4) calcar large; 5) snout in profile anteriorly inclined; 6) tympanum distinct; 7) mental gland absent; 8) dorsolateral stripe absent; 9) tarsal and FROGS OF THE HYLA BOGOTENSIS GROUP 23 anal stripes white above, brown below; 10) tadpoles having 7-8/10 tooth rows, two rows of labial papillae, and no serrations on beaks. Hyla denticulenta differs from all other members of the group by its large size, bifid and flat subarticular tubercles, large calcars, and anteriorly inclined snout; also, the presence of 7-8/10 tooth rows and absence of serrations on the beaks distinguish the tadpoles from all others. Description—N=1 ¢ (holotype), 1 ° (paratype); pertinent mea- surements and proportions given in table 1. Head as wide as body; snout moderately short, round in dorsal aspect, anteriorly inclined in profile (especially in female); nostrils three-fourths distance from eye to tip of snout, not protuberant; internarial area flat; canthus round; loreal region concave; lips thin, round; top of head flat; eyes not greatly protuberant; supratympanic fold weak, curved downward toward angle of jaw, obscuring upper part of tympa- num; tympanic ring distinct; tympanum one-half diameter of eye, separated from eye by distance equal to half again diameter of tympanum. Axillary membrane absent; forearm robust, bearing ulnar fold; prepollical tubercle large, elliptical; palmar tubercle large, bifid; distal subarticular tubercle on fourth finger flat, bifid; others round; supernumerary tubercles small, indistinct; fingers short, bearing small discs (that on third finger slightly larger than tympanum); length of fingers from shortest to longest 1-2-4-3; webbing vestigial between first and second fingers, extending from base of penultimate phalange of second to distal end of antepenultimate phalange of third, from base of penultimate phalange of third to base (male) or middle (female) of penultimate phalange of fourth finger. Hind limb moderately robust; dermal fold on knee; calcar prominent, triangular; inner tarsal fold absent; outer tarsal fold extending length of tarsus; inner metatarsal tubercle elliptical, visible from above; outer tarsal tubercle absent; subarticular and supernumerary tubercles about equal in size, small, round; toes long; length of toes from shortest to longest 1-2-3-5-4; discs slightly smaller than those on fingers; webbing extending to base of penultimate phalange of fourth toe, to discs on other toes. Anal opening directed posteroventrally at midlevel of thighs (male), posteriorly at upper level of thighs (female); skin on belly and proximal posteroventral surfaces of thighs weakly granular; skin on other surfaces smooth. Tongue broadly cordiform, shallowly notched posteriorly, barely free behind; dentigerous processes of prevomers slightly anteromedially inclined posterior to small ovoid choanae, bearing 21 (male), 20 (female) teeth; vocal slit extending from posterolateral base of tongue to angle of jaw; vocal sac sub- gular, possibly bilobate. Coloration.—In life, the male holotype had a pale green dorsum covered with many small white flecks; black flecks were present 24 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY on the head and back. The flanks were creamy yellow; the throat, chest, groin, anterior surfaces of the thighs, and ventral surfaces of the shanks were bluish green. The belly was creamy yellow, and the iris was reddish gold. The female had a green dorsum with a white margin to the lips and white anal, ulnar, and tarsal stripes. The axilla, groin, and ventral surfaces of the thighs were blue-green; the flanks were yellow-orange. The throat and belly were greenish white, and the iris was dark gold with black reticulations. In preservative, all pigment is lost, except for many small brown flecks on the head and anterior two-thirds of the back in the male and a few scattered flecks on the eyelids, head, and middorsum of the female. Also, white flecks are apparent dorsolaterally in the male. In both specimens narrow white anal, ulnar, and tarsal stripes are bordered below by a narrow line of brown pigment, which is expanded on the palm and sole. Tadpoles—Five tadpoles (KU 189531) are from the type locality. The smallest individual is in developmental stage 25 and has a body length of 17.5 mm and a total length of 51.5 mm; the largest is in stage 38 and has measurements of 22.0 and 65.5 mm. Body ovoid, four-fifths as deep as wide, widening and deepening abruptly just anterior to the eyes; nostrils midway between eyes and tip of snout; spiracle sinistral; spiracular opening directed pos- teriorly just below midline at about midlength of body; cloacal tube dextral; caudal fin not extending onto body; caudal muscula- ture robust, gradually tapering, extending nearly to tip of pointed tail; depth of musculature at one-third length of tail 57 percent of total depth of tail. Mouth large, ventral; lips deeply indented posteriolaterally, completely bordered by two rows of small papillae; seven or eight upper and ten lower rows of teeth; proximal upper row narrowly interrupted medially; upper beak slender with long, tapering lateral processes; lower beak broadly V-shaped; both beaks smooth or bearing minute serrations (Fig. 3D). In preservative, the body is dark brown above and dull gray below; the caudal musculature is cream with dense brown pigment laterally, forming distinct transverse bars, separated by narrow cream interspaces, dorsally. The fins are transparent, except for a few flecks on the dorsal fin. In life, the body is pale gray; the tail is distinctly mottled black, gray, and white, and the fins are transparent. Etymology.—tThe specific name is derived from the Latin den- ticulus meaning with small teeth, and the Latin lentus meaning full of; the name refers to the many rows of teeth in the tadpoles. Distribution—This species is known from only two localities: Charta at an elevation of 2400 m on the northern slope of the Cordillera Central, and Valdivia at an elevation of 1400 m on the northern slope of the Cordillera Central. These localities are sepa- rated by the low Rio Magdalena Valley. It is unlikely that gene FROGS OF THE HYLA BOGOTENSIS GROUP 25 flow exists between the populations; thus, it is possible that addi- tional material will reveal differences between the populations. Remarks.—At Charta the frogs were found along a shallow river south of the village. The river, a tributary of the Rio Surata, flows through pasture land with a few scattered trees. Tadpoles and a metamorphosing young having a snout-vent length of 24.6 mm and a tail stub of 10.4 mm were found in pools in the river bed. The holotype was found by day under a large rock in a small stream cascading into the river bed. The adult female was found in the town of Valdivia, where it was perched on a leaf of a large herb 2 m above the ground on a steep bank at night. The mating call is unknown. Hyla phyllognatha Melin Figure 6C Hyla phyllognatha Melin, 1941:30 [Holotype—NHMG 474 from Roque, Departamento San Martin, Peru; Douglas Melin collector]. Diagnosis —1) Webbing not extending to, or just extending to, distal subarticular tubercle on fourth finger; 2) distal subarticular tubercles on fingers single, conical; 3) ulnar and tarsal folds present; 4) calcar absent or small; 5) snout in profile round or truncate; 6) tympanum distinct; 7) mental gland present in males, present or absent in females; 8) dorsolateral light stripe absent; 9) tarsal and anal stripes absent or white; 10) tadpoles having 6/9 tooth rows, two rows of labial papillae, and small serrations on beaks. Hyla phyllognatha is like H. colymba and H. alytolylax but has less webbing and no dorsolateral stripe. In some specimens of H. phyllognatha a calcar is present, but this is smaller than the calcars in H. denticulenta, which further differs by having bifid subarticular tubercles, anteriorly inclined snout, and more webbing. The tadpoles of H. phyllognatha have 6/9 tooth rows, a character shared with H. colymba, the tadpoles of which differ from those of H. phyllognatha by not having transverse brown blotches on the dorsal caudal musculature. Coloration.—In life, the dorsum is olive-green anteriorly chang- ing to olive-tan posteriorly; the dorsal surfaces of the limbs are green. Minute black flecks are present on the head and middorsal part of the body, and small white flecks are scattered on the body and limbs. The margin of the lip is yellow, and the anal and tarsal stripes are pinkish white. The hidden surfaces of the limbs are dull green. The belly is creamy yellow, and the webbing is dull yellow. The vocal sac is green, and the lining of the mouth is bluish green. The iris is dull bronze with black reticulations. One female found by day was bright green with bluish white flecks. In preservative, the dorsum is cream to grayish brown with scattered dark flecks apparent in paler specimens. The venter and hidden surfaces of the thighs are cream. 26 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Tadpoles.—A typical tadpole in developmental stage 27 from 16.5 km NNE of Santa Rosa, Ecuador, has a body length of 16.5 mm and a total length of 50.3 mm. Body ovoid, slightly wider than deep; nostrils slightly closer to tip of snout than to eyes; spiracle sinistral; spiracular opening directed posteriorly just below midline about two-thirds length of body; cloacal tube dextral; caudal fin not extending onto body; caudal musculature robust, tapering gradually, extending nearly to tip of pointed tail; depth of musculature at one-third length of tail 60 percent of total depth of tail. Mouth large, lips indented posterolaterally, completely bordered by two rows of small labial papillae; six upper and nine lower rows of teeth; sixth upper row narrowly interrupted medially; beaks moderately slender, bearing small, blunt serrations; upper beak arched with lateral processes barely expanded distally; lower beak broadly V-shaped (Fig. 3C). In preservative, the body is dark brown above and gray below; the caudal musculature is cream with dense brown flecking laterally and dark brown transverse bars anterodorsally. The fins are trans- parent with brown flecks, except anteriorly on ventral fin. In life, the body is dark brown with small green, lichenous flecks. The caudal musculature is tan with brown blotches; the fins are clear with orange spots. The iris is dark bronze. Distribution —Hyla phyllognatha is known from elevations of 610-1740 m on the Amazonian slopes of the Cordillera Oriental of the Andes and associated ranges, such as the Cordillera del Dué and Cordillera Carabaya, from northern Ecuador to southern Peru. Remarks.—Males call from low bushes over torrential mountain streams; calling males have been found in April, July, August, and October. Gravid females have been found in October. The tadpoles live in gravel-bottomed pools in the streams. Recently metamor- phosed young have been found on vegetation along streams at night and in bromeliads by day. Four young have snout-vent lengths of 18.4-20.3 (x=19.5) mm; in life, the dorsum is green with black flecks middorsally. The mating call consists of a series of short, loud whistles, repeated at a rate of about 210 notes per minute. The notes have about 280 pulses per second; the fundamental frequency at about 2560 Hertz is dominant (Fig. 4B). _ No information has been published on Hyla phyllognatha since Melin’s (1941) original description. The holotype (NHMG 474) is an adult male having a snout-vent length of 32.3 mm and a mental gland. The specimen compares favorably with series of fresh specimens from Equador and three individuals from southern and central Pert. FROGS OF THE HYLA BOGOTENSIS GROUP 27 Hyla platydactyla Boulenger Figure 6D Hyla platydactyla Boulenger, 1905:183 [Holotype—BMNH 1904.6.30.17 (RR 1947.2.13.14) from Mérida, Estado Mérida, Venezuela; Bricefio collector]. Hyla paramica Rivero, 1961:112 [Holotype—UMMZ 59016 from Escorial, Estado Mérida, Venezuela; received from W. F. H. Rosenberg]. New synonymy. Hyla jahni Rivero, 1961:113 [Holotype—UMMZ 46465 from Escorial, Estado Mérida, Venezuela; presented by A. G. Ruthven]. New synonymy. Diagnosis—1) Webbing not extending to distal subarticular tubercle on fourth finger; 2) distal subarticular tubercles on fingers single or bifid in males, bifid and conical in females; 3) ulnar and tarsal folds absent; 4) calcar absent; 5) snout in profile truncate; 6) tympanum distinct or covered; 7) mental gland present or absent in both sexes; 8) dorsolateral stripe present or absent; 9) tarsal and anal stripes absent; 10) tadpoles having 4/5 tooth rows, one row of labial papillae, and large serrations on beaks. Hyla platydactyla is like H. bogotensis in having bifid, conical subarticular tubercles, no tarsal or anal stripes, no ulnar or tarsal folds, and tadpoles with 4/5 tooth rows and one row of labial papillae; these characters distinguish these two species from other members of the group. Hyla platydactyla can be distinguished from H. bogotensis by having less webbing and, in large adults, discrete dark spots dorsally. Also, the tadpoles of H. platydactyla have larger serrations on the beaks than do those of H. bogotensis. Coloration—In life, specimens from a locality 15-20 km NW of Mérida, Venezuela, had reddish brown dorsal surfaces with many small black spots on the back. The throat was greenish tan, and the belly was white. The iris was copper with fine black reticulations. Through the courtesy of Juan A. Rivero, I have seen colored photographs of individuals having olive-tan dorsal colora- tion and creamy yellow dorsolateral stripes. Rivero stated (pers. com.) that some individuals are dark brown above and that in most specimens the anterior and posterior surfaces of the thighs are orange. In preservative, in adults the dorsal surfaces of the head, body, forearms, and shanks are tan to brown with small dark brown to black spots present in most specimens. The dorsolateral stripe, thighs, and ventral surfaces are cream. Tadpoles.—In a series from 15-20 km NW of Mérida, Venezuela, the smallest tadpole is in developmental stage 25 and has a body length of 8.5 mm and a total length of 26.5 mm; the largest tadpole (stage 37) has measurements of 23.0 and 60.5 mm. Body ovoid, three-fourths as deep as wide; nostrils slightly closer to eyes than to tip of snout; spiracle sinistral; spiracular opening directed pos- teriorly on midline about midlength of body; cloacal tube dextral; caudal fin not extending onto body; caudal musculature robust, 28 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY tapering gradually, extending nearly to tip of pointed tail; depth of caudal musculature at one-third length of tail 80 percent of total depth of tail. Mouth moderately large; lips deeply indented postero- laterally, completely bordered by one row of labial papillae; four upper and five lower rows of teeth; fourth upper row narrowly in- terrupted medially; fifth lower row short; upper beak moderately massive, arched, with robust lateral processes and large, pointed serrations; lower beak massive, broadly V-shaped, bearing large blunt serrations (Fig. 3A). In preservative, the body is dark brown above and paler brown below; the caudal musculature is creamy tan with a heavy suffusion of brown laterally and two or three distinct longitudinally rec- tangular blotches on the dorsum, separated by narrow cream inter- spaces. The dorsal fin and posterior one-third of ventral fin are heavily flecked with brown. Distribution—This species is known only from the Mérida Andes in western Venezuela, where it occurs at elevations from 1600 to at least 2500 m, inhabiting subtropical and temperate areas. Remarks,—Rivero (1961:115) diagnosed Hyla paramica from H. jahni by the former having a “. . . rounded snout, less defined canthus and in lacking supratympanic fold, canthal, palpebral, and supratympanic lines and closely set dark dots above.” He (1961: 116) stated that Hyla platydactyla differed from H. jahni “. . . in the shorter snout and less defined canthus, absence of a light canthal and supratympanic line, narrower interorbital space, more vertical loreal region and different coloration [no dorsal spots].” Rivero based his descriptions of H. paramica on five specimens and of H. jahni on seven, all of which are rather poorly preserved; he examined no specimens of H. platydactyla. I have examined all of the specimens studied by Rivero and the holotype of H. platydactyla, together with several series from western Venezuela (total of 59 adults and subadults). I find no consistent structural differences, such as snout shape, definition of canthal ridge, or inclination of loreal region, by which to distinguish more than one taxon in western Venezuela. The dorsolateral stripe (canthal, palpebral, and supratympanic lines of Rivero) is variable; a stripe is evident in the supratympanic region in slightly more than half of the specimens; in some of these it extends posteriorly to the axilla. Also, in some of these specimens no stripe is evident on the canthus. On the basis of preserved specimens alone, there seems to be a continuum from well-defined to no stripes. Thus, only one species, H. platydactyla, can be recognized. The mating call consists of a series of short notes repeated at a rate of 540 notes per minute; the pulse rate is about 177 pulses per second, and the fundamental frequency at about 2790 Hertz is dominant (Fig. 4A). FROGS OF THE HYLA BOGOTENSIS GROUP 29 At a locality 15-20 km NW of Mérida, Venezuela, adults were found at night on leaves of herbs on a steep bank over a roadside ditch. Tadpoles were found in quiet pools in a cascading stream. Juan A. Rivero stated (pers. com.) that adults are found in brome- liads by day and that males call from bushes over streams at night. He also noted that the frogs produce a smelly exudate when handled. Cochran and Goin (1970:256) proposed the name combination Hyla labialis platydactyla; the name was used for the population of frogs in the eastern Andes of Colombia that is referable to the highly variable Hyla labialis, a species quite distinct from H. platydactyla. RESUMEN Las ranas del grupo Hyla bogotensis se crian en los arroyos de las vertientes de los Andes en el occidente de Venezuela, Colombia, Ecuador, y Peru, y en las sierras de Costa Rica y Panama. Las ranas de este grupo se caracterizan por: 1) su tamafio moderado (longitud de cabeza-cuerpo de los machos hasta 45 mm, en las hembras hasta 53 mm); 2) el dorso de color verde opaco o carmelita con manchas pequenas oscuras espacidas en el; 3) los dedos con tres cuarta partes cubiertos por la membrana interdigital; 4) la glandula del mentdn en el apice de éste; 5) membrana _ axilar ausente; 6) el cuadratoyugal articulado con el maxilar; 7) los nasales pequenos, separados ampliamente en el medio; 8) la fon- tanela frontoparietal grande; 9) renacuajos con colas musculares, largas, la boca ventral grande, completamente bordeada por papilas labiales, con 4/5—8/10 hileras de dientes. Se analizaron 14 caracteres de ranas adultas por medio del programa de computacién de anéalisis discriminante multiple; los seis grupos resultantes fueron tratados como OTUs en los programas de NT-SYS, en el cual 11 caracteres de adultos y cinco de los renacuajos constituyeron los datos basicos para la construccién de los fenogramas. Los resultados de estos andalisis son sostenidos por el analisis de las llamadas de reclamo de tres especies. Se reconocen seis especies en el grupo: 1) H. alytolylax especie nueva, laderas del Pacifico de los Andes en Ecuador; 2) H. bogo- tensis (Peters, 1882), Andes del centro y sur de Colombia; 3) H. colymba Dunn, 1931, sierras de Costa Rica y Panama; 4) H. denticulenta especie nueva, laderas del Caribe de los Andes en el norte de Colombia; 5) H. phyllognatha Melin, 1941, laderas amazonicas de los Andes en Ecuador y Pert; 6) H. platydactyla Boulenger, 1905, Andes del occidente de Venezuela. Hyla paramica Rivero, 1961, y Hyla jahni Rivero, 1961, son colocadas en sinénimia de Hyla platydactyla Boulenger, 1905. 30 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY SPECIMENS EXAMINED Hyla alytolylax—COLOMBIA: Chocé: Camino de Yupe, 420 m, LACM 73000-1. Valle: Anchicayé, KU 148703-4. ECUADOR: Bolivar: Balsabamba, 800 m, KU 132432, 132545 (tadpoles). Cotopaxi: 20.3 km W Pilalé, 830 m, KU 142857 (tadpoles). Pichincha: 3.7 km E. Dos Rios, 1190 m, KU 142856 (tadpoles); Las Palmas, 920 m, KU 132427; Tandapi, KU 111892-903, 111904 (skeleton), 111905-10, 112357-8 (tadpoles), 112359 (young), 117982-4, 118123 (tadpoles), 120851-60, 132425-6. Hyla bogotensis—COLOMBIA: Cauca: Moscopan, 2500 m, UMMZ 121032. Cundinamarca: Bogota, 2630 m, CAS-SU 22970 (tadpole), 22971 (young), KU 125363, UMMZ 123946-8, ZMB 10209; Boquerén, 2900 m, FMNH 81916; Paramo de Palacio, Siberia, UPR-M 1800. Huila: east of Paramo de Puracé, 2500 m, LACM 50562. Hyla colymba.—COSTA RICA: Cartago: Moravia, 1200 m, KU 30886, 31864, 31865 (skeleton). PANAMA: Bocas del Toro: La Loma, 610 m, MCZ 10234-5; Rio Changena, 650 m, KU 104237 (young); Rio Changena, 830 m, KU 104236 (tadpoles). Coclé: El Valle, 560 m, AMNH 59606. Darién: Cerro Citurio, Serrania de Pirre, 1100 m, KU 116357; Cerro Mali, 1410 m, GML 4-00426; Laguna, 820 m, KU 77414; ridge between Rio Jaqué and Rio Imamadé, 730 m, KU 116779 (tadpoles), 116780-1 (young). Panamd: Altos de Pacora, 740 m, KU 95979. Hyla_ denticulenta—COLOMBIA: Antioquia: Valdivia, 1400 m, KU 133452. Santander: Charta, 2400 m, KU 133450-1, 139531 (tadpoles). Hyla phyllognatha— ECUADOR: Morona-Santiago: Macas, AMNH 33911. Napo: Bermejo No. 4 (oil well site), 15 km ENE Umbaqui, 720 m, KU 123130-2; Rio Azuela, 1740 m, KU 143199-204, 143543 (young), USNM 193349-50; Rio Salado, 1 km upstream from Rio Coca, 1410 m, KU 146794 (tadpoles); Salto de Agua, 2.5 km NNE Rio Reventador, 1660 m, KU 143205, 143544 (young), 146793 (tadpoles); San José, AMNH 22164; 16.5 km NNE Santa Rosa, 1700 m, KU 143197-8, 143542 (tadpoles). Pastaza: Mera, 1140 m, KU 121418 (tadpoles), 121419 (young); Puyo, 960 m, FMNH 172632; Rio Alpayacu, 1 km E Mera, 1100 m, KU 121040. Tungurahua: 18.5 km E Banos, 1600 m, KU 141596 (young). PERU: Junin: Perene, AMNH 17260, 17277. Puno: La Unidn, Rio Huacamayo, Carabaya, 610 m, BMNH 1907.5.7.36. San Martin: Roque, NHMG 471. Hyla platydactyla—VENEZUELA: Mérida: Camino de La Culata, UPR-M 2763, 2765; Carretera La Azulita, PUR-M 2762; Escorial, 2500+ m, FMNH 3567-8, NHMW 6397 (4), 6398, UMMZ 46465, 59016, 105439, UPR-M 3158-62; La Culata, 2700 m, AMNH 10636-40, BMNH_ 1905.5.31.83-87, NHMW 6396 (3), UPR-M 3150-2, 3932-5, 4330; La Mucuy, 2500 m, UPR-M 4321-2; Mérida, 1620 m, BMNH 1909.4.30.81-83, 1912.11.1.83, 1947.2.13.14, MCZ 2523; 15-20 km NW Mérida, road to La Azulita, 1700 m, KU 1334380-3, 139521 (tadpoles); 32 km NW Mérida, road to La Azulita, 2010 m, KU 139522 (tadpoles); Rio Albarregas, 2400 m, UPR-M 2767-70; Rio Chama, Mérida, 1620 m, UMMZ 95310 (tadpoles). Tachira: 15 km from Delicias, to Rubio, 1800 m, UPR-M 2249-50, 2766; Guacharaquita, near La Grita, UPR-M 4858. FROGS OF THE HYLA BOGOTENSIS GROUP 31 LITERATURE CITED BouULENGER, G. A. 1905. Descriptions of new tailless batrachians in the collection of the British Museum. Ann. Mag. Nat. Hist., (7)16:180-184. Cocuran, D. M. and C. J. Goin 1970. Frogs of Colombia. Bull. U.S. Natl. Mus., 288:xii+655 pp. Dixon, W. J. ( Ep.) 1971. BMD Biomedical computer programs. Univ. California Publ. Auto. Comp., 2, ed. 2:x+600 pp. DuUELLMAN, W. E. 1963. A review of the Middle American tree frogs of the genus Ptychohyla. Univ. Kansas Publ. Mus. Nat. Hist., 15:297-349. 1966. Taxonomic notes on some Mexican and Central American hylid frogs. Ibid., 17:263-279. 1970. The hylid frogs of Middle America. Monog. Mus. Nat. Hist. Univ. Kansas 1:xi+753 pp. 1972. South American frogs of the Hyla rostrata group (Amphibia, Anura, Hylidae). Zool. Meded. Leiden, 47:177-192. DvuELLMAN, W. E. and M. J. FouQuETTE, Jr. 1968. Middle American hylid frogs of the Hyla microcephala group. Univ. Kansas Publ. Mus. Nat. Hist., 17:517-577. DuELLMAN, W. E. and L. TRuEB 1966. Neotropical hylid frogs, genus Smilisca. Ibid., 17:281-375. Dunn, E. R. 1931. New frogs from Panama and Costa Rica. Occas. Papers Boston Soc. Nat. Hist., 5:385-401. 1944. Herpetology of the Bogota area. Rev. Acad. Colombiana Cien. Exact. Fisicas y Nat., 6:68-81. KuuceE, A. G. and J. S. Farris 1969. Quantitative phyletics and the evolution of Anurans. Syst. Zool., 18:1-32. MELIN, D. 1941. Contribution to the knowledge of Amphibia of South America. Goteborgs Kungl. Vetensk.-och Vitterh.-Sam. Handl., Ser. B, 1(4):1-71. PETERS, W. 1882a. Eine neue Gattung von Batrachiem, Hylonomus, aus Bogota. Sitzber. Gesell. Naturfors. Fr. Berlin, 1882(7):107-109. 1882b. Der Namen der Batrachiergattung Hylonomus in Hyloscirtus zu aindern und legte zwei neue Arten von Schlangen, Microsoma notatum und Liophis Ygraecum. Ibid., 1882(8):127-129. RivERo, J. A. 1961. Salientia of Venezuela. Bull. Mus. Comp. Zool., 126:1-207. Routr, F. J. and J. KisHpauGH 1966. NT-SYS, a system of multivariate statistical programs for use in numerical taxonomy. Dept. Entomology, Univ. Kansas. TayLor, Epwarp H. 1952. A review of the frogs and toads of Costa Rica. Univ. Kansas Sci. Bull., 35( 1) :577-942. VUILLERMIER, B. S. 1971. Pleistocene changes in the fauna and flora of South America. Sci- ence, 173( 3999 ):771-780. UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly pub- lished in the above series are now published as Occasional Papers, Museum of Natural History. The Miscellaneous Pub- lications, Museum of Natural History, began with number 1 in 1946. Longer research papers are published in that series. Monographs of the Museum of Natural History were initiated in 1970. Institutional libraries interested in exchanging publications may obtain the Occasional Papers and Miscellaneous Publica- tions by addressing the Exchange Librarian, University of Kan- sas Library, Lawrence, Kansas 66044. Individuals may pur- chase separate numbers of all series. Prices may be obtained upon request addressed to Publications Secretary, Museum of Natural History, University of Kansas, Lawrence, Kansas 66044. Editor: LinpA TRUEB Managing Editor: Wi.L1AM E. DuELLMAN PRINTED BY UNIVERSITY OF KANSAS PRINTING SERVICE LAWRENCE, KANSAS OCCASIONAL PAPERS of the Slacsncmunnaas anaes scare MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas NUMBER 12, PAGES 1-28 FEBRUARY 20, 1973 MORPHOMETRIC VARIATION AND LIFE HISTORY OF THE CYPRINID FISH NOTROPIS STRAMINEUS (COPE) By JUtwe Tanyorac' The sand shiner, Notropis stramineus (Cope) is a common in- habitant of shallow, sandy streams from Quebec and New York westward to Saskatchewan and Montana, and southward to México. Sand shiners occupy streams of diverse sizes, from springs discharg- ing only a few cubic feet per second to the largest rivers in the Mississippi system. As its vernacular name implies, N. stramineus is usually associated with sandy stream bottoms; the species has rarely been found in upland areas (Ouachita, Ozark, Cumberland, and Appalachian regions) within its otherwise extensive range. The purposes of this paper are: 1) to indicate the extent and degree of geographic correlation of variation in several morpho- logical features of N. stramineus, and 2) to describe some aspects of its life history, with primary reference to populations in the Kansas River at Lawrence, Kansas. Problems relating to the nomenclature of the sand shiner have been resolved by Suttkus (1958) and Bailey and Allum (1962:64- 68). Suttkus demonstrated that the name Moniana (= Notropis) deliciosa (Girard, 1856) is applicable not to the sand shiner but instead to the weed shiner, Notropis texanus (Girard, 1856). Suttkus concluded that the oldest name available for the sand shiner is 1 Department of Zoology, University of Ankara, Ankara, Turkey. This paper is based on research accomplished while the author was a Research Associate at the Museum of Natural History at the University of Kansas, supported by a Fellowship from the American Association of University Women and a grant from the University of Ankara. 2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Hybognathus stramineus Cope, 1864; he considered Minnilus micro- stomus Rafinesque, 1820, and Alburnus lineolatus Agassiz, 1863, to be nomina dubia. The following names were listed as chronological synonyms of stramineus by Suttkus (type localities in parentheses): Hybopsis missuriensis Cope ( Missouri River at St. Joseph, Missouri); H. scylla Cope (Red Cloud Creek, tributary to Platte River); Cliola chlora Jordan (upper Missouri region); Notropis phenacobius Forbes (Peoria, Illinois); and N. reticulatus Eigenmann and Eigen- mann (Qu’Appelle, Assiniboia [Assiniboine River in Saskatchewan, Canada] ). Bailey and Allum (1962:64-68) reported that sand shiners in South Dakota “fall into two sharply defined morphotypes” that they recognized as subspecies (N. s. stramineus and N. s. missuriensis), distinguished by differences in number of circumference scale-rows. Bailey and Allum allocated H. scylla, C. chlora, and Hybopsis mon- tana Meek, 1884, to the synonymy of N. s. missuriensis. Metcalf (1966:124-131) supported the opinion of Bailey and Allum, on the basis of his counts of circumferential and predorsal scale-rows. Pre- viously, Hubbs and Lagler (1958:37, 43) and several other authors had indicated the existence of a third subspecies, southwestern in occurrence. The southwestern populations are those with which the name deliciosa was improperly associated. GEOGRAPHIC VARIATION Methods In my analysis of geographic variation in the sand shiner, nine characters were examined on 896 specimens from 61 localities, fol- lowing procedures defined by Hubbs and Lagler (1958:19-26). Meristic characters used were number of lateral-line scales, pre- dorsal-scale rows, and circumference scale-rows. Morphometric characters used were predorsal length, length of caudal peduncle, head length, head width, length of orbit, and postorbital length of head, all measured to the nearest 0.1 mm and expressed as thou- sandths of standard length. Values obtained for each character were compared, among lo- calities, by application of a program entitled Sum of Squares Simul- taneous Test Procedures (SS-STP), adapted to an IBM 7090/94 computer. The analysis indicates which of the locality means, for each character, fail to differ from one another at the 95 percent confidence level. On the distribution maps that accompany the dis- cussion of each character, the results have been coded as follows: VARIATION AND LIFE HISTORY OF NOTROPIS STRAMINEUS' 3 solid circles indicate localities where mean values are not signif- icantly different from the lowest mean obtained at any locality; solid triangles indicate localities where means are not significantly differ- ent from the highest mean value obtained at any locality; and open circles designate the remaining localities (those having intermediate values significantly different from means of the extreme groups). The 61 localities include three in the Great Lakes drainage, 58 in the Mississippi basin, one in the Guadalupe River in Texas, and one in the Rio Grande system. All are within the presumed native range of the species. In the Mississippi basin, seven series are from the Ohio River system; 13 are from upper Mississippi system (upstream from the mouth of the Missouri River, excepting one collection from the Meramec River immediately south of the Missouri River); 26 are from the Missouri River system; six are from the Arkansas River system; and two are from the upper Red River system. Results Circumference scale-rows (Fig. 1).—The SS-STP results indicate that the distinctive sets of sample means fall into eastern and west- ern groups that correspond to the two subspecies (stramineus and missuriensis, respectively) recognized by Bailey and Allum (1962:64- 68), Metcalf (1966:124-131), and others; their interpretations were based primarily on this character. In my samples, the range of non- significant low mean numbers of circumference scales is 22.2-24.6, the range of non-significant high mean numbers 27.7-30.0. These two sets include 72 percent of the localities. Intermediate numbers of scale-rows around the body were found at 17 localities, in the Platte River system, in the Missouri River and its smaller tributaries from the mouth of the Platte eastward to the Chariton River in northern Missouri, in the upper Neosho River (Arkansas River drain- age), and in the Red River system in western Oklahoma. Postorbital length of head (Fig. 2).—This character differenti- ates approximately the same samples that were separated by circum- ference scale-counts (Fig. 1). Low mean values (0.1101-0.1202, as a proportion of standard length) are mainly eastern, and high mean values (0.1316-0.1401) mainly western. The two sets include 57 percent of the 56 samples in which data are available for this char- acter. Intermediate lengths of the postorbital part of the head are found mainly in the Platte and lower Missouri rivers, the upper Neosho River in Kansas, and the upper Red River. Intermediate values in the Rio Grande and in the Osage, Meramec, and Gasconade rivers of the Missouri system in Missouri are discordant with results t OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Circumference scale-rows A 2775-3007 O 2480-2770 @ 2220-2460 250 t5 50 100 as SCALE OF MILES Fic. 1. Geographic trends in numbers of circumference scale-rows in Notropis stramineus. Triangles represent sample-means not significantly differ- ent from the highest mean value at any locality (.05 level); solid circles indi- cate means not significantly different from the lowest mean value of any locality; open circles represent locality means intermediate between those ex- treme sets. based on circumference scales, in which these localities were a part of the eastern group. Length of orbit (Fig. 3)—Geographic variation in the orbital diameter of sand shiners is great, but individual variation within local populations is small. Therefore, the SS-STP discriminated numerous (31) combinations of localities having means not signif- icantly different—a greater number of small sets of localities than emerged for any other character. As a result, few localities are in- cluded in either the “low” or the “high” groups, whose ranges of values were 0.6210-0.6980 and 0.9088-1.0030, respectively. The mean orbital diameters at most localities lie between these extreme sets, in the group designated by open circles as “intermediate” in VARIATION AND LIFE HISTORY OF NOTROPIS STRAMINEUS 5 Postorbital length of head A |316- 140) O l203-'300 @ |I0\- 1202 105 ele) 15) 90 85 Fic. 2. Geographic trends in postorbital length of head in Notropis stramineus. Symbols as in figure 1. figure 3. The orbit is smallest in the westernmost populations, and largest (relative to standard length) in populations near the center of the range of the species (eastern Missouri and Iowa). The large- eyed populations fall within the “eastern” set as defined by scale counts. In terms of absolute values, most “intermediates” located northeastward from Iowa and Missouri have relatively large eyes, as do specimens from the Rio Grande and Guadalupe in Texas. It is noteworthy that the results using orbital diameter and postorbital length of head are not entirely complementary; populations having the largest orbits do not have minimal postorbital lengths. Predorsal scale-rows (Fig. 4)—The total extent of variation in number of predorsal scale-rows is small, relative to local (individual) variation in sand shiners. Therefore, the SS-STP did not yield dis- crete sets of localities having distinctively “low” and “high” mean values; the range of “low” means, not significantly different from 6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY \ — Length of orbit ) ae 7 A 2088-1003 Ae! re Y © 7005-8910 \ @ 6210-6980 25 50 i SCALE OF MILES 105 00 LS) 90 85 Fic. 3. Geographic trends in length of orbit in Notropis stramineus. Sym- bols as in figure 1. one another, extended from 13.86 to 15.13 scales, and the range of “high” means from 14.93 to 16.53 scales. Therefore, in figure 4, the localities plotted as having “intermediate” values (open circles) are ones that are a part of both sets, rather than ones excluded from both sets. These five localities are scattered across the range of the species, at about the same latitude. Values at other localities are geographically correlated, forming a pattern generally like that shown by the characters previously discussed. Two departures are notable: 1) greater westward extent of the “eastern” group into areas that yielded intermediate values for other characters; and 2) exceptionally high numbers of predorsal scales at the northernmost localities, causing their populations to be associated with the west- ern rather than the eastern group in this character. The mean num- bers of predorsal scale-rows at the three northeastern localities in question (Ohio, Michigan, and Minnesota) are 15.14, 15.80, and 15.20, respectively. VARIATION AND LIFE HISTORY OF NOTROPIS STRAMINEUS'- 7 Predorsal scale-rows A 1513(1493)-1653 O 14993-1513 ® |386-1490(513) = — SCALE OF MILES Fic. 4. Geographic trends in number of predorsal scale-rows in Notropis stramineus. Symbols as in figure 1. Predorsal length (Fig. 5).—Western populations of sand shiners have the shortest predorsal lengths, in relation to their standard lengths. However, the geographic pattern shown by this character does not conform well to an east-west clinal trend, because “high” and “low” values are found at adjacent localities in several parts of the range: upper Red River, Kansas River, lower Missouri River, upper Illinois River, and upper Ohio River. Predorsal length and the number of predorsal scale-rows are not positively correlated. Rather, the opposite tendency exists; those populations having the shortest bodies anterior to the dorsal fin usu- ally have the largest number of scales along the nape. Most such populations are western or northern. As in predorsal scale-counts, local variation is so great compared to inter-locality variation that no wholly distinctive sets of localities emerged from the SS-STP analysis. The “low” (0.5077-0.5201) and “high” (0.5201-0.5480) 8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Predorsal length A 52I|(520!)-5480 O 520! ® 5077-5193(520)) Fic. 5. Geographic trends in predorsal length in Notropis stramineus. Sym- bols as in figure 1. sets overlapped by one locality. That locality, in the Rio Grande drainage, is therefore the only one shown as “intermediate” by an open circle in figure 5. Head length (Fig. 6).—Although variation in head length has only limited relation to geographic region, most northern and east- ern populations have shorter heads than do southwestern popula- tions. The analysis separated localities into numerous (27) sets for this character; hence, the “high” and “low” sets contain only 23 localities, 41 percent of the total from which data were taken. The ranges of mean values within these two sets are well separated (low=0.2494-0.2596; high=0.2711-0.2857). All localities belonging to the low group lie along or east of the Mississippi River or in the Platte River system, with the exception of one locality in the Kansas River basin. All localities having high mean values lie west of the VARIATION AND LIFE HISTORY OF NOTROPIS STRAMINEUS- 9 Head length A 27\||-2857 O 2599-2709 @ 2494-2596 95 85 Fic. 6. Geographic variation in head length in Notropis stramineus. Sym- bols as in figure 1. Mississippi, except for one locality in the upper Ohio basin. Inter- mediate head-lengths are found at widely scattered localities. Head width (Fig. 7).—In general, northeastern sand shiners have relatively narrow heads, whereas southwestern sand shiners have broad heads. The distribution of high and low mean values for this character resembles that for head length, demonstrating that the total size of the head increases westward, and especially south- westward. At the 18 localities included in the “low” group in figure 7, mean head-widths varied from 0.1293 to 0.1439; mean head-widths at the 13 localities in the “high” group varied from 0.1556 to 0.1698. Length of caudal peduncle (Fig. 8).—Geographic pattern is scarcely evident in the distribution of distinctively high and low mean values for this character. The longest caudal peduncles tend to occur at high latitudes and high elevations, hence at peripheral localities. Caudal peduncles are usually shortest in the central, and 10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY \ Wee Head width Be wy, A |556-1698 : y. we yy 25 50 100 if i he J © 1446-1551 “SCALE OF MILES \ @ 1293-1439 on Bia N WA 105 100 95 90 85 Fic. 7. Geographic variation in head width in Notropis stramineus. Sym- bols as in figure 1. especially the west-central, part of the range of the sand shiner. Individual (local) variation accounts for a large part of the total variation in length of the caudal peduncle. The group of low means (0.2152-0.2348) includes 27 localities, and the group of high means (0.2360-0.2512) 21 localities; only seven localities (13%) fall into the intermediate group. Lateral-line scales (Fig. 9)—-Low numbers of lateral-line scales characterize sand shiners in the central and southern parts of the range of the species. Lateral scale-rows are more numerous at peripheral localities in the east, north, and west. Therefore, lateral- line scales vary independently of circumference scales and predorsal scales, and tend to be most numerous at high latitudes or high ele- vations. The group of low mean values (32.61-34.00) includes 25 localities, and the group of high values (34.53-36.25) includes 17 localities, leaving 18 localities (30%) in the intermediate group. VARIATION AND LIFE HISTORY OF NOTROPIS STRAMINEUS 1] Length of caudal peduncle A 2360-2512 | a Y © 2349-2359 nd SCALE OF MILES @ 2152-2348 4 105 100 95 30 85 | Fic. 8. Geographic variation in length of caudal peduncle in Notropis stramineus. Symbols as in figure 1. Character Summary and Taxonomic Conclusions The SS-STP analysis disclosed similar geographic trends in three of the nine characters. Circumference scale-rows, predorsal scale- rows, and postorbital length of head have extreme values in essen- tially the same sets of eastern and western populations. The popu- lations are segregated somewhat differently by the SS-STP results in the remaining six characters. Orbital size is smallest in the western- most populations, but largest near the center of the range of the species; populations having the largest orbits are among the eastern set distinguished by the three characters mentioned above. Predorsal length diminishes from east to west, but local variation in this char- acter is great and members of the two extreme sets are intermixed broadly through the central part of the range. Head width sorts, principally, southwestern populations from those elsewhere. Head length, caudal peduncle length, and the number of lateral line scales 12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Lateral-line scales A 3453-3625 O 3400-3450 @ 326I- 3400 95 Fic. 9. Geographic variation in lateral-line scales in Notropis stramineus. Symbols as in figure 1. vary without clear geographic pattern, although the highest values for the last two characters tend to occur at peripheral localities. In view of nonconformities in the variational patterns of these characters, recognition of subspecies may seem to be unwarranted in Notropis stramineus. However, I believe the results are sufh- ciently in accord with one another, in five traits having well-defined geographic patterns, to justify continued recognition of two taxa. Their distinguishing features, synonymies, and ranges are stated below. Notropis stramineus stramineus (Cope) Diagnosis.—Circumference scales 22-25; postorbital length of head 11-12 percent of standard length; orbital diameter more than 8 percent of standard length; head width less than 15 percent of standard length; predorsal scale-rows usually 13-15. VARIATION AND LIFE HISTORY OF NOTROPIS STRAMINEUS 13 Range.—Southern Saskatchewan to Quebec (Slastenenko, 1958), southward through Great Lakes drainage and Ohio River basin; upper Mississippi basin westward to eastern Iowa, southern Missouri and southeastern Kansas (including Osage River system); Red River in eastern Oklahoma and Texas; southwestward in Gulf-coastal streams to Rio Grande. Remarks.—On geographic grounds, the names N. phenacobius Forbes, 1885, and N. 2 *ticulatus Eigenmann and Eigenmann, 1893, apply to this subspecies. Notropis stramineus missuriensis (Cope) Diagnosis.—Circumference scales usually 27-30; postorbital length of head 13-14 percent of standard length; orbital diameter less than 8 percent of standard length; head width more than 14 per- cent of standard length; predorsal scale rows 15-17. Range.—Missouri and Arkansas River systems; intergrades with N. s. stramineus in eastern South Dakota and Nebraska (including Platte River system), small tributaries of Missouri River in north- eastern Kansas and northwestern Missouri, Neosho River in Kansas and upper Red River system in western Oklahoma. Remarks.—On geographic grounds, the names Hybopsis scylla Cope, 1870, Cliola chlora Jordan, 1878, and (according to Bailey and Allum, 1962:68) Hybopsis montanus Meek, 1884, apply to the subspecies missuriensis. LIFE HISTORY A study of some aspects of the life history of the sand shiner in the Kansas River at Lawrence was based on 3632 specimens, col- lected biweekly or monthly in 1967 (March 24, May 2, June 8 and 24, July 6, 13, and 24, August 9, September 1, and October 18). The study area extended along the north shore of the Kansas River for a distance of several hundred yards downstream from a low-flow dam in Lawrence. The Kansas River is a turbid stream having variable flow. At the gauging station nearest to my study area, discharge has varied from 185 cubic feet per second (cfs) to 483,000 cfs within the 30-year period of record. In 1967, minimum and maximum rates of dis- charge were 220 and 88,900 cfs. Flow on dates when sand shiners were collected increased from 468 cfs (March 24) to 41,700 cfs (June 24), and diminished to approximately 2,000 cfs in September. Sand shiners were collected by means of seines 6 to 15 feet long, 14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY having 1/8 to 3/16-inch woven nylon mesh. The fish were preserved in 10 percent formalin for three or more days, washed in water, and transferred to 70 percent ethyl alcohol, where they remained for at least one week before data were taken from them. The accounts of age- and size-composition and of reproduction that follow are based mainly on the collections from the Kansas River at Lawrence; observations on other populations are appended to these accounts. The latter observations are based on series of specimens deposited in the University of Kansas Museum of Natural History. The series selected are representative of the western sub- species, commonly recorded as N. s. missuriensis (from South Da- kota, Wyoming, Colorado, Nebraska, Kansas, and New Mexico), and eastern subspecies, commonly recorded as N. s. stramineus (from Kentucky, Wisconsin, Minnesota, Illinois, Iowa, Missouri, Kansas, Oklahoma [Red River], Texas, and Mexico). Series of “in- tergrades” from eastern Kansas, southeastern Nebraska, southwest- ern Iowa, and Missouri were examined also. These collections were obtained in spring or summer in various years. Age- and Size-composition Most samples of sand shiners from the Kansas River consisted of three age-groups, based on evidence obtained from opercles, scales, and length-frequency distributions. For 233 sand shiners of selected sizes, opercles and scales were removed, cleaned, and examined under magnification using transmitted light. Annuli were discernible on both structures, but usually were most obvious on the opercle, where they appeared as fine opaque lines parallel to the posterior margin of the bone. The 1967-annulus had not formed on fish taken March 24, and only a few yearlings caught on May 2 had completed that annulus. All age-I fish had an annulus by June 8, but the 1967 year-mark was not apparent on all age-II and age-III specimens until mid-July. Thus, prior to July 24, ages of individual fish were inferred from the positions of annuli (extent of growth beyond the outermost annulus), as well as the number of annuli observed on their opercles and scales. The age- and size-distributions of the 10 samples of sand shiners obtained from March 24 to October 18 are shown in figure 10. The number of fish in the samples varies greatly; on figure 10, differences in the numerical scale (vertical scale) on various dates should be noted. Age-III fish were present, but in diminishing numbers, from March 24 through July 6; none was found thereafter. Age-II fish were numerous on March 24 and May 2, but diminished in abun- VARIATION AND LIFE HISTORY OF NOTROPIS STRAMINEUS 15 ae [tena luse tl, alee Seeale — 1 =| isi0 , . _ | \ i Ae 4@ ‘ = | (2) 13 July \ 5 ‘i eal ae Ao 4a 40+ ye : me) Y; ae | ee ae eae 7 w I a “| = gop . | ad f . | 2 i = 4 40-r \ aoe) | 5 R =I ie tod \ a = ie (4 y, 2007 | 0) = . 8 9 August 4 a i 80- ts 100 el | f hve Il e = \e (1) 40 : \ | Zz © 8 June | Me 0 ~~ oH (2)-* | September ° | 40; | 20r | a T ~ (© parleuen ean 12 18 24 30 36 42 48 l2 18 STANDARD LENGTH IN MM Fic. 10. Length-frequency distribution in samples of Notropis stramineus from the Kansas River at Lawrence, obtained on various dates in 1967. Roman numerals indicate the age-groups represented in the samples, arabic numerals the number of individuals assigned to each age-group. dance later. Age-I fish dominated all samples from June through October. Nearly all growth in length by age-groups I and II was accom- plished before July 13. From that date onward, the size-range and average size of the fish in my samples remained almost constant, except for the sample obtained September 1 (Fig. 10). That sample consisted of surprisingly small fish judged to be age-I by examina- tion of opercles and scales. One possible reason for the disparate size-range in the September sample can be suggested. Deep, swift flow at the study-site made seining unusually difficult, and all speci- mens were caught from one small pool, only a few inches deep, that extended into a slight depression on a large sandbar. On other dates samples were taken from more extensive areas, and greater range of depths, in the mainstream as well as in pools along sandbars. 16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY The size-ranges of different age-groups overlapped in all sam- ples. The smallest age-III fish found was 38 mm in standard length (March 24), the largest 47 mm (July 6). The size-range in age-II was 21 mm (March 24) to 45 mm (October 18), and of age-I fish 15 mm (March 24) to 43 mm (October 18). Young-of-the-year first appeared in my collections on July 13, following disappearance of age-III fish from the samples. Special efforts to obtain young, using 1/8 inch-mesh netting as well as 3/16 inch-mesh seines, were made only on July 24 and October 18. Young as small as 12 mm in standard length following preservation were present on both of these dates. Populations of N. s. missuriensis and N. s. stramineus from other localities in Kansas and Missouri consisted of age-III and younger fish. Three-year-old sand shiners occurred as late as August, but no four-year-old fish were found among specimens from these states. The largest specimen of N. s. missuriensis in collections from 30 localities is 64.2 mm in standard length, and was obtained in the Cache la Poudre River, Colorado, in September. The largest N. s. stramineus in collections from 52 localities is 61.5 mm long; it was obtained in Black Earth Creek, Wisconsin, in June. No differences in age-composition, average size, or maximum size distinguish the two subspecies. In both, northern popula- tions seem larger in average size (standard length) than southern populations. Reproduction Sex-ratio—Male and female gonads are distinguishable by gross inspection in sand shiners more than 25 mm in standard length. For this and later analyses, the fish in each sample were grouped by 3- mm intervals of length; my discussion concerns specimens larger than 26 mm (groups 27-30 mm and longer). Sex was ascertained for 2101 sand shiners from the Kansas River, of which 1128 (53%) were males (Table 1). Males were more numerous than females in all samples except those on July 13 and 24—dates on which I think spawning was in progress. On some other dates, females out- numbered males in the smaller size-groups (Table 1), indicating that the average size attained by males slightly exceeded that of females. Nevertheless, the maximum size attained by females (46.0 mm on July 13) was as great as that of males (46.6 mm on August 9) in my samples from the Kansas River. In N. s. missuriensis from 22 other localities, males were more numerous than females in 13 cases. Males outnumbered females in 17 of 26 collections of N. s. stramineus for which sexes were deter- ly US " 4 VARIATION AND LIFE HISTORY OF NOTROPIS STRAMINE TOTS €16 S8cl G 8 LT 8h = SOT OLT LOE LOV s9 OL S dali [210.1 Ol SIT 66 T 98 68 63 OL ee le 6h 0c, oc Seke = SI Se 0b e 11 ESOP eT SG Th mcmccr LS o “ee, © ‘POST ‘ides T OCE LVI €< ae! LS ae LG Go 9€ OS 67 oo 08 ra] tol 5. OY JsNnsny 6 60T GCE 9 Lr OLI SST ie ote gE iste oe oe ly ees CL Rel Gio) eee yl Clee Gy 1S oo Sl 9S BF OCG. | Si a6 Bea Ee @ Apf pe = Ayn €T GGG OCI CET — Guy) 61 86 O€ VE Oe, Tk GE SG & °° Ajn{ 9 0G Sac Ge) ssf i! Gel teal oun{ FZ CCV GLI [SG ilsay al. OT LG vy 18 LIT eT BN oun[ g SIT cg 699 Lee & © OT GG 86 VG VG 8 ABN G YARIN FZ “LOGT “ST 20q0290O Ysnoriyy FZ yore WLo1F QOUIIME'T JB IOATY sesuey oY} UI poure}qoO sisuatunssim snauiudiys sido1joN JO saydures Ul O1WeI-xeg “[ ATAV];, lope s a STIV.LOL 9ST LOT 5 me ere SP - SP ae Ca ae CP - eH A see, SP - 6S Ole! Ol eames - 66 - 98 CR amma 96 - €€ OOn 28° Ef - 08 me ey cee Gr ic 6 P (uur) ySuUET piepurys 18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY mined. The largest individuals were males in 17 of 22 series of missuriensis and in 19 of 26 series of stramineus. Reproductive period—In males, readiness for spawning was in- ferred from the presence of nuptial tubercles on the head and pec- toral fins. No males smaller than 28 mm in standard length bore tubercles in the Kansas River population. Most males larger than 32 mm were tuberculate in all samples from May 2 through September 1. Most males smaller than 32 mm were tuberculate only in samples taken from July 13 through September 1. No tuberculate males were found on March 24 or October 18. In females, readiness for reproduction was inferred from the size (weight) of the ovaries, and the size and number of eggs therein. The weight of ovaries in individual fish, and the average diameter of the ten largest ova (or oocytes) in the same fish, are shown for various dates in figures 11 and 12. Ovaries and eggs were uniformly small in March and May, but they enlarged rapidly thereafter. I doubt that spawning occurred before June 24, when ovary-weights and egg-sizes in the largest fish still were maximal. Most fish still had not spawned by July 6, although some must have done so, judging from reduced weights of ovaries in five relatively large fish (Fig. 11). By July 13, most fish larger than 36 mm in standard length had spawned, as indicated by the smaller size of the ovaries and eggs in these large fish on that date, as compared with earlier dates (Fig. 12). In females smaller than 36 mm, enlargement of ovaries and eggs continued through July and August. I think many of these small fish spawned in late summer. Therefore, I conclude that the spawning season extended from early July until September, but that progressively smaller fish comprised the spawning popula- tion after July 6. Age-II fish contributed importantly to the spawn- ing population through mid-July. After July 24, only one age-II female (a spent fish) occurred in my samples. Probably, the entire spawning population in the latter part of the season consisted of yearlings. At localities other than the Kansas River at Lawrence, periods of sexual development and reproductive activity are as follows: Males of N. s. missuriensis usually are tuberculate from April to September. All samples taken in May, June, and July from South Dakota, Wyoming, Nebraska, and Kansas had tuberculate speci- mens. Samples taken in August and early September from Wyoming, Colorado, Nebraska, Kansas, and New Mexico still contained some tuberculate specimens. The smallest tuberculate male of N. s. mis- suriensis, 26.1 mm in standard length, was found on June 12 in the Medicine River, a tributary of the Arkansas River in Kansas. VARIATION AND LIFE HISTORY OF NOTROPIS STRAMINEUS 19 e24 March o2 May Itt tr at tt 050 O 2 fo@ © ot00 8 Boron, Of © y e8 June °24June 0° c .2OO ° esc a) Cae) SS a) 013 July WEIGHT OF GONADS IN GRAMS foe) e D6 OY @® % e I 24 26 28 SOs oe 34 36 38 40 42 44 46 STANDARD LENGTH IN MM. Fic. 11. Gonosomatic index (ovary weight/total fish weight) in female Notropis stramineus obtained from the Kansas River at Lawrence in 1967, as a function of date of capture and standard length of specimens. Males of N. s. stramineus are tuberculate from April (Oklahoma and Texas) to July (Illinois and Wisconsin) and August (Lowa, Kansas, Missouri). Specimens taken on August 25 (Minnesota) and September 1 (Missouri) had no tubercles. In populations regarded as intergrades (eastern Kansas, south- eastern Nebraska, southwestern Iowa, and northern Missouri), tu- berculate males were found from May through August. Females of N. s. missuriensis usually had maximal ovary weights in June. However, a population from Gypsum Creek, McPherson Co., Kansas, had large gonads at the end of April; also, the smallest mature female, 24.5 mm in standard length, with large eggs was found in that collection. The largest individuals in a sample from Spring Creek, Smith Co., Kansas, seemingly had spawned by June 24; smaller fish probably had not spawned by that date. In the Arkansas system in southwestern Kansas, most sand shiners may complete spawning in June, but a population from Crooked Creek, Meade Co., had ovaries heavier than 0.200 g on July 4. No females 20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY ©24 March °2 May 024 June 013 July DIAMETER OF EGGS 25 on 26 28. 30 32 34 36 38 40 425744 STANDARD LENGTH IN MM. Fic. 12. Size of ova in female Notropis stramineus from the Kansas River at Lawrence, 1967. Vertical scale in mm. Each point plotted represents the aver- age diameter of 10 of the largest eggs found in an individual fish. had ova exceeding 0.79 mm diameter later than August 29 in Kansas. One specimen from Wyoming, 46.0 mm in standard length, 24 June, had not spawned and had the largest ovaries (0.735 g) of any fish examined in my study. In a separate study of sand shiners in the Smoky Hill River in western Kansas, Summerfelt and Minckley (1969:447) reported that “Spawning probably occurred several times between June and August, but apparently most fish spawned in late July and August in both 1965 and 1966.” Those authors noted a striking year-round correlation between water temperature and egg size; mid-summer temperatures were between 20°C and 25°C in 1965 and 27-33°C in 1966 when eggs were largest. VARIATION AND LIFE HISTORY OF NOTROPIS STRAMINEUS 21 Females of N. s. stramineus in series that I examined had maxi- mal ovary weights in May (Kansas and Missouri) to July (Michigan and Wisconsin). A population from Middle Creek, Miami Co.. Kansas, probably had completed spawning by June 28, while one from Floyds Fork Creek, Jefferson Co., Kentucky, and one from the Loutre R., Montgomery Co., Missouri, had ovaries heavier than 0.200 g on June 2 and 6. None had ova exceeding 0.79 mm in diam- eter later than August 30 in Missouri. Ovary weight and maximum egg-size (0.95 mm) in stramineus are about the same as in mis- suriensis. Females in series identified as intergrades had maximal ovary weights and egg-size in May, as in the case of N. s. stramineus from the same latitude. The smallest female, 29.6 mm, with large eggs was found in May (North Fork Cottonwood River, Marion Co., Kansas. ) Seemingly, the spawning period for N. s. stramineus varies lati- tudinally (May in South to July in North), but is mainly earlier and shorter in duration than the spawning season of N. s. missuriensis in Kansas and Missouri. Fecundity—Ovaries were categorized as 1) undeveloped—gray ovaries containing nucleated cells (oocytes) and small eggs (ova) with diameters not more than 0.33 mm; 2) developed—yellow ova- ries having eggs from 0.33 mm to 0.95 mm in diameter; 3) spent— gray ovaries flaccid and containing few eggs. Fecundity was determined from females that appeared to be well developed but not to have spawned. Ordinarily eggs were counted in one of the ovaries, because the two lobes appeared to be the same size; this number was doubled to determine the total egg complement. When the two lobes appeared to differ in size, the eggs were counted in both lobes. The “diameter of ova,” as shown in table 2, represents the average diameter of the ten largest eggs in each fish; egg-diameters were determined by means of an ocular micrometer at constant magnification, and converted to mm by cali- bration to a stage micrometer. All contained eggs of varied sizes during the spawning season. Egg diameters of mature fish as given by Summerfelt and Minckley (1969:447) are smaller than those I report, because their measurements were taken from the total egg complement of each ovary. Females of N. s. missuriensis from the Kansas River had maximal egg counts from June 8 to July 6, as shown in table 2. Seven females, from 32.8 mm to 43.9 mm in standard length, contained 550 to 1533 eggs. OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 22 0996 160 a> cco'0 i an?) V LV (ie a Fe ae ee oe CS Avy OT 9ESS L8°0 c+ ES'0 mites 6 FP Dh et ie ie - Secale ABW OL FEET 6L'0 i 910 ae Ser ie sa a ee. AVN OT 9T6I £60 ae 862'0 a= V Gy eS a aay ie Rees AR OL PIES 160 _— S660 mak: STF Her. Ea ee ae ABW OT 9LOI €3°0 ai SIT‘0 Pat €'8E ie ws ee eunf, 9 9¢9 £8°0 if €L0°0 esd v'SE Ls a ee eunf 9g SNAUNUDLIS SNAUNUDAYS 810.140 NJ 80S@ S6'0 =: SOF‘0 a 6°6F 1) er ere ee cone 06E% S60 a3 E8€'0 = cP te a eo eunl 6 G9TG L8°0 on S860 ep SV eS. Sate ae ae eounf 6 cesT 160 VT 0 LEGO 8c9'T 6'€P a coe ounl 7% SETT S6'0 €1T0 LG6G0 trol TOP ae ae a ae eunf 7% IOFT 160 910 GGG 0 Ofe'T V'6E ti ay ey ee ae Ajnf g S9TT 160 eT'0 SLT'0 6L¢ 1 9°8E Lm en wy te A oe Anf 9 LI8 L8°0 9T 0 VST 0 6760 OLE ees) ae. Ajnf 9 VLE L8°0 IT0 OOT'O 663°0 Go SE ep ot ae ve ee ounl 7% OSS £8°0 O10 6L0°0 tTL'0 SCE a> oo tee. 2 oe ounf 7% SISUALINSSHUL SNAUIULD.LYS 10140 NJ BAG jo (ulul) 8AG xopuy (3) speuoy (3) YSsty (uu) YsueT os8y 9}eq Jaquinn ysos1e'T oryeulosouos) JO JYSIOM, JO JYSIOA, prepur}s LIMOsstjJ Ul Ulseq OURS OY} WOLF snauUDsys snauUDAys s1d0140N UdAeS pue sesuey Ul UISeg IOATY LMOssIpyY 94} UWlOIZ sisuatinssi snaunuDijs sido1jJON JO SaTVUIAZ QT OF vJep pozjer pue Aypunoay °*% aTavy VARIATION AND LIFE HISTORY OF NOTROPIS STRAMINEUS 23 Fecundity ranged from 2162 to 2508 eggs for three N. s. mis- suriensis females from North Fork Prairie Dog Creek, Decatur Co., Kansas; their lengths were 41.5 to 49.9 mm (Table 2). N. s. stramineus females from the Loutre and the Pomme de Terre rivers, Missouri, 35.4 mm to 47.4 mm in standard length, had 626 to 2660 eggs (Table 2). The range in egg-complement observed in N. stramineus was 550-2660. The results indicate considerable individual variation in fecun- dity but little or no difference of fecundity between the subspecies. Fecundity increased with standard length (or age) of the fish. Discussion of Life-cycle The population of sand shiners in the Kansas river, followed through the spring and summer of 1967, consisted of three age- groups. Many fish two or three years old were found in spring, but these became scarce in late summer, when the population was domi- nated by yearlings. After June 24 there was little or no increase in the modal length of individuals representing age-groups I and II. Modal lengths of each of these age-groups on June 24 exceeded the modal length of the next older age-group at the beginning of the growing season (24 March and 2 May). Either growth ceased by the end of June or the larger fish in both age-groups were depleted by mortality thereafter. Age-III fish disappeared from samples in early July. Attainment of maximal size of each age-group coincided with the onset of reproduction. Thus, the life cycle of this popu- lation of N. s. missuriensis can be summarized as shown in table 3. I think that growth by older individuals occurs in spring, but terminates as these fish reproduce and disappear in July. Growth by small yearlings continues until they too spawn, later in summer. Many of them may fail to spawn, but survive to become spawners (the two-year-old component) in the following year. This cycle has some attributes of an “annual” life-cycle, modified to provide a protracted reproductive period and consequent vari- ation in size and age of individuals belonging to the same year-class. That variation tends to protract the pericd during which maturity is attained by different individuals, and thereby sustains a prolonged reproductive season. Such a cycle may have survival advantage in streams with erratic variation in flow. Abrupt changes in waterlevel are unlikely to destroy the entire year’s production, regardless of when the changes occur. Yet the longevity of the sand shiner is short, enabling a rate of turnover in the population only slightly less rapid than that characterizing annual life cycles. 24 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY TaBLE 3. Mean standard lengths (mm) of sand shiners from the Kansas River on various dates in 1967. Age-groups Date I II Ill DAP Nianche aes tie ret: 3878 26 a2 40 Da NMayP leet tsetse ah aAccundy 26 32 40 | Period of growth: SUE. ee ere ed oe 29 38 41 | increasing biomass aa Ue ees Sn 30 42, —— | age-groups I-III Gey Cena Lh ai 35 42 45 RS Aly yee te eae ea 35 44 —— | Period of repro- sf Se, ae 32 4] —— | duction: decreasing CONGR Fa eee 33 45 —~ | biomass age-groups iG ees ao 96° Be 2 Sie SCO CES. 8 re ee 38 42, a I am uncertain whether the hypothesis above can be extended to include N. s. stramineus. There is some evidence that the period of spawning by the eastern subspecies is brief at specific localities, compared to that of N. s. missuwriensis. Maturation occurs earlier in N. s. stramineus than in N. s. missuriensis in Missouri and Kansas. Northern populations of N. s. stramineus mature and spawn later than do southern populations of that subspecies, but latitudinal variation in the spawning time of N. s. missuriensis seemed slight in the samples I examined. SUMMARY Although geographic agreement among characters is imperfect, the sand shiner, Notropis stramineus, seems logically divisible into eastern and western subspecies (N. s. stramineus and N. s. missurien- sis, respectively). The range of the nominate subspecies extends from the Rio Grande and Guadalupe rivers northeastward across the lower Red River and the southernmost tributaries of the Missouri River to the Mississippi River, eastward and northward to the head- waters of the Ohio River basin and the Great Lakes basin. The western subspecies occurs in rivers of the Arkansas and Missouri systems that extend westward across the plains to the slopes of the Rocky Mountains. Intergradation occurs in the Platte and lower Missouri river systems, the upper Neosho River of the Arkansas system, and the upper Red River. The two subspecies do not differ demonstrably in size attained (ca. 60 mm), age attained (3 years), total reproductive period (May or June through August), or egg-complement (range 550-2660). At single localities, however, the reproductive period probably is longer VARIATION AND LIFE HISTORY OF NOTROPIS STRAMINEUS 25 in N. s. missuriensis than in N. s. stramineus, and latitudinal vari- ation in the time of spawning is more pronounced in N. s. stra- mineus. At similar latitude in Kansas and Missouri, females of N. s. stramineus mature earlier (May) than females of N. s. missuriensis (June-July ). A population of western sand shiners from the Kansas River at Lawrence contained both yearling and older individuals in March, 1967, but most two- and three-year-old fish disappeared before autumn. Growth in length by the larger fish was nearly confined to spring, terminating when they spawned early in July. Small year- lings matured later in summer. Males seemed slightly more numer- ous and slightly larger than females, and appeared to be sexually active for a longer term in spring and summer. Most individuals larger than 28 mm (standard length of alcoholic specimens) were mature. ACKNOWLEDGMENTS I thank Dr. Frank B. Cross of the Museum of Natural History, University of Kansas, for his suggestions and advice throughout the course of my work, for use of specimens and facilities in that mu- seum, and especially for assistance in preparation of this manuscript. I am grateful to several graduate students in ichthyology at the museum for obtaining sand shiners from the Kansas River in 1967. Mr. T. H. Swearingen prepared final drawings of the graphs and maps. Computer time was made available by the Computation Center of the University of Kansas; Dr. A. C. Echternacht and Miss Linda Flatten aided me greatly in obtaining the SS-STP analysis of my data. My study was partly supported by a grant from the fellow- ship fund of the American Association of University Women, Educa- cational Foundation. SPECIMENS EXAMINED Forty-eight of 61 collections examined in this study are deposited in the Museum of Natural History, University of Kansas (KU), 6 in the United States National Museum (USNM), 2 in the University of Michigan Museum of Zoology (UMMZ), and 5 were borrowed from the Field Museum of Natural History (FMNH), Chicago. Some data from collections in USNM and UMMZ were obtained from an unpublished manuscript by Bernard C. Nelson. Specimens from USNM were measured and counted by Dr. Frank B. Cross of the University of Kansas in the United States National Museum. 26 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Specimens examined are listed below, together with museum catalogue number, number of individuals examined (in parentheses), range of standard length (mm), locality of capture, and year of collection. Collections are arranged by State, alphabetically; series from México (Rio Grandé drainage) conclude the list. COLORADO: KU 4751 (20), 36.5-48.5 mm, Pueblo Co.: Arkansas R. at Pueblo, 1959; KU 4797 (20), 48.3-56.4 mm, Weld Co.: South Fork Platte R., 7 mi E Greeley, 1959; KU 5566 (20), 47.2-64.2 mm, Larimer Co.: Cache la Poudre R. 2 mi S, 4 mi E Ft. Collins, 1960. ILLINOIS: FMNH 43398 (15), 42.2-54.7 mm, McHenry Co.: Conf. Rush Cr. and Kishwaukee R., 2 mi W Marengo, 1941; FMNH 60754 (15), Will Co.: N branch Hickory Cr., 2 mi NE Frankfort, 1950; FMNH 90949 (15), 41.5-54.0 mm, Will Co.: Marley Cr., below trestle at 108th Rd. above Alpine, 1954; FMNH 61053 (15), 43.5-48.5 mm, Lake Co.: Des Plaines R. 1/2 mi upstream from Deerfield Rd., 1953; FMNH 61241 (15), 42.9-55.6 mm, Kankakee Co.: Kankakee R. 4 mi downstream from Kankakee, 1955. INDIANA: UMMZ 106607 (15), 40.0-51.8 mm, Wayne Co.: E Fork Whitewater R.; USNM 36499 (15), 43.8-50.4 mm, White River, Gosport. IOWA: KU 8629 (15), 38.7-46.9 mm, Buchanan Co.: Wapsipinicon R. at Park 4.5 mi N Jesup, 1964; KU 8669 (15), 40.4-55.9 mm, Montgomery Co., Little Tarkio Cr., 1964. KANSAS: KU 3943 (15), 40.6-49.0 mm, Ford Co.: Arkansas R., 1958; KU 3973 (15), 38.0-48.5 mm, Rush Co.: Walnut Cr., Sec. 27, T18S, R18W, 1958; KU 4058 (20), 36.9-49.9 mm, Smith Co.: Spring Cr., Sec. 10, T4S, R13W, 1958; KU 4700 (20), 41.2-50.0 mm, Miami Co.: Little Wea Cr., Sec. 10, T17S, R24E (Kay Ranch), 1959; KU 7316 (20), 35.9-51.1 mm, Jefferson Co.: Honey Cr., Sec. 28, T9S, RI9E, 1963; KU 8140 (15), 39.5-46.3 mm, Marion Co.: Cottonwood R., 1964; KU 8586 (15), 37.5-44.3 mm, Comanche Co.: Salt Fork Arkansas R., 1964. KENTUCKY: KU 4083 (15), 41.5-49.5 mm, Jefferson Co.: Floyds Fork Cr. 1 1/2 mi NE Wood, 1858. MICHIGAN: UMMZ 110283 (15), 41.0-49.2 mm, Mason Co.: Bass Lake outlet, 1932; USNM 193163 (10), 39.9-47.5 mm, Antrim-Kalkoska Co’s.: Torch Lake, 1952. MINNESOTA: KU 3422 (5), 31.2-43.6 mm, Jackson Co.: Des Moines R., 1954. MISSOURI: KU 5162 (15), 37.4-48.4 mm, Osage Co.: Gasconade R. on Mo. 89, Sec. 16, T42N, RSW, 1960; KU 7815 (20), 39.5-48.4 mm, Hickory Co.: Pomme de Terre R. below dam at Pomme de Terre Reservoir, 1963; KU 7962 (10), 36.1-38.8 mm, Lincoln Co.: Cuivre R. 2.5 mi N Troy, 1962.00 9134 (10), 40.8-45.3 mm, Marion Co.: South R. 3 mi S Palmyra, 1964; KU 9190 (10), 35.0-43.8 mm, Marion Co.: South Fabius R. 7 1/4 mi NW Palmyra, 1964; KU 9625 (7), 34.4-43.1 mm, Gasconade Co.: Missouri R. at Gasconade, 1963; KU 9764 (15), Atchison Co.: Missouri R. at mouth Nishnabotna R., 1963; KU 9774 (15), 26.9-34.9 mm, Buchanan Co.: Missouri R. at St. Joseph, 1963; KU 9788 (15), 27.6-34.8 mm, Lafayette Co.: Missouri R. at Lexington, 1963; KU 10054 (10), Marion Co.: Mississippi R. near West Quincy, 1963; KU 10601 (15), 38.5-46.1 mm, Franklin Co.: Meramec R., 1963; KU 10638 (9) 41.3-50.6 mm, Phelps Co.: Dry Fork Cr., 7.5 mi SE Rolla, 1963; KU 10994 (10), 39.0-45.0 mm, Adair Co.: stream below lake 5 mi W Kirksville, 1963; KU 11058 (15), 37.4-45.9 mm, Chariton Co.: Mussel Fork Chariton R. 6 mi N Keytesville, 1963; KU 11074 (15), 39.9-53.9 mm, Atchison Co.: Tarkio Cr. 4 mi S Tarkio, 1963. NEBRASKA: KU 4829 (20), 46.0-60.2 mm, Banner Co.: Pumpkin Cr. on VARIATION AND LIFE HISTORY OF NOTROPIS STRAMINEUS 27 Nebr. 29, 1959; KU 4847 (20), 48.9-55.4 mm, Keith Co.: North Platte R. at Keystone, 1959; KU 7043 (20), 41.0-54.0 mm, Franklin Co.: Thompson Cr., Sec. 28, TZN, RI3W, 1961; KU 7100 (20), 42.0-52.4 mm, Cherry Co.: North Minnechaduza Cr. 4 mi E Valentine, 1961; KU 7130 (15), 44.3-50.3 mm, Dundy Co.: South Republican R 1/2 mi N Kan./Nebr. border on Nebr. 61, 1961; KU 7148 (20), 40.8-48.5 mm, Otea Co.: trib. North Fk. Little Nemaha R. 4.8 mi S Jct. Nebr. 50 and US 34, 1961; KU 7157 (20), 38.4-56.8 mm, Dixon/Cedar Co’s.: 1/2 mi W Concord, 1961; KU 8228 (20), 35.5-47.1 mm, Adams Co.: Little Blue R. 3.6 mi S Roseland, 1961. NEW MEXICO: KU 4231 (17), 42.0-48.4 mm, Mora Co.: Mora R. 1 mi SE Valmora, 1957. OHIO: KU 2790 (7), 31.8-46.8 mm, Ashtabula Co.: Grand R. at Mechan- icsville, 1944-1945; KU 11378 (5), 36.7-43.6 mm, Pickaway Co.: Big Daddy Dr., 1966. OKLAHOMA: KU 8054 (20), 37.4-49.1 mm, Beckham Co.: Timber Cr. 3.2 mi E Sayre, 1964; KU 8098 (15), 41.0-46.2 mm, Beckham Co.: N Fk. Red River at Sayre, 1964. SOUTH DAKOTA: KU 10175 (20), 39.5-46.5 mm, Harding Co.: Little Missouri R. 4 mi S, 7 mi W Ladner, 1961. TENNESSEE: USNM 190885 (15), 39.3-45.3 mm, Sevier Co.: Waldens Cr., T59-19 on Sevierville-Wear Valley Road, 2 1/2 mi S of US 441, 1959. TEXAS: KU 5996 (8), 32.5-40.9 mm, Hays Co.: Blanco R. at Wimberley on T-12, 1961. WEST VIRGINIA: USNM uncatalogued (8), 38.2-44.7 mm, Price Hill, Pond Fork, Rd. 23-55, 1955; USNM uncatalogued (10), 39.8-49.5 mm, Trace Fk. Mud R. 200 yds above mouth (Coal R. System), 1956; USNM uncatalogued (7), Low Gap, Swinging Bridge and RR trestle section, Rd. 19-56, 1956. WISCONSIN: KU 4923 (10), 43.5-52.6 mm, Polk Co.: St. Croix R., 1959; KU 7276 (8), 50.0-61.5 mm, Dane Co.: Black Earth Cr. 1 mi SE Mazomanie, 1962. MExIco COAHUILA: KU 2982 (3), 40.2-44.8 mm, unnamed river 1 mi NW EI- moral, 1952; KU 3006 (3), 28.9-38.5 mm, 2 mi W Jiménez, 1952; KU 3020 (4) and KU 3034 (8), 36.0-41.7 mm, 2 mi W Jiménez, Rio San Diego, 1952. 28 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY LITERATURES CITED BaILey, R. M., AND ALLUM, M. O. 1962. Fishes of South Dakota. Univ. Michigan Mus. Zool., Misc. Publ. 119:65-68. June 5. Husss, C. L., AND LAGLER, K. F. 1958. Fishes of the Great Lakes Region (revised edition). Cranbrook Inst. Sci., Bull. 26. xiii + 213 pp., 44 pls. METCALF, A. L. 1966. Fishes of the Kansas River System in relation to zoogeography of the Great Plains. Univ. Kansas Publ., Mus. Nat. Hist., 17:124-132. March 24. SLASTENENKO, E. P. 1958. The distribution of freshwater fishes in the provinces and main water basins of Canada. Bull. Shevchenko Sci. Soc., 1(6):1-11. February. SUMMERFELT, R. C. AND MINCKLEY, C. O. 1969. Aspects of the life history of the sand shiner, Notropis stramineus (Cope), in the Smoky Hill River, Kansas. Trans. Amer. Fish Soc., 98(3):444-453. SuTtTkus, R. D. 1958. Status of the nominal cyprinid species Moniana deliciosa Girard and Cyprinella texana Girard. Copeia, 4:307-318. December 22. ,irmsnor nmr \ hi I “| i 1 i OCCASIONAL PAPERS | 1 | IA DY LPIA I of the ——————— MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas NUMBER 13, PAGES 1-12 FEBRUARY 20, 1973 LIFE HISTORY AND ECOLOGY OF THE HYLID FROG OSTEOCEPHALUS TAURINUS, WITH OBSERVATIONS ON LARVAL BEHAVIOR By WILLIAM E. DUELLMAN' AND JEAN LESCURE® The large hylid frog Osteocephalus taurinus Steindachner is widely distributed in the Amazon Basin and the Guianas in South America. Despite the abundance of specimens of this species and its four congeners (O. buckleyi, leprieurii, pearsoni, and verrucigerus ) in museum collections, little information is available concerning the life history and behavior of Osteocephalus. Bokermann (1964) re- marked on the calling behavior and deposition site of O. taurinus. Trueb and Duellman (1970) presented data on the mating call and life history of O. verrucigerus (Werner) and provided the only de- scription of a tadpole of the genus. Field observations on four species of Osteocephalus in Amazonian Ecuador were summarized by Trueb and Duellman (1971). Independently, data were gathered on tadpoles of Osteocephalus taurinus by Duellman and his associates in Brasil and Ecuador, and by Lescure in French Guiana. As a result, the larval development of the species is described, together with some aspects of larval be- havior observed experimentally and in the field. In addition, the mating call of O. tawrinus is described for the first time. Tadpoles have been deposited in the collections of Museum National d’His- toire Naturelle (MNHN), Université de Paris (UP), and the Um versity of Kansas Museum of Natural History (KU). ‘Curator, Division of Herpetology, Museum of Natural History, University of Kansas. Reptiles et Poissons, Museum National d'Histoire Naturelle, 57 Rue Cuvier, Paris, and Laboratoire Arago, 66650 Banyuls-sur-Mer, France. 2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY LARVAL DEVELOPMENT The only record of egg deposition by Osteocephalus taurinus was by Bokermann (1964), who observed that this species deposits its eggs in a film on the surface of flooded depressions in the forest. Lar- vae were transformed in the laboratory by Lescure. The post-meta- morphic frogs clearly could be associated with adult O. taurinus, thus substantiating the identification of these and other larvae. Description.—A typical tadpole in development stage 37 (Gos- ner, 1960) from Kérenroch, French Guiana, has a body length of 18.3 mm and a total length of 48.5 mm. Body two-thirds as deep as wide, deepest posteriorly; in dorsal view elongately elliptical with round snout; in profile snout narrow, round. Eyes small, widely separated, directed laterally; nostrils directed anterolaterally about midway between eyes and tip of snout. Spiracle sinistral; spiracular opening directed posterodorsally just below midline at about mid- length of body. Cloacal tube short, dextral. Caudal musculature moderately slender, curved upward terminally, extending nearly to tip of narrowly rounded tail; caudal fins about equal in depth, deep- est at about two-fifths length of tail; depth of caudal musculature at midlength of tail slightly less than depth of either fin; dorsal fin not extending onto body (Fig. 1). Mouth moderately small, directed ventrally; median third of upper lip bare; rest of mouth bordered by two rows of small labial | papillae; lips indented posterolaterally. Beaks slender, bearing small, blunt serrations; upper beak forming a broad arch with long, slender lateral processes; lower beak broadly V-shaped. Three upper and five lower rows of teeth; first row composed of small teeth and broadly interrupted medially; third upper row narrowly interrupted medially; fifth lower row composed of small teeth and not extending so far laterally as other rows (Fig. 2). In life and in preservative, the tadpoles are entirely black. Ontogenetic Change.—Small tadpoles in developmental stage with total lengths of 13.0-14.2 mm (x=13.6, N=10) have less pig- ment in the fins than do larger tadpoles. Otherwise, there is no change in pigmentation during development; all tadpoles beyond Fic. 1. Tadpole of Osteocephalus taurinus, KU 148706, total length 48.5 mm. LIFE HISTORY OF OSTEOCEPHALUS TAURINUS 3 Fic. 2. Mouthparts of tadpole of Osteocephalus taurinus, KU 148706; line equals 1 mm. stage 25 are entirely black. The sizes of tadpoles at Kérenroch, as correlated with development stages, are shown in figure 3. The most obvious changes during larval development are in the 60) L mm 50 40 30 20 10 34 35 36 37 38 39 40 41 42 43 44 45 46 juv. stages Fic. 3. Sizes (total length) of 79 tadpoles as correlated with developmental stages. Each point is the mean total length for a sample in a given develop- mental stage. 4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY mouthparts. The smallest tadpoles available are in stage 25 and have only two upper and three lower rows of teeth. The third lower row is weakly developed; the outermost upper row and the two outer lower rows are absent. Larger tadpoles in stage 25 have a full complement of tooth rows. During metamorphosis the outer tooth rows are the first to degenerate. Small tadpoles in stage 25 have a few small labial papillae arranged in one row. Most tadpoles in stages 27-36 have one row of labial papillae; beginning at stage 37 the papillae become more numerous and tend to form two rows in the larger tadpoles. These ontogenetic changes are like those de- scribed for Triprion petasatus by Duellman and Klaas (1964) and for Smilisca phaeota by Duellman and Trueb (1966). Recently metamorphosed young have snout-vent lengths of 19.5- 20.5 (x=19.7, N=3). In life, the dorsum is gray with clusters of melanophores; the anterior and posterior surfaces of the thighs, flanks, and venter are black. In larger individuals the limbs and flanks become gray, and in still larger specimens the flanks and thighs become tan, the belly cream, and the dorsum tan with brown markings. Comparisons—The tadpoles of Osteocephalus taurinus differ from those of O. verrucigerus by having three (instead of two) upper rows of teeth and in having fewer labial papillae; two rows of papillae are present in all O. verrucigerus, except hatchlings. Fur- thermore, the tadpoles of O. verrucigerus are not uniform black; they have a pale transverse, crescent-shaped mark ventrally on the body. Superficially, the tadpoles of O. taurinus resemble those of Hyla geographica, which also are uniform black, but which have two upper and four lower rows of teeth and deeper caudal fins (Bokermann, 1963). LARVAL ECOLOGY Lescure observed tadpoles of Osteocephalus taurinus in French Guiana near an ORSTOM hydrological research station at Kéren- roch (5° 5’ N, 53° 2’ W, 5 m elev.), on the left bank of the Sin- namary River, 45 km upstream (south) from its mouth. Behind the station two rivers, of which the Crique Grégoire is the largest, flow into the Sinnamary at the same point. The soil of the granite basin crossed by these rivers is a clayish sand with coarse sand dominant. During heavy rains the rivers rise rapidly and overflow, so as to form one wide expanse of water flooding the forest for a width of several hundred meters. The current is very strong near the Sinnamary. LIFE HISTORY OF OSTEOCEPHALUS TAURINUS 5 The vegetation consists of tropical rainforest having an abundance of large lianas. On three occasions (February 1969, February and June 1971) large aggregations of tadpoles of O. tauwrinus were found in flooded areas of the forest where the current was weak and the depth of the water was 0.5 to 1 m. The tadpoles swam along the new banks amongst dead leaves or grazed on moss on logs in the water. Each aggregation probably contained more than 1,000 tadpoles. One catch consisted of 296 tadpoles after many had escaped. Within the aggregations most tadpoles were in stages 36 and 37 of development; six were in stages 29 and 30, and one was in stage 25 (Fig. 4). Others were more advanced, for shortly after the capture eight in- dividuals left the water to complete their metamorphosis. Of 79 tad- poles collected on the same day, more than half were in stages 37-39. The tadpoles in a given aggregation do not metamorphose at the same time. Those in stages 44 and 45 leave the water one at a time and climb onto vegetation, whereas those in stage 43 remain swim- ming with the group. Thus, there is no metamorphic aggregation as there is in Scaphiopus ( Bragg, 1950) or in Bufo marinus (Lescure, 80 70 60 50 40 30 20 10 am 29.30 33.34 37-38 41-42 45.46 stages 2? 26 7.28 31.32 35-36 39.40 43.44 Fic. 4. Population structure (number of individuals in different develop- mental stages) with respect to developmental stages of 296 tadpoles removed from one aggregation. 6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY per. obser.) where tadpoles in advanced stages aggregate and re- main immobile in shallow water. On 18 June 1970, Martha L. Crump observed five aggregations of tadpoles of O. taurinus in a shallow stream on the Ducke Reserve near Manaus, Amazonas, Brasil. The stream, about 1 m wide and no more than 25 cm deep, flowed slowly through young second- growth rainforest. Parts of the stream were in direct sunlight, but tadpoles were observed only in shaded areas along about 20 m of the stream. Four aggregations containing about 100 small tadpoles each; a sample of one of these consisted only of individuals in stage 25. A fifth aggregation contained an estimated 500 tadpoles; a sample consisted of individuals in stages 33-36. A small series of tadpoles in stages 35-39 was seined from a swampy pool with no current at Veracruz, Pastaza Province, Ec- uador, 950 m, on 4 July 1968, by John D. Lynch. These were part of an aggregation of “several hundred” tadpoles. On 6 July 1968, Lynch obtained one tadpole in stage 42 in quiet water at the edge of the Rio Puyo, 3 km S Puyo, Pastaza Province, Ecuador, 980 m. LARVAL BEHAVIOR Observations made at Kérenroch, Veracruz, and at the Ducke Reserve indicate that tadpoles of Osteocephalus taurinus form large aggregations and that this behavior occurs throughout larval de- velopment. The following observations and experiments were made by Lescure at Kérenroch, French Guiana. Observations.—The tadpoles in an aggregation are close together and frequently touch one another. When the group is stationary, it assumes a spherical shape. Tadpoles in the middle of the mass are immobile and not oriented in any particular direction; those on the periphery typically wriggle and seem to push towards the middle. As a group moves along the margin of a bank or log, the tadpoles along the leading edge of the group and those in the middle slow their movements, apparently in order to feed. The tadpoles in the rear of the group bump into those ahead of them but continue to advance by moving along the outer edge of the group. Apparently they cannot penetrate the middle of the group, so they move to the front, where they graze or move back along the log towards the middle. This circulation of tadpoles within the group continues with the result that tadpoles from the rear keep moving around the periphery to the front even though food remains behind. Thus, the entire aggregation progresses forward in a manner analagous to a slowly rolling ball (Fig. 5). LIFE HISTORY OF OSTEOCEPHALUS TAURINUS ~l Fic. 5. Schematic diagram of movement of a group of tadpoles along a log; solid dots = A, stars = B, open circles = C. I. Disposition of tadpoles within group at position one. I’. Tadpoles C advance by moving along edge of group. II. Tadpoles C are in front of the group; tadpoles B begin to advance. III. Tad- poles B are in front; tadpoles A begin to advance. IV. Tadpoles A are in front (same disposition as in position one). The arrow indicates direction of move- ment of the aggregation; small arrows within ellipses indicate movements of tadpoles within the aggregation. Occasionally the group branches out in another direction, but either the tadpoles soon return to the aggregation, or the entire group follows the new direction. When an individual tadpole leaves the group, it does not wander far and soon returns to the group. Experiments——Two hundred tadpoles were removed from an aggregation and placed in a rectangular tank, 8 x 1 m with water 20-30 cm deep. During the several days that the tadpoles were kept in the tank they demonstrated no gregarious behavior. However, when they were replaced in the river, they formed aggregations. The tadpoles were released in three groups at 5 m intervals in the river. Forty minutes later the group that was close to the bank stopped at a submerged twig. Immediately the tadpoles began to feed voraciously with their heads down and tails quivering. The two other groups, which were several meters away, immediately began to swim faster. In a few moments they reached the feeding group and joined to form one active mass of tadpoles around the twig. Several minutes later the excitement subsided, and the whole aggregation moved slowly along the bank. Discussion.—The method of progression of aggregations and the general lack of orientation of tadpoles within aggregations of tad- poles of Osteocephalus taurinus differ from schooling behavior in fishes. The tadpoles do not swim in parallel formations, and instead of maintaining a constant distance from one another, the tadpoles tend only to remain close together. However, a certain coordination of movement exists when a group swims in one direction. The tight grouping probably is maintained by the continual contact among individuals. The lack of contact on the edges of the group may provoke the movements toward the middle when the group is sta- tionary or the lateral movements when the group is in motion. Pos- sibly the need for oxygen or food initiated the movement of tadpoles from below to swim around the edge of the group to the surface. 8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Movements upward through the middle, as observed in Rana tem- poraria by Savage (1961), were not observed. Feeding activity seems to be an important factor in the gre- gariousness of the tadpoles of O. taurinus. Living in temporarily flooded areas, the tadpoles do not have permanent grazing grounds. Thus, the discovery of a supply of food by one group of tadpoles may normally result in frenzied feeding activity which attracts other groups of tadpoles. The grouping seems to be a lasting one, because the group moves as a whole even when not feeding. The presence of one unlimited dimension in the water may be a requirement for the maintenance of an aggregation. No gregarious- ness was observed in tadpoles kept in a rectangular tank. The same phenomenon occurs in fish that normally live in schools; Schaffer (1955) noted that schools of young herring broke up in quadran- gular tanks but remained aligned in annular ones. Obviously, neither tactile nor visual stimuli can account for the regrouping witnessed in the second experiment. Possibly chemical sensations were involved, but because of the frantic movements of the tadpoles that discovered the food, it may be assumed that the resulting vibrations in the water attracted the distance groups. Never- theless, visual stimuli are involved in some movements by the tad- poles. The shadow cast by the observer on a group of tadpoles near the surface results in the group plunging downward. This collective reaction is not the only kind of protective reaction by a group. If an object is dropped or pushed into a group, the tadpoles scatter in small groups, but in the absence of further disturbance, they regroup within 15 min. Wassersug and Hessler (1971) demonstrated that the stimulus for aggregation is visual in Xenopus laevis (Daudin). The presence of a current in the water may affect the aggrega- tional behavior of the tadpoles, but one aggregation has been ob- served in a swampy pond. Furthermore, tadpoles in the quiet water in the tank did not aggregate. Bragg (1968) defined two kinds of aggregations in tadpoles: 1) social, in which the stimulus for aggregation is generated by the tadpoles, and 2) asocial, in which the stimulus is from environmental factors. Bragg concluded that aggregations of plankton-feeding tad- poles were social aggregations, but that groups that formed at concentrations of food were asocial aggregations. Tadpoles of O. taurinus aggregate at concentrations of food but also are gregarious when not feeding. The aggregations have a social aspect, because when moving the tadpoles remain close together and react defen- sively as a group. These are not chance aggregations. They are ele- LIFE HISTORY OF OSTEOCEPHALUS TAURINUS 9 mentary groups, as defined by Lescure (1968). The accumulated observations and results of the experiments suggest that tactile stimuli may be the primary factor for maintenance of aggregations in tadpoles of O. taurinus, but the vibrations caused by tadpoles that find concentrations of food may initiate the formation of an aggre- gation. Chemo-stimuli remain an unstudied possibility. We have no data on temperature responses of the tadpoles; Brattstrom (1962) demonstrated that tadpoles of Hyla regilla Baird and Girard agere- gated in response to thermal gradients. MATING CALL We have one lengthy recording of Osteocephalus taurinus re- corded at Limoncocha, Napo Province, Ecuador, on 30 June 197], by W. Ronald Heyer, who informed us that the call was typical of others that he heard at the same locality. This individual produced two kinds of notes—short melodious notes (herein termed type A) and longer, poorly modulated tones (type B). In a two-minute sequence, type B notes were always pre- ceded by two or three type A notes, but in other cases the latter were produced without subsequent type B notes. Thus, any given call group might consist of N A or N B. The number of type A notes per call group is 2 or 3 (x=2.4, N—87), and the number of type B notes per call group is 0-4 (x=—0.46, N37). In a given sequence of calls the frequency of notes and combinations of types of notes in individual call groups was (frequency of occurrence in parentheses; not sequential as produced): 2A (18), 2A + 1B (4), 3A (9), 3A + 1B (2), 3A + 2B (2), 3A + 3B (1), 3A + 4B (1). The rate at which call groups are produced is 27.5 per min, and note repetition rate is 83.5 notes per min. Pulse rates of type A and B notes are about 160 and 95 pulses per sec, respectively. In se- quences of type A notes, the first and third (if present) notes have an equal harmonic arrangement, whereas the harmonics are higher pitched in the second note (Fig. 6A-B). The first harmonic is domi- nant, and each successive higher harmonic has less energy. The first harmonic is at 570-625 (x=598, N=10) Hertz in the first note in a sequence, and in the second note the first harmonic is at 770-805 (X=788, N=10) Hertz. The duration of type A notes is 0.03-0.05 (x=0.042, N=10) sec; the duration of type B notes is 0.36-0.65 (x=0.47, N=5) sec. Type B notes generally have poor modulation (Fig. 6C). The call of Osteocephalus taurinus can be described as “boop- boop-boop,” followed or not by “worrr.” The call differs notably 10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Kilohertz Time in Seconds Fic. 6. Mating call of Osteocephalus taurinus showing different sequences of notes. A. Two type A notes. B. Three type A notes. C. Three type A notes followed by two type B notes. KU Tape 1204; Limoncocha, Napo Province, Ecuador; 30 June 1971; 24.6° C; effective band width 40 Hz. from that of O. verrucigerus, which consists of a series of well- pulsed, low, guttural notes (Trueb and Duellman, 1970). The mating call of Osteocephalus taurinus is highly complex by comparison with known calls of other hylids. The type A and B notes are not comparable to the primary and secondary notes pro- duced by many small hylids; in these notes the acoustical differences are in duration and phase (Duellman, 1970), whereas the type A and B notes differ in several acoustical parameters and apparently differ in information content. ACKNOWLEDGMENTS We express our gratitude to Mr. Brugiere, Director of the Cen- tral Offce de la Recherche Scientifique et Technique Outre-Mer of Cayenne for his hospitality extended to Lescure at the hydrological station ORSTOM at Kerenroch and to Duellman’s field associates, Martha L. Crump and John D. Lynch, for providing specimens and field observations. We are indebted to W. Ronald Heyer for the tape LIFE HISTORY OF OSTEOCEPHALUS TAURINUS | recording of O. taurinus and to Linda Trueb for the illustrations depicted in figures | and 2. RESUME Le tétard d’Osteocephalus taurinus (Hylidae) est enti¢rement noir, il a une petite bouche ventrale avec trois rangées supérieures de dents et cinq rangées inférieures. Les lévres ont une rangée de papilles excepté dans la partie médiane de la lévre supérieure; chez les plus grands individus, les papilles plus nombreuses sont disposées en deux rangées. Des groupements d’environ 1000 tétards furent observés dans des parties inondées de la forét ou le courant est faible. Les obser- vations et les résultats des expériences suggerent que les contacts entre les tétards sont le principal stimulus de ce comportement erégaire. Le stimulus déclencheur de la formation du groupe serait les vibrations de l'eau provoquées par l’agitation des tétards dé- couvrant de la nourriture. L’appel sexuel d’O. taurinus est complexe et consiste en deux catégories de notes. La production d'une longue note a peine modulée parait dépendre d’une premiere série de deux ou trois notes courtes et plus modulees. RESUMEN El renacuajo del hilido, Osteocephalus taurinus, es completa- mente negro, tiene boca ventral moderademente pequena, con tres hileras de dientes en el labio superior y cinco en el inferior. La parte media del labio superior carece de papilas, el resto de los labios posee una sola hilera; sin embargo, en los individuos de mayor tamafio hay proliferacién de las papilas en dos hileras. Se encontraron renacuajos en todos los estadios de desarrollo en agregaciones hasta 1000 individuos colectados en las partes anegadas con agua en movimiento, cercanas a los rios. Las observaciones y los resultados experimentales realizados sugieren que el estimulo principal en la formacién de las agregaciones es tactil, aunque el estimulo inicial tal vez sea la presencia de comida. El complejo canto nupcial de O. taurinus consiste de dos tipos distintos de notas; la produccidén de una nota larga, poco modulada aparentemente depende de la producién previa de dos o tres notas melodiosas cortas. 12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY LITERATURE CITED BOKERMANN, W. C. A. 1963. Girinos de anfibios brasileiros—1l. (Amphibia-Salientia). An Acad. Brasileira Cien., 35:465-474. 1964. Field observations on the hylid frog Osteocephalus taurinus Fitz. Herpetologica, 20:252-255. Brace, A. N. 1950. Mass movement at metamorphosis in the savannah spadefoot, Sca- phiopus hurterii Strecker. Proc. Oklahoma Acad. Sci., 31:26-27. 1968. The formation of feeding schools in tadpoles of spadefoots. Was- mann Jour. Sci., 26(1):11-16. BRATTSTROM, B. H. 1962. Thermal control of aggregational behavior in tadpoles. Herpeto- logica, 18:38-40. DUELLMAN, W. E. 1970. The hylid frogs of Middle America. Monog. Mus. Nat. Hist. Univ. Kansas, 1: xi+753 pp. DUELLMAN, W. E. AND L. T. KLaas 1964. The biology of the hylid frog Triprion petasatus. Copeia, (3):308- Oat. DUELLMAN, W. E. AND L. TRUEB 1966. Neotropical hylid frogs, genus Smilisca. Univ. Kansas Publ. Mus. Nat. Hist., 17:281-375. Gosnepr, K. L. 1960. A simplified table for staging anuran embryos and larvae, with notes on identification. Herpetologica, 16:183-190. LEscurE, J. 1968. Le comportement social des batraciens. Rev. Comportement Anim., 2(4) :1-33. SAVAGE, M. R. 1961. The ecology and life history of the common frog Rana temporaria. Pitman, London, 221 pp. SCHAFFER, W. 1955. Uber das Verhalten von Jungerheringsschwarmen in Aquarium. Arch. Fischereiwiss., 6:276-287. TRUEB, L. AND W. E. DUELLMAN 1970. The systematic status and life history of Hyla verrucigera Wemer. Copeia, (4):601-610. 1971. A synopsis of Neotropical hylid frogs, genus Osteocephalus. Occas. Papers Mus. Nat. Hist. Univ. Kansas, 1: 1-47. WaASSERSUG, R. AND C. M. HESSLER 1971. Tadpole behaviour: Aggregation in larval Xenopus laevis. Anim. Behav., 19:386-389. JrV\utl ACADEMY OF SCIENCE & Oo |] Wi Yuvili | : | A i ) 4 7 t ey oo 4) 1 / A lk Wu" i ga Ver iw Jd J 1 in TAN | if ( RNIA OCCASIONAL PAPERS of the The University of Kansas Lawrence, Kansas NUMBER 14, PAGES 1-8 FEBRUARY 20, 1973 A PENNSYLVANIAN DISSOROPHID AMPHIBIAN FROM KANSAS By THEeoporE H. Eaton’ Among rhachitomous labyrinthodont Amphibia of the Late Paleozoic, the Dissorophidae and Trematopsidae currently are con- sidered to be separate but closely related families. They are gen- eralized in most features but show trends toward terrestrial life. Commonly recognized distinctions between the two families include the great enlargement of the external nares in the trematopsids, whereas the dissorophids have: 1) transverse plates of bony armor along the back usually attached to the neural spines; 2) broad, rounded interpterygoid vacuities; and 3) palatine bone forming part of the lower rim of the orbit. Apart from the limitations imposed by incomplete specimens, these features are neither consistently present in the appropriate group nor mutually exclusive between the two. For instance, DeMar (1966a) described a Permian genus Longiscitula with enlarged nares; yet because of other characters such as dorsal armor, he placed it in the Dissorophidae. Vaughn (1969) showed that another Permian genus, Ecolsonia, has broad, rounded interpterygoid fenes- trae such as characterize the dissorophids, but because of the pres- ence of large nasal openings, Vaughn referred the genus to the Trematopsidae. It is not known whether or not armor was present in Ecolsonia. Amphibamus from the Middle Pennsylvanian Allegheny beds of * Curator, Vertebrate Paleontology, Museum of Natural History, University of Kansas. 2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Linton, Ohio, is the oldest dissorophid; fortunately it can be studied from a number of specimens (Carroll, 1964). Watson (1940), who investigated Amphibamus in reference to the origin of frogs, noted that it is probably more aquatic and more primitive in some ways (e.g., exclusion of the frontal from the rim of the orbit by the pre- frontal and postfrontal) than any other dissorophid. The genus described here from Kansas is from the Late Penn- sylvanian. Although in most respects it is an unspecialized dissoro- phid, it does have an enlarged nasal opening. The other known genera are variably characterized by small or large nostrils, presence or absence of armor, and wide or narrow palatine fenestrae; all are of Permian age. It is a part of the thesis of this paper that we should no longer recognize two families in this group, but only one, for which the older name, Dissorophidae, would be used. Two other valid families can be included in a superfamily, Dissorophoidea; these are the Zatracheidae and Doleserpetontidae. Both are closely related to dissorophids. Doleserpeton is a minute amphibian, prob- ably terrestrial, from the Lower Permian fissure deposits at Fort Sill, Oklahoma. This animal was described by Bolt (1969) as a possible “protolissamphibian.” Its structure, apart from the pedicellate, bi- cuspid teeth and large pleurocentra, differs remarkably little from that of Amphibamus. The specimen named and described here is a nearly complete skull and skeleton of a small labyrinthodont amphibian, order Temnospondyli, suborder Rhachitomi, family Dissorophidae, taken from the Rock Lake Shale member of the Stanton Limestone, Mis- sourian series, Upper Pennsylvanian, approximately 6 mi NW of Garnett, Kansas. The locality is that from which Petrolacosaurus, Hesperoherpeton, Edaphosaurus ecordi, and other members of the Garnett fauna were obtained. The dissorophid is in a limy shale bed containing plant fragments and numerous bones of the primitive pelycosaur Clepsydrops. Actiobates new genus Type species.—Actiobates peabodyi new species. Holotype—University of Kansas Museum of Natural History (VP) 17941; locality, age, and formation as given in preceding para- graph. Diagnosis—Skull broad, shallow, comparable to Amphibamus and Ecolsonia; cranial roofing bones thin, with pit-and-ridge orna- mentation externally; basipterygoid joint moveable; frontals entering margin of orbits; otic notch moderate, its posterior opening slightly PENNSYLVANIAN DISSOROPHID AMPHIBIAN 3 narrowed; marginal teeth uneven, those of anterior end of max- illary large. Actiobates differs from Amphibamus in having a greatly enlarged nasal opening and from Ecolsonia in having the frontal and parietals of nearly equal size, and the postparietals narrow and elongate later- ally. Diagnosis of the species is the same as that of the genus. This specimen is mentioned by Estes (1965) and Estes and Reig (1973). Etymology.—tThe generic name is derived from the Greek actios meaning shore, and bates meaning one who walks. The specific patronym is in remembrance of Frank E. Peabody, who initiated the extensive study of the Garnett fauna. DESCRIPTION AND DISCUSSION The skull is flattened by pressure, but has sustained little damage and slight displacement of bones (Fig. 1). Its medial length is 59 mm, but the lateral edges have been pressed out a few millimeters, making the skull a little broader than it was in life. At the same time the nasal and premaxillary parts were spread laterally, leaving a gap between. In addition there is a small round fenestra between the tips of the nasals, which is commonly present in dissorophids. Ventrally, the mandibles were pressed against the lateral parts of the palate and upper jaw; a full description of this area must await further preparation. The pterygoid, parasphenoid and otic parts are remarkably well preserved. In the ventral view (Fig. 1b) the stapes and opisthotic are shown on the left side of the drawing as they are in the specimen; on the right, they are placed in their original positions relative to one another and the fenestra ovalis. The postcranial skeleton and details of the skull not yet available will be described in a subsequent paper. The vertebral column is nearly complete, but scattered, because of the rhachitomous division of each vertebra. The forelimbs, including the girdle and feet, are complete; similarly, most of one hind limb is complete. A few scat- tered chevron bones and part of the pelvis are present. Four small bones are present anteriorly that look like double-ended paddles; these are probably units of the dorsal armor, but they lack attach- ment to the neural spines. One curious feature is the presence of an occipital (one basi- and two exoccipitals in one unit) of a small individual of Clepsy- drops inside the mouth cavity and pressed up into the orbit. A few other associated fragments suggest that this was not a chance re- lationship, but instead, that the amphibian was feeding on the pelycosaur. Figure 1 shows most of the visible features of the skull, but some + OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Fic. 1. Skull of Actiobates peabodyi, KU (VP) 17941, actual size. A. Dor- sal view. B. Ventral view. Abbreviations: cl, edge of clavicle; f, frontal; ju, jugal; la, lacrimal; mx, maxillary; na, nasal; op, opisthotic; or, orbit; pa, pari- etal; pm, premaxillary; po, postorbital; pp, postparietal; pof, postfrontal; prf, prefrontal; pro, prootic; ps, parasphenoid; pt, pterygoid; gj, quadratojugal; qu, quadrate; spt, supratemporal; sq, squamosal; st, stapes; ta, tabular. PENNSYLVANIAN DISSOROPHID AMPHIBIAN 5 items need to be discussed. Comparison of the expanded nasal opening with those of most dissorophids and other labyrinthodonts shows that the anterior part (one-third) has not been changed by the enlargement, but that the posterior two-thirds resulted from reces- sion of the edges of the lacrimal and prefrontal bones and some of the lateral edge of the nasal. The lacrimal and prefrontal are left as little more than a cross-bar separating the orbit and nostril. What- ever the reason for this expansion may have been, it need not have affected the original function of the anterior portion, which was for the passage of air. The more lateral and anterior parts of the palate are concealed beneath by the lower jaws which are crushed against them; it may be possible to reveal these parts of the palate by further preparation while reconstructing the lower jaw at the same time. Much of the pterygoid is visible; its joint with the basisphenoid was evidently moveable. The expanded ventral surface of the pterygoid and that part of the parasphenoid which is between the basipterygoid processes are closely scattered with minute (0.25 mm diameter or less), round, flattened, dome-like teeth. Each tooth occupies, but may not fill, a round, shallow alveolus. Some of the teeth are much smaller than the alveolar opening and deeply placed in it; these are apparently in early stages of eruption. Other teeth nearly fill the alveolus to the rim, and the largest appear to have fused with the margin. A few have their rounded peaks worn off exposing the inner cavity. The basisphenoid is exposed on either side of the converging, ventralmost part of the parasphenoid; the bone bears grooves for the interal carotids. The medial edges of these grooves are bounded by the parasphenoid except in their anterior parts, where the basi- pterygoid processes project laterally. The cultriform process of the parasphenoid is slender and crushed against the skull roof; thus, it is not possible to see a cranial cavity or foramina. There is no indi- cation of a sphenethmoid. No palatine, ectopterygoid, or vomerine teeth can be seen, be- cause these bones are covered. The few visible teeth of the jaws are shown in figure 1b. There appears to be space for about 30 teeth in each jaw. Apparently, the largest teeth are two located on the for- ward end of the maxilla. Morphologically, the otic region is most interesting. The otic capsule is represented mainly by the prootic, and is bounded later- ally by the quadrate process of the pterygoid, anteriorly by the basal process, medially by the parasphenoid plate, and posteriorly by the 6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY anterior edge of the clavicle. At the posterolateral corner of the otic capsule is the fenestra ovalis, with the stapes slightly displaced, as seen on the left side of figure 1b. Beyond its rounded footplate, the stapes has a foramen in its shaft; distal to the foramen, the shaft curves laterally. The shaft lies dorsal to the quadrate process of the pterygoid, beyond which its tip shows. Posterior to the foot of the stapes, lies a short bone which evidently forms a portion of the wall of the fenestra ovalis. I assume that this bone is the opisthotic, which is considerably reduced and bears a short paroccipital process laterally. On the right side of figure 1b, the stapes and opisthotic are shown in their proper positions relative to the fenestra ovalis. The resemblance of this apparatus to those of frogs and sala- manders is at once obvious. If we allow a little more reduction of the opisthotic until it is a small, flat plate, and eliminate its ossifi- cation (as is usual for the otic region among modern amphibians), the opisthotic becomes quite precisely the operculum of the otic capsule. Primitively, the paroccipital process was for muscle attach- ment. When the bone was reduced, the muscle (opercularis) re- mained inserted on the operculum; it extends back to the anterior edge of the scapula in the modern Amphibia. It may have done so in dissorophids in the early stages of terrestrial life. This alone is not enough to demonstrate conclusively that Actiobates is directly an- cestral to Lissamphibia, because we do not yet know the relationship of the expanded nasal opening to the reduction of the skull in frogs, salamanders, and caecilians. On the other hand, the condition of the ear in Actiobates approaches that of the modern Amphibia re- markably closely. Elements of the hyobranchial apparatus, although not figured, are present. They are somewhat displaced against the otic region of the left side of the skull. SKULL CHARACTERS AND RELATIONSHIPS The question whether dissorophids and trematopsids represent one family or two depends on the occurrence of differential charac- ters or combination thereof. But interpretation of these, in tum, requires viewing a given character, or characters, as a stage or “state” in the evolution of certain morphological features. For instance, among the Rhachitomi a moveable basipterygoid articulation is clearly more primitive than a fixed articulation. The moveable articulation occurs in the two Pennsylvanian genera, but not, so far as is known, in those of the Permian. On the other hand, union in a fixed joint could have occurred more than once; if it did, then PENNSYLVANIAN DISSOROPHID AMPHIBIAN ji possession of a fixed joint is not, by itself, evidence of a direct rela- tionship among genera. Likewise, there is no doubt that the small external nares charac- teristic of most labyrinthodonts are primitive with respect to large external nares. The former occur in Amphibamus, Dissorophus and most other genera in the Dissorophidae, but not in Actiobates, Ecol- sonia, Longiscitula, or the genera usually placed in the Trema- topsidae. Again, the enlarged condition, being specialized, could have originated more than once. The size of the external nares is not a simple character, and the modification of at least three bones in a particular area of the skull in the same way in more than one evolutionary line seems improbable. Such an interpretation would require independent evidence from other characters that genera having enlarged nares represent separate phyletic lines. Morphologically it is not possible to determine the relative prim- itiveness of wide, rounded interpterygoid vacuities as compared with narrow or small interpterygoid vacuities. However, within the scope of dissorophids and trematopsids, we can suppose that the wide, rounded state is primitive because it is present in Amphibamus and Actiobates of the Pennsylvanian in addition to other dissorophid genera. The fenestrae are narrow and relatively small in several trematopsids. If these taxa are closely related and form a natural group (whether one family or two), then there is a high probability that the narrowed condition is secondary. In spite of the fact that the most primitive labyrinthodonts had no interpterygoid vacuities, it is reasonable to suppose that rounded interpterygoid vacuities were present in primitive dissorophids, because they are known in Amphibamus. None of the features of the skull that has been discussed, nor the occurrence of dorsal armor, seem to be suitable to distinguish the two families. In the case of each character there is at least one taxon for which a character state cannot be determined. Furthermore, the distribution of known characters is discordant. Thus, any arrange- ment of the genera in separate groups requires that some incon- sistency in overlapping or contradictory characters be introduced in their diagnosis. It seems more satisfactory, therefore, to recognize only one family, the Dissorophidae, for all the animals hitherto placed in this and the Trematopsidae. 8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY SUMMARY A virtually complete skeleton of a new dissorophid labyrintho- dont, Actiobates peabodyi, is reported from the Rock Lake Shale member of the Stanton Limestone, Missourian Series, near Garnett, Kansas. The skull is characterized by an expanded narial opening, characteristic of Permian trematopsids and seen here for the first time in a Pennsylvanian amphibian. In other respects, the skull is that of a generalized dissorophid. The distribution of characters among dissorophids and trematopsids no longer seems to warrant recognition of more than one family, the Dissorophidae, for both. LITERATURE CITED Bout, J. R. 1969. Lissamphibian origins: possible Protolissamphibian from the Lower Permian of Oklahoma. Science, 166:888-891. CaRRoLE, R. L. 1964. Early evolution of the dissorophid amphibians. Bull. Mus. Comp. Zool., Harvard, 131(7 ):163-250. DeEManr, R. E. 1966a. Longiscitula houghae, a new genus of Dissorophid Amphibian from the Permian of Texas. Fieldiana Geology, 16(2) :45-53. 1966b. The phylogenetic and functional implications of the armor of the Dissorophidae. Fieldiana Geology, 16(3):55-88. Estes, RICHARD 1965. Fossil salamanders and salamander origins. Amer. Zool., 5:319- 334. EstTEs, RICHARD AND OSVALDO REIG 1973. The early fossil record of frogs: a review of the evidence. In J. L. Vial, Evolutionary biology of the Anurans. University of Missouri Press, Columbia (in press). VAUGHN, P. P. 1969. Further evidence of close relationship of the Trematopsid and Dis- sorophid Amphibians with a description of a new genus and spe- cies. Bull. So. Calif. Acad. Sci., 68(3):121-130. Watson, D. M. S. 1940. The origin of frogs. Trans. Roy. Soc. Edinburgh, 60:195-231. hissllNe 070. (5 re MAY 1 4 1973 LIBRARY OCCASIONAL PAPERS of the MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas NUMBER 15, PAGES 1-43 MAY 2, 1973 POPULATION STUDIES OF THE CAVE BAT (MYOTIS VELIFER): REPRODUCTION, GROWTH, AND DEVELOPMENT By Tuomas H. Kunz! An integral part of the ecology of an animal is the manner in which it successfully adapts to its environment. Under conditions of thermal seasonality in the temperate zones, insectivorous bats have evolved strategies that compress reproduction and develop- ment into a relatively short time period in the warm season, when food is abundant and temperature conditions are optimal for rapid growth of young. Seasonal reproductive patterns of temperate insectivorous bats (including autumn copulation, spring ovulation and gestation, and summer parturition) have been documented for several species. Much of the available information on reproduction has been sum- marized by Asdell (1964), Barbour and Davis (1969), and Carter (1970). Wimsatt (1945, 1960a) gave a thorough account of breed- ing behavior, pregnancy, and parturition of several vespertilionids and also discussed aspects of reproduction relating to hibernation (Wimsatt, 1960b, 1969). Orr (1970) summarized studies on growth and development. There are a number of important studies detailing aspects of natural history, population dynamics and behavior of Myotis velifer (J. A. Allen) (Twente, 1955a, 1955b; Tinkle and Milstead, 1960; Tinkle and Patterson, 1965; Dunnigan and Fitch, 1967; and Hay- ward, 1970). Other important contributions are those of Vaughan 1 Assistant Professor, Department of Biology, Boston University, Boston Massachusetts 02215. This paper represents part of a Ph.D. dissertation sub- mitted to the Department of Systematics and Ecology, University of Kansas. CALIFORNIA. ACADEMY OF SCIENCES 2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY (1959) on functional morphology and Constantine (1957) on pelage and molt. However, these studies do not assess an entire seasonal cycle, and little is known with respect to the reproduction, growth and development of M. velifer. In this study I have considered the seasonal chronology of reproduction and examined factors relating to growth and development in two populations occupying physically and thermally contrasting nurseries in Kansas. The ecological and geographic distribution of M. velifer typi- cally is patchy and extends northward from Honduras into the cen- tral Great Plains. The study area lies within the Gypsum Hills and adjacent areas of south-central Kansas. A complete discussion of the study area was given by Kunz (1973). See the appendix for place names, coordinates, and counties of each locality mentioned in text. Of the four races of M. velifer currently recognized, the northern- most race is now known as Myotis velifer grandis Hayward (1970) and occurs in parts of Texas, Oklahoma and in south-central Kansas. Previously, the name Myotis velifer incautus (J. A. Allen) was used for these bats (Miller and Allen, 1928). Climatically, the study area is typical of the southwestern plains. Most of the area falls within the O° C January isotherm and above the 26° C July isotherm (U.S. Department of Commerce, Climatol- ogical Data). The length of the average frost-free period varies from 180 to 190 days. The last vernal frost occurs in April and the earliest frost in autumn usually is in late September or early October. The summers are long, hot, and dry; it is not uncommon for temperatures of 39° C to prevail for several days. The warmest months are July and August. Winters generally are mild, although temperatures may remain below freezing for short periods. The coldest period usually occurs in February and early March, although this may vary con- siderably from year to year. Precipitation varies locally and season- ally and months may pass with little measurable amounts recorded. Mean annual precipitation is 16-28 in. (406-711 mm). Acknowledgments Several persons contributed to various aspects of this study. For field assistance I acknowledge Joseph T. Collins, Lawrence M. Cavin, John D. Lynch, Eric M. Rundquist, and Larry C. Watkins. Ronald E. Domsch wrote programs for sorting and tabulating growth and weight data. Arlin Pound and Stanley D. Roth were particularly helpful in providing maps, and introducing me to a number of caves in the area. The cooperation of land owners, ranch- ers, and county extension agents is greatly appreciated. I am grate- ful to representatives of the National Gypsum Company, Medicine Lodge, Kansas, and Buffalo, New York, for permission to study at the National Gypsum Mine, Sun City, Kansas, and to Mr. and Mrs. Fay Smith of Wilmore for their hospitality during my stay in the field. | SSS ES POPULATION STUDIES OF THE CAVE BAT 3 Appreciation is extended to Frank B. Cross, Robert S. Hoffman, and J. Knox Jones, Jr., who read and criticized an earlier version of this manuscript. I am also grateful to M. Brock Fenton and Stephen R. Humphrey who offered several helpful suggestions. I am particu- larly indebted to Frank B. Cross who made available financial and travel support, both for field and laboratory studies, through the State Biological Survey of Kansas. Computer time was secured through the Department of Systematics and Ecology and University of Kansas Computation Center. METHODS AND MATERIALS Field studies were begun in March 1968 and continued through January 1971. Data for mark-recovery analyses include all recov- eries through June 1970. All bats were marked using size 2b, lipped, aluminum bat bands (U.S. Fish and Wildlife Service, Washington, D.C.). Some bands were color-anodized (blue, green, and red) for marking juveniles occupying different nurseries. Four “seasons,” corresponding to Summer (June-August), Autumn (September- October), Winter (November-March), and Spring (April-May), were selected to best represent the seasonal changes in population behavior. Although this scheme does not correspond exactly to the seasonal activity of the entire population, it does represent a general pattern and probably includes most populations in the study area. Environmental Measurements Lost Colony and Wilmore were selected for intensive study in order to assess the influence of ambient temperature and relative humidity on reproduction, growth and development. These two nursery roosts, a cave and a barn respectively, were selected mostly because populations of approximately the same size occupy them in the summer. Also, the roosts invite comparative study because their environments contrast sharply with respect to physical and thermal properties. Temperature and relative humidity recordings were taken in roost areas using seven-day recording hygrothermographs and a Bendix Psychron. Twenty-four hour recordings of micro-roost temperatures (including body-surface temperatures), were obtained using a 12 channel, Model 44, YSI telethermometer with general purpose ther- mister probes positioned in roost areas at the Wilmore barn. Periodic recordings were made from April through October (Kunz, 1973). Similar attempts were made to measure micro-roost temperatures and body-surface temperatures of bats occupying Lost Colony but these efforts were unsuccessful. Whenever thermisters were posi- tioned in an area that had been occupied by bats they would either abandon that specific site or would move to a different part of the cave. Temperature and relative humidity measurements also were taken at other roosts. 4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Population Estimates, Age, Sex, and Reproductive Condition Methods used to estimate population size of active bats de- pended on the number and size of entrances used by the bats. Where single entrances to caves or buildings were used, flight counts were employed (see Dwyer, 1966; Humphrey, 1971) and _ this method probably provided the most accurate estimate. Flight counts were not practical at sites with multiple-use entrances; in such situ- ations, estimates were often based on the size of the stained area in a roost, the quantity of the accumulated guano, or number of neo- nates present after adults had departed. In a few instances, esti- mates were based on observations of clusters occupying ceiling areas. Sex ratios were determined mostly from trap samples (Tuttle, ms.) taken at entrances during emergence and return activity, over streams, or by taking hoop-net or hand samples within a roost. The two latter methods were avoided during the nursery period (June- July) in order to reduce disturbance, and were used selectively at other times. Young bats born in summer could be distinguished from adults at least through August. Generally, after this time, closure of the epiphyseal region of the phalanges occurred and many young bats developed adult-like pelage, and some were nearly equivalent to adults in body weight (Kunz, 1971). After August, age determi- nation was based on the condition of teats in females and size of testes in males. Young females usually have smaller, non-pendulant teats in contrast to those of post-lactating adults, and young males have smaller testes than adults. However, this aging criterion for males becomes less reliable in late September and October when testicular regression occurs in adults. Reproductive condition of females (pregnancy, lactation, and post-lactation) was determined by palpation. In April and early May pregnant females can not be distinguished from those that may be non-reproductive. In summer, non-pregnant females were con- sidered to be lactating if milk could be extruded from the mam- mary tissue, or white areas were notably visible beneath the skin. When milk no longer could be obtained, or mammary tissue was visibly pink and undergoing regression, individuals were considered to be post-lactating. Judgments of reproductive activity were based solely on the presence or absence of a discernible fetus in the latter two or three weeks of gestation and the presence of milk in the first week of lactation. When all individuals have given birth, the diffi- culty of estimating the proportion of reproductive females in the population was compounded by the fact that some females may have aborted in late term or lost their offspring in early days follow- ing parturition. These individuals may appear to be non-reproduc- tive upon gross examination so the category is probably appropriate POPULATION STUDIES OF THE CAVE BAT ) for them. Reliable estimates of non-reproductive females in a pop- ulation also are contingent on the fact that they, too, reside in nurseries. Establishment of a seasonal reproductive pattern for adult males was based mostly on linear measurements of testes and the cauda epididymides from preserved samples. Field observations on the changes in the size and position of testes and epididymides provide a general measure of seasonal activity; however, I think that tem- poral aspects of the reproductive pattern of adult males can be extrapolated from the pattern observed in preserved specimens. Growth and Development Studies of prenatal growth were based on preserved fetuses, which were removed from autopsied females and weighed to the nearest 0.01 gram using a Mettler top-loading analytical balance. Linear measurements of the width and length of the uterine horns and crown-rump length were taken from preserved specimens. The field methods utilized in studies of post-natal growth were selected primarily to minimize disturbance to nursery colonies. Data obtained from any one nursery were limited by the location and type of structure, the size of the resident colony, and its relative sus- ceptibility to disturbance. Studies at Lost Colony were conducted mostly for the assessment of morphological growth, whereas those at Wilmore were conducted for the purpose of recording behavioral observations. Preliminary observations at Lost Colony and at other nurseries suggested that an interval from one to one and a half hours was per- missible to capture, process, and return young to their roosts before adults returned from foraging. During this period some behavioral observations were made on social organization and age distribution of young within the roost. The procedure of sampling and observing young while adults were foraging was advantageous in reducing dis- turbance to the colony. However, the recovery rate of known age individuals was low once young became volant. In spite of this disadvantage, the procedure was preferable to sampling bats in the daytime when excessive disturbance to adults may result in high mortality and/or abandonment. Captures of newborn (or very young) bats were essential for a complete analysis of post-natal growth. Young bats having attached umbilical cords were assumed to be approximately one day old. This assumption has been used by others (see Pearson et al., 1952; Davis, 1969b), although it may not always be valid. When the rela- tive humidity of roosts is high, the umbilical cord may remain at- tached to a neonate longer than one day. Young bats were sexed, banded, weighed, and measurements of the forearm and fifth finger 6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY were taken. Color-anodized bands were used in roost areas for dis- tinguishing young from adults once the former were fully grown. At approximately 10-day intervals, newly captured neonates and young that had been previously marked were captured and processed. By the beginning of the fourth week, several young that had been marked in the first period had become volant. Throughout the summer unmarked samples of young bats that were trapped at colony entrances or in roost areas were processed as described above. Periodic samples of unmarked young corresponding to sizes of known-age bats, were collected from several different colonies. These bats were killed and preserved in 10 percent formalin for studies of pelage, dentition, and food consumption. Drawings of wing areas were made before preservation for the purpose of study- ing age-related changes in wing-loading. Wing areas were traced on paper in the field and surface area determinations were made fol- lowing the methods described by Davis (1969a). Linear and wing- area measurements, were taken using a metric dial caliper (0.1 mm) and a polar planimeter (0.1? inches), respectively; the latter meas- ures were converted to metric units (cm?). Binoculars (7 x 35) were used for making observations at Wil- more. At this site ambient light conditions were usually sufficient for making diurnal observations from a distance of approximately five meters without noticeably disturbing the colony. On periodic visits to each nursery, I examined areas beneath roosts for evidence of juvenile mortality. However, this search was hindered at Lost Colony because small young became buried in the wet guano after falling to the cave floor. This problem was not encountered at building nurseries where guano usually dried rapidly under conditions of low relative humidity. Growth Analysis A FORTRAN program, fitting properties of the von Bertalanffy growth model applicable to mark-recovery data (Fabens, 1965), was used in the analysis of post-natal growth. The only basic input requirements of this program are sets of linear or weight data (ar- ranged as paired measurements, and corresponding capture intervals that need not be equal) and a separate input that includes linear size or weight of some known-age individuals (preferably those taken at or near birth). In the analysis that follows, linear growth is expressed by the general formula X=a(1-be*t) and weight by the formula W=[a(1-be*™*) ]*, where X represents the computed length of the forearm or fifth finger, and W is the 2/3 power of the body weight. The quantities a, b, and k in both equations are parameters: a is a measure of the asymptotic size; b is a measure related to the size of the animal at birth; and k is a measure of the intrinsic rate of growth. The values e and t represent the base of the natural loga- rithm and the number of days of growth, respectively. POPULATION STUDIES OF THE CAVE BAT yi A thorough account of the mathematical and physiological the- ory of the model was presented by von Bertalanffy (1960); he cau- tioned that at present quantitative analysis of growth in mammals must be considered provisional. Laird et al. (1965) and Laird (1966) used models based on the Gompertz equation for studying growth of body weight in mammals. However, von Bertalanfly argued that models based on this equation are not only unsuitable for analyses of linear measurements but also lack a physiological basis for appli- cation to study of growth. Curve-fitting of growth data is useful to summarize data for de- scriptive purposes. If a growth curve is derived from a theoretical model, so that parameters have some additional meaning, independ- ent investigations then can be conducted to test the theory on which the model is based. The von Bertalanffy growth curve of the decay- ing exponential type (von Bertalanffy, 1960) has been derived from a model that has physiological significance and provides parameters that are useful for independent testing. The underlying assumption of this model is that growth, in the biological sense, is intimately related to metabolism and is characterized by the highest rates in the early stages of development, decreasing exponentially with time. RESULTS Nursery Roost Environments Nursery “caves” seldom showed significant daily fluctuations in temperature or relative humidity. Hygrothermograph recordings taken at Lost Colony in late spring and summer 1969 revealed little daily change in ambient temperature or relative humidity. A repre- sentative record in the second week of July showed that the cave temperature remained near 20° C and the relative humidity near 100 percent (Fig. 1). Periodic measurements of temperature and relative humidity in the nursery area at the National Gypsum Mine suggest a similar pattern. Much of the gypsum substrate becomes saturated with ground-water following spring and early summer rains at these sites. The accumulation of this water, the presence of seasonally intermittent streams or standing water, and the relatively low air circulation in summer reduce evaporative conditions and contribute to the high relative humidity. Building nurseries, in contrast to cave nurseries showed marked daily variations of temperature and relative humidity. A comparison of temperature recordings and relative humidity at Lost Colony and Wilmore in the second week of July (Fig. 1) demonstrates these striking differences. In this period temperatures at Wilmore varied from 25-35° C, and relative humidity ranged from 40-100 percent. Because Wilmore was occupied by spring transient and a resident summer population, the recordings shown in figure 2 were selected to depict the roost conditions during this period. It is clear that bats 8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY LOST COLONY 7-14 JUL 1969 WILMORE SslSSULIIE9 Fic. 1. A comparison of hygrothermograph records during the nursery period (second week of July) at Lost Colony and Wilmore. On both records the temperature measurements (upper) and relative humidity measurements (lower) represent ambient conditions in the colony irrespective of the thermal influence or vapor pressure changes brought about by the presence of the bats. The diagonal lines indicate the 2400 hour (midnight) of each day. occupying this roost potentially were exposed to widely fluctuating conditions. There is a general seasonal pattern of increasing mean ambient temperatures and decreasing relative humidity from early spring to late summer. Environmental fluctuations may vary considerably depending on the physical structure of the roost, the location of the building or cave with respect to topography, surrounding vegetation, and the facing direction of the entrance. Ambient temperature in small buildings usually fluctuated more than in large structures. Nursery sites in caves, in contrast to those in buildings, were usually located in areas where little, if any, light penetrated beyond the entrance area. In most buildings, however, light was sufficiently bright dur- ing parts of the day to permit observation without the use of arti- ficial sources. At Lost Colony and the National Gypsum Mine no light was evident in the nursery areas but at Wilmore bats occupied dimly lighted areas and the darkest areas were not occupied. Clustered bats modify the microclimate of their nursery roosts. Tendencies of bats to cluster, coupled with the physical features of ee POPULATION STUDIES OF THE CAVE BAT 9 Oh aaa | | | 26 MAR- 2 APR 1970 20 %RH TEMP °C oO Aas eo oO TEMP ?¢ ND oO 25-30 MAY II969 Oo %o RH 20 TENMP=<'C 19-26 JUL 1969 Yo RH 20 | 7-14 AUG 1969 TEMR2 %1C %o RH Fic. 2. Hygrothermograph records showing seasonal pat- terns of ambient temperature and relative humidity at a barn occupied by transient and nursery populations of M. velifer. The diagonal lines indicate the 2400 hour (mid- night) of each day. the roost, combine as important factors in maintaining high roost temperatures. Microenvironmental temperatures recorded in roosts at Wilmore in late June often reached daytime levels exceeding 35° C (Fig. 3). To assess the effect of bats on their roost microclimate, temperature differentials between test probes and control probes were determined (see insert Fig. 3). I have assumed that these differences represent the thermal influence that bats had on their roost microclimate. On 29-30 June these differences varied from 4- 10 12° C depending on the time of day and the number of bats present at any one time. Much of the loft floor at Wilmore was covered with loose straw and broken hay bales to a depth ranging from two to 40 centimeters. The accumulated litter may have been important in providing insulation by trapping heat that was generated by bats OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY \ AMBIENT \ 30 : \S ¢ NES INSS eo eVEIL CINE “(C, pe) O De) O O TEMPERATURE <¢ O 1200 1600 2000 2400 0400 0800 TIME Fic. 3. Microenvironmental temperatures recorded at the Wilmore nursery on 29-30 June 1969 (after most females had given birth). Data are based on hourly temperatures recorded along a vertical gradient in a joist crevice from thermisters positioned at 6 and 18 cm from the ceiling (see Kunz, 1973 for further explanation). One daily record of a control probe (6 cm) and the outdoor ambient temperature are shown for comparison. The insert shows temperature differentials between control sites (bats absent) and _ test sites (bats present). The solid and open triangles at the bottom of the figure denote sunset and sunrise time, re- spectively. roosting on the joists supporting the loft floor. POPULATION STUDIES OF THE CAVE BAT 11 The daily and seasonal fluctuations of relative humidity in build- ing roosts also were subject to modification by bats. In spite of the pattern of relative humidity recorded in the Wilmore barn (Figs. 1 and 2), the presence of clustered bats in the daytime tends to modify the vapor pressure in roosts when, otherwise, it is low. In June and July 1969 and 1970, roosts were examined periodically after adults had departed to feed. These observations revealed that the wood substrate of joists on which the bats roosted was saturated with urine. Instruments were not available that would permit remote micro-determinations, but measurements of relative humidity using a Bendix Psychron positioned at the opening of joist crevices re- vealed substantially higher readings than from the hygrothermo- graph recordings taken in open areas near the joist crevice. It is quite probable that as these wood joists become saturated during periods of occupancy, increases in vapor pressure become compara- ble to the 100 percent saturation recorded in cave nurseries. In early spring, the thermal environment at Lost Colony ap- peared to be unsuitable for occupancy; temperatures varied from a low of 8° C in mid-April to 13° C in mid-May, and little daily variation was evident. These temperatures were considerably lower than those recorded at other spring roosts and that were occupied by transient populations (Kunz, 1971). When individuals first arrived at Lost Colony in early May, they usually selected the warmest sec- tions available. In contrast, Wilmore was subjected to the environ- mental fluctuations of the outside environment. In early spring, roost temperatures occasionally dropped below 0° C but by May daily highs reached 30° C (Fig. 2). Adult Sex Composition of Nursery Roosts Adult male M. velifer accompany females to nursery roosts; this phenomenon is apparently uncommon in other members of the genus. The sexes occur in nearly equal numbers in spring transient roosts and in nurseries until parturition begins (Table 1). Monthly comparisons of the proportion of adult sexes occupying Lost Colony and Wilmore showed that the males occupying these two sites de- clined measurably in May and June. Proportionately fewer males remained in the barn at Wilmore from May through August than occupied Lost Colony during the same period. Samples of bats taken at the National Gypsum Mine in July and August, 1969, revealed that a comparatively large proportion of males occupied this site. Here, many males occupied cool caves and were solitary or roosted in small groups. On 19 August several bats were found scattered throughout the mine at considerable distances from the nursery area; of 39 individuals captured, 38 were adult males. A trap sample on the same date during emergence activity yielded 406 bats, of which 296 were males. 12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY TABLE 1. Seasonal comparisons of adult male Myotis velifer occupying nurseries at Lost Colony (cave) and Wilmore (barn). Lost Colony Wilmore Number of Pooled Percent Numberof Pooled Percent Month samples N Males males samples N Males males Mar a ne nies cea | AQ 23 47.0 5:0) page a 1 4 be 50.0 a 112 sy 45.6 Mays 22555 2 216 106 49.0 4 304 87 28.6 | Ane iy eee 4 696 106 15:2 ys 343 28 G2 | (0) Nees eae 3 574 194 33.8 5 305 24 7.9 Ae ~ 2s 4 687 sy 20.0 4 93 10 LOY Sep. ast = 3 241 48 20.0 Zz 15 bd 13. Oct) 2.28 1 68 65 95.4 I 20 20 ~=100.0 Reproduction MALES Testes and Epididymides.—Seasonal changes in size of testes and epididymides of adults show that testes increase in size follow- ing departure from hibernacula with maximum hypertrophy occur- ring in late August (Fig. 4). These observations concur with those of Krutzsch (1961) and Hayward (1970) based on samples from Texas and Arizona, respectively. Increase in the size of testes corre- sponds to the period of spermatogenesis in summer reported for other vespertilionids (Wimsatt, 1960b, 1969). In early autumn when testes decrease in size, the epididymides become distended with spermatozoa. Maximum engorgement of the cauda epididymides occurs in October and is followed by a decrease in size throughout the winter. Testes are abdominal throughout hibernation and early spring, but in mid-summer (July) they descend into the pigmented sheaths adjacent to the tail in the uropatagium. FEMALES Age at Sexual Maturity—Female M. velifer apparently reach sexual maturity in their first year of life. Thirty-nine females were recovered from 21 May through 21 June 1970, that had been marked as juveniles in the preceding summer; all were either pregnant or lactating. It was not determined whether or not yearling females are inseminated in spring or at other times. Ovulation and Fertilization—Wimsatt (1944, 1945) suggested that ovulation in M. lucifugus probably occurs between the time of departure from hibernation and arrival at nurseries in spring. If fertilization in M. velifer occurs at this time, then this would include the period from late March to late April for populations in this study. In 1970, some females had already departed from hiber- POPULATION STUDIES OF THE CAVE BAT 13 . # eS ies Patio: a+ aN S) lO lO 10 9 10 lO. 9 8 MILLIMETERS O at ver t MONTHS Fic. 4. Seasonal changes in size of testes and cauda epididymides of adult male M. velifer. Dice grams indicate mean, range and 2 SE. Sample sizes are shown between the two graphs. nacula and had arrived at spring roosts by 25 March, although the median departure date from hibernacula was estimated to be mid- April (Kunz, 1971). Gestation and Prenatal Development.—The means for length and width of the right uterine horn (Table 2) are plotted to portray the general period of gestation and pattern of prenatal growth (Fig. 5). Prenatal growth follows a sigmoidal pattern with a character- istic lag phase in the early stages. Throughout this early period of growth, the length and width of the uterine horn were nearly pro- portional. Most of the growth up to this time mainly reflected the increase in the size of the uterus, the placenta, and extraembryonic membranes. Once these tissues are well developed, embryos grow rapidly as evidenced by their elongation, change in the relative pro- portions of the linear dimensions of the uterine horn, and increase in embryo size and weight. Mean weight of the fetus at birth was approximately 3.0 grams; on 29 May, about three weeks before 14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY TABLE 2. Prenatal growth of Myotis velifer, where growth is expressed as the length and width (+ 0.1 mm) of the right uterine horn, weight (+ 0.01 gm) and crown-rump length (+ 0.1 mm) of the fetus (means + SE and ranges are given). Length of right Width of right uterine horn (mm) uterine horn (mm) Date N x SE Range X SE Range 24 Apr ___ 2 286 014 419-40) 19 069 Tee 8 Mays... 1D, ALO F025 ( 2.9- 6.3) 36. 029 ( 2.3- 6.1) 29 Nay ass 9 6 0:99 ( 9.1-17.4) 10.1 0.89 ( 6.4-14.4) 1S Fon 2. fo, 2 Oa (18.8—24.1 ) 136 046 (12.5-14.8 ) ir lame 1 SLR) 1.92 (16.3-26.4 ) 15.8 . 0:61 (14.5-16.9) 20. Fame 22 OF Zo.n 10°94 (20.4-28.0 ) 16.1 (0:50 (14.0-18.2) Weight of embryo (gm) Crown-rump length (mm) Date N x SE Range X SE Range 24 Apr 12 8 May: ____ 12 Bes a _ =e es mes 29) May” —._ 9 043 0.073 (0.05-0.73 ) 10.3 0.98 ( 6.2-15.9) il Akon! ee 15 1.66 0.068 (1.20-2.25 ) 20.7 0.43 (17.9-23.5) li6: jun = te 223 -OEL76 (1.10-3.30 ) 23.6 0.78 (16.4—27.1) Ale qu 2 8 2.80 0.251 (1.90-3.92) 23.1 072 (204260) parturition, mean embryo weight was 0.43 grams, representing only 14.3 percent of the fetal weight at birth. It is clear from these data that most of the prenatal growth occurs in the last three weeks of gestation. The sex ratio of fetuses did not differ significantly from unity. Before late May sex determination was not possible by external macroscopic examination of the fetus. Sex determinations made on and following 29 May revealed 26 males and 22 females (Chi- square, 0.31, P > 0.05). The sexes of five fetuses taken on 29 May could not be determined owing to their exceedingly small size and relatively underdeveloped condition. Samples from various nurseries were taken from late May through August to assess the reproductive condition of females. The data summarized in table 3 represent the number and percentage of pregnant, lactating, and post-lactating females captured in this in- terval. Data for Lost Colony and Wilmore are plotted separately (Fig. 6) and form the basis for the following discussion. A median parturition date was estimated when 50 percent of the reproductive females sampled at each location had produced young. In 1969 this date was estimated to be on 21 June both at Lost Colony and Wilmore. If one assumes that 15 April was the median date for fertilization and 21 June for the median parturition date, then the gestation period for M. velifer is approximately 67 days. If individuals that ——S SS SS ————————— a Sea See SS a POPULATION STUDIES OF THE CAVE BAT 15 30 RIGHT UTERINE HORN ae ta ul LENGTH Li = Be a = 10 O APR MAY JUN Fic. 5. Period of gestation and pattern of prenatal growth of M. velifer expressed as the mean increase in length and width of the right uterine hom (also see table 2). arrived at Wilmore on 26 March were indeed pregnant and partu- rition began soon after 11 June, this would suggest a gestation period of approximately 77 days, assuming that these individuals were the first to give birth. Based on the available data, the most reliable estimate probably lies somewhere between 60 and 70 days. Wimsatt (1945) estimated that the gestation period in M. lucifugus ranged from 50 to 60 days and Pearson et al. (1952) reported the gestation period for Plecotus townsendii in California as 56 to 100 days. Parturition—tThe parturition period at Lost Colony and Wil- more extended over a period approximating four weeks in 1969 and corresponded closely to the range of departure dates from hiber- nacula. Periodic examination of the number of young occupying these two nurseries in June and July supported the contention that most of the young are born within a short period. Nearly 80 percent of the births occurred in a two-week period from 14 to 28 June. On 8 June 1969 no young were present in the nursery at Lost Colony, although approximately 100 and 400 neonates were present on 16 and 17 June, respectively. These data indicate that approximately 300 young were born within a 24-hour period between the nights of 16 and 17 June. Ten days later, on 27 June, there were nearly 4500 16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY TABLE 3. Seasonal changes in the reproductive condition of adult female Myotis velifer occupying nursery colonies in Kansas.2 Pregnant Lactating Post-lactating Date N Percent WN _ Percent WN _ Percent Location 22-23 May 1970 48 100.0 Wilmore 28 May 1969 230 100.0 Englewood» 28-29 May 1969 69 100.0 Wilmore 30-31 May 1969 96 100.0 Lost Colony 8-9 Jun 1969 124 100.0 Lost Colony 9-10 Jun 1969 10° 91.0 1 9.0 Nat'l Gypsum Mine 10-11 Jun 1970 42 100.0 Wilmore 16-17 Jun 1969 So.) 6470 1d Ss Lost Colony 18 Jun 1969 56 96.5 2 oD Nat’] Gypsum Mine? 18-19 Jun 1969 blo S162 VIG= 236 Belvidere 19-20 Jun 1970 147 666 68 33.4 Wilmore 21 jun 1970 266° “7451 = 90) 72533 Lost Colony» 21-22 Jun 1970 AB: 50.0 " 4p. 50:0 Nat'l Gypsum Mine 27-28 Jun 1969 Is 12:0 95 “68:0 Lost Colony 28-29 Jun 1969 € 1.0" “92 "930 Wilmore 29-30 Jun 1969 210 100.0 Wilmore 7-8 Jul 1969 122 100.0 Lost Colony 8-9 Jul 1969 107 100.0 Wilmore 9-10 Jul 1969 27 100.0 Wilmore 18-19 Jul 1969 153 100.0 Lost Colony 19-20 Jul 1969 10 100.0 Wilmore 27-28 Jul 1969 78 742 27 25.8 Lost Celom 28-29 Jul 1969 5 83.4 1 16.6 Wilmore 31 Jul 1969 101 60.4 66 39.6 Double Entrance S» 31 Jul — 1 Aug 1969 196 77.5 57 22.5 Protection 6-7 Aug 1969 42 37.5 70 62.5 Lost Colony 7-8 Aug 1969 3 15.7 16 84.3 Wilmore 8-9 Aug 1969 6 109 49 89.1 Wilmore 10 Aug 1969 31 14.9 177 85.1 Lost Colony» 16-17 Aug 1969 93 100.0 Lost Colony 17-18 Aug 1969 18 100.0 Wilmore 19 Aug 1969 1 0.4 210 99.6 Natl Gypsum Mine 20 Aug 1969 189 100.0 Protection” 27-28 Aug 1969 11 100.0 Wilmore 31 Aug — 1 Sep 1969 136 100.0 Lost Colony 2 Non-reproductive females were excluded because they usually represented less than one per cent of any sample. b Roost sample. POPULATION STUDIES OF THE CAVE BAT Li Boor COLONY LACTATION = 43 DAYS Sen lOO Y i Wf = PREGNANT LACTATING POST -LACTATING Li] YT ys Grae N=319 as LJ QO. O JUN JUL AUG WILMORE LACTATION = 43. DAYS lOO _ = PREGNANT POST -LACTATING O N=95 fe 50 LJ fle JUN JUL AUG Fic. 6. A comparison of the summer reproductive patterns of female M. velifer expressed as the percentage of pregnant, lactating, and post-lactating individuals sampled at Lost Colony and Wilmore. The median parturition date is indicated when 50 percent of the reproductive females had already produced young. The interval between the median parturition date and the median cessation date of lactation represents an estimated individual lactation period. young ranging from one day to nearly two weeks of age. If 50 per- cent of the young were born by 21 June this would mean that in the four-day period from 18 to 21 June, an average of more than 500 young were born per day. Similar observations were made at Wil- more. A trap sample taken at an entrance to the National Gypsum Mine on the night of 22 June 1970 (45 pregnant and 45 lactating females) revealed a comparable median parturition date. Some authors have suggested that yearling females of some spe- 18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY cies give birth later in the season than older bats (see Griffin, 1940; Pearson et al., 1952; Davis and Hitchcock, 1965). To test the null hypothesis that there was no difference in parturition dates of year- ling or adult M. velifer, I examined females that had been marked at Lost Colony and Wilmore in 1969 and recovered there between 19 and 21 June 1970. Of 22 adults and 28 yearlings recovered in this period, six adults (27.2%) had already given birth or were lactating and only four yearlings (14.8%) had given birth. A ¢ test for the equality of these two percentages was non-significant (t=1.14, P> 0.05) thereby supporting the null hypothesis. Actual parturition was not observed in the field, although studies conducted in captivity (unpublished observations) concur with those of Twente (1955b) and Wimsatt (1945, 1960a) for M. velifer and M. lucifugus, respectively. Fresh paper was placed beneath nursery areas at Wilmore in an attempt to determine if the placentae were dropped or eaten. The fact that few were found beneath roosts occupied by several hundred young is circumstantial evidence that placentaphagia is practiced by M. velifer. Nursing—Soon after parturition, females begin to nurse their young. A noticeable decrease in the intensity of audible vocaliza- tions was observed between 0800 and 1000 and again between 1500 and 1800 in the Wilmore barn. I have interpreted this to indicate two nursing periods; one follows the return of females in morning and another occurs in late afternoon. The evidence is circumstantial, but it seems reasonable that a reduction in vocalization would occur when young are nursing. Also, between 1100 and 1300 hours I ob- served a large proportion of adult females occupying open-faced joists adjacent to the areas mostly occupied by both young and adults at other times. Based on samples of young and the amount of milk found in their stomachs during the daytime, Davis et al. (1962) suggested that female Tadarida brasiliensis have two daily nursing periods. In the first weeks following parturition, many female M. velifer return to their young within an hour or two after emergence; whether or not nursing takes place at this time was not established. When females departed in the evening, little milk could be ex- pressed from the mammary tissue, whereas females returning to the roost after 0400 appeared to have fully distended mammary glands. Little is known concerning the length of time females nurse their offspring. The lactation period may be approximately 43 days (or six weeks) if one assumes that the time interval between the esti- mated median parturition date and the median cessation date of lactation represents the average for an individual (Fig. 6). Most young begin to forage well before the age of six weeks (see section on Growth and Development). And, evidence based on captures of POPULATION STUDIES OF THE CAVE BAT 19 TaBLeE 4. Annual reproductive effort of female Myotis velifer. Percentage of Reproductive reproductive Date Females females® females 20-31 May 443 443 100.0 1-10 Jun 180 77 98.4 11-20 Jun 242, 240 99.2 21-30 Jun 861 861 100.0 Total 1.726 1,721 99.9 a Pregnant and lactating females (table 3) sampled from late May through June; reproductive activity (pregnancy and lactation) was determined by pal- pation. foraging young suggests that they continue to nurse after they have become volant. Reproductive effort—Yearling and adult females captured at all nurseries were pooled at 10-day intervals from 20 May through 30 June for an analysis of annual reproductive effort (Table 4). These data, coupled with the fact that only a few barren females were found, indicate that most adult female M. velifer in the population are reproductive and also that females reach sexual maturity in their first year. If non-reproductive females remain solitary or occupy other sites separate from nurseries then the annual reproductive ef- fort of 99.9 percent may be an overestimate. Some pregnant females, which had aborted or lost their offspring at birth, may have been considered as non-reproductive individuals upon external examina- tion. If these individuals were counted as non-reproductives, but in reality had been parous, then the overestimate resulting from the possibility of a greater proportion of non-reproductives at other locations is offset. Growth and Development GROWTH PATTERNS Growth curves and descriptive parameters based on 90 paired records for body weight, forearm, and fifth finger are shown in fig- ure 7. Relative growth is expressed as the percentage of body weight and size of adults from samples taken in mid-August (Fig. 9). For linear measures (forearm and fifth finger), relative growth is based on curves shown in figure 7. Relative changes in body weight were based on sample means shown in figure 8 and tables 5 and 6. Data for both sexes were combined in the analysis because there were no significant differences in weight or linear measures in the early stages of post-natal development. Unless otherwise indicated, linear measurements are based on values extrapolated from the computed curves. It should be understood that these curves repre- 20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY TABLE 5. Body weight, wing size, and wing-loading of known age, pre-volant juvenile Myotis velifer (means and standard errors ) .2 Forearm Fifth finger Age (mm) (mm ) Weight (gm) Wing area (cm?) wing loading (days) N» x SE x X X (gm/cm?) 1-2 so. 17-0 0:09. 19:0 O11) 332 (03> 1075. Olas .298 Let? 3 26," 30:5. “021 .32:55 026 76 0:12) wl 00 Oso 245 21-22 26 413 029 485 039 106 0.21 82.58 1.38 128 a Samples are based on recaptured individuals that were initially marked when approximately one to two days old. b Each sample size for wing area was 10. sent best fit approximations to the observed data, and that variation due to errors in measurement and stochastic properties are inherent. Bats that had been marked as neonates seldom were recovered in the roost area more than twice. Of 175 neonates marked at Lost Colony in 1969, 90 recoveries of 64 individuals were made. Loss of bands may account for a low recovery owing to the fact that bands may slip over the wrist of neonates before wing dimensions reach sufficient size to hold them in place. However, only two lost bands later were found beneath the roost. Apparent “loss of bands” may occur if mothers move their young following disturbance (see Roer, 1968; Fenton, 1969a; Davis, 1970), and account for a low recovery of non-volant young. After young became volant, the probability of recovering them (while foraging or at entrances) is even lower. On the day of birth the calculated lengths of the forearm and fifth finger were 16.0 and 18.0 mm, respectively. (The forearm meas- ured approximately 34% of the adult size, whereas the fifth finger was only 29%.) By three weeks of age the forearm exceeded 80 per- cent of the adult size and by the fourth week, when most of the young had begun to forage, the forearm (43.0 mm) had reached 90 TABLE 6. Seasonal changes in body weight of volant, juvenile Myotis velifer (means and standard errors ). Males Females Body weight Body weight Date N x SE N X SE 18 Jul 1969 17 10.4 0.26 16 10.4 0.34 27 Jul 1969 49 10.7 0.12 58 10.7 6 pa 6 Aug 1969 59 9.8 Or 40 10.2 0.15 16 Aug 1969 34 10.2 0.13 38 10.4 0.14 31 Aug 1969 36 11.8 0.11 53 ii) 0.09 7 Sep 1969 35 12.0 O- Ta 52 12.4 0.07 18 Oct 1969? 55 14.7 0.17 = =e es IS 2Oct, 1969 30 15.3 0.25 4 Taken in a hibernaculum. POPULATION STUDIES OF THE CAVE BAT 21 60 X= 61.5(1-0.725e7 9.0902) 5O op) Ge Lu ce f 40 X= 47.4 (1- 0.685 e~ 9-0689t) a = S Ze (op) z S I 30 0.05), the above data suggest that gestation may have been further delayed at Lost Colony. It is of interest that the population densi- ties at each nursery in 1970 were reduced from their 1969 levels; the estimated population at Lost Colony in June 1970 was approximately 2000 adult females, compared to approximately 5000 in 1969 (Kunz, 1971). At Wilmore, the 1970 population also was lower than in 1969 but the reduction was less than at Lost Colony. Considering the fact that bats can increase the micro-temperature of their own roost and that populations in 1970 were reduced from the previous year, the size of the resident population may influence the duration of gestation. Smith (1956) reported significant differences in the weight gains 32 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY of pregnant Myotis lucifugus in two different years. The fact that she found no correlation between ambient temperature and weight gain in the two years suggests that the observed difference in weight gain may be accounted for by the difference in population size in the two years of study (see Smith, 1954). Clustering behavior and maintenance of high metabolic levels usually are characteristic of females late in pregnancy (Smith, 1956; Stones and Wiebers, 1967). Herreid (1967) has shown that energy cost is less among individuals of Tadarida brasiliensis occupying large clusters than those in small clusters or singles. Perhaps when a nursery colony is reduced in size, and fewer bats are present to cluster, more energy is expended in maintaining homeothermy, and proportionately less is available for development. Additional studies involving examination of preg- nant females in spring are needed to fully assess the influence of clustering behavior, population size, and ambient temperature on rates of gestation. Daily production of milk probably reaches maximal levels in the second or third week following parturition and corresponds to the increasing size of the young and the increased food consumption at this time (Kunz, 1973). Davis et al. (1962) suggested that female Tadarida brasiliensis may produce an amount of milk equivalent to 16 per cent of their lactating body weight within a 24-hour period. Based on the 12 gram average weight of lactating Tadarida (Short, 1961), daily milk production per female should be in excess of 1.9 grams. Since Davis et al. (1962) also reported an average weight increase from daily food consumption of only one gram, there is little doubt that their estimation of milk production was too high or, alternatively, that their evaluation of food consumption was too low. Although I have no comparable data on milk production for M. velifer, my estimates on daily food consumption for lactating females (Kunz, 1973) suggest that their estimate of food consumption for Tadarida (a bat comparable in size to M. velifer) was much too low. Growth and Development Few studies provide data on growth of known-age bats under natural conditions. Because the forearm has been the most consist- ently measured element in growth studies, comparisons of the rela- tive stages of development of several species were made at birth and at the age of six weeks to illustrate the similarity in growth. The forearm of M. velifer at birth and at six weeks are 34 and 96 percent, respectively, of adult size. Proportions based on data given for Antrozous pallidus (Davis, 1969) at birth and at six weeks are 32 and 96 percent, respectively. Corresponding values for Plecotus townsendii are 38 percent at birth and 98 percent at the age of six weeks (Pearson et al., 1952). Similar proportions based on data presented by Kratky (1970) for M. myotis are 37 percent at birth POPULATION STUDIES OF THE CAVE BAT 33 and 98 percent at six weeks. This evidence illustrates the similarity of developmental stages at the time of birth and at the time when young bats probably are weaned. Pearson et al. (1952) reported an absolute growth rate of 0.85 mm per day for the forearm of Plecotus townsendii in the first 16 days of growth, and Davis et al. (1968) reported that some indi- vidual Eptesicus fuscus showed increases in forearm length in early stages of growth up to 2.6 mm per day. Based on data extrapolated from growth curves presented by the above authors, I have com- puted for comparison absolute growth rates for the forearm in the first 10 days. Rates of 1.20 mm per day for Plecotus townsendii and 1.45 for Eptesicus fuscus generally are comparable to the value of 1.56 determined for M. velifer. At birth, the proportion of body weight of young to the weight of post-parturiant female M. velifer (25.8%) closely approximates the proportions for M. lucifugus (24.6%) and M. nigricans (25.6%) recorded by Wimsatt (1960b) and Wilson (1971), respectively. In the first two months of post-natal life, maximum body weight of M. velifer is attained at the age of three to four weeks. Loss of weight after this time seems to be common as it has been observed both in field and laboratory studies (Short, 1961; Kleiman, 1969; Davis, 1969; Kratky, 1970). Short suggested that this decrease may be accounted for by loss of body fats after young have become volant. The fact that similar decreases have been observed under conditions of laboratory confinement (Kleiman, 1969) suggests that nutritional conditions at time of weaning are important. Thus weight loss under natural conditions in this period may be accounted for by a combination of factors related to the weaning period and early flight activity, including reduced milk consumption, expendi- ture of accumulated fats, and inefficient insect capture (Kunz, 1973). Perhaps the deposition of fat in early developmental stages, and prior to the initiation of flight, is an important strategy that helps sustain juveniles through the weaning period. Davis and Hitchcock (1965) observed that juvenile M. lucifugus continued to nurse after they had become volant and similar obser- vations have been reported for M. mysticinus and M. daubentoni (Nyholm, 1965). The fact that volant young M. velifer contained both milk and the remains of insects (Kunz, 1973) suggests that weaning does not take place abruptly at the time young initiate foraging activity. Vocalizations that Gould (1971) recorded suggest that weaning may be initiated by the withdrawal behavior of young (weaning in M. lucifugus and Eptesicus fuscus apparently took place at the age of three weeks under conditions of laboratory con- finement). Kleiman (1969) observed that weaning of Eptesicus serotinus and Pipistrellus pipistrellus in captivity began in the third or fourth week but was not completed until two to three weeks later. 34 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Unusually long nursing periods have been reported for some bats reared in captivity (Eisentraut, 1936; Kleiman, 1969; and others). Differences in weaning time between captive bats and those under natural conditions may be due to artifacts resulting from nutritional deficiences existing under laboratory confinement. Juvenile M. velifer began feeding on insects near the time when permanent teeth were fully erupted (approximately 3-4 weeks); however, adult-like feeding behavior and success (Kunz, 1973) was not achieved until permanent teeth were fully developed (approxi- mately 7-8 weeks). These observations are in contrast to those of Kratky (1970) who reported that permanent teeth in M. myotis were fully erupted in 30 days and that they did not begin to feed on insects until the age of 40 days (approximately 7 weeks). Con- tinued linear growth and stabilization of body weight soon after foraging is initiated in M. velifer, results in a low wing-loading, and probably is important in leading to the development of successful feeding behavior. Two generations of hair in the juvenile pelage were evident— over-hair, that began development in the prenatal period, and under- hair which appeared at approximately one week. A second gener- ation apparently began as early as seven days as evidenced by darkening of the skin (melanin deposition). A juvenile molt was initiated at approximately six weeks (when young bats had become nutritionally independent) and was not completed until September (10 weeks). These observations corre- spond to those reported by Mazak (1963) who noted that termina- tion of hair growth in juvenile M. myotis coincided with the time that external measurements reached adult size. Although parturition in this study occurred over a period of four weeks, there was no evidence of age segregation beyond the fact that the young less than two days old did not join older bats in forming clusters. The fact that juveniles less than two days of age were found scattered in peripheral areas suggests that either females intentionally deposit newborn young away from the densest part of the colony, or that the older young are more mobile and have a strong tendency to cluster. It seems likely that mothers would be able to locate their newborn easily if the latter were solitary, espe- cially before the young have developed a reciprocal vocal response to their mothers’ calls. Dwyer (1963) suggested that the behavior of females to deposit young on the periphery of nursery clusters may reduce the mortality of neonates by decreasing the probability of being dislodged by larger, more active individuals. Gould (1971) theorized that if bats remained organized into different age groups corresponding to the time of their birth, such behavior also may facilitate mothers recognizing their young as they returned from feeding. POPULATION STUDIES OF THE CAVE BAT 35 Environment and Growth Nursery sites that can be maintained at temperatures near ther- mal neutrality probably are important to the promotion of rapid growth in young. Also, less energy would be required for the main- tenance of homeothermy at such temperatures. The selection of warm nursery roosts along with clustering behavior probably con- tribute measurably to a reduction in energy cost to individuals (Herreid, 1963, 1967). Cluster formation in nurseries not only pro- vides a potential avenue for conductive and convective heat ex- change for the enhancement of growth (Dwyer and Hamilton-Smith, 1965) but also in promoting efficient digestion and assimilation of food (Twente, 1955b; Davis et al., 1962). One of the initial designs of this study was to examine the re- lationships between growth and roost microclimate at Lost Colony and Wilmore. Efforts to obtain comparative temperature data at Lost Colony, however, were not completely successful. Measure- ments that were comparable are those from hygrothermograph rec- ords (Fig. 2). These data demonstrate that cave temperatures at Lost Colony remained near 20° C. Because bats at Lost Colony maintained high activity levels in daytime, I assumed that cluster temperatures (body surface temperatures ), at least during and sub- sequent to parturition, would be similar to those recorded at Wil- more. If this assumption is valid, and individuals at Lost Colony had body temperatures in daytime between 32 and 37° C, body temperature-ambient temperature differentials would be expected to vary from 12-17° C. At Wilmore daytime differentials ranged from 5-12° C in the nursery period (see insert Fig. 3). If, on the average, higher temperature differentials are needed to maintain cluster tem- peratures at Lost Colony, then the energy cost of occupying Lost Colony would be greater than for occupying Wilmore. And, any difference in energy cost in occupying these two sites may be re- flected in different growth rates, reproductive periods, or quantities of daily food consumption. In spite of the apparent thermal differences at Lost Colony and Wilmore a seasonal comparison of reproductive parameters at these two sites (Fig. 6) suggests that there were no differences in repro- ductive chronology. The fact that parturition began at approxi- mately the same time, the median parturition dates were the same and the lactation periods were comparable further suggests that post-natal growth rates may have been similar. If early flight of young bats can be used as a measure of com- parable growth, then I would suggest that there was no measurable difference in growth rates at these two nurseries in 1969. On 7-8 July and 8-9 July 1969, when young bats were first captured while departing to forage, approximately the same number of individuals was trapped at entrances in a comparable period or time; five were 36 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY taken at Lost Colony and six at Wilmore. Because both nurseries were occupied initially by the same number of young (approxi- mately 5000), these data suggest that the same proportion of indi- viduals in these two colonies reached flight stage at the same time. Mean forearm lengths of the earliest volant young trapped at Lost Colony (43.6 + 0.56) and Wilmore (43.2 + 0.70) are not signif- icantly different (t=0.54, P> 0.05). Based on observations of Pearson et al. (1952) that Plecotus townsendii grew more rapidly at warmer temperatures, and the fact that Herreid (1963) recorded increases in cave temperatures throughout the nursery period, Herreid (1967) postulated that bats born in the early part of the parturition period would grow more slowly than those born later. To test this hypothesis groups of neo- nates that were born at Lost Colony both in the early and late stages of the parturition period were individually marked. One group con- sisted of individuals taken on 17 June, when approximately 300 (16% of the total juvenile population) had been delivered. The second group was taken on 27 June, after approximately 4500 (more than 90%) young were present. If growth rates differed according to the time of birth, I expected to find differences in the first three weeks since the most rapid growth occurs during this period (Fig. 8). In- dividuals marked as neonates on the above dates were recovered at intervals of 10 and 20 days after initial marking. Of all paired com- parisons made (using a ¢ test) no significant differences in linear size or weight were found (P> 0.05). Although differences in growth may be evident in later periods, sample sizes (less than five on any one capture date) at these times were inadequate for making valid comparisons. Although these data fail to support the hypoth- esis presented by Herreid (1967), they do not disprove the premise that temperature may have a measurable effect on development. Additional studies are needed to examine the influence of en- vironmental factors and roost temperatures on growth and develop- ment of bats. Studies using the von Bertalanfly growth analysis (Fabens, 1965) may be useful for comparing growth patterns at dif- ferent environmental temperatures. According to von Bertalanffy (1960), growth occurs most rapidly where protein turnover is high. Interspecific and intraspecific growth comparisons could be made by comparing catabolic constants in roosts subject to different thermal environments—the higher the catabolic constant, the faster the growth rate. The catabolic constant is determined as 3k, where k is the intrinsic rate of growth (see Methods and Materials). An ana- bolic constant also may be computed (Fabens, 1965), but because it is a function of body weight, and computed curves may not corre- spond to observed measures, this determination may be less useful. POPULATION STUDIES OF THE CAVE BAT 37 Geographic Variation Bats occurring at northern latitudes and higher altitudes usually give birth later than do those in more southerly areas and at lower elevations. Differences in temperature, abundance of resources, and variation in periods of hibernation all have been implicated as in- fluencing factors. Few studies on M. velifer are available that per- mit a broad geographic comparison of reproduction. Hayward (1970) assumed that the average birth date of populations in Ari- zona was 17 June, although he presented little evidence to substan- tiate his assumption. Davis et al. (1962), however, reported that parturition in M. velifer occurred several weeks before parturition of Tadarida brasiliensis in southern Texas. Examination of dates re- corded for Tadarida would suggest that most of the births for M. velifer occurred in early June. This and the fact that young report- edly were flying in Texas by late June suggest that the average parturition period for that latitude was two to three weeks sooner than in Kansas. Owing to the wide distribution of M. velifer it would be of interest to compare reproduction in the northern popu- lations with those occurring at lower latitudes, especially in areas where the availability of resources may be less influenced by ther- mal seasonality. SUMMARY A study detailing reproduction, growth and development of the cave bat, Myotis velifer, was conducted in south-central Kansas from March 1968 through January 1971. Reproductive and growth pat- terns were examined at two physically and thermally contrasting nurseries (barn and cave) that included a temporal analysis of ges- tation, parturition and lactation. Post-natal growth was determined from mark-recapture data. Reproductive activity of males was ex- amined throughout an annual cycle. Adult males and females occurred in approximately equal num- bers at transient and nursery roosts in spring. By early June the adult sex composition at the nursery sites was predominantly female. A comparison of the sex composition of adults occupying a cave and a building revealed that proportionately fewer males occupied the building. These differences may be related to the intolerance of males for high roost temperatures in buildings. Testes began increasing in size in June and reached maximum hypertrophy in late August. After late August and early September the testes decreased in size and remained in an arrested state until the following summer. In early autumn the epididymides increased in size and became maximally distended with spermatozoa by mid- October. Most copulations occurred in October. Ovulation probably occurs in April when bats depart from hiber- 38 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY nacula. The gestation period was 60 to 70 days and most of the pre- natal growth occurs in the latter three weeks of gestation. In 1969, parturition began in the second week of June and extended for approximately four weeks with a median parturition date of 21 June. Nearly 80 percent of all young were born in a two week period from mid to late June. Females apparently are reproductive in their first year with no significant tendency for them to give birth later than older females. No apparent differences in growth rates were evident in bats occupying a cave and a building nursery. A comparison of reproductive parameters at two sites in 1969 and 1970 revealed that parturition in 1970 was later than in 1969 suggesting that cooler temperatures and a reduced population size may have contributed to this observed difference. Post-natal growth followed a decaying exponential model. The forearm and fifth finger lengths at birth were 34 and 29 percent of adult size, respectively. Growth occurred rapidly in the first three weeks in which time the above linear dimensions exceeded 80 per- cent of adult size. By the fourth week when young bats began to forage, the forearm had reached 90 percent of adult size. The fifth finger did not reach equivalent proportions until the fifth or sixth week. Adult linear size was achieved at the age of 9 to 10 weeks (late August and early September). Young bats at birth weighed 3.0 grams (25.8% of adults); by the time they were three weeks of age they exceeded 85 percent of adult weight. By the fourth week their weight stabilized between 82 and 86 percent of adult weight until fat deposition began in late August. Flight was initiated within the confines of the nursery roost at the age of three weeks. When bats began to forage permanent teeth were fully erupted. Weaning occurred at approximately six weeks when linear dimensions had reached 90 percent of adult size. Juve- nile pelage increased in density until the age of three to four weeks. A juvenile molt was initiated at the age of six weeks (when young became nutritionally independent) and was completed near the time that external measurements reached adult size. The importance of clustering behavior and the maintenance of roost temperatures were discussed with respect to successful repro- duction and growth. POPULATION STUDIES OF THE CAVE BAT 39 APPENDIX Name Coordinate (N, W) County Belvidere (Barn) ..._______ geek ae 37°30'35",, “99°038'56” Kiowa Double Entrance S Cave __........ Si 0430? 9920225" Comanche Emglewood (Bar) 31-07302, 100°03'03”” Clark MostsColony Cave: 37°0PAG6”, 98°59'06” Barber National Gypsum Mine _.....- 37°21'46”, 98°56’47” Barber Protecttome (Bai) 5 31 20325 9928 53" Comanche Wymimere (iparl) 2 ot 19°37", 99°09’ 16” Comanche 40 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY LITERATURE CITED ASDELL, S. A. 1964. Patterns of mammalian reproduction. Cornell Univ. Press, Ithaca, New York, 2nd ed., 670 pp. Barsour, R. W., and W. H. Davis 1969. Bats of America. Univ. Press of Kentucky, Lexington, 286 pp. CarTER, D. C. 1970. Chiropteran reproduction. Pp. 233-246, in About bats (B. H. Slaughter and D. H. Walton, eds.), Southern Methodist Univ. Press, Dallas. CHRISTIAN, J. J. 1956. The natural history of a summer aggregation of the big brown bat, Eptesicus fuscus fuscus. Amer. Midland Nat., 55:66-95. CONSTANTINE, D. G. 1957. Color variation and molt in Tadarida brasiliensis and Myotis velifer. J. Mamm., 38:461-466. Davis, R. 1969. Growth and development of young pallid bats, Antrozous pallidus. J. Mamm., 50:729-736. 1970. Carrying of young by flying female North American bats. Amer. Midland Nat., 83:186-196. Davis, R. B., C. F. Herre, II, and H. L. SHort 1962. Mexican free-tailed bats in Texas. Ecol. Monogr., 32:311-346. Davis, W. H., and H. B. Hrrcucock 1965. Biology and migration of the bat, Myotis lucifugus, in New Eng- land. J. Mamm., 46:296-313. Davis, W. H., R. W. Barsour, and M. D. HAssELL 1968. Colonial behavior of Eptesicus fuscus. J. Mamm., 49:44-50. Dunnican, P. B., and J. H. Fircu 1967. Seasonal movements and population fluctuations of the cave bat (Myotis velifer) in south-central Kansas. Trans. Kansas Acad. Sci., 70:210-218. Dwyer, P. D. 1963. The breeding biology of Miniopterus schreibersi blepotis (Tem- minck ) (Chiroptera) in northeastern New South Wales. Australian J. Zool., 11:219-240. 1966. The population pattern of Miniopterus schreibersii (Chiroptera) in north-eastern New South Wales. Australian J. Zool., 14:1073-1137. Dwyer, P. D., and E. HAMILTON-SMITH 1965. Breeding caves and maternity colonies of the bent-winged bat in south-eastern Australia. Helictite, 4:3-21. EISENTRAUT, M. 1936. Zur Fortpflanzungsbiologie der Fledermause. Zeit. Morph. Okol. Tiere, 31:27-63. 1937. Die Wirkung niedriger temperaturen auf die Embryonalent Wi- klung bei Fledermausen. Biol. Sentralblatt. Leipzig, 57:59-74. FABENS, A. J. 1965. Properties and fitting of the von Bertalanffy growth curve. Growth, 29 :265-289. FENTON, M. B. 1969a. The carrying of young by females of three species of bats. Cana- dian J. Zool., 47:158-159. 1969b. Summer activity of Myotis lucifugus (Chiroptera: Vespertilionidae ) at hibernacula in Ontario and Quebec. Canadian J. Zool. 47:597- 602. POPULATION STUDIES OF THE CAVE BAT 4] 1970. The deciduous dentition and its replacement in Myotis lucifugus (Chiroptera: Vespertilionidae ). Canadian J. Zool., 48:817-820. Grover, T. D., and D. H. Youne 1963. Temperature and the production of spermatozoa. Fertil. Steril., 14:441-450. GouLp, E. 1971. Studies of maternal-infant communication and development of vocalizations in the bats Myotis and Eptesicus. Comm. Behav. Biol. (Part A), 5:263-313. GriFFIN, D. R. 1940. Notes on the life history of New England bats. J. Mamm., 21:181- 187. GUTHRIE, M. J. 1933. The reproductive cycle of some cave bats. J. Mamm., 14:199-216. Haywaprp, B. J. 1970. The natural history of the cave bat Myotis velifer. Res. Sci., West- ern New Mexico Univ., 1:1-74. HeERRED, C. F., II 1963. Temperature regulation of Mexican free-tailed bats in cave habitats. J. Mamm., 44:560-573. 1967. Temperature regulation, temperature preference and tolerance, and metabolism of young and adult free-tailed bats. Physiol. Zool., 40:1-22. HuMPHREY, S. R. 1971. Population ecology of the little brown bat Myotis lucifugus, in Indiana and north-central Kentucky. Ph.D. dissertation, Oklahoma State Univ., 138 pp. KLEMAN, D. G. 1969. Maternal care, growth rate and development in the noctule (Nycta- lus noctula), pipistrelle (Pipistrellus pipistrellus) and serotine (Ep- tesicus serotinus) bats. J. Zool., 157:187-211. Kratky, J. 1970. Postnatale Entwicklung des Grossmausohrs, Myotis myotis ( Bork- hausen, 1797). Vest. Ceskoslov. Spol. Zool., 33:202-218. Krutzscu, P. H. 1961. The reproductive cycle in the male vespertilionid bat Myotis veli- fer. Anat. Rec., 139:309 (abstract ). Kunz, T. H. 1971. Ecology of the cave bat, Myotis velifer, in south-central Kansas and northwestern Oklahoma. Ph.D. dissertation. The University of Kansas, 148 pp. 1973. Feeding ecology of a temperature insectivorous bat (Myotis velifer). M.S. submitted for publication. Larrp, A. K. 1966. Postnatal growth of birds and mammals. Growth, 30:349-363. Lairp, A. K., S. A. TYLER, and A. D. BARTON 1965. Dynamics of normal growth. Growth, 29:233-248. Mazak, V. 1963. Hair growth in large mouse-eared bat, Myotis myotis myotis Bork- hausen, 1797 (Mammalia, Chiroptera), during its praenatal and early postnatal life. Vest. Ceckoslov. Spol. Zool., 27:234-242. MILLER, G. S., Jn., and G. M. ALLEN 1928. The American bats of the genera Myotis and Pizonyx. Bull. U.S. Natl. Mus., 144:viiit218 pp. 42 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY MILLER, R .E. 1939. The reproductive cycle in male bats of the species Myotis lucifugus and Myotis grisescens. J. Morph., 64:267-295. NyYHoLM, E. S. 1965. Zur Okologie von Myotis mystacinus (Leisl.) und M. daubentoni (Leisl.) (Chiroptera). Ann. Zool. Fenn., 2:77-123. Orn, RoE: 1970. Development: Prenatal and Postnatal. pp. 217-231, in Biology of bats (W. A. Wimsatt, ed.), Vol. 2, Academic Press, New York. PEARSON, O. P., and M. R. Korrorp, and A. K. PEARSON 1952. Reproduction of the lump-nosed bat (Corynorhinus rafinesquei) in California. J. Mamm., 33:273-320. Racey, P. A. 1969. Diagnosis of pregnancy and experimental extension of gestation in the pipistrelle bat. J. Reprod. Fertil., 19:465-474. Rice, D. W. 1957. Life history and ecology of Myotis austroriparius in Florida. J. Mamm., 38:15-32. Roker, H. 1968. Nehmen die Weibchen des Mausohrs, Myotis myotis (Borkhausen ) , ihr Neugeborenes auf ihren Nahrungsflugen mit? Zeit. Tier- psychol., 25:701-709 (English summary ). Roer, H., and W. EcsBAEK 1969. Uber die Balz der Wasserfledermaus (Myotis daubentoni) (Chiro- ptera) in Winterquartier. Lynx, 10:85-91 (English summary). SHort, H. L. 1961. Growth and development of Mexican free-tailed bats. Southwest- ern Nat., 6:156-163. SLUITER, J. W. 1961. Sexual maturity in males of the bat Myotis myotis. Proc. Kon. Ned. Akad. Wet., Amsterdam, C, 64:243-249. SMITH, E. 1954. Studies on the life history of non-cave dwelling bats in northeastern Ohio. Ohio J. Sci., 54:1-12. 1956. Pregnancy in the little brown bat. Amer. J. Physiol., 185:61-64. STONES, R. C., and J. E. WIEBERS 1967. Temperature regulation in the little brown bat. Myotis lucifugus. Pp. 97-109, in Mammalian hibemation III (K. C. Fisher et al., ed.), American Elsevier, New York. TINKLE, D. W., and W. W. MILsTEAD 1960. Sex ratios and population density in hibemating Myotis. Amer. Midland Nat., 63:327-334. TINKLE, D. W., and I. G. PATTERSON 1965. A study of hibernating populations of Myotis velifer in northwest- ern Texas. J. Mamm., 46:612-633. TWENTE, J. W., JR. 1955a. Aspects of a population study of cavern-dwelling bats. J. Mamm., 36 :379-390. 1955b. Some aspects of habitat selection and other behavior of cavern dwelling bats. Ecology, 36:706-732. VAUGHAN, T. A. 1959. Functional morphology of three bats: Eumops, Myotis, Macrotus. Univ. Kansas Publ., Mus. Nat. Hist., 12:1-153. 1970. Flight patterns and aerodynamics. Pp. 195-216, in Biology of bats (W. A. Wimsatt, ed), vol. 1, Academic Press, New York. POPULATION STUDIES OF THE CAVE BAT 43 VON BERTALANFFY, L. 1960. Principles and theory of growth, Pp. 137-259, in Fundamental as- pects of normal and malignant growth (W. W. Nowinski, ed.), Elsevier Publ. Co., Amsterdam. WIixson, D. E. 1971. Ecology of Myotis nigricans (Mammalia: Chiroptera) on Barro Colorado Island, Panama Canal Zone. J. Zool., 163:1-13. WIMsaTT, W. A. 1944. Growth of the ovarian follicle and ovulation in Myotis lucifugus lucifugus. Amer. J. Anat. 74:129-159. 1945. Notes on breeding behavior, pregnancy, and parturition in some vespertilionid bats of the eastern United States. J. Mamm., 26:23- 33. 1960a. An analysis of parturition in Chiroptera, including new observations on Myotis I. lucifugus. J. Mamm., 41:183-200. 1960b. Some problems of reproduction in relation to hibernation in bats. Bull. Mus. Comp. Zool., 124:249-267. 1969. Some interrelations of reproduction and hibernation in mammals. Pp. 511-549, in Dormancy and survival (H. W. Woolhouse, ed.), Academic Press, New York. UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly pub- lished in the above series are now published as Occasional Papers, Museum of Natural History. The Miscellaneous Publica- tions, Museum of Natural History, began with number 1 in 1946. Longer research papers are published in that series. Monographs of the Museum of Natural History were initiated in 1970. All manuscripts are subject to critical review by intra- and extra- mural specialists, final acceptance is at the discretion of the publications committee. Institutional libraries interested in exchanging publications may obtain the Occasional Papers and Miscellaneous Publica- tions by addressing the Exchange Librarian, University of Kan- sas Library, Lawrence, Kansas 66044. Individuals may pur- chase separate numbers of all series. Prices may be obtained upon request addressed to Publications Secretary, Museum of Natural History, University of Kansas, Lawrence, Kansas 66044. Editor: LinpA TRUEB Managing Editor: Wi1Lu1AM E, DUELLMAN PRINTED BY UNIVERSITY OF KANSAS PRINTING SERVICE LAWRENCE, KANSAS y) } A Y 22 ff Vide A) CC Ct CALIFORNIA ‘| ACADEMY OF SCIENCES SEP 19 1973 sural ~— 1D. f OCCASIONAL PAPERS of the MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas NUMBER 16, PAGES 1-66 SEPTEMBER 7, 1973 A REVIEW OF THE CENTROLENID FROGS OF ECUADOR, WITH DESCRIPTIONS OF NEW SPECIES By Joun D. Lyncu' AND WILLIAM FE. DUELLMAN* Since the description of Centrolene geckoideum by Jiménez de la Espada (1872), there have been few contributions to our knowl- edge of Ecuadorian centrolenid frogs. Boulenger (1882, 1898, 1899) named three Ecuadorian species as Hyla or Hylella. Noble (1924) recorded another species. Goin (1961) named three new species, and Goodman and Goin (1970) commented on additional Ecua- dorian specimens of Centrolene geckoideum. The generic status of these frogs has been reviewed by Noble (1920), Taylor (1951), Goin (1964) and Savage (1967). Taylor (1951) perceived the distinctive features of the assemblage and pro- posed the recognition of the Centrolenidae, a family occurring from southern México to Peri and Surinam and with an additional pro- liferation of species in southeastern Brasil. The Middle American species have been reviewed by Taylor (1949, 1952, 1958), Duellman and Tulecke (1960), Savage (1967), and Savage and Starrett (1967). Taylor and Cochran (1953) sum- marized the Brasilian species; Cochran and Goin (1970) accounted for the known Colombian species, and Rivero (1968) published on the Venezuelan centrolenids. Additional species have been named from Pert: ( Boulenger, 1918), Surinam (Goin, 1966), Brasil (Lutz and Kloss, 1952), and Guyana (Goin, 1968). * Associate in Herpetology, Museum of Natural History, University of Kan- sas, and Associate Professor, Department of Zoology, University of Nebraska, Lincoln, Nebraska 68508. * Curator, Division of Herpetology, Museum of Natural History, University of Kansas. 2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Currently the Centrolenidae is composed of two genera—Cen- trolene Jiménez de la Espada, 1872, and Centrolenella Noble, 1920; approximately 50 species are recognized in the family. Eight species have been recorded from Ecuador: Centrolene geckoideum and Centrolenella buckleyi, cochranae, fleischmanni, griffithsi, ocellifera, parabambae, and petersi. Field work in Ecuador from 1967 through 1972 has resulted in the accumulation of nearly 200 specimens, representing all of the previously known Ecuadorian species of Centrolenella, two species heretofore unknown from Ecuador, and eleven new species named herein. We have examined the type specimens of all Ecuadorian taxa. Our investigations show the presence of 19 species of centro- lenids in Ecuador. We present data on all the Centrolenella but include Centrolene geckoideum only in the key. Additionally, we treat the Colombian Centrolenella medemi, although the species has yet to be found in Ecuador. Most areas of Ecuador have been explored in the course of our field work. Duellman worked mostly in the Amazonian lowlands and slopes, whereas Lynch spent most of his time at high elevations in the Andes and on the Pacific slopes of the Andes. Both of us spent limited time on the mesic Pacific lowlands. Based on our ex- perience, we have noted a high degree of endemism in the faunas on the Pacific and Amazonian slopes and in some semi-isolated mountain ranges in Ecuador. We suspect that the centrolenid fauna is much larger than the 19 species now known from Ecuador.? Ac- cordingly, in this paper we examine the current state of our knowl- edge of Ecuadorian centrolenids. Acknowledgments For the loan of specimens or for provision of working space in their respective institutions, we are grateful to James R. Dixon, Alice G. C. Grandison, M. S. Hoogmoed, Konrad Klemmer, Charles W. Myers, Giinther Peters, Douglas A. Rossman, Charles F. Walker, and George R. Zug. We are grateful to the following persons whose efforts in the field enhanced our collections of centrolenid frogs: Joseph T. Collins, Martha L. Crump, Stephen R. Edwards, Thomas H. Fritts, Marsha Lynch, Bruce MacBryde, John E. Simmons, Ger- ald R. Smith, Linda Trueb, and Charles F. Walker. Field work in Ecuador was partially supported by Watkins Museum of Natural * Subsequent to the completion of this paper, we received a specimen of an apparently unnamed species of Centrolenella, obtained at the Estacién Biolégica Rio Palenque, Provincia Los Rios. The specimen would be identified in our key as C. cochranae. CENTROLENID FROGS OF ECUADOR 3 History Grants, University of Kansas (Duellman and Lynch), So- ciety of Sigma Xi (Lynch), Penrose Fund (Grant no. 5549) of the American Philosophical Society (Lynch), and the Committee on Systematics and Evolutionary Biology, University of Kansas (Lynch). Duellman’s study of specimens in European museums was made possible by a grant (no. 5063) from the Penrose Fund of the Amer- ican Philosophical Society. Abbreviations for collections used throughout the text are: AMNH_ American Museum of Natural History BMNH British Museum ( Natural History ) CAS California Academy of Sciences KU University of Kansas Museum of Natural History LSU Louisiana State University RMNH_ Rijksmuseum van Natuurlijke Historie SMF Senckenbergische Museum Frankfurt TCWC Texas Cooperative Wildlife Collection UMMZ University of Michigan Museum of Zoology USNM_ United States National Museum (National Museum of Natural History ) ZMB Zoologisches Museum Berlin TAXONOMIC CHARACTERS With the exception of Centrolene geckoideum, living centro- lenids usually are small, slender-limbed, green frogs. Most preserved examples seem to offer a limited suite of characters useful in their classification. The apparent paucity of characters is in part illusory, for there has been a multiplicity of names applied to comparatively few species in some areas; furthermore, there has been a limited amount of well-preserved material with detailed records of color- ation of living frogs. In the following paragraphs we discuss those taxonomic characters found to be useful by us and contemporary investigators (Savage, 1967; Savage and Starrett, 1967). Prevomerine teeth.—Prevomerine teeth and dentigerous proc- esses are absent in 9 species: anomala, buckleyi, fleischmanni, grif- fithsi, megacheira, munozorum, pellucida, peristicta, and pipilata. Judgment on the absence of teeth should be held in abeyance for anomala and pellucida, inasmuch as we have a single specimen of each. Centrolenella pellucida is a member of the fleischmanni group, and if it does have prevomerine teeth, it would be the only species of the group to have them. Centrolenella anomala seems to be allied most closely to C. cochranae, some specimens of which lack prevomerine teeth. Prevomerine teeth usually are absent in grandisonae. The teeth usually are present in cochranae, flavopunc- tata, ocellifera, prosoblepon, and siren, and are found in all ex- 4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY amples examined of audax (4), medemi (1), midas (11), and resplendens (2). Color of bones.—In living and recently preserved specimens of many centrolenids, the bones are green. The color is seen most easily on the ventral surface of the shank. We have seen living ex- amples of all species discussed here except medemi. Color of the bones was recorded for most specimens after field work in 1967; our only material of two species was collected in 1967 (ocellifera and resplendens ), and no notations of bone color were made. Based on color notes of living frogs, the following taxa have green bones: audax, buckleyi, cochranae, flavopunctata, grandisonae (pale), griffithsi (pale), megacheira, midas, peristicta (pale), pipi- lata, prosoblepon, and siren; white bones were observed in anomala, fleischmanni, munozorum, and pellucida. Some specimens of green- boned species collected in 1967 still retain green bones. Dissection of preserved specimens of medemi, ocellifera, and resplendens, re- vealed no green pigment in the bones. We are tempted to argue that the bones of these species were white in life; however, we must point out that the pale green bones of griffithsi quickly fade to white in preservative. Humeral spines—On some centrolenids, a hook-like process ex- tends anteriorly from the deltoid ridge of the humerus. This fea- ture was used initially to distinguish Centrolenella and Cochranella but is no longer regarded as adequate grounds for generic distinc- tion (Goin, 1964). Humeral spines usually are present only in males, and the spines appear to become more pronounced in larger, and presumably, older individuals. The spine is usually absent in fe- males (Eaton, 1958, reported small spines in female prosoblepon). The spines are present in audax, buckleyi, grandisonae, medemi (small), peristicta, pipilata, and prosoblepon. Normally griffithsi lacks humeral spines, but one individual (KU 142649) has small humeral spines. The distal end of the humerus of males is pro- vided with expanded lateral flanges in some of our examples of grandisonae, peristicta, and pipilata; sexual dimorphism of this sort is well-known in some Eupsophus and some Leptodactylus (Lynch, 1971). Snout shape.—The shape of the snout in dorsal view (Fig. 1) varies from subacuminate (fleischmanni) to round (audax, buckleyi, flavopunctata, grandisonae, munozorum, ocellifera, pellucida, peri- sticta, prosoblepon, and resplendens) or truncate (anomala, coch- ranae, griffithsi, medemi, megacheira, midas, and siren). Truncate snouts (in dorsal view) result from the protuberant nostrils lending an angular appearance to the top of the snout. Protuberant nostrils CENTROLENID FROGS OF ECUADOR 5 Fic. 1. Heads of Centrolenella: Dorsal views. A. C. fleischmanni, KU 146606. B. C. audax, KU 143290. C. C. anomala, KU 143299. Lateral views. D. C. resplendens, KU 118053. E. C. munozorum, KU 118054. F. C. griffithsi, KU 118040. are also found among some of those species with round snouts (dor- sal view) but the nostrils do not extend as far anteriad. In lateral view (Fig. 1), the snout profile varies from strongly sloping (resplendens) to a weakly sloping or rounded (buckleyi), to rounded flavopunctata, grandisonae, munozorum, pellucida, peri- sticta, and in some female prosoblepon), to truncate (anomala, audax, cochranae, weakly in fleischmanni, griffithsi, medemi, mega- cheira, midas, ocellifera, pipilata, most prosoblepon, and siren). The laterally truncate snouts again reflect the presence of protuberant nostrils (more so than any differences in shape of the underlying skeleton ). An additional feature of the snout is the angularity of the canthus and loreal region. The canthus rostralis is scarcely evident in fleisch- manni, munozorum, and pellucida, whereas it is much more distinct in those frogs having more vertical loreal regions (all other species, except grandisonae, peristicta, and pipilata). Tympanum. —Tympana are present in all centrolenids we ex- amined. The tympana are concealed beneath the skin in medemi, munozorum, and pellucida, but at least partially visible in others (lower one-fourth visible in buckleyi to entirely visible in some fleischmanni, peristicta, and pipilata). The upper edge of the tym- panum usually is obscured by a supratympanic fold or may be simply covered by a thickening of the skin in the absence of a dis- tinct supratympanic fold. The extent of the exposed portion of the tympanum is probably not a reliable feature for species recognition. 6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY The tympanic region is directed strongly dorsolaterally (as op- posed to being nearly vertical in most frogs) in fleischmanni, muno- zorum, and pellucida; this feature, in combination with the weakly marked canthus, seems to give these frogs a flatter head than that seen in the other Ecuadorian species. These species also have a strong dorsolateral orientation to the tympanum. The tympana are strongly oriented laterally (little or no dorsal or posterior inflection) in grandisonae, ocellifera, and peristicta. The tympana are oriented posterolaterally with little or no dorsal inflection in anomala, buck- leyi, midas, pipilata, and siren. The tympana are oriented dorso- laterally, but much less so than in fleischmanni and its allies, in audax, cochranae, griffithsi, megacheira, prosoblepon, and resplen- dens; in these frogs there may be little to moderate posterior in- flection of the tympana. Skin texture —The skin of the venter is coarsely areolate in all species examined. The skin of the dorsal surfaces varies in texture and is useful in species recognition. The dorsal skin of medemi is smooth, unlike any other Ecuadorian centrolenid. Most species have finely shagreened skin on the dorsum without warts or spinules (audax, flavopunctata, fleischmanni, grandisonae, griffithsi, midas, munozorum, ocellifera, peristicta, pellucida, prosoblepon, and siren). Shagreened skin with scattered enameled‘ warts is found in pipilata and resplendens; resplendens has many more warts (Plate 2C). Centrolenella cochranae has scattered warts on a shagreened dor- sum; C. anomala has a shagreened dorsum with more numerous warts, and C. megacheira has pustular skin on the dorsum. The skin on the dorsum of buckleyi is shagreened with spinules, which are most numerous laterally. Dermal ornamentation—Few species of the family have what Taylor and Cochran (1953) termed “decoration” or tarsal and ulnar folds. We find that many of the folds and tubercles are enameled and thus appear to be distinctive. Centrolenella grandisonae has low tubercles along the outer edge of the forearm and tarsus; the tubercles are present distally along the lateral edge of the hand and foot to the digital pads. A similar arrangement of more pronounced tubercles is found in C. peristicta (Fig. 3) and C. pipilata. Centro- lenella resplendens has a scalloped fringe along the outer edge of the hand and arm (Fig. 3), and foot and tarsus (Fig. 4); the fringe continues around the heel. A thin fringe, without warts or scallop- ing, occurs along the hand and forearm and the foot and tarsus of C. pellucida. *“The term “enameled” is used throughout to denote the shiny white ele- vations found in the skin of some centrolenids. CENTROLENID FROGS OF ECUADOR 7 Fic. 2. Anal ornamentation in Centrolenella. A. Ventral view of posterior ventrum of C. siren, KU 146621. B. Posterior view of anal region of C. pellu- cida, KU 143298. C. Dorsal view of C. resplendens, KU 118053. Most of the Ecuadorian frogs have two to four large, round subanal tubercles (Fig. 2). Five species have more extensive anal ornamentation. Six Ecuadorian species do not have subanal tuber- cles (anomala, fleischmanni, medemi, munozorum, ocellifera, and pellucida). The small tubercles lateral and posteroventral to the vent in grandisonae, peristicta, and pipilata are enameled and seem to rep- resent a rudimentary character-state for the “anal decoration” char- acter-state reported by Taylor and Cochran (1953); none of these species has enlarged para- or postanal warts or folds. Centrolenella pellucida has a transverse fold below the vent (Fig. 2); the fold is enameled. The “anal decoration” is most extensive and unique in C. resplendens (Fig. 2). A similar feature appears to be present in Taylor and Cochran’s (1953) surda of southeastern Brasil. In re- splendens, a pair of thick folds extends laterally from the vent onto the posterior surface of the thighs; the folds are connected at their lateral extent by a semicircular fold extending medio-ventrally be- neath the anus. These folds and the area between them are orna- mented with enameled warts and short ridges. 8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Fic. 3. Palmar views of hands of Centrolenella. A. C. griffithsi, KU 118040. B. C. grandisonae, KU 118047. C. C. flavopunctata, KU 121050. D. C. muno- zorum, KU 118054. E. C. resplendens, KU 118053. F. C. peristicta, KU 118051. Hands and feet.—The first finger is as long as or longer than the second in all Ecuadorian centrolenids. In two Peruvian species we have examined, the first finger is shorter than the second. The pal- mar and subarticular tubercles are difficult to see on most preserved examples and are not accorded significance here. The fingers bear discs that are wider than long but vary between being more nearly round or truncate. The species with more rounded finger discs are buckleyi, fleischmanni, griffithsi, munozorum, pellucida, pipilata, resplendens; megacheira is intermediate between rounded and trun- cate discs include anomala, audax, cochranae, flavopunctata, gran- CENTROLENID FROGS OF ECUADOR 9 disonae, medemi, midas, ocellifera, peristicta, prosoblepon, and siren. The finger discs are larger than those of the toes in all species examined, but in several species (anomala, audax, fleischmanni, me- demi, megacheira, munozorum, pellucida, pipilata, prosoblepon, and resplendens) the discs are more nearly equal in size than in cochranae, flavopunctata, grandisonae, griffithsi, midas, ocellifera, peristicta, and siren. The fingers bear narrow lateral fringes in ano- mala, audax, buckleyi, fleischmanni, grandisonae, megacheira, muno- zorum, pellucida, peristicta, pipilata, prosoblepon, and resplendens but not in cochranae, flavopunctata, griffithsi, medemi, midas, ocelli- fera, or siren. The extent of webbing of the hands and feet has been used commonly in order to distinguish species of centrolenids. In record- ing the extent of webbing we have followed Savage and Heyer (1967). The least finger webbing is seen in anomala, cochranae, griffithsi, megacheira, and siren (Fig. 3); buckleyi grandisonae, and pipilata have only slightly greater webbing of the fingers (Fig. 3). In these eight taxa the webbing does not enclose the distal sub- articular tubercle. The most extensive webbing seen in Ecuadorian Centrolenella was found in the holotype of petersi and an example referred to fleischmanni (KU 121052). In these specimens the web reaches the disc of the fourth finger and nearly to the disc of the third (III1%s-OIV ); the modal webbing for fleischmanni is TI114-21V. The webbing completely encloses the distal subarticular tubercle of fingers III and IV in flavopunctata, medemi, peristicta, and resplen- dens (Fig. 3) and is only slightly less extensive in fleischmanni, midas, ocellifera, and prosoblepon (Fig. 3). The distal subarticular tubercle of IV, but not III, is free of webbing in audax and pellucida. Most species studied have a small outer metatarsal tubercle; none was found in audax, flavomaculata, fleischmanni, medemi, midas, munozorum, ocellifera, pellucida, resplendens, or siren. The extent of webbing on the feet parallels that of the fingers, but species group less well. The least webbing is seen in cochranae, griffithsi, and siren, and with only slightly more webbing in anomala, buckleyi, megacheira, and pipilata (Fig. 4). The webbing formulae for these seven species range from 12-24I11%-3-T112--31V3-2V of cochranae and siren to the comparatively well-webbed pipilata (11-2*111-2*111-24%1V2%-1V ). Nine species have more webbing than pipilata but less than the extensive webbing of medemi and re- splendens (audax, flavopunctata, fleischmanni, grandisonae, midas, ocellifera, pellucida, peristicta, and prosoblepon). Toe webbing is most extensive in medemi where the web reaches the discs on all but the fourth toe. 10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Fic. 4. Plantar views of feet of Centrolenella. A. C. griffithsi, KU 118040. B. C. grandisonae, KU 118047. C. C. resplendens, KU 1180653. D. C. flavo- punctata, KU 121048. Peritoneal coloration—tThe parietal peritoneum is white (heart not visible in life) in all Ecuadorian Centrolenella except fleisch- manni, munozorum, and pellucida; in these three species the heart is visible through the skin of the chest. The visceral peritoneum is white in fleischmanni, munozorum, pellucida, and resplendens. The latter is the only Ecuadorian species with both white parietal and visceral peritonea. In living frogs, if the heart is visible, the parietal peritoneum is clear; if the heart is not visible, the parietal peritoneum is opaque CENTROLENID FROGS OF ECUADOR ot and white. The character-states of the visceral peritoneum are more difficult to observe in living frogs. If the intestine is dark (as seen through the skin of the venter), the visceral peritoneum is clear; if the intestine is yellow or white, the visceral peritoneum is opaque and white. In preserved examples, inspection of peritoneal pigmentation is best made by cutting into the abdomen and recording whether the intestine is white or not (visceral peritoneum) and whether the in- side of the belly beneath the liver is white or not (parietal peri- toneum). Three peritoneal pigmentation patterns are presently known: 1) clear parietal, white visceral (fleischmanni and pulverata groups and at least C. albotunica, eurygnatha, and vanzolinii of southeastern Brasil); 2) white parietal, clear visceral (prosoblepon group); and 3) white parietal, white visceral (antioquiensis and resplendens ). Ground color in preservative —The ground color of preserved frogs is cream to creamy-white in fleischmanni (including the holo- type of petersi), munozorum, and pellucida. The ground color of anomala is pale brown. The ground color of flavopunctata, medemi, and pipilata is gray to slate-gray. That of grandisonae, griffithsi, ocellifera, peristicta, and resplendens is pale to dull lavender and contrasts with the darker lavender ground color of audax, buckleyi, cochranae, megacheira, midas, prosoblepon, and siren. Color patterns—The most simple color patterns are those of audax, flavopunctata, midas, and siren (white flecks on ground color), griffithsi (lavender to black flecks on ground color), buck- leyi (moderately distinct white labial stripe continuing onto flanks separating lavender or purple dorsum from cream venter), and fleischmanni and munozorum (faint peppered reticulation of dorsal surfaces ). No color pattern is seen in pellucida. The color pattern of resplendens consists of many small white spots (enameled warts) on a dull lavender ground color. Centro- lenella medemi has a simple pattern of large cream spots on the dorsal surfaces, unlike the pattern of any other centrolenid. The other centrolenids with non-ocellated color patterns include grandisonae, peristicta, and pipilata with small, diffuse black (or lavender) spots and equally diffuse white spots on the dorsal sur- faces and megacheira and prosoblepon with discrete dark lavender spots on the dorsum and limbs (spotting reduced or absent in prosoblepon). The other three species known from Ecuador (anomala, coch- ranae, and ocellifera) and the Peruvian ocellata have a dorsal pat- tern of ocelli. The ocelli are large in ocellata but small in the three 12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Fic. 5. Dorsal patterns of ocellated Centrolenella. A. C. anomala, KU 143299. B. C. cochranae, KU 121035. C. C. ocellata, LSU 25990>) BG: ocellifera, KU 118046. Ecuadorian ocellated Centrolenella (Fig. 5). The ocelli of anomala are black with white centers; those of cochranae are deep lavender with white centers, and those of ocellifera faint lavender without white centers. The pattern of ocellifera also includes small non- ocellated white spots (Fig. 5). KEY TO ECUADORIAN CENTROLENID FROGS 1. Discs on fingers as large as eye _______. Centrolene geckoideum Discs on fingers less than half size of eye ___.___= = === 2 2. Dorsum tan with black ocelli enclosing cream (orange in life) SPOUS: et ee eee oak oe een ee a Centrolenella anomala Dorsum white, cream, gray, or lavender (green in life) with or without spots or ocelli _.3 2. eee 3 3. Heart visible in life; bones white; dorsum cream or white in preservative ocelli absent .00..._. 4 Heart not visible in life, bones green (unknown in medemi, ocellifera, and resplendens); dorsum lavender or slate gray in preservative; ocelli and spots present or absent 6 4) lympanum visiple 22 = Centrolenella fleischmanni Tympanum concealed __________ Eee 5 5. Forearm and tarsus each bearing a dermal fold on ventrolateral edge; transverse dermal fold below anus __.----------___-_____ 2.835} in) ipenrd Siete set ene Centrolenella pellucida 10. 1a 12. 13. 14. 15. 16. CENTROLENID FROGS OF ECUADOR 13 Forearm and tarsus lacking dermal folds; no transverse dermal nO CMMCIOW "AMUS Centrolenella munozorum Webbing extending no more than midway between basal and distal subarticular tubercles on fourth finger _._-_____________ {i Webbing extending at least to base of distal subarticular tuber- remOUmenOUItEL WOOT ona 10 Wersalskin pustular Centrolenella megacheira Mioneleskin shagreened, 4)40) 2 bese, heen hs oe pee ee 8 Dorsum with distinct small black ocelli enclosing pale spots Pe ot Oe Centrolenella cochranae Dorsum with white (yellow in life) flecks __ Centrolenella siren Dorsum with or without diffuse black flecks ee as Oe Ns et Se FN, Sel RS Centrolenella griffithsi Broad, scalloped dermal fringe on ventrolateral edges of fore- arm and foot; U-shaped anal fold; snout sloping anteroventrally; dorsum with white warts Centrolenella resplendens Niumilcariveso fancy (os. sat Mee Rs ot, oe 11 Dorsum of body and limbs slate gray with large cream spots; webbing extending to discs on all but fourth toe _--- NE once eT eae Centrolenella medemi Dorsum of body and limbs unicolor, ocellated, or flecked; web- PpiMmeaglessyextenSive. On Meet 6.0 6 te 1! Dorsum with dark ocelli Centrolenella ocellifera Worstm: without ocelli 4... 42s ee 13 Webbing extending beyond distal subarticular tubercle of ROERURIMS ENIAC cot ati Le Be eee 14 Webbing not extending beyond distal subarticular tubercle of HCG LS wk NST) ai men lek Mae a ie celle else aly A 16 Dorsum unicolor or with distinct dark flecks; no pale flecks Centrolenella prosoblepon Dorsum with pale flecks; diffuse dark flecks present or not —. 15 Dorsum of body slate gray (dark green in life) with white (yellow in life) flecks ........______ Centrolenella flavopunctata Dorsum of body pale lavender (pale green in life) with dark lavender spots and few white flecks Centrolenella peristicta Dorsum with pale flecks, no dark flecks; prevomerine teeth FOROS Ny wtteloemat tees ey ent is Yale peek Kee an a Sly oe Ls ees Lif 14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Dorsum unicolor with dark and pale flecks; prevomerine teeth absent = PY oes yt ee a 18 17. Humeral spine present in males; pale flecks numerous on dor- STU aie en CS Oe ee Centrolenella audax Humeral spine absent in males; pale flecks few, principally late rallye Aus viet. 2 bee ee ee Centrolenella midas iS we Dorsumiunicolor: Ce. Centrolenella buckleyi Dorsum with dark and pale flecks __. _.._- = 34a 19 19. Snout truncate in dorsal and lateral profiles ___ "7s 3: BEL eee a cine be Re ane a See a Centrolenella pipilata Snout round in dorsal and lateral profiles. _.__ - =a = Ln Centrolenella grandisamee ACCOUNTS OF SPECIES In the following accounts we attempt to make comparable state- ments in the diagnoses and descriptions. In order to facilitate com- parisons of primary character states among species, we have num- bered what we consider to be the primary character in the diagnoses. The webbing formula in the diagnosis of each species is the modal formula; variation is given in the description. Measurements and proportions are for Ecuadorian specimens only, unless otherwise noted. Specimens designated as paratypes and paratopotypes (in addi- tion to holotypes) are those on which descriptions of new species were based. In some cases measurements of referred specimens have been incorporated into statements of ranges of variation. Colored photographs of 15 species are reproduced on Plates 1 and 2 (following pages 16 and 26). Taxonomic changes and their justifications, comments on type specimens and literature, and observations on behavior and ecology are given in the “Remarks” section of each account. All specimens examined are listed in a terminal section of the paper. Centrolenella anomala new species Plate 2H Holotype —KU 143299, an adult male, 24.1 mm, from the Rio Azuela, 1740 m, Quito-Lago Agrio road, Provincia Napo, Ecuador, obtained on 23 October 1971, by William E. Duellman. Diagnosis.—1) prevomerine teeth absent; 2) bones white; 3) parietal peritoneum white; visceral peritoneum clear; 4) color in CENTROLENID FROGS OF ECUADOR 15 life tan with black ocelli with orange-tan centers; in preservative, brown with black ocelli with white spots; 5) webbing between outer fingers III3*-2%IV; 6) webbing on foot 12--24I11-241IT1-2*1V 2%- #V; 7) snout truncate in dorsal and lateral profiles; 8) dorsal skin shagreened with minute spicules and elevated warts corresponding to ocelli; 9) arms and legs lacking dermal fringes; 10) humeral spine absent in male; 11) lower two-thirds of tympanum visible, directed posterolaterally with dorsal inclination. Centrolenella anomala differs from all other centrolenids by be- ing tan instead of green; it further differs from other species having ocellated patterns (cochranae, ocellata, and ocellifera) by having more ocelli and scattered black flecks between ocelli. Moreover, anomala differs from ocellifera by having less webbing between the outer fingers, from cochranae by lacking prevomerine teeth, and from ocellata by having the snout truncate and lower two-thirds of the tympanum visible, whereas in ocellata the snout is round and the tympanum is concealed. Description.—Adult male moderate-sized, 24.1 mm in snout-vent length; females not known. Head slightly wider than body; width of head 33.2 percent of snout-vent length; snout short, truncate in dorsal and lateral profiles (Fig. 1); canthus round; loreal region concave; lips not flared; nostrils nearly terminal on snout, protuber- ant, directed dorsolaterally; internarial area depressed. Eye large, protuberant, directed anterolaterally. Supratympanic fold very weak, barely covering upper edge of tympanum; tympanum directed posterolaterally with dorsal inclination. Prevomerine dentigerous processes and teeth absent; choanae small, ovoid, near margin of mouth; tongue not notched posteriorly, barely free behind; vocal slits extending from lateral base of tongue to angles of jaws. Humeral spine absent; ulnar fold and tubercles absent; first finger longer than second; fourth finger nearly as long as third; lat- eral fringes present on fingers; webbing absent between first and second fingers, vestigial between second and third; webbing formula for outer fingers III3*-2%IV; discs broad, truncate; subarticular tu- bercles large, round, simple; supernumerary tubercles absent; pal- mar single, ovoid; nuptial excrescences absent. Hind limbs slender; length of tibia 54.4 percent of snout-vent length, tarsal folds and tubercles absent; inner metatarsal tubercle large, elongate; outer metatarsal tubercle small, round; subarticular tubercles large, round; supernumerary tubercles absent; feet about two-thirds webbed; webbing formula [2-24111-24TII1-2*1V2%-14V; discs on toes slightly smaller and more nearly round than those on fingers. 16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Skin on dorsal surfaces of head, body, forearms, and shanks with many minute spicules and larger spicules corresponding to ocelli; skin of belly and ventral surfaces of thighs granular; other surfaces smooth; anal opening directed posteroventrally at midlevel of thighs; anal folds and tubercles absent. Color in preservative: dorsal surfaces of head, body, forearms, fourth fingers, thighs, shanks, feet, and fourth and fifth toes brown with dark brown flecks and head, body, forearms, and shanks with black ocelli enclosing white spots (Fig. 5); other surfaces creamy tan. Color in life: dorsum tan with small black ocelli enclosing orange-tan spots; chest white; heart not visible; throat, visceral peri- toneum, and ventral surfaces of limbs lacking pigment; bones white; iris bronze with black reticulations. Distribution —This distinctive species is known only from the type locality, a rivulet flowing into the Rio Azuela on the east slope of Volcan Reventador at an elevation of 1740 m (Fig. 6). The small stream is about 50 m north of the bridge over the Rio Azuela on the Quito-Lago Agrio road. In this area there is a narrow relatively gentle slope between the steep slopes of Volcaén Reventador to the west and the chasm of the Rio Coca to the east. Remarks.—The single specimen was kept alive for two days and two nights, during which time it was observed frequently. At no time did it take on any greenish color; it remained tan with no noticeable change in pigmentation. After 18 months in alcohol the dorsum has a faint lavender tint. The holotype was on a mossy limb of a bush about 1.5 m above a cascading rivulet at night. Hyla phyllognatha and four other spe- cies of Centrolenella (megacheira, pellucida, pipilata, and siren) were found in the same stream and in other small streams nearby in the cloud forest, but no other individuals of C. anomala were observed. Etymology.—The specific name is from the Greek anomalos meaning unusual, deviating from the general rule; the name is used in allusion to the distinctive tan color of this species. Centrolenella audax new species Holotype —KU 146624, an adult male, 23.0 mm, from Salto de 35 PLATE 1. A. Centrolenella fleischmanni, KU 146607; B. C. munozorum, KU 123225; C. C. pellucida, KU 143298 (holotype); D. C. midas, KU 123219 (holotype); E. C. siren, KU 143293; F. C. flavopunctata, KU 123224; G. C. cochranae, KU 123216; H. C. anomala, KU 143299 (holotype). All x 2. CENTROLENID FROGS OF ECUADOR i 600 M. CONTOUR OVER 3000M. OVER 5000M. O 40 100 KILOMETERS 76 Fic. 6. Distribution of Centrolenella anomala (square), C. cochranae (circles), C. ocellifera (triangles), C. resplendens (hexagons). Agua, 2.5 km NNE of Rio Reventador on Quito-Lago Agrio road, 1660 m, Provincia Napo, Ecuador, obtained on 7 April 1972, by William E. Duellman. Paratypes—KU 143290 and 143292, adult males, from 16.5 km NNE of Santa Rosa on Quito-Lago Agrio road, 1700 m, Provincia Napo, Ecuador, obtained on 18 October 1971, by Joseph T. Collins and William E. Duellman. Diagnosis ——1) prevomerine teeth 2-4; 2) bones green; 3) pa- rietal peritoneum white; visceral peritoneum clear; 4) color in life pale green with small yellow flecks; in preservative, lavender with white flecks; 5) webbing between outer fingers III2*-2IV; 6) web- bing on foot I1-2-I11-2‘1II1-2*1V2*-1V; 7) snout round in dorsal view, truncate in lateral profile; 8) dorsal skin shagreened; 9) arms 18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY and legs lacking dermal folds; 10) humeral spine present in males; 11) lower four-fifths of tympanum visible, directed dorsolaterally with slight posterior inclination. The coloration of audax is like that of flavopunctata, midas, and siren, but the males of these species lack humeral spines. No other species having humeral spines has a green dorsum with yellow flecks. Description—Adult males moderate-sized, 23.0-23-6 mm (x=23.3, N=3) in snout-vent length; females not known. Head much wider than body, width of head 34.8-43.5 percent (x=37.8, N=3) of snout-vent length; snout short, round in dorsal view, truncate in lateral profile (Fig. 1); canthus round; loreal region barely concave; lips not flared; nostrils four-fifths distance from eyes to tip of snout, slightly protuberant dorsolaterally; internarial area depressed. Eye moderately large, directed anterolaterally. Supra- tympanic fold absent; lower four-fifths of tympanum visible, di- rected dorsolaterally with slight posterior inclination. Prevomerine dentigerous processes posteromedially inclined, narrowly separated medially, between longitudinally elliptical choanae, bearing 2-4 teeth; tongue nearly round, barely free behind; vocal slits extending from midlateral base of tongue to angles of jaws. Humeral spine large, parallel to humerus; ulnar fold and tuber- cles absent; first finger longer than second; third finger slightly shorter than fourth; lateral fringes present on fingers; webbing ab- sent between first and second fingers, vestigial between second and third; webbing formula for outer fingers III(2+-2%)-(2--2')IV; discs broad, truncate; subarticular tubercles small, round, simple; super- numerary tubercles absent; palmar tubercles rectangular, simple; nuptial excrescences absent. Hind limbs moderately slender; length of tibia 51.3-55.6 percent (x=54.0, N=3) of snout-vent length; tar- sal folds and tubercles absent; inner metatarsal tubercle small, ellip- tical, outer metatarsal tubercle absent; subarticular tubercles small, round; supernumerary tubercles absent; feet about two-thirds webbed; webbing formula I1-(2--2*)IT1-(2*-24)ITI1-2‘1V2'-1V; discs on toes slightly smaller than those on fingers. Skin on dorsal surfaces shagreened; skin on belly and proximal ventral surfaces of thighs granular; other surfaces smooth; anal open- ing directed posteroventrally at upper level of thighs; pair of large tubercles ventral to anus. Color in preservative: dorsal surfaces of head, body, forearms, thighs, and shanks lavender with small white dots; hands, feet, and ventral surfaces cream. Color in life: dorsum green with gold flecks; fingers and toes pale yellow; chest white; heart not visible; visceral peritoneum and CENTROLENID FROGS OF ECUADOR 19 600 M. CONTOUR [| OVER 3000M. | 3] OVER 5000M. ODN4A08 E00 KILOMETERS Fic. 7. Distribution of Centrolenella audax (triangles), C. grandisonae (square), C. medemi (hexagon), and C. prosoblepon (circles). ventral surfaces of limbs unpigmented; bones green; iris pale bronze with black reticulations. Distribution —This species presently is known from only two localities at elevations of 1660 and 1700 m on the Amazonian slopes of the Cordillera Oriental of the Andes (Fig. 7). The type of locality is a deeply recessed waterfall in a nearly vertical cliff at a point 2.5 km north-northeast of the bridge over the Rio Reventador on the Quito-Lago Agrio road. Remarks.—The holotype was found in a bromeliad on a cliff below and in the spray zone of the waterfall. In another bromeliad a metamorphosing young having a snout-vent length of 13.5 mm and a tail 19.5 mm in length was found. Two individuals were found at night on vegetation over a stream 16.5 km north-northeast of Santa 20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Rosa; three other species of Centrolenella were found there—mega- cheira, pipilata, and siren. Etymology.—The specific name is Latin, meaning daring, and is used in allusion to the precipitous regions inhabited by the species. Centrolenella buckleyi (Boulenger) Plate 2E Hylella buckleyi Boulenger, 1882:420 [Syntypes —BMNH 78.1.25.16 from In- tac, Provincia Imbabura Ecuador; BMNH 80.12.5.201 from “Paitanga” (=Pallatanga ), Provincia Chimborazo, Ecuador]. Hyla purpurea Nieden, 1923:267 [Substitute name for Hyllela buckleyi Bou- lenger]. Cochranella buckleyi—Taylor, 1951:35. Centrolenella buckleyi—Goin, 1964:6. Diagnosis—1) prevomerine teeth absent; 2) bones green; 3) pa- rietal peritoneum white; visceral peritoneum clear; 4) color in life dark green; in preservative, purple; 5) webbing between outer fin- gers III24-2*1V; 6) webbing on foot 11%-2IT1-2*1I14-2kTV3*-1zV; 7) snout round in dorsal view, in profile round above and slightly sloping anteriorly; 8) dorsal skin shagreened with minute spinules; 9) arms and legs lacking dermal folds; 10) humeral spine present in males; 11) lower one-fourth to one-half of tympanum visible, di- rected posterolaterally with slight dorsal inclination. Centrolenella buckleyi is like many specimens of griffithsi and some prosoblepon in being uniform lavender above in preservative, but it differs from both of these by having an inclined snout in lat- eral profile, spinules in the dorsal skin, and only the lower one-fourth to one-half of the tympanum visible. It further differs from griffithsi by having more webbing on the hand and humeral spines in males, and from prosoblepon by having less webbing on the hand and lack- ing prevomerine teeth. Description—Adults large, snout-vent length 28.4-29.5 mm (x=29.0, N=2) in males, 29.8-34.4 mm (x=31.7, N=5) in females. Head not as wide as body; width of head 32.5-38.6 percent (x=34.7, N=7) of snout-vent length; snout short, round in dorsal view, round above and slightly sloping anteriorly in profile; canthus round; loreal region concave; lips flared; nostrils two-thirds distance from eyes to tip of snout, barely protuberant laterally; internarial area flat. Eyes moderately large, directed anterolaterally. Supratympanic fold mod- erately heavy; lower one-fourth to one-half of tympanum visible, directed posterolaterally with slight dorsal inclination. Prevomerine dentigerous processes and teeth absent; choanae small, ovoid; tongue cordiform, shallowly notched behind; free posteriorly for about one-fourth of its length; vocal slits extending posterolaterally from midlateral base of tongue to angles of jaws. CENTROLENID FROGS OF ECUADOR 21 Humeral spine short, pointed, nearly parallel to humerus; ulnar fold and tubercles absent; first finger equal in length to second; fourth finger slightly shorter than third; lateral fringes present on fingers; webbing absent between first, second, and third fingers; webbing formula for outer fingers III(2*-24)-(24-2%)IV; discs broadly rounded; subarticular tubercles moderately large, conical, simple; distal tubercle on fourth finger weakly bifid; supernumerary tuber- cles small, numerous on basal segments; palmar tubercle large, ovoid; nuptial excrescences absent. Hind limbs moderately robust; length of tibia 49.4-56.4 percent (x=51.2, N=7) of snout-vent length; tarsal folds and tubercles absent; inner metatarsal tubercle large, ovoid; outer metatarsal tubercle small, round; subarticular small, round; supernumerary tubercles small, present basally; toes about three-fourths webbed; webbing formula I(1%-1%)-(2-2*)II(1-1*)- 2*TTT(1-1%s)-(2+-3*)TV(2%-3*)-(14-2*)V; discs on toes slightly smaller than those on fingers. Skin on dorsum shagreened with minute spinules; skin on belly and proximal posteroventral surfaces of thighs granular; other sur- faces smooth; anal opening directed posteroventrally at midlevel of thighs; several small and four large tubercles below anus. Color in preservative: dorsal surfaces, except fingers and toes, purple; other surfaces cream; narrow cream stripe on edge of upper lip in some specimens. Color in life: dorsal surfaces bright to dark green, sharply de- marcated laterally from white lower flanks; chin and most of venter pale green; parietal peritoneum yellow; heart not visible; edge of upper lip, outer edge of tarsus, and anal stripe white; bones green; iris pale copper flecked with black. Distribution —Centrolenella buckleyi occurs from the Mérida Andes in western Venezuela southward through the Andes and in- terandean valleys to southern Ecuador (Fig. 8). Although the spe- cies has been recorded at elevations as low as 1500 m, authenticated records in Colombia and Ecuador are all above 2000 m; we have found the species at elevations up to 3000 m. Remarks.—We have found individuals in terrestrial bromeliads and in vegetation in ciénegas, the quaking meadows in many high Andean localities. Francisco Leén of the Universidad Catolica in Quito collected two specimens from bromeliads in a Eucalyptus grove near Machachi. Thomas H. Fritts obtained one from a bro- meliad on an island in Laguna Cuicocha. Centrolenella buckleyi may breed in the same situations as other centrolenids—rapid, mountain streams—but buckleyi frequents non-stream situations (Laguna Cuicocha, the bog-like ciénegas) where trees, if present, 22 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 81 80 79 78 77 76 600 M. CONTOUR [|] OVER 3000M. Oo | OVER 5000M. OV40 100 ot KILOMETERS 8] 80 79 78 TEGE 76 Fic. 8. Distribution of Centrolenella buckleyi (circles), C. peristicta (square), and C. pipilata (triangles); open symbols are literature records. are small and rarely overhang water. Because of the absence of streams, we suspect that the tadpoles may develop in bromeliads or in ciénegas, a radical departture from the adaptive zone of centro- lenid frogs. As noted by Goin (1961:101), one of the syntypes (BMNH 80.12.5.201) is now represented by disarticulated bones and the other (BMNH 78.1.25.16) is missing. Rivero (1968) named Centrolenella buckleyi venezuelensis from the Mérida Andes of western Venezuela. We have not been con- cerned with northern populations which may or may not be conspe- cific with buckleyi; thus, we use the binomial for the populations in Ecuador. CENTROLENID FROGS OF ECUADOR 23 Centrolenella cochranae (Goin) Plate 1G Cochranelle cochranae Goin, 1961:97 [Holotype-—BMNH 1912.11.1.68 from El Topo, Rio Pastaza, Provincia Tungurahua, Ecuador]. Centrolenella cochranae Goin, 1964:6. Diagnosis——1) prevomerine teeth 0-3; 2) bones green; 3) pari- etal peritoneum white; visceral peritoneum clear; 4) color in life pale green with minute purple ocelli with red centers; in preserva- tive, lavender with purple ocelli with white centers; 5) webbing be- tween fingers III3-2%1V; 6) webbing on foot 12-24111%-2#II12-31V3+- 1%V; 7) snout truncate in dorsal and lateral profiles; 8) dorsal skin shagreened with elevated warts corresponding to ocelli; 9) arms and legs lacking dermal folds; 10) humeral spine absent in males; 11) lower three-fourths of tympanum visible, directed dorsolaterally. The first impression of the dorsal pattern of cochranae is of black flecks on a green ground color; however, upon closer inspection it is evident that the markings actually are small ocelli (Fig. 5). Centro- lenella cochranae differs from other species having ocellated dorsal patterns (anomala, ocellata, and ocellifera) by having smaller ocelli and prevomerine teeth (absent in some cochranae). Furthermore, the species differs from anomala by having a green instead of brown dorsum, from ocellifera by having less webbing between the outer fingers, and from ocellata by having the snout truncate and the lower three-fourths of the tympanum visible, whereas in ocellata the snout is round and the tympanum concealed. Description——Adults moderately large; snout-vent length 23.8- 26.7 mm (x=25.0, N=6) in males, 30.0 mm in single female. Head noticeably wider than body; width of head 33.0-35.3 percent (x=34.0, N=7) of snout-vent length; snout short, truncate in dorsal and lateral profiles; canthus round, loreal region concave; lips moderately flared; nostrils nearly terminal, directed anterolaterally; internarial area depressed. Eye large, protuberant, directed anterolaterally. Supra- tympanic fold absent; lower three-fourths of tympanum visible, di- rected dorsolaterally. Prevomerine dentigerous processes absent in one specimen, present in six; processes transverse between choanae, bearing 1-3 teeth; choanae small, ovoid, near edge of mouth; tongue cordiform, free posteriorly; vocal slits extending from midlateral base of tongue to angles of jaws. Humeral spine absent; ulnar fold and tubercles absent; first fin- ger longer than second; fourth finger slightly shorter than third; lateral fringes absent on fingers; webbing absent between first and second fingers, vestigial between second and third; webbing formula for outer fingers III(3--3+)-(2%-3)IV; discs broader than deep, 24 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY rounded; subarticular tubercles small, round, simple; supernumerary tubercles absent; palmar tubercle large, elliptical; nuptial excres- cences absent. Hind limbs slender; length of tibia 55.7-61.8 percent (x=58.7, N=7) of snout-vent length; tarsal folds and tubercles absent; inner metatarsal tubercle large, flat, elongate; outer metatarsal tu- bercle small, conical; subarticular tubercles small, round; super- numerary tubercles absent; feet about two-thirds webbed; webbing formula [2-(2*-2%)II(14-1%)-(2--2%) I1I2-- (3--3* )IV(3--3*)-(2-2-)V; discs on toes smaller and more nearly round than those on fingers. Skin on dorsal surfaces smooth with scattered spicules; skin on belly and ventral surfaces of thighs granular; other surfaces smooth; anal opening directed posteroventrally at midlevel of thighs; pair of large tubercles below anus. Color in preservative: dorsal surfaces of head and body lavender with small black ocelli enclosing minute white dots or some solid black flecks; other dorsal surfaces creamy tan with black flecks on thighs and shanks of some specimens. Color in life: pale green with black or purple ocelli enclosing pink or red dots; chest white; heart not visible; visceral peritoneum and ventral and concealed surfaces of limbs unpigmented; bones green; iris pale gray with black flecks. Distribution —Most localities from which Centrolenella coch- ranae is known are in the lower valley (1100-1300 m) of the Rio Pastaza on the eastern face of the Andes (Fig. 6); additionally the species is known from an elevation of 1150 m in the Cordillera del Dué above the Rio Coca (+ 180 km NNE of the Rio Pastaza val- ley). On the basis of these few localities it seems that cochranae, along with flavopunctata, inhabits intermediate elevations between the ranges of the lowland species (medemi, midas, munozorum, and resplendens) and the numerous species in the higher cloud forests (anomala, audax, megacheira, pellucida, pipilata, and siren). Remarks.—Goin (1961:97) described cochranae as lacking pre- vomerine teeth, but he (1964:2) noted the presence of teeth in six other specimens. Six of our seven specimens have prevomerine teeth, and one lacks not only teeth but dentigerous processes. We compared two of our specimens (KU 121033, 123217) with the holotypes of cochranae (BMNH 1912.11.1.68) and C. ocellata (Boulenger, 1918) (BMNH 1912.11.1.19) from Huancabamba, De- partamento Pasco, Peri. Our specimens compare favorably with the holotype of cochranae but are different from ocellata in several characters. Centrolenella ocellata differs from C. cochranae as fol- lows: 1) prevomerine teeth absent; 2) ocelli much larger (Fig. 5); 3) snout truncate; 4) first finger shorter than second. CENTROLENID FROGS OF ECUADOR 25 All of our specimens were obtained at night from vegetation overhanging small mountain streams. The call consists of a single, high-pitched note. Other than the calls, no evidence of reproduction was noted while we were collecting in the range of cochrane ( April, June, July, August). Individuals seemed to be sparsely distributed in the habitat. Centrolenella flavopunctata new species Plate IF Holotype -—KU 121048, an adult male, 21.6 mm, from Mera, Provincia Pastaza, Ecuador, obtained on 14 July 1968, by John D. Lynch. Paratopotypes—KU 121041, 28 June 1968, John D. Lynch; 121043-46, 2 July 1968, John D. Lynch and Gerald R. Smith; 121049, 14 July 1968, John D. Lynch; 121050-51, 24 July 1968, John D. Lynch. Diagnosis.—1) prevomerine teeth 0-3 on low processes; 2) bones green; 3) parietal peritoneum white; visceral peritoneum clear; 4) color in life green with pale yellow flecks; in preservative, slate gray with white flecks; 5) webbing between outer fingers III2--14IV; 6) webbing on foot [1-14I10-2-III1-2 TV2-1V; 7) snout round in dorsal and lateral profiles; 8) dorsal skin shagreened; 9) arms and legs lacking dermal folds; 10) humeral spine absent in males; 11) lower three-fourths of tympanum visible, directed dorsolaterally with slight posterior inclination. Centrolenella flavopunctata is like midas and siren in having a lavender dorsum with white flecks (in life, green with gold flecks ) and no black flecks, but it differs from both species by having a rounded, instead of truncate, snout and slightly more webbing on the hands and feet. Centrolenella siren further differs from flavo- punctata by having nearly the entire tympanum visible and strongly inclined posteriorly, whereas only about three-fourths of the tym- panum is visible and directed dorsolaterally in flavopunctata. Description—Adults rather small; snout-vent length 20.6-23.2 mm (x=21.6, N=7) in males, 24.1-25.7 mm (x=24.9, N=3) in females. Head slightly wider than body; width of head 34.6-38.3 percent (x=36.2, N=10) of snout-vent length; snout short, round in dorsal and lateral profiles; canthus round; loreal region barely concave; lips not flared; nostrils nearly terminal on snout, not protuberant, directed antero-dorsolaterally; internarial area barely depressed. Eye moder- ately large, directed anterolaterally. Supratympanic fold absent; lower three-fourths of tympanum visible, directed dorsolaterally with slight posterior inclination. Prevomerine dentigerous processes 26 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY small, low, widely separated between longitudinally rectangular choanae, bearing 0-3 teeth; tongue broadly cordiform, barely free posteriorly; vocal slits extending from midlateral base of tongue to angles of jaws. Humeral spine absent; ulnar fold and tubercles absent; first fin- ger longer than second; fourth finger much shorter than third; fringes absent on fingers; webbing absent between first and second fingers; webbing formula for other fingers IT1-(2%-3*)IIT(14-2)-(1-1%)IV; dises truncate, subarticular tubercles small, round, simple; supernumerary tubercles absent; palmar tubercle large, elliptical; nuptial excres- cences absent (Fig. 3). Hind limbs moderately robust; length of tibia 55.3-62.0 percent (x=58.0, N=10) of snout-vent length; tarsal folds and tubercles absent; inner metatarsal tubercle small, flat, elliptical; outer metatarsal tubercle absent; subarticular tubercles small, round; supernumerary tubercles absent; feet about three- fourths webbed; webbing formula I(0-1)-(1-1%)II(0-1)-(0-2) III(0-1)- (14-2+)TV(2--2*)-(0-1)V; discs on toes smaller and more nearly round than those on fingers ( Fig. 4). Skin on dorsal surfaces shagreened; skin on belly and proximal posteroventral surfaces of thighs granular; other surfaces smooth; anal opening directed posteriorly at upper level of thighs; pair of large tubercles below anus. Color in preservative: dorsal surfaces of head, body, and limbs lavender or slate gray with many minute white flecks; other surfaces dull creamy tan. Color in life: dorsal surfaces of head, body, and limbs pale green with numerous minute yellow flecks on body and limbs; edge of upper lip pale yellow; fingers and toes yellow; chest white; visceral peritoneum unpigmented; throat pale bluish green; bones green; iris pale grayish white, with or without golden tint, with dark gray or brown flecks or fine reticulations. Distribution.—Most specimens are from elevations of 1000-1800 m in the Pastaza Valley in the eastern slope of the Andes; the species is also known from an elevation of 720 m in the Serrania de Um- baqui, 200 km NNE of the former locality (Fig. 9). One faded specimen with no visible yellow flecks from San José Abajo, Provin- cia Napo (AMNH 22187) tentatively is referred to this species. The locality is between 700 and 1000 m on the eastern slope of Volcan Sumaco, about 130 km NE of the Pastaza Valley. Centrolenella > PLATE 2. A. Centrolenella megacheira, KU 143245 (holotype); B. C. grif- fithsi, KU 121039; C. C. resplendens, KU 118053 (holotype); D. C. pipilata, KU 143278 (holotype); E. C. buckleyi, KU 144131; F. C. prosoblepon, KU 146609; G. C. peristicta, KU 121053. All x 2. —_— ae PlATE «2 CENTROLENID FROGS OF ECUADOR 27 8] 80 LY) 78 HU 76 ae 600 M. CONTOUR [] OVER 3000M OVER 5O00M. QO 40 100 KILOMETERS 81 80 Te, 78 77 76 Fic. 9. Distribution of Centrolenella flavopunctata (circles), C. griffithsi (hexagons), C. megacheira (solid squares), C. midas (triangles), and C. siren (open squares); half closed squares are localities for both C. megacheira and C. siren. flavopunctata, together with C. cochranae, inhabits intermediate elevations between the ranges of the lowland species (medemi, mi- das, munozorum, and resplendens) and the cloud forest inhabitants at higher elevations (anomala, audax, megacheira, pellucida, pipilata, and siren). Although cochranae and flavopunctata occur at about the same elevations on the eastern slopes of the Andes, the two species have not been found in sympatry. Remarks.—One was found on a bush in forest by day; most were collected in a deep ravine. Males were perched on small herbs in the spray-zone of a small waterfall. No eggs were observed although one gravid female was found. At several small streams between Mera and the Rio Alpayacu, males were calling on 2, 14, and 24 July 28 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 1968. Recently metamorphosed young having snout-vent lengths of 14.5 and 16.0 mm were found on 28 June and 6 July 1968. Etymology.—The specific name is a combination of the Latin flavus, meaning golden yellow, and the Latin punctatus, meaning dotted, and is used in reference to the dorsal coloration. Centrolenella fleischmanni (Boettger) Plate 1A Hylella fleischmanni Boettger, 1893:251 [Holotype—SMF 3760 from San José, Provincia San José, Costa Rica]. Hylella cappellei Lidth de Jeude, 1904:94 [Holotype—RMNH 4463 from Saramacca, Surinam; synonymy fide Goin, 1964:1]. Centrolenella fleischmanni—Noble, 1924:67. Centrolenella cappellei—Noble, 1926:18. Cochranella fleischmanni—Taylor, 1951:34. Cochranella petersi Goin, 1961:96 [Holotype—BMNH 1902.5.27.24 from Rio Durango, Provincia Esmeraldas, Ecuador]. New synonym. Centrolenella fleischmanni—Goin, 1964: 1. Centrolenella petersi—Gooin, 1964.6. Diagnosis—1) prevomerine teeth absent, 2) bones white; 3) parietal peritoneum clear; visceral peritoneum white; 4) color in life pale green with pale yellow spots; in preservative, cream with faint dark flecks; 5) webbing between outer fingers III2-14IV; 6) web- bing on foot [1-2-I10-2I111-2‘1V2*-1V; 7) snout subacuminate in dorsal view, round in lateral profile; 8) dorsal skin shagreened; 9) arms and legs lacking dermal folds; 10) humeral spine absent in males; 11) tympanum almost entirely visible, directed dorsolaterally with slight posterior inclination. Centrolenella fleischmanni differs from other Ecuadorian centro- lenids that have the heart visible in life and a white dorsum in preservative (munozorum and pellucida) by having the snout sub- acuminate in dorsal view and truncate in lateral profile, and the tympanum distinct. The other species have round snouts and the tympanum concealed. Furthermore, pellucida differs from fleisch- manni by having ulnar, tarsal, and anal folds. Centrolenella oro- costalis from Venezuela differs from fleischmanni by having minute enamel (yellow in life) flecks on the dorsum. Description.—Adult males small; snout-vent length 19.2-21.2 mm (x=20.4, N=5). Head wider than body; width of head 38.3-40.6 per- cent (x=39.2, N=4) of snout-vent length; snout short, shallow, roundly subacuminate in dorsal view, round in lateral profile (Fig. 1); canthus round; loreal region shallowly concave; lips not flared; nostrils three-fourths distance from eye to tip of snout, barely pro- tuberant, directed dorsolaterally; internarial area slightly depressed. Eye large, protuberant, strongly oriented anteriorly. Supratympanic CENTROLENID FROGS OF ECUADOR 29 600 M. CONTOUR OVER 3000M OVER 5000M. O 40 100 KILOMETERS 76 Fic. 10. Distribution of Centrolenella fleischmanni (circles), C. munozorum (triangles), and C. pellucida (square); open symbol is a literature record. fold absent; tympanum small, about one-fourth diameter of eye, nearly entirely visible, directed dorsolaterally with slight posterior inclination. Prevomerine dentigerous processes and teeth absent; choanae small, round, near margin of mouth; tongue ovoid, free pos- teriorly for about one-fourth of its length; vocal slits extending from midlateral base of tongue to angles of jaws. Humeral spine absent; ulnar folds and tubercles absent; first finger longer than second; fourth finger nearly as long as third; lateral fringes present on fingers; webbing vestigial between first and second and second and third fingers; webbing formula for outer fingers III(0- 2)-(1%-1%)IV; discs moderate, rounded, subarticular tubercles small, round, simple; supernumerary tubercles absent; palmar tubercle small, ovoid; nuptial excrescences absent. Hind limbs slender; 30 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY length of tibia 51.9-57.3 percent (x=55.1, N=4) of snout-vent length; tarsal folds and tubercles absent; inner metatarsal tubercle low, flat, elliptical; outer metatarsal tubercle absent; subarticular tubercles small, round; supranumerary tubercles absent; feet about three- fourths webbed; webbing formula I(0-1)-( 14-2) II(0-1)-(2-2*)III(1- 2*)-(2-2*)TV(2-2+)-1V; discs of toes round, slightly smaller than those on fingers. Skin on dorsal surfaces of head and body shagreened; skin on belly and ventral surfaces of thighs granular; other surfaces smooth; anal opening directed posteriorly at upper level of thighs; anal folds and tubercles absent. Color in preservative: dorsum creamy white with scattered black flecks visible under microscope; eyelids and hepatic peritoneum white; skin elsewhere transparent. Color in life: dorsum pale green with pale yellow or yellowish green spots, so large in some individuals so as to give appearance of a pale frog with darker green reticulations; heart visible; belly white; tips of digits yellow; other surfaces unpigmented; bones white; iris white to pale yellow. Distribution.—This is the most widespread species of Centro- lenella, ranging from Veracruz and Guerrero, México to Ecuador and Surinam (Goin, 1964; Savage, 1967). In Ecuador it occurs on the Pacific lowlands and to an elevation of 1460 m at Tandapi on the Pacific slopes of the Cordillera Occidental of the Andes (Fig. 10). Remarks.—Numerous males were calling over small streams in April at the Estacién Biolégica Rio Palenque north of Quevedo. The single specimen from Tandapi was on a leaf of an herb about 2m above the ground at night. Goin (1961:96) diagnosed petersi as differing from fleischmanni by having more extensive webbing on the hand. One specimen (KU 121052) is like the holotype in having a webbing formula for the outer fingers III0-1%IV, whereas other Ecuadorian specimens have III(1%-2)-14IV. Examination of series of specimens from Costa Rica and Panama reveals that most fall within the range of variation in the webbing of the hand as displayed by the three Ecuadorian speci- mens, but three have only one free digit on the third finger and about one and one-half digits free on the fourth finger. Due to the lateral fringes on the fingers, the determination of the point of de- parture of the web is highly subjective in some specimens. We have observed living fleischmanni in México, Guatemala, Costa Rica, and Panama, as well as in Ecuador. Comparison of colored photographs of living individuals from throughout this range reveals that north- CENTROLENID FROGS OF ECUADOR 31 ern frogs tend to be less distinctively marked than southern ones, but the reticulate pattern is evident in some Panamanian specimens. Despite the wide geographic range of the species as outlined by Goin (1964:4), he was reluctant to accept the occurrence of fleisch- manni in Ecuador; he stated: “I have examined the specimen (USNM 60520) that Noble (1924:67) recorded as fleischmanni from Guevedo [=Quevedo, 56 km south of a locality where we obtained four specimens], Ecuador, and agree with him that this individual looks like typical fleischmanni from along the north coast of South America. It seems improbable to me, however, that this species actually occurs in Ecuador.” We have examined the type specimens of all of the nominal species included in the foregoing synonymy and have compared our findings with data obtained from series of living and preserved frogs. We conclude that the holotype of Cochranella petersi and KU 121052 represent the extreme in variation of webbing in Cen- trolenella fleischmanni and that Cochranella petersi Goin, 1961, is a junior synonym of Centrolenella fleischmanni (Boettger, 1893). Centrolenella grandisonae Cochran and Goin Centrolenella grandisonae Cochran and Goin, 1970:513 [Holotype-—BMNH 1910.7.11.68 from Pueblo Rico, Departamento Caldas, Colombia]. Diagnosis.—1 ) prevomerine teeth usually absent; 2) bones pale green; 3) parietal peritoneum white; visceral peritoneum clear; 4) color in life green with minute yellow, white, and black flecks; in preservative, pale lavender with small gray spots and white flecks; 5) webbing between outer fingers III2%-2*IV; 6) webbing on foot 11-2-I11-2-11T1-21V2*14V; 7) snout round in dorsal and lateral pro- files; 8) dorsal skin shagreened; 9) row of low, indistinct tubercles on ventrolateral edges of forearm and tarsus; 10) humeral spine present in males; 11) tympanum entirely visible, directed laterally with posterodorsal inclination. Two other Ecuadorian species are lavender (in preservative) with dark spots and white flecks. Of these, pipilata differs from grandisonae by having ulnar and tarsal folds, truncate snout, and incised webbing. Centrolenella peristicta is nearly identical to grandisonae in coloration and structural features, except that peri- sticta has more webbing on the hand (1% phalanges free on fourth finger; 2* free in grandisonae) and is smaller (mean snout-vent length 19.7 mm; 24.8 in grandisonae ). Description—Adults moderately large; snout-vent length in males 23.7-25.8 mm (x=24.8, N=4); females not known. Head about as wide as body; width of head 24.6-33.3 percent (X=31.1, N=4) of 32 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY snout-vent length; snout short, rounded in dorsal view, in lateral view inclined above, truncate below, giving a rounded appearance; canthus rounded; loreal region barely concave; lips not flared; nos- trils nearly terminal on snout, slightly protuberant dorsolaterally; internarial area slightly depressed. Eye large, directed anterolater- ally. Supratympanic fold weak; entire tympanum visible, directed laterally with posterodorsal inclination. Prevomerine dentigerous processes and teeth absent; choanae small, round; tongue cordiform, notched behind, free posteriorly for about one-fourth of its length; vocal slits extending from midlateral base of tongue to angles of jaws. Humeral spine blunt, perpendicular to humerus; row of low tubercles on ventrolateral edge of forearm; first finger longer than second; fourth finger much shorter than third; lateral fringes present on fingers; webbing absent between first and second fingers, vestigial between second and third fingers; webbing formula for outer fingers IIT ( 2%-2%)-(2--2+)IV; discs truncate; subarticular tubercles small, round, simple; supernumerary tubercles absent; palmar tubercle ovoid, simple; nuptial excrescences absent (Fig. 3). Hind limbs slender; length of tibia 54.9-57.3 percent (x=55.8, N=4) of snout-vent length; row of low, indistinct tubercles on ventrolateral edge of tarsus; inner metatarsal tubercle small, elliptical; outer metatarsal tubercle small, ovoid; subarticular tubercles small, round; super- numerary tubercles absent; toes about two-thirds webbed; webbing formula I(0-1%)-1%-2*) II (0-1) -(1-2*) TI (1-14)-( 2-2) IV (2-2) -(1- 1%)V; discs smaller and more nearly round than those on fingers (Fig. 4). Skin on dorsal surfaces shagreened; skin on belly and proximal posteroventral surfaces of thighs granular, other surfaces smooth; anal opening directed posteriorly at upper level of thighs; pair of large tubercles below anus. Color in preservative: dorsal surfaces lavender with large dark smudges and small white flecks; other surfaces cream. Color in life: green with minute yellow and white flecks and dark green and black spots on body and limbs; tips of digits pale yellow; humeral spine bluish green, vocal sac green; chest white; heart not visible; visceral peritoneum transparent; bones green; iris pale golden bronze flecked with black. Distribution —Centrolenella grandisonae occurs at moderate ele- vations on the Pacific slopes of the Cordillera Occidental of the Andes from southwestern Colombia to northwestern Ecuador (Fig. 7). In Colombia it is known from Pueblo Rico, 1540 m and Santa CENTROLENID FROGS OF ECUADOR 33 Leticia, 2000 m, Departamento Caldas, and in Ecuador from Tan- dapi, 1460 m, Provincia Pichincha. Remarks.—Three calling males were found along the small stream in cloud forest at Tandapi in July 1967. With the exception of one metamorphosing young found along the Rio Tandapi, gran- disonae was observed only along the one stream, where males were calling from leaves of bushes and trees by a waterfall. None of the other four species of Centrolenella known from Tandapi was found along this stream. The metamorphosing young has a snout-vent length of 13.5 mm and a tail stub of 2 mm. The holotype of grandisonae is the largest known specimen of the species (27.4 mm) and is the only one having prevomerine teeth. Centrolenella griffithsi (Goin) Plate 2C Cochranella griffithsi Goin, 1961:99 [Holotype—BMNH 1940.2.20.4 from Rio Saloya, Provincia Pichincha, Ecuador]. Centrolenella griffithsi Goin, 1964:6. Diagnosis——1) prevomerine teeth absent; 2) bones pale green; 3) parietal peritoneum clear; visceral peritoneum white; 4) color in life yellowish green with or without dark flecks; in preservative, dull lavender; 5) webbing between outer fingers III3-2%IV; 6) webbing on foot I2-24II11-2*III1-241V2%-1V; 7) snout truncate in dorsal and lateral profiles; 8) dorsal skin shagreened; 9) arms and legs lacking dermal folds; 10) humeral spine absent in males; 11) lower three- fourths of tympanum visible, directed dorsolaterally with posterior inclination. The diagnostic characters of griffithsi resemble those of three other Centrolenella—buckleyi, megacheira, and prosoblepon. Of these, megacheira is much larger and has pustular dorsal skin. Cen- trolenella buckleyi and prosoblepon have more webbing, and the males have humeral spines. Description—Adults moderately large; snout-vent length 19.7- 96.1 mm (x=24.1, N=14) in males, 21.6-24.8 (x=23.4, N=3) in fe- males. Head slightly wider than body; width of head 30.6-34.7 per- cent (x=32.3, N=17) of snout-vent length; snout short, truncate in dorsal and lateral profiles (Fig. 1); canthus round; loreal region concave; lips not flared; nostrils nearly terminal on snout, slight protuberant dorsolaterally; internarial area barely depressed. Eye moderately large, directed anterolaterally. Supratympanic fold barely evident; lower three fourths of tympanum visible, directed dorsolaterally with posterior inclination. Prevomerine dentigerous processes and teeth absent; choanae large, quadrangular; tongue 34 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY cordiform, distinctly notched behind, barely free posteriorly; vocal slits extending from midlateral base of tongue to angles of jaws. Humeral spine absent; ulnar fold and tubercles absent; first fin- ger longer than second; fourth finger noticeably shorter than third; lateral fringes absent on fingers; webbing absent between first, sec- ond, and third fingers; webbing formula for outer fingers III3-24IV; discs broad, rounded; subarticular tubercles small, round, simple; supernumerary tubercles absent; palmar tubercle large, elliptical, simple; nuptial excrescences absent (Fig. 3). Hind limbs slender; length of tibia 52.4-62.3 percent (X=55.2, N=17) of snout-vent length; tarsal folds and tubercles absent; inner metatarsal tubercle small, ovoid; outer metatarsal tubercle absent; subarticular tubercles small, round; supernumerary tubercles absent; feet about one-half webbed; webbing formula I(2-2*)-(2*-2%)II(1*-1%)-(24-3-)ITI14-(2%-3-)IV(2%- 3-)-2-V; discs on toes smaller and more nearly round than those on fingers (Fig. 4). Skin on dorsal surfaces shagreened; skin on belly and postero- ventral surfaces of thighs granular; other surfaces smooth; anal opening directed posteriorly at upper level of thighs; pair of large tubercles below anus. Color in preservative: dorsal surfaces of head, body, forearms, shanks, and feet dull lavender with or without minute black flecks; other surfaces cream. Color in life: dorsal surfaces pale yellowish green with or with- out dark green flecks; tips of digits pale yellow; chest white; heart not visible; visceral peritoneum transparent; bones pale green; iris whitish bronze. Distribution—Centrolenella griffithsi occurs on the Pacific slopes of the Cordillera Occidental of the Andes from southwestern Colum- bia to northwestern Ecuador (Fig. 9); the species inhabits cloud forests at elevations of 1200-2170 m. Remarks.—Most of our specimens were obtained from low vege- tation in cloud forest at Tandapi. Males were calling from leaves of herbs and bushes over cascading streams. One was found in the axil of an elephant-ear plant by day. Calling males were observed by Lynch on each of his five trips to Tandapi (March 1968, June 1968, July 1967, 1968, and 1970). Sparsely distributed and sporadically calling males were heard every evening at Tandapi, but the breed- ing season clearly is in March, when most available sites were occu- pied by calling males, and vegetation overhanging streams was festooned with egg masses. Eggs were not observed in June or July. In July 1968, while we were collecting astroblepid catfishes in the gravel at the bottom of a small stream, two centrolenid tadpoles CENTROLENID FROGS OF ECUADOR 35 were found. These may be the tadpoles of griffithsi, the most abun- dant of the five species of Centrolenella known from Tandapi; only griffithsi adults were found along the stream. Structurally identical tadpoles were collected at Pilalé, Provincia Cotopaxi, 2500 m. Goin (1969:99) gave the type locality as “Rio Saloya, Ecuador, 4000 feet.” This locality probably is the point on the Rio Saloya where it is crossed by the road from Chillogallo to Santo Domingo de los Colorados, the only road west from Quito in 1940, when the specimens were collected. The point where the road crosses the Rio Saloyo is about 1200 m. The species is abundant in the next valley south, that of the Rio Pilatén. Centrolenella medemi Cochran and Goin Centrolenella medemi Cochran and Goin [Holotype—USNM 152277 from Puerto Asis, Comisaria Putumayo, Colombia]. Diagnonis.—1 ) prevomerine teeth 2-3; 2) bone color unknown; 3) heart apparently not visible; 4) color in life unknown; in pre- servative, dark gray with many cream spots; 5) webbing between outer fingers III2--14IV; 6) webbing on foot 10-0I10-0II10-11V1-0V; 7) snout truncate in dorsal and lateral profiles; 8) dorsal skin smooth; 9) arms and legs lacking dermal folds; 10) small humeral spine present in males; 11) tympanum concealed. Centrolenella medemi is unique among known centrolenids by having a dark gray dorsum with many large, round spots on all dor- sal surfaces. The only other Amazonian species having humeral spines, audax and pipilata, have distinct tympana, less webbing, and a pale green dorsum with small yellow flecks (audax) or dark green with black spots and yellow flecks (pipilata). Description—Adult male moderate in size, 25.3 mm in snout- vent length; head much wider than body; width of head 39.5 percent of snout-vent length; snout moderately short, truncate in dorsal and lateral profiles; canthus round; loreal region barely concave; lips not flared; nostrils nearly terminal, not protuberant, directed dorso- laterally; internarial area flat. Eye moderately large, directed antero- laterally. Supratympanic fold absent; tympanum concealed. Pre- vomerine dentigerous processes short, transverse between choanae, bearing 2-3 teeth; choanae small, ovoid, near margin of mouth; tongue ovoid, barely free posteriorly; vocal slits extending from midlateral base of tongue towards angles of jaws. Humeral spine curved, not projecting; ulnar folds and tubercles absent; lateral fringes absent on fingers; webbing formula 12-24%]12-- S*12-141IV; discs large, truncate; subarticular tubercles large, round, simple; supernumerary tubercles absent; palmar tubercle 36 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY large, simple, ovoid; nuptial excrescences absent. Hind limbs robust; length of tibia 68.8 percent of snout-vent length; tarsal folds and tubercles absent; inner metatarsal tubercle low, elongate; outer meta- tarsal absent; subarticular tubercles small, round; supernumerary tubercles absent; feet essentially fully webbed; webbing formula 10-0110-O1T10-11V1-0V; discs on toes slightly smaller and more nearly round than those on fingers. Skin on belly granular; skin on other surfaces smooth; anal open- ing directed posteriorly at upper level of thighs; anal folds and tubercles absent. Color in preservative: all dorsal surfaces slate gray with many round cream spots; ventral surfaces cream. Color in life unknown. Distribution Known only from the type locality at an elevation of about 280 m on the Rio Putumayo in Amazonian Colombia (Fig. 7). Remarks.—This distinctive species is included here in the antic- ipation that it will be found in Ecuador; the type locality is only 20 km north of the Ecuadorian border. Centrolenella megacheira new species Plate 2A Holotype—KU 143245, an adult male, 27.1 mm, from a stream 16.5 km NNE of Santa Rosa, 1700 m. on Quito-Lago Agrio road, Provincia Napo, Ecuador, one of a series obtained on 17 October 1971, by Joseph T. Collins and William E. Duellman. Paratopotypes—KU_ 143246-72, BMNH_ 1971.1854-55, CAS 135498-9, UMMZ 131668 (3) obtained on 17-19 October 1971, by Joseph T. Collins and William E. Duellman. Diagnosis—1) prevomerine teeth absent; 2) bones green; 3) parietal peritoneum white; visceral peritoneum clear; 4) color in life green with black spots; in preservative, lavender with black spots; 5) webbing between outer fingers III3--2%IV; 6) webbing on foot [2--2*111-241111%-2%1V3--1%V; 7) snout truncate in dorsal and lat- eral profiles; 8) dorsal skin pustular; 9) arms and legs lacking dermal folds; 10) humeral spine absent in males; 11) lower three-fourths of tympanum visible, directed dorsolaterally with slight posterior in- ination. Centrolenella megacheira is distinguished from all other Andean species by its large size, pustular dorsal skin, and small amount of webbing on the hands and feet. The presence of black flecks on the dorsum also is characteristic of prosoblepon, males of which have humeral spines. CENTROLENID FROGS OF ECUADOR 37 Descriptions —Adults large; snout-vent length 27.1-32.8 mm (x=28.2, N=20) in males, 32.1-32.8 mm (x=32.5, N=3) in females. Head much wider than body; width of head 32.8-37.4 percent (x=34.3, N=23) of snout-vent length; snout short, truncate in dorsal and lateral profiles; canthus rounded; loreal region slightly concave; lips barely flared; nostrils four-fifths distance from eye to tip of snout, slightly protuberant dorsolaterally; internarial area depressed. Eye large, directed more anteriorly than laterally. Supratympanic fold weak; lower three-fourths of tympanum visible, directed dorso- laterally with slight posterior inclination. Prevomerine dentigerous processes and teeth absent; choanae small, ovoid; tongue cordiform, barely free posteriorly; vocal slits extending from midlateral base of tongue to angles of jaws. Humeral spine absent; ulnar fold and tubercles absent; hand large; first finger equal in length to second; fourth finger slightly shorter than third; lateral fringes present on fingers; webbing absent between first and second fingers; webbing formula for other fingers II ( 2*-2% ) -( 34-3%) TI ( 24-3-)-(24-3)1V; discs broader than deep, rounded; subarticular tubercles large, ovoid, simple; supernumerary tubercles absent; palmar tubercle large, ovoid, simple; nuptial ex- crescences absent. Hind limbs slender; length of tibia 53.7-62.3 per- cent (x=56.9, N=23) of snout-vent length; tarsal folds and tubercles absent; inner metatarsal tubercle large, elliptical; outer metatarsal tubercle small, ovoid; subarticular tubercles small, round; super- numerary tubercles absent; feet about one-half webbed; webbing formula I(2-2)-(2*-2%) IT (1-1%)- (24-27) TI (14-14) -( 24-3") TV ( 2%- 3-1)-(14-2-)V; discs on toes slightly smaller and more nearly round than those on fingers. Skin on dorsal surfaces pustular; skin on belly and ventral sur- faces of thighs granular; other surfaces smooth; anal opening di- rected posteriorly at upper level of thighs; pair of large tubercles below anus. Color in preservative: dorsal surfaces of head, body, hind limbs, and forearms lavender with small, round, black dots; hands, feet, upper arms, and ventral surfaces cream. Color in life: dorsum green with black dots; flanks cream; chest white; heart not visible; visceral peritoneum unpigmented; othey ventral surfaces green; iris pale grayish bronze; bones green. Distribution —tThe species currently is known at two localities at elevations of 1700 and 1740 m on the Amazonian slopes of the Cor- dillera Oriental in Ecuador (Fig. 9). The type locality is a smahk stream which drains into a tributary of the Rio Salado. This stream 38 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY is crossed by the Quito-Lago Agrio road at a point 16.5 km north- northeast of the village of Santa Rosa. Remarks.—Individuals were found at night on leaves and stems of bushes and trees overhanging streams in cloud forest. Males were calling in October 1971. At the type locality, audax, pipilata, and siren were found with megacheira; at the Rio Azuela, anomala, pel- lucida, pipilata, and siren occurred along the same streams with megacheira. Etymology.—tThe specific name is from the Greek megas, mean- ing large, and the Greek cheiros, meaning hand; the name is used in reference to the exceedingly large hands of this species. Centrolenella midas new species Plate 1D Holotype —KU 123219, an adult male, 19.2 mm, from Santa Cecilia, 340 m, Provincia Napo, Ecuador, obtained on 22 June 1968, by Linda Trueb. Paratypes—KU 107026, 23 November 1966, William E. Duell- man; KU 146625, 2 April, Martha L. Crump; KU 150622, 25 June 1971, Philip S. Humphrey; KU 150623, 28 August 1971, Martha L. Crump, all from the type locality. KU 125334, 23 May 1969, Thomas H. Fritts, and UMMZ 129314, 6 May 1969; Charles F. Walker from Lago Agrio, 330 m, Provincia Napo, Ecuador. Diagnosis —1) prevomerine teeth 1-3; 2) bones green; 3) pari- etal peritoneum white; visceral peritoneum clear; 4) color in life dark green with yellow flecks; in preservative, lavender with white flecks; 5) webbing between outer fingers IJI2-2- IV; 6) webbing on foot I1-2IT1-21111-2*1V2*-1V; 7) snout truncate in dorsal and lateral profiles; 8) dorsal skin shagreened; 9) arms and legs lacking dermal folds; 10) humeral spine absent in males; 11) lower two-thirds of tympanum visible, directed dorsolaterally with strong posterior in- clination. Centrolenella midas is like flavopunctata and siren in having a lavender dorsum with white flecks (in life, green with gold flecks) and no black flecks; it differs from flavopunctata by having a trun- cate, instead of round snout and slightly less webbing on the hands and feet. Centrolenella siren differs from midas by having less web- bing and a more prominent tympanum oriented posterolaterally, instead of dorsolaterally. Description.—Adults small; snout-vent length 17.4-19.2 mm (x=18.4, N=3) in males, 20.6-25.6 mm (x=22.7, N=7) in females. Head wider than body; width of head 33.3-39.1 percent (x=36.0, N=10) of snout-vent length; snout short, truncate in dorsal and lateral profiles; CENTROLENID FROGS OF ECUADOR 39 canthus round; loreal region concave; lips rounded; nostrils nearly terminal on snout, slightly protuberant dorsolaterally; internarial area depressed. Eye moderately large, directed anterolaterally. Supratympanic fold absent; lower two-thirds of tympanum visible, directed dorsolaterally with strong posterior inclination. Dentigerous processes of prevomers small, low, widely separated between mod- erately large, round choanae, bearing 1-3 teeth; tongue broadly cordiform, barely free posteriorly; vocal slits extending from mid- lateral base of tongue to angles of jaws. Humeral spine absent; ulnar fold and tubercles absent; first fin- ger longer than second; fourth finger noticeably shorter than third; webbing vestigial between first and second fingers; webbing formula for other fingers II(2-2*)-(3'-3%) IIT (2--2+ ) -( 1%-2*)IV; discs truncate; subarticular tubercles small, round, simple; supernumerary tubercles absent; palmar tubercle large, elliptical; nuptial excrescences absent. Hind limbs moderately slender; length of tibia 53.1-62.0 percent (x=57.7, N=10) of snout-vent length; tarsal folds and tubercles ab- sent; inner metatarsal tubercle elongate, rounded in section; outer metatarsal tubercle absent; subarticular tubercles small, round; supernumerary tubercles absent; feet about two-thirds webbed; web- bing formula I(0-1%)-(2--2*)II(%-1)-(2-2*) ITT1-( 2-2% )IV (2-2*)-IV; discs on toes more nearly round than those on fingers. Skin on dorsal surfaces shagreened; skin on belly and proximal posteroventral surfaces of thighs granular; other surfaces smooth; anal opening directed posteriorly at upper level of thighs; pair of large tubercles below anus. Color in preservative: dorsal surfaces of head, body, and limbs lavender with few small white flecks on body; other surfaces creamy white. Color in life: dorsum of head, body, and limbs dark green with a few small yellow flecks dorsolaterally on body; hands and feet dull greenish yellow; chest white; heart not visible; visceral peritoneum unpigmented; iris silvery bronze with black reticulations; bones green. Distribution—This species is known from three localities at ele- vations of 330-570 m along the Rio Aguarico in the upper Amazon Basin in Ecuador (Fig. 9). In this area it occurs in sympatry with munozorum and resplendens. Remarks.—Individuals have been found throughout the year on leaves of herbs and trees along small rivulets in rainforest. The call consists of three short notes. Etymology.—tThe specific name is that of a king in Greek my- thology, at whose touch everything turned to gold. The name is 40 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY associated with this frog known along the Rio Aguarico, meaning rich water, in reference to gold found in the river, and in allusion to the gold flecks on the frogs. Centrolenella munozorum new species Plate 1B Holotype—KU 118054, an adult male, 20.2 mm, from Santa Cecilia, 340 m, Provincia Napo, Ecuador, obtained on 18 June 1967, by John D. Lynch. Paratypes—KU 105251, 13 July 1966, Charles M. Fugler; KU 123225, 13 July 1968, William E. Duellman; KU 150620, 13 August 1971, Martha L. Crump; KU 150621, 10 October 1971, Martha L. Crump, all from Santa Cecilia, and UMMZ 129313 from Lago Agrio, 330 m, 14 km E Santa Cecilia, Provincia Napo, Ecuador, 6 May 1969, Charles F. Walker. Diagnosis—1) prevomerine teeth absent; 2) bones white; 3) parietal peritoneum clear; visceral peritoneum white; 4) color in life pale green with pale greenish yellow spots; in preservative, creamy white with pale gray reticulations; 5) webbing between outer fingers III1%-14IV; 6) webbing on foot I0-1110-1%III1-21V2-- 1V; 7) snout round in dorsal and lateral profiles; 8) dorsal skin shagreened; 9) arms and legs lacking dermal folds; 10) humeral spine absent in males; 11) tympanum concealed; strongly directed dorsolaterally. Two other Ecuadorian species have the heart visible in life and a white dorsum in preservative. Centrolenella pellucida differs from munozorum by having ulnar, tarsal, and anal folds, and fleischmanni differs by having the snout subacuminate in dorsal view and truncate in lateral profile and the tympanum is distinct; in munozorum and pellucida the snout is round, and the tympanum is concealed. The Venezuelan orocostalis has enamel white (yellow in life) flecks on the dorsum. Description Adults small; snout-vent length in males 18.8-20.5 mm (X=19.7, N=5), in one female 20.7 mm. Head wider than body; width of head 37.7-40.7 percent (x=38.4, N=6) of snout-vent length; snout short, shallow, round in dorsal and lateral profiles (Fig. 1); canthus round; loreal region shallowly concave; lips slightly flared; nostrils about four-fifths distance from eye to tip of snout, barely protuberant, directed dorsolaterally; internarial area slightly de- pressed. Eye moderately large, protuberant, directed anterolaterally. Supratympanic fold absent; tympanum concealed, strongly directed dorsolaterally. Prevomerine dentigerous processes and teeth absent; choanae small, ovoid, near margin of mouth; tongue ovoid, barely CENTROLENID FROGS OF ECUADOR 4] free posteriorly; vocal slits extending from posterolateral base of tongue to angles of jaws. Humeral spine absent; ulnar fold and tubercles absent; first finger longer than second; fourth finger nearly as long as third; lateral fringes present on fingers; webbing vestigial between first and second and second and third fingers; webbing formula for outer fingers III (1%- 2-)-(1-14)IV; discs small, rounded, subarticular tubercles small, round, simple; supernumerary tubercles present on proximal segments of first and second fingers; palmar tubercle small, round; nuptial ex- crescences absent (Fig. 3). Hind limbs slender; length of tibia 57.5- 59.6 percent (x=58.4, N=5) of snout-vent length in males, 54.1 per- cent in one female; tarsal folds and tubercles absent; inner meta- tarsal tubercle small, elongate; outer metatarsal tubercle absent; subarticular tubercles small, round; supernumerary tubercles absent; fringe on inner edge of first toe; toes about three-fourths webbed; webbing formula I0-(1--%)II(0-1)-(1%-2-)III(0-1)-(2--2* )IV(2--2)- (0-1)V; discs on toes round, slightly smaller than those on fingers. Skin on dorsal surfaces of head and body shagreened; skin on belly and proximal ventral surfaces of thighs weakly granular; other surfaces smooth; anal opening directed posteriorly at upper level of thighs; anal folds and tubercles absent. Color in preservative: dorsal surfaces of head, body, forearms, thighs, and shanks creamy white with many minute black flecks giving appearance of gray with unpigmented spots; eyelid and he- patic peritoneum white; other surfaces unpigmented. Color in life: dorsum pale green with pale yellow or yellowish green spots; limbs pale green, with slightly darker crossbars in one; thighs unpigmented; iris pale gold. Distribution—Currently this species is known from only two localities along the Rio Aguarico in the Amazonian lowlands of Ecuador (Fig. 10). Remarks.—All individuals were found in lowland rainforest. At Santa Cecilia the frogs were found on leaves of bushes and trees at night: one over a pond, one away from water in primary forest, one on a palm frond 2 m above a stream, and one on an herbaceous leaf more than 2 m above a stream. The specimen from Lago Agrio was obtained from the foliage of a large tree that was felled during the clearing of primary forest. Etymology.—tThe specific name is a patronym for Ing. Ildefonso Munoz B. and Sra. Blanca Munoz, our congenial hosts at Santa Cecilia. 42 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Centrolenella ocellifera (Boulenger) Hyla ocellifera Boulenger, 1899:277 [Holotype—BMNH 98.5.19.3 from Pa- ramba, Provincia Imbabura, Ecuador]. Cochranella ocellifera—Taylor, 1951:35. Centrolenella ocellifera—Goin, 1964:6. Diagnosis——1) prevomerine teeth absent; 2) bones white (?); 3) parietal peritoneum white; visceral peritoneum clear; 4) color in life green with yellow spots; in preservative, pale lavender with faint purple ocelli enclosing white spots; 5) webbing between outer fin- gers III2-14IV; 6) webbing on foot I1-2I11-2-III1-21V2‘-1V; 7 snout round in dorsal view, truncate in lateral profile; 8) skin shagreened; 9) arms and legs lacking dermal folds; 10) humeral spine absent in males; 11) lower two-thirds of tympanum visible, directed laterally with slight posterodorsal inclination. Centrolenella ocellifera has shagreened skin and few large ocelli on the dorsum. Other species having ocellated dorsal patterns (ano- mala, cochranae, and ocellata) have minute spicules in the dorsal skin and less webbing between the outer fingers. Furthermore, ocellifera differs from anomala by having a green instead of brown dorsum, from ocellata by having the lower two-thirds of the tym- panum visible (concealed in ocellata), and from cochranae by havy- ing larger ocelli, snout round in dorsal view, and prevomerine teeth absent (cochranae has small ocelli, snout truncate in dorsal view, and prevomerine teeth usually present). Description Adults moderate-sized; snout-vent length 20.0 mm in male, 26.7 mm in female. Head as wide as body; width of head 32.9-37.0 percent (x=35.0, N=2) of snout-vent length; snout short, round in dorsal view, truncate in lateral profiles; canthus round; loreal region concave; lips slightly flared; nostrils four-fifths distance from eye to tip of snout, not protuberant, directed laterally; inter- narial area flat. Eye large, protuberant, directed anterolaterally. Supratympanic fold not evident; lower two-thirds of tympanum vis- ible, directed laterally with slight posterodorsal inclination. Prevo- merine teeth absent in one male, 2-3 on transverse processes between choanae in one female; choanae small, rectangular; tongue ovoid, slightly free posteriorly. Humeral spine absent; ulnar folds and tubercles absent; first fin- ger larger than the second; fourth finger nearly as long as third; fingers extensively webbed; webbing formula 1[2%-2112-3III2-(1%- 1%)IV; fringe on outer edge of fourth finger; discs moderately broad, truncate; subarticular tubercles small, round, simple (distal tubercle on fourth fingers bifid in female); supernumerary tubercles absent; palmar tubercle single, ovoid; nuptial excrescences absent. Hind CENTROLENID FROGS OF ECUADOR 43 limbs long, slender; tibia length 52.4-59.5 percent (x=55.9, N=2) of snout-vent length; tarsal folds and tubercles absent; inner metatarsal tubercle moderately small, elliptical; outer metatarsal tubercle ab- sent; subarticular tubercles small, round; supernumerary tubercles absent; feet about three-fourths webbed; webbing formula I1-2IT1- 2-III(1--1)-21V2+-1V; fringe on inner edge of first toe; discs on toes smaller and more nearly round than those on fingers. Skin on dorsal surfaces shagreened, that on belly and ventral surfaces of thighs weakly granular; other surfaces smooth; anal opening directed posteriorly at upper level of thighs; anal folds and tubercles absent. Color in preservative: dorsal surfaces of head, body, and shanks pale lavender, scattered dark brown flecks; three or four thin purple ocelli with cream centers in scapular region (Fig. 5); chest and belly cream; other surfaces transparent. Color in life: body green with pale yellow spots and dark green flecks on body; limbs paler green; chest white; heart not visible; visceral peritoneum transparent; iris gray-bronze. Distribution—Centrolenella ocellifera inhabits the Pacific slopes of the Andes (Fig. 6). In addition to the holotype from Paramba (777 m), we have a specimen from Tandapi (1460 m) and have seen a specimen in the Gustavo Orcés-V. collection in the United States National Museum from Pilalé (2320 m). Remarks.—Our specimen was found at night on a fern in cloud forest. The fern was at the edge of a cliff about 10 m above the Rio Pilaton. Centrolenella pellucida new species Plate 1C Holotype —KU 143298, a gravid female, 22.0 mm from the Rio Azuela, 1740 m, Quito-Lago Agrio road, Provincia Napo, Ecuador, obtained on 20 October 1971, by William E. Duellman. Diagnosis—1) prevomerine teeth absent; 2) bones white; 3) parietal peritoneum clear; visceral peritoneum white; 4) color in life pale green with diffuse yellow spots; in preservative, uniform cream; 5) webbing between outer fingers III2*-2IV; 6) webbing on foot 11-2*111-1411T1-2‘1V2'-1V; 7) snout round in dorsal and lateral pro- files; 8) dorsal skin shagreened; 9) unscalloped dermal fold on outer edge of hand and forearm and on foot and tarsus; transverse dermal fold below anus; 10) humeral spine absent in males; 11) tympanum concealed. Among the Ecuadorian species (fleischmanni, pellucida, and munozorum) having the heart visible in life and the dorsum white 44 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY in preservative, C. pellucida is unique in possessing ulnar, tarsal, and anal folds. Moreover, in C. fleischmanni the snout is subacuminate in dorsal view and truncate in lateral profile, and the tympanum is distinct; in C. pellucida and C. munozorum the snout is round, and the tympanum is concealed. The Venezuelan orocostalis, a member of the fleischmanni group, also lacks dermal folds. Description—Males unknown; adult female small, 22.0 mm in snout-vent length; head slightly wider than body; width of head 36.4 percent of snout-vent length; snout short, round in dorsal and lateral profiles; canthus round; loreal region barely concave; lips not flared; nostrils nearly terminal, directed laterally, not protuberant; inter- narial area flat. Eye large, protuberant, strongly oriented anteriorly. Supratympanic fold absent; tympanum concealed. Prevomerine dentigerous processes and teeth absent; choanae large, oval; tongue ovoid, barely free posteriorly. Humeral spine absent; narrow, unscalloped dermal fold on ven- trolateral edge of forearm and outer edge of hand; fringes absent on fingers; first finger longer than second; fourth finger slightly shorter than third; fingers extensively webbed; webbing formula 1[2-2*II2-- 3ITI2*-21V; discs moderately large, rounded; subarticular tubercles small, low; supernumerary tubercles absent; palmar tubercle small, indistinct. Hind limbs slender; tibia length 56.4 percent of snout- vent length; narrow, unscalloped dermal fold along outer edge of tarsus and fifth toe; inner metatarsal tubercle small, ovoid; outer metatarsal tubercle absent; subarticular tubercles small, low; super- numerary tubercles absent; feet about three-fourths webbed; web- bing formula [1-21T1-14III1-2*1V2*-IV; discs on toes round, slightly smaller than those on fingers. Skin on dorsum of head and body shagreened; skin on other sur- faces smooth; anal opening directed posteriorly at upper level of thighs; transverse dermal fold below anus at posteroventral edge of thighs (Fig. 2). Color in preservative: dorsum creamy white with minute purple flecks visible under magnification; hepatic peritoneum white; skin on ventral surfaces transparent. Color in life: dorsum pale green with diffuse yellow spots; venter and hidden surfaces of limbs lacking pigment; fingers and toes yel- low; parietal peritoneum clear; heart visible; bones white; iris pale silvery bronze. Distribution—This small species is known only from the type locality on the east slope of Volcan Reventador on the Amazonian slopes of the Andes (Fig. 10). Remarks.—The holotype was on the leaf of an herb over a small CENTROLENID FROGS OF ECUADOR 45 stream at night. See the account of anomala for a detailed descrip- tion of the type locality and comments on associated species. Etymology.—The specific name is Latin meaning transparent and is applied to this species having a transparent parietal peritoneum. Centrolenella peristicta new species Plate 2G Holotype.—KU 118051, an adult male, 20.6 mm, from Tandapi, 1460 m, Provincia Pichincha, Ecuador, obtained on 23 July 1967, by John D. Lynch. Paratopotypes—KU _ 118052, 24 July 1967, Marsha Lynch; 121053, 28 July 1968, Gerald R. Smith. Diagnosis—1) prevomerine teeth absent; 2) bones pale green; 3) parietal peritoneum white; visceral peritoneum clear; 4) color in life pale green with minute yellow, white, and black flecks; in pre- servative, pale lavender with dark lavener spots and few white flecks; 5) webbing between outer fingers IHI2--14IV; 6) webbing on foot [1-2-I11-21IT1-2*1V2-1V; 7) snout round in dorsal and lateral profiles; 8) dorsal skin shagreened; 9) row of low tubercles on ven- trolateral edges of forearm and tarsus; 10) humeral spine present in males; 11) tympanum entirely visible, directed laterally with slight dorsal inclination. Two other Ecuadorian species are lavender (in preservative ) with dark spots and white flecks. Of these pipilata differs from peristicta by having ulnar and tarsal folds, truncate snout, and in- cised webbing. Centrolenella grandisonae is nearly identical to peristicta in coloration and structural features, except that grandi- sonae has less webbing on the hand (2* phalanges free on fourth finger; 1% free in peristicta) and is larger (mean snout-vent length 24.8 mm; 19.7 mm in peristicta). Description —Adults small; snout-vent length 18.7-20.6 mm (x=19.7, N=2) in males, 20.5 mm in one female. Head no wider than body; width of head 32.5-32.7 percent (x=32.6, N=3); snout short, round in dorsal and lateral profiles; canthus round; loreal region concave; lips rounded; nostrils nearly terminal on snout, slightly protuberant dorsolaterally; internarial area depressed. Eye moderate sized, directed more laterally than anteriorly. Supratympanic fold weak; tympanum entirely visible, directed laterally with slight dor- sal inclination. Prevomerine dentigerous processes and teeth absent; choanae small, ovoid; tongue cordiform, shallowly notched behind, barely free posteriorly; vocal slits extending from midlateral base of tongue to angles of jaws. Humeral spine curved, terminus parallel to humerus; row of low 46 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY tubercles on ventrolateral edge of forearm; first finger longer than second; fourth finger much shorter than third; lateral fringes present on fingers; webbing vestigial between first and second fingers; web- bing formula for other fingers II ( 2-2%)-3*III2--14IV; discs truncate; subarticular tubercles small, round; distal subarticular tubercle on fourth finger bifid; supernumerary tubercles small, round, present on proximal segments of digits 2-4; palmar tubercle large, ovoid; nup- tial excrescences absent (Fig. 3). Hind limbs slender; length of tibia 52.2-58.8 percent (x=54.8, N=3) of snout-vent length; row of low tubercles on ventrolateral edge of tarsus; inner metatarsal tubercle elliptical; outer metatarsal tubercle small, ovoid; subarticular tu- bercles small, round; supernumerary tubercles absent; feet about two-thirds webbed; webbing formula J1-(1%-2)ITI-(2--2*)ITI1-(2--2*) IV(2-2*)-1V; discs smaller and more nearly round than those on fingers. Skin on dorsum shagreened; skin on belly and proximal postero- ventral surfaces of thighs granular; other surfaces smooth; anal opening directed posteriorly at upper level of thighs; pair of large tubercles below anal opening. Color in preservative: dorsal surfaces lavender with dark spots and scattered cream flecks; other surfaces cream. Color in life: dorsal surfaces pale to medium green with dark green spots and white and yellow flecks; flanks yellow-green; belly yellow; vocal sac green; tips of digits pale yellow; heart not visible; bones green; iris grayish bronze with brown or copper ring around pupil. Distribution—This species is known only from Tandapi (for- merly Cornejo Astorga ), a village at the point where the Quito-Santo Domingo de los Colorados road crosses the Rio Pilatén at an eleva- tion of 1460 m on the Pacific slopes of the Cordillera Occidental of the Andes ( Fig. 8). Remarks.—All three individuals were obtained in July on vege- tation in cloud forest at night. One was on an elephant-ear leaf; one was on a fern in the spray zone of a waterfall, and one was on a bush over a waterfall. None was calling. Etymology.—tThe specific name is derived from the Greek peristiktos, meaning dappled, and refers to the spotted color pattern. Centrolenella pipilata new species Plate 2D Holotype —KU 143278, an adult male, 22.9 mm, from a stream 16.5 km NNE of Santa Rosa, 1700 m, on Quito-Lago Agrio road, CENTROLENID FROGS OF ECUADOR 47 Provincia Napo, Ecuador, obtained on 17 October 1971, by William E. Duellman. Paratopotypes—KU 143279-83, 17-18 October 1971, William E. Duellman and Joseph T. Collins. Diagnosis ——1) prevomerine teeth absent; 2) bones green; 3) parietal peritoneum white; visceral peritoneum clear; 4) color in life dark green with diffuse black spots and pale yellow flecks; in pre- servative, gray with dark and pale flecks; 5) webbing between outer fingers II124-2%I1V; 6) webbing on foot indented I1-2*II1-2*III1- 41V2%-IV; 7) snout truncate in dorsal and lateral profiles; 8) dorsal skin shagreened with elevated warts corresponding to pale flecks; 9) unscalloped fringe on outer edge of hand, forearm, and foot; fringe with low scallops on tarsus; 10) humeral spine present in males; 11) tympanum entirely visible, directed posterolaterally with no dorsal inclination. Centrolenella pipilata resembles two other Ecuadorian species in coloration, but both grandisonae and peristicta differ in having ulnar and tarsal tubercles instead of folds and in having round, in- stead of truncate, snouts. Two other Ecuadorian species have ulnar and tarsal folds; pellucida is white in preservative and in life lacks markings and a white peritoneum, whereas resplendens is a much larger, fringe-limbed frog. Males of both of those species lack hu- meral spines. Description—Adults moderately small; snout-vent length 19.5- 22.9 mm, (x=21.5, N=10) in males, 21.8-22:1 mm,(x=21-9) N=2) in females. Head noticeably wider than body; width of head 32.9-38.9 percent (x=35.2, N=12) of snout-vent length; snout extremely short, truncate in dorsal and lateral profiles; canthus round; loreal region barely concave; lips not flared; nostrils three-fourths distance from eye to tip of snout, protuberant anterodorsolaterally; internarial area slightly depressed. Eye large, protuberant, directed more anteriorly than laterally. Supratympanic fold weak; tympanum entirely visible, directed posterolaterally with no dorsal inclination. Prevomerine dentigerous processes and teeth absent; choanae small, ovoid; tongue cordiform, shallowly notched behind, barely free posteriorly; vocal slits extending from midlateral base of tongue to angles of jaws. Humeral spine blunt, oriented at about 30° from humerus; un- scalloped dermal fringe on ventrolateral edge of forearm and outer edge of hand; first finger longer than second; fourth finger notice- ably shorter than third; lateral fringes on fingers; webbing absent between first, second, and third fingers; webbing formula for outer fingers III (2*-2%)-(2-2*)IV; webbing incised; discs broad, rounded; subarticular tubercles moderately large, subconical, simple; super- 48 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY numerary tubercles small, present on proximal segments of all digits; palmar tubercle large, ovoid; nuptial excrescences absent. Hind limbs moderately slender; length of tibia 55.7-61.9 percent (x=59.7, N=12) of snout-vent length; scalloped dermal fold on ventrolateral edge of tarsus; unscalloped dermal fold on outer edge of foot; inner metatarsal tubercle large, ovoid; outer metatarsal tubercle small, round; subarticular tubercles small, round; supernumerary tubercles minute, present on proximal segments of digits; toes about two- thirds webbed; webbing formula I( 1-12) -(2-2+)II1-(2-2%) III (1-14)- (2*-2% ) TV ( 245-2 )-(1-1%)V; discs on toes slightly smaller than those on fingers. Skin on dorsal surfaces shagreened with elevated warts corre- sponding to white spots; skin on belly and ventral surfaces of thighs granular; other surfaces smooth; anal opening directed posteroven- trally at midlevel of thighs; many small and two large tubercles below anus. Color in preservative: all dorsal surfaces lavender with dark spots and white dots; ventral surfaces cream. Color in life: dorsum dark green with diffuse black flecks and pale yellow flecks; flecks on side of head silvery white; chest white; heart not visible; other ventral surfaces pale green; discs pale yel- low; bones green; iris pale bronze with black reticulations. Distribution —In Ecuador, C. pipilata is known from two locali- ties (1700 and 1740 m) on the Amazonian slopes of the Cordillera Oriental of the Andes (Fig. 8). Remarks.—All individuals were found at night on vegetation along cascading mountain streams in cloud forest. Both females were in amplexus; one deposited a clutch of 18 eggs having clear jelly and pale green yolks. At the Rio Azuela, pipilata was found in sympatry with anomala, megacheira, pellucida, and siren, and at 16.5 km north-northeast of Santa Rosa, it was found with audax, megacheira, and siren. At first we thought our specimens might be Centrolenella johnelsi Cochran and Goin, but the much larger size of johnelsi ( ¢ 29.3 mm) and certain structural differences negated this assignment. Further- more, johnelsi is known only from San Pedro, Departamento An- tioquia, in northern Colombia. See the account of megacheira for a description of the type locality. Etymology.—The specific name is an adjectival derivative of the Latin verb pipila, meaning to peep, and refers to the characteristic call of this and many other centrolenid frogs. CENTROLENID FROGS OF ECUADOR 49 Centrolenella prosoblepon (Boettger) Plate 2F Hyla prosoblepon Boettger, 1892:45 [Syntypes—SMF 3756 and ZMB 28019 from “Plantago Cairo” (La Junta), near Limén, Provincia Limén, Costa Rica]. Hyella puncticrus Boulenger, 1896:431 [Syntypes—BHNH 96.10.8.70-71 from La Palma, Provincia San José, Costa Rica]. Hyla parabambae Boulenger, 1898:125 [Holotype—BMNH 98.4.28.163 from Paramba, Provincia Imbabura, Ecuador]. New synomym. Centrolene prosoblepon—Noble, 1924:66. Centrolene parambae (emendation )—Dunn, 1933:73. Cochranelle parambae—Taylor, 1951:35. Cochranella parabambae—Taylor, 1951:35. Centrolenella prosoblepon—Goin, 1964:5. Centrolenella parabambae—Goin, 1964:6. Diagnosis.—1) prevomerine teeth 0-4; 2) bones green; 3) pari- etal peritoneum white; visceral peritoneum clear; 4) color in life green, usually with black dots; in preservative, lavender, usually with dark lavender dots; 5) webbing between outer fingers III2- 2LV; 6) webbing on foot [1-2I11-2*II11-21V2*-1V; 7) snout round in dorsal view, truncate in lateral profile; 8) dorsal skin shagreened; 9) arms and legs lacking dermal folds; 10) humeral spine present in males; 11) lower two-thirds of tympanum visible, directed dorso- laterally with posterior inclination. This species most closely resembles megacheira and _ griffithsi, both of which have less webbing and lack humeral spines in males. Centrolenella megacheira further differs from prosoblepon by hav- ing much larger hands and pustular, instead of shagreened, skin on the dorsum. Unspotted prosoblepon are colored like buckleyi, which also has humeral spines in males, but buckleyi has less webbing, an inclined snout, and only the lower one-fourth of the tympanum visible. Description—Adults large, snout-vent length 21.7-25.6 mm (x=24.1, N=5) in males, 25.4-27.2 mm (x=26.4, N=3) in females. Head slightly wider than body; width of head 33.1-35.6 percent (x=34.1, N=8) of snout-vent length; snout moderately short, round in dorsal view, truncate in lateral profile; canthus round; loreal re- gion concave; lips slightly flared; nostrils three-fourths distance from eyes to tip of snout, slightly protuberant dorsolaterally; internarial area slightly depressed. Eye moderately large, directed anterolater- ally. Supratympanic fold weak; lower two-thirds of tympanum vis- ible, directed dorsolaterally with posterior inclination. Prevomerine dentigerous processes posteromedially inclined, narrowly separated medially between moderately large, ovoid choanae, bearing 0-4 teeth; tongue broadly cordiform, shallowly notched behind, barely 50 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY free posteriorly; vocal slits extending from posterolateral edges of tongue to angles of jaws. Humeral spine pointed, oriented about 30° anterior to humerus; ulnar fold and tubercles absent; first finger longer than second; fourth finger slightly shorter than third; lateral fringes present on fingers; webbing absent between first and second fingers, vestigial between second and third; webbing formula for outer fingers III (2-- 2*)-(1-1%)IV; discs broad, truncate; subarticular tubercles small, round, simple; supernumerary tubercles absent; palmar tubercle large, ovoid, simple; nuptial excrescences absent. Hind limbs mod- erately slender; length of tibia 51.4-57.6 percent (x=54.6, N=8) of snout-vent length; tarsal folds and tubercles absent; inner metatarsal tubercle small, flat, elliptical; outer metatarsal tubercle small, ovoid; subarticular tubercles small, round; supernumerary tubercles absent; feet about two-thirds webbed; webbing formula I(1-1%)-(2*-2*)IT1- (2-2*)IIT1-(2--2*)IV(2-2*)-1V; discs truncate, slightly smaller than those on fingers. Skin on dorsal surfaces of head, body, forearms, and shanks shagreened; skin on belly and posteroventral surfaces of thighs gran- ular; other surfaces smooth; anal opening directed posteroventrally at upper level of thighs; pair of large tubercles below anus. Color in preservative: dorsal surfaces, exclusive of two inner fingers and first three toes lavender, usually with numerous small dark lavender dots; other surfaces cream. Color in life: dorsal surfaces green with or without black flecks; tips of digits pale yellow; chest white; heart not visible; throat green; bones green; iris grayish white to pale bronze with brown or gray flecks. Distribution.—This species occurs to elevations of about 1200 m from lower Central America (Caribbean and Pacific slopes in Costa Rica and Panama) southward on Pacific slopes and lowlands to western Eicuador, where it has been taken at elevations of 220 to 800 m (Fig. 7). We have examined two specimens from the Pacific versant of Colombia. Cochran and Goin (1970:508) listed C. para- bambae from Medellin, Rio Mecaya, and Serrania de Macarena, Colombia. The last two localities are east of the Andes, so it is doubtful if the specimens from there actually are prosoblepon. Remarks.—The identity of Centrolenella parabambae has been greatly confused in the literature. As noted by Savage (1967:330): “Apparently the name [C. parabambae] has been applied to any population of small, uniformly lavender (in preservative) centro- lenids with vomerine teeth from Panama, Colombia, and Ecuador.” Savage noted that Panamanian frogs referred to “parambae” by —— CENTROLENID FROGS OF ECUADOR 51 Dunn (1933) are spinosa. Two characteristics of parabambae (as given in the type description by Boulenger, 1898) are responsible for the confusion: 1) absence of a humeral spine; the holotype is a female; 2) uniformly lavender dorsum, except for dark flecks on hind limbs. We compared a nearly uniformly lavender female (KU 121055) from Santo Domingo de los Colorados, Ecuador, with the holotype and noted that structurally the two frogs were identical; the holo- type has small indefinite dark flecks on the thighs, shanks, and feet, whereas KU 121055 has one fleck on one shank and two and three flecks on the feet. Subsequent comparison of KU 121055 with two females and five males of prosoblepon from Ecuador revealed that the only differences were in the numbers and disposition of dark flecks on the dorsum. The numbers of flecks in the nine specimens are (average value for limbs; means in parentheses after ranges) : head 0-38 (11.0), body 0-58 (23.1), forearm 0-11 (6.1), thigh 0-10 (5.2), shank 0-23 (9.4), foot 2-11 (5.1). In those specimens having many flecks on the body (49 and 58), the flecks are present over the entire dorsum; in those having fewer flecks (19, 21, 24, 32), the flecks are present only laterally. One individual has only five flecks (all laterally), and two lack flecks on the body. Two individuals have three and five small white flecks dorsally. One specimen from Colombia has reduced flecking, whereas the other is normal. Cursory examination of series of prosoblepon from Costa Rica and Panama revealed considerable variation in the num- ber of flecks on the dorsum, but no specimens lacked flecks. Taylor (1952:772) mentioned a specimen from Moravia, Costa Rica in which the spotting was “. .. sparse and confined largely to limbs and pos- terior part of dorsum.” The apparent continuum of variation in dorsal pattern and the absence of structural features to distinguish flecked and plain individuals necessitates the placement of Centro- lenella parabambae (Boulenger, 1898) in the synonymy of Centro- lenella prosoblepon ( Boettger, 1892). Three of our specimens were found on low vegetation in forest; four others were calling from vegetation over a cascading mountain stream at Balzapamba in July 1970. Tadpoles agreeing with the description of prosoblepon by Star- rett (1960:12) were collected late at night as they swam on the bottom of silt-bottomed pools in streams at Balzapamba. Centrolenella resplendens new species Plate 2C Holotype —KU 118053, an adult male, 27.3 mm, from Santa 52 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Cecilia, 340 m, Provincia Napo, Ecuador, obtained on 14 June 1967, by John D. Lynch. Diagnosis.—1) prevomerine teeth 2 on low processes; 2) bones white (?); 3) parietal and visceral peritonea white; 4) color in life dark green with white to bluish green flecks; in preservative, dull lavender with white flecks; 5) webbing between outer fingers I1I2--1IV; 6) webbing on foot I1-1%I10-2*11T1-2I1V2--0V; 7) snout round in dorsal view and gradually inclined anteroventrally in lat- eral view; 8) dorsal skin shagreened with elevated warts correspond- ing to white flecks; 9) scalloped dermal fold on outer edge of hand, forearm, and elbow, on heel, tarsus, and foot; U-shaped anal fold; 10) humeral spine absent in males; 11) lower four-fifths of tym- panum visible, directed dorsolaterally. Centrolenella resplendens is unlike any other Centrolenella known from South America; euknemos and pulverata from Central America are like resplendens in having scalloped dermal folds on the limbs. Centrolenella pulverata differs from the other two species by having the heart visible in life, dorsum white in preservative, snout short and slightly inclined anteroventrally, and dorsal skin uniformly granular. In contrast, in euknemos and resplendens the heart is not visible in life, the dorsum is lavender in preservative, the snout is long and gradually inclined anteroventrally, and the dorsal skin is shagreened with scattered elevations. Centrolenella re- splendens differs from euknemos by having a deeper snout that is round instead of subacuminate in dorsal view, more extensive anal folds, and more extensive webbing. The modal webbing formulae in euknemos are: hand—I[3-3113-3%III2-21V, foot I1-1%I11-28ITT1- 21V2-1V (Savage and Starrett (1967:606). Description—Adult male large, 27.3 mm in snout-vent length; females not known. Head slightly wider than body; width of head 34.8 percent of snout-vent length; snout long, round in dorsal view, gradually sloping from nostrils to tip in lateral profile (Fig. 1); canthus round; loreal region barely concave; lips slightly flared; nos- trils about two-thirds distance from eyes to tip of snout, not pro- tuberant, directed dorsolaterally; internarial area barely depressed. Eye large, directed more laterally than anteriorly. Supratympanic fold absent; tympanum directed dorsolaterally. Prevomerine den- tigerous processes low, short, transverse between choanae, each bearing two teeth; choanae large; longitudinally rectangular; tongue ovoid, barely free posteriorly; vocal slits extending from postero- lateral base of tongue to angles of jaws. Humeral spine absent; scalloped dermal fold around elbow, along ventrolateral edge of forearm and outer edge of fourth finger; CENTROLENID FROGS OF ECUADOR 53 first finger longer than second; fourth finger slightly shorter than third; lateral fringes on fingers; webbing absent between first and second fingers, vestigial between second and third; webbing formula for outer fingers III2--1IV; discs large, nearly round; subarticular tubercles small, round, simple; supernumerary tubercles absent; palmar tubercle simple, large, ovoid; nuptial excrescences absent (Fig. 3). Hind limbs moderately robust; length of tibia 53.5 percent of snout-vent length; scalloped fold around heel, along ventrolateral edge of foot and outer edge of fifth toe; inner metatarsal tubercle large, triangular; outer metatarsal tubercle absent; subarticular tu- bercles small, round; supernumerary tubercles absent; feet nearly fully webbed; webbing formula I1-1%II0-2‘HI1-21V2--0V; discs on toes round, slightly smaller than those on fingers (Fig. 4). Skin on dorsal surfaces of head, body, and limbs, and lateral surface of head and flanks shagreened with small tubercles corre- sponding to white spots; belly and proximal posteroventral surfaces of thighs granular; other surfaces smooth; anal opening directed posteriorly at upper level of thighs, bordered laterally and ventrally by fleshy tuberculate U-shaped fold; two conical tubercles below fold (Fig. 2). Color in preservative: head, flanks, and dorsal surfaces of body and limbs (except thumb and toes 1-3) dull lavender with many small, white spots; venter and posterior surfaces of thighs cream. Color in life: dorsum dark green with white to bluish green spots; fringes on forearm and foot and fold around anus white; venter yellow; iris gray. Distribution—Known only from the type locality and Santa Maria de Sucumbios, Colombia, about 40 km NNW of Santa Cecilia (Fig. 6). Remarks.—The holotype was perched on a leaf of a bush about 30 cm above the ground during light rain at night in primary rain- forest. Subsequent work at Santa Cecilia from 1967 through 1972 has resulted in the collection of thousands of frogs, but no additional specimens of resplendens have been found. The Colombian specimen (AMNH 88083) is a juvenile having a snout-vent length of 16.5 mm. The white dermal folds and flecks are essentially the same as those in the holotype, but the webbing, as typical of juveniles, is less extensive. Etymology.—The specific name is derived from the Latin verb resplendo meaning to glitter, and is used in allusion to the jewel-like appearance of the living frog. 54 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Centrolenella siren new species Plate 1E Holotype —KU 146610, an adult male, 20.7 mm, from a small tributary of the Rio Salado, about 1 km upstream from the Rio Coca, 1410 m, Provincia Napo, Ecuador, one of a series obtained on 7 April 1972, by William E. Duellman and John E. Simmons. Paratopotypes.—KU 146611-23, same date and collectors. Diagnosis ——1) prevomerine teeth 0-2; 2) bones green; 3) pari- etal peritoneum white; visceral peritoneum clear; 4) color in life green with yellow flecks; in preservative, lavender with white flecks; 5) webbing between outer fingers III2%-2%IV; 6) webbing on foot 12-2411 135-27s1111s-3-1V3-2-V; 7) snout truncate in dorsal and lateral profiles; 8) dorsal skin shagreened; 9) arms and legs lacking dermal folds; 10) humeral spine absent in males; 11) lower four-fifths of tympanum visible, directed posterolaterally with slight dorsal in- clination. Centrolenella siren is like flavopunctata and midas in having a lavender dorsum with white flecks (in life, green with gold flecks) with no black flecks, but it differs from both of those species by having much less webbing on the hands and feet and a more promi- nent tympanum oriented posterolaterally, instead of dorsolaterally. Centrolenella flavopunctata further differs from siren by having a rounded, instead of truncate, snout. Description—Adults moderately small; snout-vent length 19.8- 22.0 mm (x=20.8, N=14) in males; females unknown. Head much wider than body; width of head 30.0-37.0 percent (x=34.7, N=14) of snout-vent length; snout extremely short, truncate in dorsal and lat- eral views; canthus round; loreal region barely concave; lips not flared; nostrils nearly terminal on snout, not protuberant, directed laterally; internarial area flat. Eye moderately large, directed antero- laterally. Supratympanic fold absent; lower four-fifths of tympanum visible, directed posterolaterally with slight dorsal inclination. Pre- vomerine dentigerous processes small, low, widely separated, be- tween small round choanae, bearing 0-2 teeth; tongue round, barely free posteriorly; vocal slits extending from midlateral base of tongue to angles of jaws. Humeral spine absent; ulnar fold and tubercles absent; first fin- ger about equal in length to second; fourth finger noticeably shorter than third; fringes absent on fingers; webbing absent between first and second fingers, vestigial between second and third; webbing formula for outer fingers III ( 2%-2% )-( 24-24 )IV; discs truncate; sub- articular tubercles small, round, simple; supernumerary tubercles absent; palmar tubercle large, elliptical; nuptial excrescences absent. $$ = CENTROLENID FROGS OF ECUADOR 55 Hind limbs slender; length of tibia 54.8-60.1 percent (x=57.3, N=14) of snout-vent length; tarsal folds and tubercles absent; inner meta- tarsal tubercle small, ovoid; outer metatarsal tubercle absent; sub- articular tubercles small, round; supernumerary tubercles absent; feet about one-half webbed; webbing formula I2-(2*-24)II(14-1%)- (27-2% ) III 1s- ( 24-3 )1V3-(1%-2)V; discs on toes smaller and more nearly round than those on fingers. Skin on dorsal surfaces shagreened; skin on belly and ventral surfaces of thighs granular; other surfaces smooth; anal opening di- rected posteriorly at upper level of thighs; pair of large tubercles below anus (Fig. 2). Color in preservative: dorsum of head, body, and limbs lavender with many small, white flecks on head and body; other surfaces pale cream. Color in life: dorsum green with gold flecks; fingers and toes pale yellow; chest white; heart not visible; other ventral surfaces pale dull green; bones green; iris pale bronze with fine black re- ticulations. Distribution —Centrolenella siren occurs in cloud forest on the Amazonian slopes of the Andes, where it is known from elevations of 1410-1740 m (Fig. 9). Centrolenella siren occurs in sympatry with anomala, audax, megacheira, pellucida, and pipilata. Remarks.—All individuals were found at night on low vegetation along small mountain streams in cloud forest. The type locality is a small stream at the south edge of the Papallacta-Lago Agrio road about 300 m west of the bridge across the Rio Salado and approxi- mately 1 km up the Rio Salado from its confluence with the Rio Coca. Etymology.—The specific name refers to the nymphs of Classical mythology, who by their sweet singing enticed seafarers to destruc- tion; the name is used here in allusion to the fact that the call of these small frogs entices collectors to the nocturnal perils of streams. DISCUSSION Savage (1967) recognized three species groups of Centrolenella in Central America. He characterized the fleischmanni group as having white bones in life, a colorless parietal peritoneum, a white visceral peritoneum, a white ground color in preservative, and in lacking prevomerine teeth and dentigerous processes. The pro- soblepon group was characterized as having green bones in life, a white (opaque) parietal peritoneum, a colorless visceral peritoneum, lavender ground color in preservative, and prevomerine teeth and 56 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY dentigerous processes. He recognized a third group for the some- what annectant pulverata. Three Ecuadorian species (fleischmanni, munozorum, and pellu- cida) clearly fit into Savage’s fleischmanni group. With the excep- tions of anomala, medemi, ocellifera, and resplendens, the remaining Ecuadorian Centrolenella agree with Savage’s definition of the pro- soblepon group. The presence of prevomerine teeth and processes is variable in the Ecuadorian representatives of the prosoblepon group (sensu Savage) but is usually constant within species. How- ever, a single specimen of cochranae and one of prosoblepon lack teeth and processes, whereas all other specimens of these species examined have them. Except for the absence of prevomerine teeth and _ processes, buckleyi, grandisonae, griffithsi, megacheira, peristicta, and pipilata agree with the definition of the Central American prosoblepon group. However, the agreement is weakened by the pale-green (contrasted with green) bones of grandisonae, griffithsi, and peristicta (a character-state of the pulverata group). Savage characterized the pulverata group as having pale green bones, a colorless parietal peritoneum (heart visible), opaque visceral peritoneum, yellowish- white ground color in preservative, and prevomerine teeth and processes. On examining the holotype of pulverata (ZMB 7842) in 1969, Duellman noted the ground color was lavender. Aside from having pale green bones, grandisonae, griffithsi, and peristicta do not agree with the definition of the pulverata group. We know of no Ecuadorian Centrolenella assignable to the pulverata group. The Venezuelan Centrolenella antisthenesi may be allied with pulverata; antisthenesi has a clear parietal peritoneum, opaque visceral peri- toneum, prevomerine teeth, lavender dorsum, green bones, and sub- anal tubercles and lacks a humeral spine. Three Ecuadorian species (and possibly the Colombian medemi) depart from the fleischmanni-prosoblepon arrangement. Centro- lenella anomala is prosoblepon-like, except in lacking prevomerine teeth, having white bones, and in lacking a lavender ground color. The opaque parietal peritoneum and brown ground color exclude it from the homogeneous fleischmanni group. Centrolenella ocellifera and resplendens presumably have white bones (green bones were not recorded in their color descriptions in life). Both have prevo- merine teeth and low dentigerous processes, an opaque parietal peritoneum, and lavender ground color. Centrolenella resplendens differs further in having white visceral peritoneum (a character- state of the fleischmanni and pulverata groups). The combination : CENTROLENID FROGS OF ECUADOR 57 of characteristics cited above does not recommend them for any of Savage's (1967) groups. We noted the absence of large subanal tubercles in anomala, fleischmanni, medemi, munozorum, ocellifera, and pellucida, and their presence in the remaining Ecuadorian Centrolenella. Subanal tubercles are absent in most Central American centrolenids; we found them in ilex, prosoblepon, and pulverata and in antioquiensis (Colombia) and antisthenesi (Venezuela). The absence of subanal tubercles in the frogs of the fleischmanni group and in anomala, medemi, and ocellifera (enigmatic in having white bones) thus may not be significant. Tubercles are present in resplendens (also enig- matic in presumably having white bones) but these may not be homologous to those in the frogs lacking the elaborate post- and para-anal ornamentation of resplendens. Humeral spines are not known in any frog of the fleischmanni group. Goin’s (1964) and Savage’s (1967) conclusion that presence or absence of the spines cannot be used to support generic sepa- ration of Centrolenella and Cochranella is substantiated here in that an otherwise “normal” griffithsi has humeral spines. This specimen may prove to be distinct from griffithsi but at present no features other than the humeral spine support that argument. Accordingly, the presence of humeral spines possibly may be intraspecifically variable, at least in griffithsi. Small spines occur on the humeri of medemi but not in anomala, ocellifera, or resplendens (species with presumed or known white bones). The presence of humeral spines in audax, buckleyi, grandisonae, medemi, peristicta, pipilata, and prosoblepon does not suggest to us that these frogs are more closely related to one another than they may be to species lacking humeral spines. This assemblage seems to be heterogeneous: three have pre- vomerine teeth and processes, two have pale green as compared to green bones, and one lacks subanal tubercles. Centrolenella antioquiensis and resplendes differ from all other known Centrolenella by having opaque parietal and visceral peri- tonea. Otherwise they share few characters; antioquiensis has a humeral spine and green bones (absent and white in resplendens) and lacks dermal ornamentation and prevomerine teeth (both pres- ent in resplendens ). The Ecuadorian Centrolenella thus far described do not readily fit into the arrangement of Central American forms proposed by Savage (1967). Inspection of Taylor and Cochran’s (1953) account of southeastern Brasilian centrolenids reveals comparable lack of fit with Savage's three groups in Central America. Because we antic- ipate the discovery of many undescribed Centrolenella in north- 58 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY western South America, at this time we prefer not to alter Savage's groupings or define new species groups based on our study of Ecua- dorian species. We offer only the following comments on putative relationships of the 19 species, as described below. Centrolenella fleischmanni, munozorum, and pellucida constitute a well-circumscribed group of species having white bones and a colorless parietal peritoneum (heart visible) in life, a white visceral peritoneum, a white ground color in preservative, and in lacking distinct canthi, subanal tubercles, humeral spines, and prevomerine teeth and processes. Centrolenella audax, cochranae, flavopunctata, midas, prosoble- pon, and siren constitute a group having green bones and an opaque parietal peritoneum (heart not visible) in life, a clear visceral peri- toneum, a very pale to dark lavender ground color in preservative, and in having distinct canthi, subanal tubercles, and prevomerine teeth and processes. Two of these (audax and prosoblepon) have humeral spines. Centrolenella buckleyi, grandisonae, griffithsi, megacheira, peri- sticta, and pipilata fall into a somewhat heterogeneous group agree- ing in most respects with that listed above, except in the uniform absence of prevomerine teeth and processes. The bones are pale green in grandisonae, griffithsi, and peristicta; griffithsi and mega- cheira lack humeral spines. Centrolenella anomala differs markedly from all other centro- lenids in the combination of white bones and an opaque parietal peritoneum in life, clear visceral peritoneum, a brown ground color in life and in preservative, distinct canthi, and in lacking prevo- merine teeth and processes, subanal tubercles, and humeral spines. Its ocellated color pattern on a shagreened dorsum with warts and the reduced webbing of the fingers lend it a striking similarity to cochranae; the similarity is out-weighed by the numerous differences between the two species. Centrolenella ocellifera also differs markedly from other centro- lenids. The presumably white bones in life, opaque parietal peri- toneum, clear visceral peritoneum, and absence of subanal tubercles, coupled with the pale lavender ground color in preservative, distinct canthi, prevomerine teeth and processes, ocellated color pattern, and the absence of humeral spines suggests some relationship with ano- mala or with the group of species listed above, which included audax and cochranae. Centrolenella resplendens appears to be trenchantly different from other Ecuadorian Centrolenella chiefly because of its dermal ornamentation (scalloped limb fringes, para- and postanal ornamen- ne ee CENTROLENID FROGS OF ECUADOR 59 Pdramo and Subparamo BS buckleyi Bel buckley/ ie, Os) x Cloud Forest e ‘at Ky janomala 2000 3 54 d “ w d S S 2 8 2 dS 6mos. >12mos. >18mos. \S24ines: Sceloporus variabilis 604 70(11.6%) 0 0 0 Anolis humilis 305 6(1.9%) 1(.33%) 0 0 Anolis cupreus 1482 48(3.24%) 3(.20%) 0 0 Anolis tropidolepis 316 27(8.45%) 7(2.22%) 0 0 Basiliscus vittatus 90 ' 6(6.66%) 4(4.45%) 2(2.23%)” “2022 e7 POPULATION STRUCTURE AND SURVIVORSHIP 31 size at sexual maturity, larger adult female size, slightly smaller number of eggs per clutch, and a higher ratio of oviparous to vivip- arous species. Of the 15 tropical species discussed here, only one, Ctenosaura similis, is late-maturing and single-brooded, whereas nine are early maturing and multiple-brooded. The other four are somewhat intermediate; the viviparous Sceloporus malachiticus is single-brooded, and Basiliscus basiliscus, B. vittatus and Anolis tropidolepis have somewhat delayed maturity. As to size, most of the 15 species are small, but the average is raised by the relatively gigantic Ctenosaura similis and also by the two species of Basiliscus. With few exceptions, the number of eggs is small. The clutch consists of only one egg in the six species of anoles and the gecko, Gonatodes albogularis. In the two teiids and the skink, the average clutch size is between 2 and 3. The average clutch size is 3.0 in Sceloporus variabilis, 3.9 in Basiliscus vittatus, 4.6 in Sceloporus malachiticus, 6.0 in Basiliscus basiliscus. Only Ctenosaura similis has large clutches of more than 20 eggs. Of the 15 species, only Sceloporus malachiticus is viviparous; of the 58 species of lizards recorded from Costa Rica, there are only three that are known to be viviparous. Tinkle, et al. (1970) noted the absence of a significant correla- tion between clutch size and body size at maturity in tropical lizards. However, such correlation definitely does exist in the 15 species that I investigated. Doubtless the lack of correlation found by Tinkle et al. resulted, at least in part, from the fact that insular species (having few natural enemies and reduced clutch size) were included along with mainland kinds. Also, their list included species of Draco, which parallels Anolis in specialization for an active, arboreal existence, including reduction in clutch size. If the insular species and the species of Draco are excluded, the remaining 12 examples of Tinkle et al. do show high correlation of body size and number of eggs per clutch. A much different combination of ecological traits from those hypothesized for tropical lizards by Tinkle et al. (1970) would be expected from concepts of selection proposed or developed by Dobzhansky (1951), MacArthur and Wilson (1967), Pianka (1970) and also MacArthur (1972). They recognized two basic types of selection to which populations are subject, depending on the type of environment. “K-selection” is most characteristic of en- vironmental type best exemplified by a tropical rain forest. In this type, physical factors are relatively benign and stable. Weather is predictable. The populations of a given organism tend to be stable, and often in equilibrium near the carrying capacity. Inter- specific and intraspecific competition is usually keen, and mortality is density-dependent. Animals are highly adapted to their com- munities and are “fit” so that they have notable longevity, with slow development, delayed maturity, large body size, few young 32 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY per clutch, repeated reproduction and parental care (occasionally expressed in viviparity). Conversely, “r-selection” is most charac- teristic of an environment typified (in extreme form) by an Arctic tundra. There, environmental conditions are unstable and often harsh. Weather is subject to extremes, which may bring about catastrophic decimation of animal populations in a density-inde- pendent fashion. As a result populations are unstable, are often below the carrying capacity, and competition may be lax. Selection favors a short life span, rapid development, early maturity, small size, energy channeled into a single, early reproductive effort, with a large number of offspring, and lack of parental care. Most terrestrial environments fall somewhere between these two extremes. Presumably the assemblage of tropical lizard species dealt with in the present study represents environments that are most likely to show K-selection. Table 2 is designed to test con- formity of each species with the two types of selection on the basis of eight separate criteria. A species that conformed with r-selection in all respects would have a rating of 8, one that conformed with K-selection would have a rating of 24 and one that was intermediate would have a rating of 16. Assuming that the criteria used are appropriate and that the ratings have been made correctly, it ap- pears that none of the species conforms entirely with either type, but that 12 of the 15 fit r-selection better than K-selection. “Coarse-grained” and “fine-grained” environments have been recognized by several recent authors as producing different types of selection, the latter permitting more specialization and closer adap- tation. According to Levins (1968) a species “loses fitness in a heterogeneous environment. It would be better off specialized, but the uncertainty of the environment forces niche expansion.” Compared on this basis, the 15 species of this study seem to have members in each type of environment and some that are inter- mediate. Basiliscus basiliscus is perhaps the best example of a species living in a coarse-grained environment. Within the area where it occurs it is not generally distributed but is limited to watercourses and their vicinity, and along them it is irregularly distributed with concentrations where there are log jams, uprooted large trees, or undermined banks with dense thickets. These alter- nating features are accentuated by seasonal change; a stream that is a rushing torrent during the rainy season may be reduced to a series of puddles in the dry season, and the log jams and thickets that are used as shelter may be drastically altered or swept away completely in time of flood. Basiliscus vittatus and Anolis lionotus are also streamside species subject to some of the same coarse- grained features. Ctenosaura similis and Sceloporus malachiticus are usually associated with irregularly distributed environmental features such as rock outcrops, buildings, or hollow trees, and their TABLE 2.—-r-selection and K-selection in Some Costa Rican Lizards POPULATION STRUCTURE AND So | Slows BUI UOTI[IS ouy—¢ 9} VIPSULIoJUI=F 9SIVOO= | AJULIOFIUN }EPIQVET qqvis=E O}CIPOULI9}UI=F [P10sS=T AVIGeyS JeUGVH ‘sIA Z UBY} BIOWU=—E sIvak G-[=G 1e9k-[—7T I9AOUIN} uone[ndog [Ss V-G—G d10UL 10 G=T spooiq Jo 1aquinyy (EIS orc 6 91OUL 10 J=T SUNOA 10 $939 JO Ioquiny euou 10 8} =E uesi0 Av[dstp Io /pue 10j0oo=% IO[OD puke 9ZIS=T tustydiour4rp jenxas GI uUeY} s10U=¢ OL956 ool “SOUL UT A}LIN} UL 0} OUT], d10UI 10 OOS =F OST-OL—6 09-S€=1 ( UIUI UI A-S ) OZI§ ise) re Ameiva quadrilineata Anolis cupreus Anolis humilis ix re Mil CD) CD) CD T= SH Seq nee Anolis intermedius Anolis limifrons Anolis lionotus Basiliscus basiliscus Anolis tropidolepis Basiliscus vittatus Sy eq or | 1 3 Cnemidophorus deppii Ctenosaura similis laliee) mee 1 i 2 i Gonatodes albogularis Leiolopisma cherriei Te) re Sceloporus malachiticus Sceloporus variabilis SURVIVORSHIP i) o 34 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY environments are likewise coarse-grained. In contrast the species of leaf litter in rain forest (Anolis humilis, Leiolopisma cherriei) or those of tree-trunks (Gonatodes albogularis, Anolis tropidolepis ) have fine-grained environments. Again, there is no definite corre- lation between these environmental types and population structure or survivorship. CONCLUSIONS Four types of population structure were found to result from the varying climates and reproductive strategies in the 15 species of tropical lizards studied in Costa Rica. Type 1 is a stable popula- tion with all ages constantly represented, from the most numerous and youngest to the relatively scarce oldest lizards; type 2 also has all age groups constantly present, but their ratios are constantly changing, with an older group sometimes better represented than a younger one reflecting fluctuation in the level of reproduction. Type 3 results from seasonality of breeding, and changes through- out the annual cycle, from relative homogeneity (with all adults) to heterogeneity with immatures of all sizes as well as adults. Type 4 consists of a structured population of many successive an- nual cohorts, and results from a short annual breeding season and delayed maturity, paralleling trends in the kinds of lizards that occur farthest from the equator. Small body size, rapid population turnover, frequent reproduc- tion, oviparity, small clutch-size, type 2 or 3 population structure, and sexual dimorphism (in color or display organs or body size or all three) are characteristic of the populations studied with few exceptions. Although occurring in communities that would be expected to have mainly K-selection, the species do not conform consistently to the criteria expected in products of K-selection. The majority of the species fit better with the criteria of r-selection, but each species shows some mixture of the two sets of traits, and no two species are alike in this respect. In contrast with lizard populations of the temperate zones, those of the tropics are less restricted to seasonal schedules. Living in more constant environments, they are able to distribute their ac- tivities, such as those concerned with reproduction, over a greater part of the annual cycle. The iguanid anoles and the sphaerodac- tyline geckos, two groups that are highly successful in the Neo- tropical region, have progressed farthest in this regard; in rain forests and cloud forests their reproductive effort is uniformly distributed over the entire year. The female lays one egg at a time, the left and right ovaries alternating, so that, usually carrying two eggs in different stages of growth, she is not much handicapped and continues her normal activities. Aside from its various other implications, the one-egg clutch may be regarded as an extreme POPULATION STRUCTURE AND SURVIVORSHIP 35 specialization for an active way of life. Each egg may be left in a different place (though both groups are known to use communal nests at times). Eggs are deposited in damp sheltered places, but are not buried in special nest burrows. Therefore production of eggs and oviposition involve a minimum of stress. Dispersal of eggs in time and space provides security from: a) catastrophic events such as flood or drought which might catch the eggs or young at a vulnerable stage, b) concentrated predation, c) intense intraspecific competition for food and space, such as might occur when many young emerge simultaneously from the same nest. Climates in the tropics that have strong seasonality impose annual cycles on their lizard populations. In areas that have a severe dry season, the lizards’ annual cycles are comparable with those that occur in species of the temperate zones except that there is no hibernation period. There is a breeding season of varying length but confined to one part of the year and presumably ad- justed to avoid the worst rigors of the year-round climate and ex- ploit the benefits, such as adequate moisture for the incubating egg, and an abundant supply of suitable food (usually insects) for the breeding adults and/or the hatchlings. Even in the wet climate of the Caribbean lowlands some lizards, especially those of seral situations, such as Anolis limifrons, Basiliscus vittatus and Leiolo- pisma cherriei alter the level of their reproductive activity in re- sponse to wetter or drier weather at certain times of year. In gen- eral, heavy precipitation is correlated with heightened reproduc- tion, whereas drought is correlated with slowing or cessation of reproductive activity. The long and severe dry season of western Costa Rica (Guana- caste and northern Puntarenas provinces, where monthly precipi- tation in December, February, March, and early April is usually less than 25 mm) limits reproduction mainly to the wetter half of the year in most species. An extreme case is Ctenosaura similis which is limited to a short and concentrated breeding season, with large clutches of eggs, relatively small hatchlings, and delayed maturity as facets of its reproductive strategy. In ctenosaurs, the young differ from adults in their behavior, microhabitats ( Hender- son, 1973), size and type of food items (Montanucci, 1968) hence intraspecific competition is minimized. In Basiliscus basiliscus, Iguana iguana and Anolis cupreus there is a less well-defined segre- gation of young and adults. SUMMARY Free-living populations of 15 species of lizards were studied at 14 localities in Costa Rica; the study sites include a wide range of altitudes, habitats and climates. Mark and recapture studies were 36 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY carried out over a three-year period for most of the species. At most of the localities two or more species were studied simultane- ously; thus the results provide a basis for both intraspecific and interspecific comparisons. Unlike lizards of the Temperate Zone, the tropical species in- cluded in this study all are active throughout the year; however most of them show some seasonality in their annual cycles. Their different population structures result largely from the timing of reproduction and rate of development and survivorship of the cohorts of young produced. In some kinds, population structure changed markedly in response to seasonal changes in the weather. Recognizable components of individual populations were adult males and females, and immatures at various stages of development. Snout-vent length is highly correlated with age in immatures; there is less correlation in adults because growth continues after attain- ment of maturity. On the basis of their population structures, reproduction and survivorship, the species studied can be arranged in a graduated series from those of small size, early maturity, high production, and rapid turnover, to those that are large and have delayed ma- turity, low production and slow turnover. Four fairly distinct but overlapping types of population structure can be recognized within this range. Type 1 is confined to climates lacking seasonal contrast and results from year-round reproduction at a uniform level. The population structure is stable and includes individuals of all ages in unchanging ratios. This type is best exemplified by Anolis tropidolepis of montane cloud forest, and less perfectly by A. hu- milis, probably A. lionotus, and by Gonatodes albogularis, all of the Caribbean rain forest. Type 2 also has year-round reproduction with all ages represented in the population at all times, but the level of reproduction and the ratios of different age groups undergo constant seasonal change. This type occurs in wet climates that have moderate seasonal change and was found developed to various degrees in Ameiva quadrilineata, Anolis limifrons, Basiliscus vit- tatus, and Leiolopisma cherriei. Type 3 is characteristic of areas having a long dry season, where lizards generally confine their reproduction to the wetter part of the year. At the beginning of the breeding season the population consists essentially of reproductive adults, but subsequently successive increments of young are added until all classes from hatchlings to adolescents are found along with the adults (by then reduced to relatively low numbers) at the end of the breeding season. Type 3 prevails in northwestern Costa Rica and was found in Anolis cupreus, A. intermedius, Basiliscus basiliscus, Cnemidophorus deppii, Gonatodes albogularis, Scelop- orus variabilis and S. malachiticus. The latter species is transitional to type 4 which otherwise was found only in the large iguanid POPULATION STRUCTURE AND SURVIVORSHIP 37 Ctenosaura similis. In type 4 there is a relatively short annual breeding season and delayed maturity resulting in a sharply struc- tured population that consists of several or many discrete annual age groups at least two of which are immatures. Survivorship was judged from the rate at which marked individ- uals disappeared from the populations on the small study areas, but some of the turnover was caused by shifts of individuals and this spatial displacement could not be evaluated accurately. In most species, population turnover was rapid, with most individuals re- placed in the course of a year. It was especially rapid in the smaller species of anoles, in Cnemidophorus deppii, Ameiva quadrilineata and Sceloporus variabilis. In contrast to these species the montane Anolis tropidolepis and the two species of basilisks are notably longer lived and individuals often survive for more than a year. Ctenosaura similis clearly is in a class by itself surviving longer than any of the smaller kinds of lizards studied. Previous findings and predictions about populations of tropical lizards are partially borne out by the present findings. Conforming with trends indicated by Tinkle et al. (1970), most of the tropical lizards included in this study have relatively small clutches, and have a low ratio of viviparous to oviparous kinds compared with some from the temperate zone. However, the lack of correlation between size of female and number of eggs per clutch reported by Tinkle et al. is not borne out. Most of the species were found to conform to the strategy of early maturity and multiple broods. Only one of the 15, Ctenosaura similis, definitely conforms to the opposite strategy of late maturity and single broods. Appraised by some of the main criteria of r-selection and K-selection, no species of the 15 is fully committed to either type; instead all are inter- mediate in some degree, with a majority favoring r-selection. RESUMEN Poblaciones salvajes de 15 especies de lagartos fueron estudiadas en 14 localidades en Costa Rica; los lugares de estudio incluyen un amplio rango de altitudes, habitats y climas. Estudios de marca y recaptura fueron Ilevados a cabo por un periodo de tres afos para la mayoria de las especies. En la mayoria de las localidades dos o mas especies fueron estudiadas simultaneamente; por lo tanto los resultados proporcionan una base para comparaciones in- traespecificas e interespecificas. A diferencia de los lagartos de las zonas templadas, las especies tropicales incluidas en este estudio son activas durante todo el ano; sin embargo, casi todas muestran periodicidad en sus ciclos anuales. Las diferentes estructuras de las poblaciones resultan mayormente de la regulacién del tiempo de reproduccién y la tasa de desarrollo y sobrevivencia de las cohortes de jévenes pro- 38 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY ducidas. En algunos tipos la estructura de las poblaciones cam- biaba marcadamente en respuesta a cambios periddics en el clima. Componentes reconocibles de poblaciones individuales fueron machos y hembras adultos, e inmaduros a varios niveles de de- sarrolo. La distancia (cabeza-tronco) esta altamente correla- cionada con la edad en los inmaduros; hay menor correlacién en los adultos porque el crecimiento continua despues de la madurez. En base a las estructuras de sus poblaciones, reproduccién y supervivencia, las especies estudiadas pueden ser agrupadas en una serie gradual de aquellos de tamafo pequeno, temprana ma- durez, alta produccién, y rapido reemplazo, a aquellos que son grandes, y que tienen madurez tardia, baja produccién y reemplazo lento. Cuatro tipos de estructuras de poblaciones bastante difer- enciables pero sobrepon‘bles pueden ser reconocidas en este rango. El tipo 1 esta confinado a climas sin contrastes de estaciones y resulta de reproduccién durante todo el afio a un nivel uniforme. La estructura de la poblacién es estable e incluye individuos de todas las edades en proporciones invariables. Este tipo esta bien representado por Anolis tropidolepis de selvas nubladas de mon- tafia, y menos perfectamente por A. humilis, probablemente A. lionotus, y por Gonatodes albogularis, todos de las selvas lluviosas del Caribe. E] tipo 2 tambien tiene reproduccién durante todo el ano con todas las edades representadas en la poblacion durante todo el tiempo, pero el nivel de reproduccién y las proporciones de diferentes grupos de edades camb‘an constantemente con los cam- bios de estaciones. Este tipo ocurre en climas himedos que tienen un cambio de estaciones moderado y fué encontrado desarrollado en varios grados en Ameiva quadrilineata, Anolis limifrons, Basilis- cus vittatus, y Leiolopisma cherriei. E] tipo 3 es caracteristico de areas que tienen una larga estacién seca, donde los lagartos con- finan su reproduccién a las partes mas hiimedas del afio. Al co- mienzo del periodo de cria la poblacién consiste esencialmente de adultos en capacidad de reproducirse, pero incrementos subsecuentes de los j6venes son afadidos hasta que todas las clases desde reciente nacido hasta adolescentes son hallados junto con adultos (entonces disminuidos a pequefios nimeros) al comienzo de cada periodo de cria. E] tipo 3 prevalece en el noreste de Costa Rica y se encuentra en Anolis cupreus, A. intermedius, Basiliscus basiliscus, Cnemi- dophorus deppii, Gonatodes albogularis, Sceloporus variabilis, y S. malachiticus. Esta ultima especie es una transicién con el tipo 4 que de otra forma fué hallado solamente en la iguanida grande, Ctenosaura similis. En el tipo 4 hay un periodo relativamente corto de cria y madurez tardia resultando en una marcada estructura de poblacién que consiste de varios 0 muchos grupos discretos de edades al menos dos de los cuales son inmaduros. La supervivencia fué estimada de la razon a la cual los individuos marcados desapare- POPULATION STRUCTURE AND SURVIVORSHIP 39 cian de las poblaciones en las pequefas areas estudiadas, pero parte del reemplazo fué causado por desplazamientos de individuos, y este desplazamiento de espacio no pudo ser evaluado con precisién. En la mayoria de las especies el reemplazo de la poblacién fué rapido, con casi todos los individuos reemplazados en el curso de un ano. Fué esencialmente rap'do en las especies mas pequefnas de Anolis, en Cnemidophorus deppii, Ameiva quadrilineata, y Scel- oporus variabilis. En contraste con estas especies la montafiosa Anolis tropidolepis y las dos especies de basiliscos son notablemente de vida mas larga e individuos frecuentemente sobreviven por mas de un ano. Ctenosaura similis esta claramente en una clase por si misma sobreviviendo por mayor tiempo que cualquiera de las especies mas pequenas de lagartos estudiadas. Anteriores resultados y predicciones sobre poblaciones de la- gartos tropicales son parcialmente anulados por estos resultados. De acuerdo a tendencias indicadas por Tinkle et al. (1970) los lagartos tropicales incluidos en este estudio tienen mayormente nidadas pequefias, y una proporcién pequena de tipos viviparos a Oviparos comparados con algunos de las zonas templadas. Sin embargo, la falta de correlacién entre el tamafo de la hembra y el numero de huevos por nidada reportados por Tinkle et al. no fué anulado. Se hall6 que la mayoria de las especies se adhieren a la estrategia de temprana madurez y multiples camadas. Solo una de las 15, Ctenosaura similis, se adhiere definitivamente a la estra- tegia opuesta de tardia madurez y una Unica camada. Ectimadas por algunos de los criterios fundamentales de seleccion-r y seleccién- K ninguna de las 15 especies puede ser catalogada como una u otra, pero todas fueron intermedias en algin grado, con una mayoria favoreciendo seleccién-r. LITERATURE CITED AtcaLaA, A. C., and W. C. Brown 1967. Population ecology of the tropical scincoid lizard, Emoia atroco- stata, in the Philippines. Copeia, 1967 (3):596-604. ASANA, J. J. 1931. The natural life history of Calotes versicoor (Boulenger) the com- mon blood-sucker. Jour. Bombay Nat. Hist. Soc., 34:1041-1047. BAKER, J. R. 1947. The seasons in a tropical rain-forest. Part 6. Lizards (Emoia). Jour. Linn. Soc. London, 41 (279) :243-247. Buair, W. F. 1960. The rusty lizard, a population study. University of Texas Press, Austin, xvi+185 pp. BusTARD, H. R. 1968. The ecology of the Australian gecko Heteronotia binoei in northern New South Wales. Jour. Zool., 156(4):483-497. 1969. The population ecology of the gekkonid lizard Gehyra variegata (Duméril and Bibron) in exploited forests in northern New South Wales. Jour. Animal Ecol., 38:35-51. AO OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY CARPENTER, C. C. 1960. Parturition and behavior at birth of Yarrow’s spiny lizard (Scelop- orus jarrovi). Herpetologica, 16(2):137-138. DoBzHANSky, T. 1951. Genetics and the Origin of Species. Columbia Univ. Press, New York. FircH. HwS; 1954. Life history and ecology of the five-lined skink, Eumeces fasciatus. Univ. Kansas Publ. Mus. Nat. Hist., 8:1-156. 1956. An ecological study of the collared lizard (Crotaphytus collaris). Ibid., 8:213-2T4. 1958. Natural history of the six-lined racerunner (Cnemidophorus sex- lineatus). Ibid., 11:11-62. 1970. Reproductive cycles of lizards and snakes. Misc. Pub. Mus. Nat. Hist. Univ. Kansas, 52:1-247. 1972. Ecology of Anolis tropidolepis in Costa Rican cloud forest. Herpe- tologica, 28:10-21. 1973. A field study of Costa Rican lizards. Univ. Kansas Sci. Bull., 50:39-126. Fircu, H. S., and H. W. GREENE 1965. Breeding cycle in the ground skink. Lygosoma laterale. Univ. Kan- sas Publ. Mus. Nat. Hist., 15:565-575. GREENE, H. W. 1969. Reproduction in the Middle American skink, Leiolopisma cherriei (Cope). Herpetologica, 25:55-56. Harr, J. 1971. Ecology of a population of the Great Plains skink (Eumeces obso- letus). Univ. Kansas Sci. Bull., 59:357-388. Harris, V. A. 1964. The life of the rainbow lizard. Hutchison Tropical Monographs, Hutchison and Co., Ltd., London, 174 pp. HENDERSON, R. W. 1973. Ethoecological observations of Ctenosaura similis (Sauria: Iguani- dae) in British Honduras. Jour. Herpetology, 7(1):27-33. Hints, HLF. 1963. The ecology of two lizards on a tropical beach. Ecol. Monogr., 33:83-112. IncER, R. F., and B. GREENBERG 1966. Annual reproductive patterns of lizards from a Bornean rain forest. Ecology, 47(6):1007-1021. Levins, R. 1968. Evolution in changing environments. Monographs in population biology, No. 2. Princeton Univ. Press, New Jersey, ix+120 pp. MacArrtuur, R. H. 1972. Geographical ecology: Patterns in the distribution of species. Harper & Row, Inc., New York, xviii+269 pp. MacArrTuur, R. H., and E. O. Witson 1967. The theory of island biogeography. Princeton Univ. Press, Prince- ton, New Jersey. Manion, K. R. and O. J. SExTon 1971. The reproductive cycle of the lizard Sceloporus malachiticus in Costa Rica. Copeia, 1971(3):517-526. MarsHAL., A. J., and R. Hoox 1960. The breeding biology of equatorial vertebrates: reproduction of the lizard Agama agama lionotus Boulenger at Lat. 0° 01’ N. Proc. Zool. Soc. London, 134:197-205. POPULATION STRUCTURE AND SURVIVORSHIP 4] MayHew, W. W. 1964a. Photoperiodic responses in three species of the lizard genus Uma. Herpetologica, 20:95-113. 1964b. Reproduction in the sand-dwelling lizard Uma inornata. Ibid., 21:39-55. 1966a. Reproduction in the psammophilous lizard Uma scoparia. Copeia, 1966(1):¥14-122. 1966b. Reproduction in the arenicolous lizard Una notata. Ecology, 47(1): 9-18. MILER, M. R. 1951. Some aspects of the life history of the yucca night lizard, Xantusia vigilis. Copeia, 1951(2):114-120. MOontTANUwCcCcI, RICHARD R. 1968. Comparative dentition in four iguanid lizards. Herpetologica, 24:305-315. Panna, S. K., and J. P. THAPLIyAL 1967. Egg laying and development in the garden lizard, Calotes versi- color. Copeia, 1967(1):121-125. PIANKA, E. R. 1970. On r and K selection. American Naturalist, 100:592-597. 1971. Ecology of the agamid lizard Amphibolurus isolepis in western Australia. Copeia, 1971(3) :527-536. SanyAL, M. K., and M. R. N. Prasap 1967. Reproductive cycle of the Indian house lizard, Hemidactylus flavi- viridis Riippell. Copeia, 1967 (3) :627-633. SExTon, O. W., H. F. HEATWOLE, and E. H. MesErH 1963. Seasonal population changes in the lizard, Anolis limifrons, in Panama. Amer. Midl. Nat., 69:482-491. SEXTON, O. W., E. P. ORTLEB, L. M. HATHAWay, R. E. BALLINGER, and P. Licut 1971. Reproductive cycles of three species of anoline lizards from the Isthmus of Panama. Ecology, 52:201-215. Smitu, R. E. 1968. Studies on reproduction in Costa Rican Ameiva festiva and Ameiva quadrilineata (Sauria: Teiidae). Copeia, 1968:236-239. TINKLE, D. W. 1967. The life and demography of the side-blotched lizard, Uta stans- buriana. Misc. Publ. Mus. Zool. Univ. Michigan, 132:1-182. TINKLE, D. W., H. M., Wixpur, and S. G. TILLey 1970. Evolutionary strategies in lizard reproduction. Evolution, 24(1): bast. TuRNER, F. B., J. R. LANNOM, Jr., P. A. MEpIcA, and G. A. HoDDENBACH 1969. Density and composition of fenced populations of leopard lizards (Crotaphytus wislizenii) in southern Nevada. Herpetologica, 25 (4) :247-256. Wityort, D. C. 1963. Reproduction in the tropical Australian skink, Leiolopisma rhom- boidialis. Amer. Midl. Nat., 70(2):442-461. Wituort, D. C., and E. O. REerrer 1965. Sexual cycle of the lizard, Leiolopisma fuscum, a tropical Aus- tralian skink. Jour. Morph., 116(3) :379-387. i ‘oe UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly pub- lished in the above series are now published as Occasional Papers, Museum of Natural History. The Miscellaneous Publica- tions, Museum of Natural History, began with number 1 in 1946. Longer research papers are published in that series. Monographs of the Museum of Natural History were initiated in 1970. All manuscripts are subject to critical review by intra- and extra- mural specialists, final acceptance is at the discretion of the publications committee. Institutional libraries interested in exchanging publications may obtain the Occasional Papers and Miscellaneous Publica- tions by addressing the Exchange Librarian, University of Kan- sas Library, Lawrence, Kansas 66045. Individuals may pur- chase separate numbers of all series. Prices may be obtained upon request addressed to Publications Secretary, Museum of Natural History, University of Kansas, Lawrence, Kansas 66045. Editor: Linpa TRUEB Managing Editor: W1LL1AM E. DuELLMAN PRINTED BY UNIVERSITY OF KANSAS PRINTING SERVICE LAWRENCE, KANSAS NOE SF a “CALIFORNIA cor | ACADEMY OF SCIENCE : NOV 16 497° i OCCASIONAL PAPERS || NOV 1 @ 1/3 | | LIBRA RY of the RAD ee MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas NUMBER 19, PAGES 1-47. NOVEMBER 9, 1973 SYSTEMATICS OF THE GENUS RHOGEESSA (CHIROPTERA: VESPERTILIONIDAE ) By RicuArp K. LAVAt' The several species of Neotropical bats of the vespertilionid genera Rhogeessa and Baeodon include some of the smallest of living mammals, although one species approaches in size the com- mon big brown bat, Eptesicus fuscus, of North America. Most of the species occur at low elevations, in habitats ranging from desert scrub to rain forest, and appear to be insectivorous. They range from the lowlands of northern México to southern Brasil, but are poorly represented in collections from s uth America south of Venezuela and Colombia. Allen (1866) described the genus Rhogeessa including two spe- cies, R. parvula and R. tumida. He implied close relationships of Rhogeessa with Nycticejus [—Nycticeius], Nyctinomus [—Ta- darida|, and with the Noctilionidae. Dobson (1878) referred Rho- geessa to a subgenus of the genus Vesperugo [=Vespertilio]. How- ever, Thomas (1892) retained Rhogeessa as a full genus, and stated that it was most closely related to Nycticejus [—Nycticeius|. Sub- sequently, Miller (1897) and others have followed Thomas. Miller (1906) erected a new genus, Baeodon, for Rhogeessa alleni Thomas. Although Simpson (1945) included Baeodon in Rhogeessa, other recent authors, including Tate (1942), Hall and Kelson (1959), and Koopman and Cockrum (1967) have retained them as separate genera. The eight nominal species of Rhogeessa were described in the following sequence: R. parvula (Tres Marias Islands, Nayarit) and 1 Adjunct Professor, Department of Systematics and Ecology, and Research Associate, Museum of Natural History, University of Kansas. (Present address: Organization for Tropical Studies, Universidad de Costa Rica, Ciudad Uni- versitaria, Costa Rica, C.A. ) 2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY R. tumida (Veracruz) by H. Allen in 1866; R. alleni (Jalisco) by Thomas in 1892; R. minutilla (Margarita Island) and R. gracilis (Puebla) by Miller in 1897; R. io (Venezuela) and R. velilla (Ecua- dor) by Thomas in 1903, and R. bombyx (Colombia) by Thomas in 1913. Hall (1952) reduced R. tumida to a subspecies of R. parvula, an arrangement followed by Hall and Kelson (1959) and several authors prior to 1959. No new taxa were described in the genus Rhogeessa until Goodwin’s (1958) revision, in which he named three subspecies, R. tumida major (Oaxaca), R. tumida riparia (Venezuela), and R. parvula aeneus (Yucatan). At the same time he included R. bombyx as a subspecies of R. tumida, and reduced R. minutilla, R. io, and R. velilla to subspecies of R. parvula. The distributional ranges Goodwin gave for the seven subspecies he recognized suggest a montage of isolated subspecies, some sepa- rated by gaps of over 1500 miles. On the basis of my preliminary examination of specimens in the Museum of Natural History at the University of Kansas, I concluded that his taxonomic arrangement failed to elucidate the actual intra- and interspecific relationships within the genus. Comments made by Alvarez and Avia (1965) and by Jones, et al. (1971) also suggest the need for a revision of the genus Rhogeessa. Therefore, with the 650 specimens now avail- able, I have undertaken a new revision, employing multivariate statistical analyses. MATERIALS AND METHODS A total of 653 specimens of Rhogeessa was examined. Included were 8 alleni (2 bacula), 350 R. tumida (43 bacula), 160 R. parvula (28 bacula), 112 R. minutilla (11 bacula), 9 R. gracilis (4 bacula) and 14 (1 baculum) of an unnamed species. Although a substantial number of juvenal specimens was encountered, no more than one per locality was included in the total number of specimens exam- ined. Because many important specimens are preserved in alcohol, skulls from these specimens were removed and cleaned. I examined the following holotypes: R. gracilis; R. tumida; R. tumida major; R. tumida riparia; R. parvula; R. parvula aeneus; R. minutilla. Holotypes of R. alleni, R. bombyx, R. io, and R. velilla were examined for me by J. E. Hill of the British Museum of Natural History. C. O. Handley of the National Museum of Natural History generously made available to me his notes on the holotypes of R. io, as compared with various specimens in the NMNH. Topo- types of R. velilla and R. parvula were examined by me. I recorded all available label and field note information, plus three external measurements, length of fur, nine cranial measure- ments, three bacular measurements, color of dorsum and of venter, presence or absence of sagittal crest, age (adults: phalangeal SYSTEMATICS OF THE GENUS RHOGEESSA 3 epiphyses ossified; juveniles: epiphyses not ossified) and toothwear (1—none; 2—present but scarcely noticeable; 3—easily noticeable on canines and molars, but little cusp detail obscured; 4—heavy, much cusp detail obscured). Measurements were taken following in general the methods outlined by Handley (1959), with excep- tions or additions as noted below: forearm (FA); third metacarpal (3MC); tibia (from center of knee joint to point of attachment of calcar to ankle); fur (greatest length in center of dorsum); greatest length of skull (GLS—including incisors); depth of braincase (DB); postorbital width (POW); mastoid width (MW); width across second upper molars (M2-M2); width of second upper molar (M2—as measured with ocular micrometer); maxillary toothrow (MAX); mandibular toothrow (MAND); measurements of bacular length, depth, and width follow LaVal (1973, Fig. 1). A set of nine specimens (Table 1) was chosen as color standards and matched to named colors from Ridgway (1912). Unfortunately certain specimens, notably those from northwestern Venezuela, fell outside the limits of variation delimited by the color standards; these were compared directly with Ridgway (1912). TaBLE 1. Colors from Ridgway (1912) applied to specimens used as color standards (tip color only). Color Specimen no. no. Dorsum Venter TCWC 19756 1 Fuscous-Black Buffy Brown KU 102619 2 Hair Brown Cartridge Buff KU 29886 3 Tawny-Olive Light Ochraceous-Buff KU 107494 4 Buffy Brown Cartridge Buff KU 105565 5 Buckthorn Brown Light Ochraceous-Buff KU 90754 6 Pinkish Cinnamon Cinnamon Buff UNM 27545 7 Warm Buff Light Ochraceous-Buff KU 97050 8 ca. Light Ochraceous-Buff Pinkish Buff KU Gly 9 Dresden Brown Light Ochraceous-Buff Karyotypes were not examined by me, but I have seen partial results of a study of karyotypic variation in the genus now being conducted by R. J. Baker (pers. com.) and his students. Baker and Patton (1967) documented the differences in karyotypes between R. tumida from Chiapas and R. parvula from Sonora and Nayarit. Specimens identified by Baker and Patton as R. gracilis are not of that species (Baker, pers. com.) and are almost certainly R. parvula, collected at the opposite end of the long cline beginning in Sonora. Scanning electron micrographs were made of hairs from the mid-dorsal region of specimens of each species. Because the appear- ance of Rhogeessa hairs varies substantially depending on the angle of view and the portion of the hair photographed, all photographs were made of the mid-section of each hair; the hair was oriented 4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY so that the side of the scales bearing the V-shaped depression in the rim would face the film. Polaroid negatives (4 & 5 in) were exposed for 70 seconds on a ETEC Autoscan scanning electron microscope, made available to me by the Department of Ento- mology, Kansas State University, Manhattan. Penes were removed from 89 of the males examined, and cleared and stained to reveal the bacula, using Anderson’s (1960) method. Bacula were measured with an ocular micrometer. All data recorded were punched on IBM cards, and statistical analyses performed on the GE 635 computer at the University of Kansas Computation Center. Although t-tests performed between the sexes indicated a significant level of sexual dimorphism for some measurements at some localities, the small sample sizes prevalent in this genus made separation of the sexes impractical. Samples usually contained a fairly well-balanced proportion of the two sexes, which would tend to offset any possible bias due to sexual dimorphism. Gabriel’s Sum of Squares Simultaneous Testing Procedure (SS- STP) was performed, first with all species in the same analysis, and then each species was analyzed separately. An SS-STP program available at the University of Kansas Museum of Natural History (as modified by Gary Powers) gives 13 standard statistics, calcu- lates and lists non-significant subsets, and prints these in graphic form, with means in descending sequence (Figs. 10, 11, 13). The SS-STP procedure, as related to systematics, is discussed by Smith (1972), who lists some of the more important literature references. The University of Kansas Numerical Taxonomy Program (NT- SYS) developed by E. J. Rohlf, J. Kishpaugh, and R. Bartcher was utilized to give the following statistical information: 1) standard- ized data correlations between localities and a two-dimensional phenogram based on these; 2) cophenetic correlation, in which cophenetic values are plotted on scatter diagrams against the cor- relation coefficients used in the phenogram; 3) standardized data distances between localities, and a two-dimensional phenogram based on these; 4) a second cophenetic correlation, plotting co- phenetic values against distance coefficients; 5) a principal com- ponent analysis in which the first five principal components are calculated; 6) a Varimax Factor Rotation by the Kaiser Method; and 7) the variance-covariance matrices projected onto the principal components (both rotated and non-rotated), and two-dimensional scattergrams plotting each of the principal components against each of the others. A stepwise discriminant analysis was also carried out, using the University of California BMDO07M program; this program identi- fies each individual specimen based on the criteria derived from the analysis. It computes canonical correlations and coefficients SYSTEMATICS OF THE GENUS RHOGEESSA 5 for canonical variables, and plots the first two canonical variables on a two-dimensional scattergram. All of the multivariate analyses were computed using the same nine characters, all of which are skin and skull measurements made by myself. These were chosen for several reasons, one of which dealt with problems resulting from missing data. Discriminant function analysis indicated that all contributed significantly to variation observed among the samples analyzed. Other characters examined either did not contribute to variation among samples, or contributed so strongly to singling out one or more species or samples that I felt they would tend to obscure the close relation- ships which are typical of species within this genus. Further, I wanted to see if a multivariate analysis based on a few measure- ments made of morphological characters, which I would expect to be highly correlated, would support suspected relationships based on other kinds of characters (such as distribution of hair on uro- patagium, size of i3, presence of cingual cusps, bacula, hair, etc. ). Blackith and Reyment (1971) stated that “Where the organisms are fairly closely related, and all the measurements are quantita- tive, .... Satisfactory analyses have been made with as few as three to six characters, although ten might be regarded as more Optimal... .” In the analyses performed by the NT-SYS program, only sample means were utilized. To help assure that means used would be representative of the geographic area which they represented, several small samples often were pooled (for example, several samples of one or two specimens each from various localities on the Caribbean slope of Honduras were combined). In other cases, large samples from a single locality were utilized. The areas from which each of the samples was taken, along with the abbreviations used in this paper, are as follow: Tamaulipas, coast (Tamp 1); Tamaulipas, interior (Tamp 2); San Louis Potosi (SLP); Veracruz, northern (VC-N); Veracruz, central (VC-C); Veracruz, southern (VC-S); Campeche (Camp); Yucatan and northern Quintana Roo (Yuc); Chiapas, Pacific Coast (Chiapas); Oaxaca, Pacific slope (Oax); Guerrero (Guer); Michoacan (Mich); Colima (Colima); Jalisco (Jal); Nayarit (Nay); Sinaloa, southern (Sin-S); Sinaloa, central (Sin-C); Sonora (Son); Guatemala, Pacific Coast (Guat); El] Salvador (El Salv); Honduras, Caribbean slope (Hond); Nica- ragua, Caribbean slope (Nic); Costa Rica, Interior (CR); Panama, Pacific slope (Pan); Colombia, north (Colomb); Venezuela, north- west (Venez-NW, 1-5); Venezuela, coast (Venez-C, 1-2); Vene- zuela, interior (Venez-int); Venezuela, northeast (Venez-NE); Trinidad (Trin); Guyana (Guyana). All specimens of R. gracilis, R. alleni, and the unnamed species are pooled into single samples. Fewer samples, each representing more individual samples com- bined into larger pooled samples, were used in the stepwise dis- 6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY criminant analysis, because the kind of output generated by that program would be unintelligible if a very large number of indi- vidual samples were chosen. These samples, along with their ab- breviations, are as follows: Southwestern México—Oaxaca and Guerrero (SWMEX); Michoacan (MICH); West-central Méexico— Jalisco, Colima, Nayarit, southern Sinaloa (WCMEX); Northwest- ern México—northern Sinaloa and Sonora (NWMEX); Eastern Meéxico—Tamaulipas, San Luis Potosi, and Veracruz (EMEX); Northern Yucatan Peninsula (YUC); Central America—Tabasco through Panama, excluding Caribbean slope from Nicaragua to Panama (CENTAM); Caribbean slope from Nicaragua to Panama (NICPAN ); Northern Colombia (COLOMB); Northern Venezuela (VENEZ); Arid northwestern Venezuela and extreme northeastern Colombia (DRYVEN); Guyana (GUYANA); Trinidad (TRIN); all R. gracilis (GRACIL); all R. alleni (ALLENTI); all specimens of the unnamed species (MIRA). Although sexual dimorphism in size was detected within some samples, t-tests did not show it to be consistently significant. On the average, females are as much as 4% larger than males. However, in R. minutilla from Venezuela, males are slightly larger on the average. ACKNOWLEDGMENTS I wish to thank the following persons and institutions for per- mission to examine specimens in their care. Abbreviations as used in lists of specimens examined are given in parentheses. Listed alphabetically. Ticul Alvarez, Escuela Nacional de Ciencias Bio- logicas, Mexico City (ENCB); Robert J. Baker, Texas Tech Uni- versity (TTU); Alberto Cadena, Universidad de los Andes, Bogata (UAB); E. Lendell Cockrum, University of Arizona (UA); James S. Findley, Museum of Southwestern Biology, University of New Mexico (UNM); Charles O. Handley, Jr., National Museum of Natural History (NMNH); Robert S. Hoffmann, Museum of Natural History, University of Kansas (KU); Emmet T. Hooper, Museum of Zoology, University of Michigan (UM); Karl F. Koopman, Ameri- can Museum of Natural History (AMNH); Barbara Lawrence, Museum of Comparative Zoology, Harvard University (MCZ); George H. Lowery, Jr., Museum of Zoology, Louisiana State Univer- sity (LSU); Helen Matuskowitz, Academy of Natural Sciences of Philadelphia (ANSP); Donald R. Patten, Los Angeles County Mu- seum of Natural History (LACM); James L. Patton, Museum of Vertebrate Zoology, University of California, Berkeley (MVZ); Randolph L. Peterson, Royal Ontario Museum (ROM); José Ramirez-P., Instituto de Biologia, Universidad Nacional Aut6noma de México (UNAM); David J. Schmidly, Texas Cooperative Wild- life Collection, Texas A&M University (TCWC); Luis de la Torre, Field Museum of Natural History (FMNH). SYSTEMATICS OF THE GENUS RHOGEESSA fl J. E. Hill of the British Museum of Natural History (BMNH) and C. O. Handley of the National Museum of Natural History are due a debt of gratitude for making available to me notes and meas- urements of those holotypes on deposit in the British Museum. I thank Robert J. Baker, Brent L. Davis, and V. Rick McDaniel for showing me unpublished results of their study of karyotypic varia- tion in Rhogeessa. Robert S. Hoffmann kindly provided editorial assistance. My study was financed, in part, by Penrose Fund Grant 6320 from the American Philosophical Society, by Biomedical Sci- ences Support Grant RR-07037, and by computing funds allocated through the Division of Biological Sciences, College of Liberal Arts and Sciences, University of Kansas. ANALYSIS OF CHARACTERS Goodwin (1958), in his revision of the genus Rhogeessa, stated that the genus (exclusive of Baeodon) could be separated into three species: “a big eared species with thin flying membranes [gracilis], a large species with short ears and thick membranes [tumida], and a small species [parvula].” He further stated that “The propor- tionate length of the forearm and color of the pelage cannot be relied on as determining factors except in a very general way. Cranial measurements can also be confusing unless accompanied by comparative material.” As might be anticipated from these intro- ductory remarks, Goodwin did not state clearly how to distinguish among parvula, tumida, and the various subspecies into which he divided them. As suggested by Goodwin, skin and skull measurements are in- adequate for distinguishing among species, except in the case of R. alleni, which is larger in most cranial variates than any other species examined, and the new species, R. mira, which is smaller than any specimens of R. parvula from Michoacan or adjoining states. However, in the multivariate analyses, notably the principal component analysis, the canonical variate analysis, and the dis- criminant function analysis, the measurements taken were adequate to discriminate many of the samples from each other (see table 4, and Figs. 4, 7, 8, 12). The discriminant function analysis indi- cated that greatest length of skull and forearm were the best and second best discriminators, respectively. Thus, I chose these two variables to demonstrate geographic variation by means of uni- variate analyses. Color of fur.—Variation in relative positions of light and dark bands seems to separate alleni and gracilis from all other species. Individual variation in contrast between bases and tips renders pelage coloration of limited usefulness in identification of individual specimens of the other four species, but differences are evident when series are compared. Molting individuals take on a grayish 8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY cast because dark gray hairs grow up throughout the old pelage before it is shed. Thus, newly molted individuals are much darker in color than individuals in old pelage. In the species which have pale bases and darker tips, individual color variation is accounted for primarily by differences in tip color, whereas color differences among samples and species typically result from variation in basal color. Length of fur—Although some individual variation in fur length occurs, this character is sufficiently constant within a population to separate certain species, and to illustrate geographic variation among populations in a single species. Distribution of fur on uropatagium.—This character evidences so little individual variation that it is a fairly reliable means of separating one of the species from the other five species. Degree of toothwear.—Toothwear was negligible in about 90 percent of specimens examined, except in the case of R. minutilla, in which more than 35 percent of the specimens had moderate to heavy toothwear. Lingual cingulum of C1.—In all species except gracilis and mira two (rarely one) cusps are present on the lingual cingulum of the Cl. These cusps are variable in development, from higher and sharper than the example shown in figure 1A, to lower and more rounded. They tend to be obscured in individuals with heavy toothwear. In the species lacking cusps, the cingulum is perfectly smooth and straight, lacking even the slightest swelling (Fig. 1B). Bacula.—With the exception of R. alleni, bacula of the various species are not sharply differentiated from each other. Although Fic. 1. Upper right canine of two species of Rhogeessa, lingual view. A. R. parvula, UA 10319, Jalisco; B. R. mira, UNAM 8594, Michoacan. x25. SYSTEMATICS OF THE GENUS RHOGEESSA 9 bacular shape shows substantial geographic and individual varia- tion within species, it also seems to differ among species in areas of sympatry and near sympatry (Fig. 2). Single bacula of parvula, tumida, gracilis, and alleni were illustrated and described by Brown, et al. (1971), but these authors did not examine bacular variation. Microstructure of hair—Benedict (1957) examined hairs from SUK < a ae ae A B D E allent gracilis UG mira m= <3 G H | ’ ee ere ee ee | —— LUO C)2>4>1, 1 and 4 rudimentary, 3 much the longest, all margined + OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY with fleshy web; palms smooth, subarticular tubercles absent. Foot with only four toes evident, small but distinct disks at tips, 4>3>5>2, the normal first toe not developed externally; soles smooth except for a feeble, rounded inner metatarsal tubercle. Tongue elongate, oval, extensively free posteriorly; two low, rounded, transverse palatal dermal ridges, the posterior the longer. Color dark brown above and below, flecked with small white spots, more numerous and larger ventrally than dorsally. Snout-vent length 12.3, tibia 6.0, foot 4.9, head width 4.2, tympanum 0.8 mm. Variation.—The largest specimen, an adult female paratopotype, is 13.2 mm in length; the male allotype is 11.2 mm in length with well developed testes, and shows no external sexual dimorphism in structure or color. No vocal sac is apparent nor are there any aper- tures present indicative of an internal sac. The membrane covering the testes is lightly pigmented. The vertebral number, as shown by X-ray photographs, is con- stant throughout the series of 13 specimens, as is also some degree of fusion of the first two vertebrae. In those individuals in which the fusion of vertebrae I and II is complete (Fig. 1), there are only six functionally independent presacral elements. Thus the reduc- tion of the column has been achieved in part by fusion, a frequent occurrence among anurans, and in part by elision, a much rarer event. The partial fusion of vertebrae V and VI of the figured specimen represents an individual variation. The coccygeal proc- esses vary from mere vestiges to the maximum shown in the figure, which is approached by only one other individual. The forward tilt of the processes supplies evidence that the shortening of the column has been achieved by the incorporation of a former verte- bral element into the coccyx, concurrent with the transfer of the sacral function from vertebra IX to vertebra VIII. A parallel situa- tion has been described by Tihen (1960) in the African bufonid genus Mertensophryne, and may be inferred in certain species of the American bufonid genus Rhamphophryne (Trueb, 1971) which exhibit seven presacral vertebrae with no evidence of fusion. The coccygeal processes of Syncope are thus not strictly homologous with those of discoglossids and ascaphids. Their presence is to be interpreted as a derived rather than primitive condition. It has been remarked frequently that departures from the standard foot structure, four fingers and five toes, are rare among anurans. Most of the exceptions seem to occur among diminutive species. In the African bufonid genus Didynamipus, with a maxi- mum recorded length of 18 mm, only three toes are apparent ex- MICROHYLID FROG FROM ECUADOR 5 ternally. Still smaller is the Neotropical brachycephalid Psyllo- phryne didactyla ( 2 10.2 mm, ¢ 8.6 mm) recently described by Izecksohn (1971), in which only two fingers and three toes are obvi- ous. The phalangeal formula of these genera is not of record. In Syncope the phalangeal formula reveals that suppression of the inner toes has not been complete, nor has it in Geobatrachus, a genus of disputed familial relationship, which also lacks an ex- ternally evident inner toe and shares with Syncope the pedal for- mula: 1, 2, 3, 4, 3. Dwarfing seems to have been accompanied by paedomorphosis affecting the foot structure independently in these four unrelated lines. Although patently specialized in most respects, Syncope retains two primitive character states, distinct tympanum and _ posterior prevomer. Derived states include loss of a presacral vertebra, re- duction of clavicle, absence of palatine, quadratojugal, externally evident inner toe, and diminutive size. Examination of Table 1, in which the principal character states of the American microhyline genera are presented, shows clearly that Syncope could not have evolved from any existing genus. Presumably it has arisen from a TABLE 1. Occurrence of Character States among American Genera of Microhylid Frogs B not to scapula complete, incomplete Included taxa Presacral vertebrae Tympanum Clavicle A to scapula Posterior prevomer Palatine Maxillary arch Premaxillary notch Coccygeal process Arcovomer* Chiasmocleis 1 Ctenophryne Dasypops Dermatonotus Elachistocleis Gastrophryne Glossostoma Hamptophryne Hypopachus 1 Hyophryne Mvyersiella** Otophryne Relictovomer Synapturanus Stereocyclops Syncope lel a5 be RR eR DR eS Re DS OUD BR ~1H CH CH HTM OM WH MH CH HH OH HH OT OT OT OT OF OL OT UT UT OT OT OT OT U1 OT OT} No. of toes | | Drow | we | se sear | | | ee (Gl (Ql @) No el@i@i@uer(@al(ey(@) a slate cleats ale atctale ale ated lt te et oiet | lara >| we >| ww | ++ 141 | + | * Unique in its T-shaped terminal phalanges ** Presacral vertebra VIII procoelous. 6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY relatively primitive stock prior to the loss of the tympanum and posterior prevomer. This same stock may have given rise sub- sequently to Chiasmocleis, Elachistocleis, Hamptophryne, and other more advanced genera. In Parker’s (1934) monograph, which relies heavily on condi- tion of the pectoral girdle, the new species keys out to Chiasmocleis, and that structure indeed is so similar to the girdle of Chiasmocleis albopunctata (Parker, 1934:117) as to obviate the need of an addi- tional figure. In Carvalho’s (1954) key to the American genera, one is balked at the first couplet because the combination of a dif- ferentiated tympanum and a reduced clavicle was unknown to that author. Etymology—The generic name, a word of Greek origin, is common to English and most European languages (with only minor variations in orthography) as a figure of speech denoting a short- ening by elision. In the present context it alludes to the loss of one presacral element from the vertebral column. The specific name acknowledges the notable contributions of Antenor Leitao de Car- valho to our knowledge of the morphology and biology of the American microhylids. Remarks——The specimens in the Kansas collection, 124001- 124011, were taken at Puerto Libre on the north bank of the Rio Aguarico, just below the mouth of the Rio Cofanes. All were col- lected at night, from litter on the forest floor or from leaves of low herbs and ferns. The two frogs in the Michigan collection, UMMZ 131699, were flushed from a large patch of low terrestrial brome- liads in humid forest about 9 km E of Puyo. Several small groups of eggs in early stages of development were found in these brome- liads. An effort to rear some of these eggs was unsuccessful, and no evidence exists that the eggs were indeed those of Syncope other than the treacherous evidence of propinquity of eggs and adults. In two mature, unspawned females the complement of well-yoked follicles is 3 and 4 respectively. One ovulated egg, free in the coelom, is about 1.2 mm in diameter and is heavily pig- mented. Acknowledgments.—Thanks are due Alice G. C. Grandison, British Museum (Natural History) and Richard Zweifel, American Museum of Natural History for the loan of three species not other- wise available to me, and to Martha Lackey for her painstaking care with the figures. Resumen.—Syncope antenori, se llama el nuevo genero y especie de ranas microhylid del este Ecuatoriano. Syncope se diferencia de MICROHYLID FROG FROM ECUADOR 7 otros microhylids americano porque tiene siete vertebras pre- sacrales, el coxis articulado con el sacro por dos condilos angostos y separados, un timpano pequeno claramente visible, y tiene solo cuatro dedos en las patas posteriores. Material Examined.—Arcovomer passarellii: KU 93232-6, 92237 (cl. & st.), Brasil. Chiasmocleis albopunctata: UMMZ 63972, 104324, Bolivia; C. bassleri: UMMZ 68197, Bolivia; C. hudsoni: BMNH 1947.2.11.93, Guyana; C. panamansis: UMMZ 126775, Panama; C. ventrimaculata: KU 126668, Ecuador. Ctenophryne geayi: AMNH 44787, Brasil; 42887, Peri. Dasypops schirchi: UMMZ 115660, Brasil. Dermatonotus muelleri: UMMZ 105318, 105321, Brasil. Elachistocleis bicolor: UMMZ 66534, Bolivia. Gastrophryne olivacea: UMMZ S-2643 (skeleton), Texas; G. usta: UMMZ S-1068 (skele- ton), Guatemala. Glossostoma aterrimum: UMMZ 123591, Costa Rica; G. aequatoriale: AMNH 17553, 17562, Ecuador. Hamptophryne _boliviana: UMMZ 63966, Bolivia. Hypopachus aquae: UMMZ S-1032 (skeleton), Guate- mala; H. oxyrrhinus: UMMZ 128077 (skeleton), 128550 (cl. & st.), Nayarit, México; H. cuneus: UMMZ S-2314 (skeleton), Tamaulipas, México; H. inguinalis: UMMZ S-2927 (skeleton), Guatemala. Hyophryne histrio: none examined; data from Carvalho (1954). Myersiella subnigra: KU 93264 (cl. & st.), Brasil. Otophryne robusta: UMMZ 85137-40, Venezuela. Relictovomer pearsei: UMMZ 45569, 45573, Colombia. Synapturanus microps: UMMZ 85143, Venezuela. Stereocyclops incrassatus: UMMZ 115659, Brasil; cranial data from Carvalho (1948). Syncope antenori: KU, UMMZ, type material, listed above, Ecuador. LITERATURE CITED Carva.uHo, A. L. DE 1948. Sobre a validez de Stereocyclops incrassatus Cope 1871 e Hypo- pachus miilleri (Boettger) 1885. Bol. Mus. Nacional, Rio de Janeiro, Zoologia, 84:1-13, 6 figs. 1954. A preliminary synopsis of the genera of American microhylid frogs. Occ. Papers Mus. Zool. Univ. Michigan, 555:1-19, 8 figs. IZECKSOHN, E. 1971. Novo genero e nova especie de Brachycephalidae do estado do Rio de Janeiro, Brasil. Bol. Mus. Nacional, Rio de Janeiro, Zoologia, 280:1-12, 16 figs. Parker, H. W. 1934. A monograph of the frogs of the family Microhylidae. London: British Museum. xiii + 208 pp., 67 figs. TIHEN, J. A. 1960. Two new genera of African bufonids, with remarks on the phy- logeny of related genera. Copeia, 1960(3):225-233. TRUEB, L. 1971. Phylogenetic relationships of certain Neotropical toads with the description of a new genus. Los Angeles County Museum Con- tributions in Science, 216:1-40, 11 figs. UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly pub- lished in the above series are now published as Occasional Papers, Museum of Natural History. The Miscellaneous Publica- tions, Museum of Natural History, began with number 1 in 1946. Longer research papers are published in that series. Monographs of the Museum of Natural History were initiated in 1970. All manuscripts are subject to critical review by intra- and extra- mural specialists, final acceptance is at the discretion of the publications committee. Institutional libraries interested in exchanging publications may obtain the Occasional Papers and Miscellaneous Publica- tions by addressing the Exchange Librarian, University of Kan- sas Library, Lawrence, Kansas 66045. Individuals may pur- chase separate numbers of all series. Prices may be obtained upon request addressed to Publications Secretary, Museum of Natural History, University of Kansas, Lawrence, Kansas 66045. Editor: LinpA TRUEB Managing Editor: WiLL1AM E, DUELLMAN PRINTED BY UNIVERSITY OF KANSAS PRINTING SERVICE LAWRENCE, KANSAS fv Oe “ fy» Aid K/33 UL Mo- U2. >/ TT ences mien et = C-uEORNIS ACADEMY OF SvIENCES JAN 1 41974 LIBRARY . of the — MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas OCCASIONAL PAPERS | NUMBER 21, PAGES 1-54 DECEMBER 21, 1973 NEW NORTH AMERICAN ERINACEINE HEDGEHOGS (MAMMALIA: INSECTIVORA ) By THoMAS H. V. Ricu’ AND DONALD L. RASMUSSEN” Modern hedgehogs, members of the subfamily Erinaceinae, first appeared in the fossil record of Asia and Europe approximately 35 million years ago during the Oligocene. Today the subfamily con- tinues to thrive on those two continents as well as Africa, where they are known to have occurred first 20 million years ago near the beginning of the Miocene. Although now extinct in North America, members of the sub- family first appeared on that continent approximately 23 million years ago near the beginning of the Miocene. At that time, three erinaceine species, belonging to as many different genera, made their appearance in the fossil record. Two of the species are closely related to forms known in Oligocene deposits of Asia and Europe and hence, apparently immigrated into North America shortly after their appearance there in the fossil record. No known erinaceids in the pre-Miocene fossil record of either the Eastern or Western Hemisphere are closely related to the third species. Therefore the question of whether this third species is a newly arrived immigrant into North America or a descendant from an Oligocene endemic form remains unresolved. The Erinaceinae persisted in North America until the latter part of the Miocene about 10 million years ago. During the interval of their presence on that continent, only one additional erinaceine * Department of Geology, Columbia University; Department of Vertebrate Paleontology, the American Museum of Natural History. Present address: The Museum, Texas Technological University, Lubbock, Texas 79409. * Department of Geology and Museum of Natural History, The University of Kansas, Lawrence, Kansas 66045. 2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY genus and species appeared after the initial, nearly simultaneous appearance of the three forms that marked the beginning of the episode. Otherwise, the group did not undergo any noticeable change in North America other than the extinction of its members. Previously, the North American history of the Erinaceinae had been documented by the description of fewer than two dozen speci- mens of which less than half were correctly recognized as erina- ceines. In an attempt to further elucidate the history of the Erina- ceinae, this report will describe a similar number of new North American erinaceine specimens and review those previously men- tioned ones which are referred to the same taxa. ACKNOWLEDGMENTS For the loan of specimens we are deeply indebted to Mr. Morris F. Skinner and Dr. Malcolm C. McKenna, Department of Vertebrate Paleontology, and Dr. Sydney Anderson, Department of Mammal- ogy, American Museum of Natural History; Dr. Larry D. Martin, Museum of Natural History, University of Kansas; Dr. Robert W. Wilson, Museum of Geology, South Dakota School of Mines and Technology; Dr. Peter Robinson, University of Colorado Museum; Dr. Robert W. Fields, Department of Geology, University of Mon- tana; the late Dr. Claude W. Hibbard, Museum of Paleontology, University of Michigan; and Dr. Elwyn L. Simons, Peabody Mu- seum of Natural History, Yale University. We also thank Drs. Craig C. Black, Malcolm C. McKenna, and Larry D. Martin for criticizing the manuscript and making several suggestions for improvement. Field work in Montana by Rasmussen was supported by: a) National Science Foundation Grant GP-1553 to Dr. Robert W. Fields, principal investigator; b) Division of Vertebrate Paleontol- ogy, Museum of Natural History, University of Kansas; and c) Watkins Fund, Museum of Natural History, University of Kansas. Ranchers Dean Tavenner and Marvin Radtke graciously gave permission to collect specimens on their properties. We thank Mrs. Mary Lee Vickers and Mrs. Geraldine J. Ras- mussen who typed this manuscript. METHODS AND ABBREVIATIONS The primary geochronologic units employed in this report are the North American and European Land-Mammal Ages. Also given are the approximate epoch and radiometric age equivalents based on data summarized in Berggren (1971) and Van Couvering (1972). Figure 1 illustrates the method employed for making measure- ments of teeth discussed in this report, and figure 11 illustrates how angular measurements of lower jaws were taken. Museum abbreviations used with catalogue numbers are as follows: NORTH AMERICAN ERINACEINE HEDGEHOGS 3 AMNH Department of Vertebrate Paleontology, American Museum of Natural History AMNH (M) Department of Mammalogy, American Museum of Natural History F:AM Frick American Mammals, Department of Vertebrate Pa- leontology, American Museum of Natural History KU Museum of Natural History, University of Kansas MPUM Department of Geology, University of Montana SDSM South Dakota School of Mines and Technology UCM University of Colorado Museum UMMP Museum of Paleontology, University of Michigan YPM Yale Peabody Museum HISTORICAL RESUME Established by Koerner in 1940, Parvericius montanus was the first erinaceine species to be recognized and named in North Amer- ica. The new species was based on a single specimen, consisting of a maxillary fragment with M!* and the buccal regions of P**, from the Miocene Deep River Formation of Meagher County, Mon- tana. Koerner regarded the species as an erinaceine and considered it to be more closely related to Erinaceus than to any other member of that subfamily, devoting the generic diagnosis to an enumeration Fic. 1. Occlusal view of erinaceine dentition showing points between which tooth measurements were taken. Abbreviations: a, anterorposterior diameter; ¢t, transverse diameter; tl w, talonid width; tr lI, trigonid length; tr w, trigonid width. For I'-P*, M*, L-P., and Ms, the transverse diameter, t, is defined as the greatest distance from the buccal to the lingual borders of the tooth measured perpendicular to the anteroposterior diameter, a, but not necessarily on a single line. For M’, the anteroposterior diameter, a, is defined as the greatest distance from the anterior to the posterior borders of the tooth measured perpendicular to the transverse diameter, t, but not necessarily on a single line. am OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY of eleven differences he considered significant between the two (Koerner, 1940:841). Although Butler (1948:490) suggested that Parvericius might belong to the tribe Erinaceini (subfamily Erinaceinae), he tenta- tively placed it in the Brachyericini (subfamily Neurogymnurinae ) in his classification of the Erinaceidae (1948:488). Butler (loc. cit.) regarded the M! as more transverse and the M® more reduced in Parvericius montanus than in Amphechinus edwardsi. Examination of Koerner’s illustration of the upper dentition of P. montanus (see also Fig. 12a, this paper) and Hiirzeler’s of A. edwardsi reveals that the M® transverse width is equal to the distance from the buccal edge of the M? opposite the metacone to the buccal side of the hypocone on both specimens (Koerner, 1940:PI. 1, Fig. la; Htirze- ler, 1944: Fig. 4). Direct examination of the type of P. montanus (YPM 13956) confirms this relationship in that specimen. In the same way, the anteroposterior dimensions of the M* on the two specimens appear to be proportionally the same, and morphologi- cally the two teeth are quite similar. Therefore, it cannot be said generally that P. montanus has a more reduced M? than is found on A. edwardsi. In 1960, R. W. Wilson identified an isolated lower molar from the Hemingfordian lower Pawnee Creek Formation of northeastern Colorado as an Mz of Metechinus marslandensis. This specimen now appears to be an M, of either Parvericius or Stenoechinus, new genus. Friant (1961) placed Parvericius in her broadly defined Erina- ceinae. Van Valen (1967:273) synonymized Parvericius with Amphech- inus for three reasons. At the time, Val Valen regarded it likely that material referred to Metechinus marslandensis was the other- wise unknown lower dentition of Parvericius montanus. On this basis, he assumed that Parvericius was similar to Amphechinus in possessing an enlarged lower incisor. Subsequently, however, the material referred to M. marslandensis has been shown to be the lower dentition of Brachyerix macrotis, an erinaceid quite unlike P. montanus (Rich and Rich, 1971:12-13). Secondly, Van Valen thought that both Metechinus (a genus he thought to have been derived from Parvericius) and Amphechinus had foreshortened skulls. However, comparison of figures of the skulls of Amphechinus and other erinaceines such as Erinaceus, reveals that the skull of Amphechinus is as elongated, if not more so, than those of other members of the Erinaceinae (cf. Viret, 1938: Fig. 1). Finally, Van Valen regarded the teeth of Amphechinus and Parvericius as quite similar. Although the available teeth of Parvericius do resemble those of Amphechinus, they show equally strong resemblances to other known erinaceines. NORTH AMERICAN ERINACEINE HEDGEHOGS 5 Fic. 2. Stenoechinus tantalus, n. gen. and n. sp. A. Lateral view; B. Occlu- sal view; C. Lingual view of holotype, KU 18001, right mandible, from the late Arikareean, upper Cabbage Patch beds, Tavenner Ranch locality 2 (KU-Mt- 21), NW 4, NW kh, SW 4%, SW &%, sec. 11, T 8 N, R 10 W, Powell Co., Montana (= locality 1, Wood and Konizeski, 1965:462). D. Occlusal view of KU 18406, left mandible, from the medial Arikareean, middle Cabbage Patch beds, Cabbage Patch locality 13 (KU-Mt-46), CSW %, SE %, sec. 1, T 10 N, R 12 W, Granite Co., Montana (= MV6547, Rasmussen, 1969:132). x6. 6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Fic. 3. Stenoechinus tantalus, n. gen. and n. sp., KU 18002, left mandible. A. Lateral view. B. Occlusal view. C. Lingual view. From the late Arikareean, upper Cabbage Patch beds, Tavenner Ranch locality 2 (KU-Mt-21), NW 4%, NW %, SW 4, SW 4%, sec. 11, T 8 N, R 10 W, Powell Co., Montana (= locality 1, Wood and Konizeski, 1965:462). x6. Mandibles of Parvericius montanus described here for the first time support the placement of this species in a genus other than Amphechinus. Although the two genera share the elongate I, that has so impressed previous workers, this new material demonstrates that the M, trigonid of Parvericius is anteroposteriorly compressed rather than being anteroposteriorly expanded as in Amphechinus. Although but a single difference, in a group as highly uniform as the Erinaceinae, we consider it of generic significance. Palaeoerinaceus minimus Bohlin (1942) was described first from Taben-buluk, Kansu, China; recently, additional material from Nareen Bulak, Mongolia, has been assigned tentatively to that species by Sulimski (1970) under the designation Amphechinus (Palaeoerinaceus) cf. minimus. Both localities have yielded faunas regarded as somewhat younger than that from the Hsanda Gol Formation of Mongolia and medial or late Oligocene in age (Mel- lett, 1968:9; Sulimski, 1970:69). Although P. minimus apparently has the enlarged I, characteristic of all members of Amphechinus, the M, trigonid is not anteroposteriorly expanded; therefore, this species is transferred to Parvericius with which it agrees in both characters. Furthermore, the few mandibles of Parvericius minimus NORTH AMERICAN ERINACEINE HEDGEHOGS 7 can be distinguished from the few of Parvericius montanus only by the greater depth of the latter, a difference probably due to onto- genetic stage of development. Therefore, P. minimus is regarded here as a junior synonym of P. montanus. Despite the removal of Parvericius montanus from Amphech- inus, the latter genus still is known in North America because Palaeoerinaceus horncloudi J. R. Macdonald (1970) from the early Arikareean Monroe Creek Formation of South Dakota may be as- signed to Amphechinus following Butler’s action synonymizing the two genera (Butler, 1948:473). J. R. Macdonald (1970:21) dis- tinguished A. horncloudi from other species of the genus by the talonid having a greater width than the trigonid on P, and the shorter trigonid on M,. However, the latter character does not appear valid when figures of other species of Amphechinus are com- pared to the type specimen of A. horncloudi. (For example, com- pare figures of the following species of Amphechinus: 1) A. horn- cloudi, Macdonald, 1970: Fig. 5; this paper: Fig. 9; 2) A. edwardsi, Hiirzeler, 1944: Fig. 13; 3) A. intermedius, Viret, 1938: Fig. 9; and 4) A. arvernensis, Viret, 1929: Pl. 28, Fig. lb.) Relative to either the width of the trigonid or the length of the tooth, the anteroposterior length of the M, trigonid is as great or greater on A. horncloudi as on any other species in the genus. From the same lithic unit and general area that yielded the type specimen of Amphechinus horncloudi, L. J. Macdonald (1972) recently has reported the presence of additional specimens of that species (under the name Metechinus marslandensis) together with specimens of Parvericius montanus. SYSTEMATIC ACCOUNTS Class MAMMALIA Linnaeus, 1758 Order INsectivora Illiger, 1811 Superfamily ERINACEOIDEA Fischer von Waldheim, 1817 Family Ertnacemae Fischer von Waldheim, 1817 Subfamily ERINACEINAE Fischer von Waldheim, 1817 Stenoechinus new genus Type Species.—Stenoechinus tantalus new species. Known Distribution—Late Arikareean (22.5-21 my, early Mio- cene ), North America. Diagnosis —Distinguished from Aethechinus, Amphechinus, Atelerix, Dimylechinus, Erinaceus, Gymnurechinus, Hemiechinus, Mioechinus, Paraechinus, Parvericius, and Postpalerinaceus by the greater width of the M! relative to its length; from Amphechinus, Dimylechinus, Mioechinus, Palaeoscaptor, and Postpalerinaceus by an anteroposteriorly compressed M, trigonid; from Aethechinus, Amphechinus, Atelerix, Erinaceus, Hemiechinus, Mioechinus, Para- 8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Fic. 4. Stenoechinus tantalus, n. gen. and n. sp., occlusal view. A. KU 18359, right M*. B. KU 18098, right M*. Same locality as in figure 3. x12. echinus, Parvericius, and Postpalerinaceus by the presence of a prominent postcingulum on M3; from Palaeoscaptor by a talonid on Mz; reduced to a prominent postcingulum; and from Dimylech- inus by presence of Ms. Etymology.—Stenos, Greek, meaning narrow, referring to the proportions of the M!, M?, M, trigonid, and M3; talonid; echinos, Greek, meaning hedgehog. Stenoechinus tantalus new species Figures 2-4 Holotype.—KU 18001, right manible fragment with M,-s, lacking ascending ramus and horizontal ramus anterior to M, except for the medial wall of the horizontal ramus that has traces of four alveoli immediately anterior to M;. Type Locality and Stratigraphic Position—Upper Cabbage Patch beds, Tavenner Ranch locality 2 (KU-Mt-21), NW %, NW 4, SW, SW 4, sec. 11, T 8 N, R 10 W, Powell Co., Montana (= Lo- cality no. 1, Wood and Konizeski, 1965:462 ). Diagnosis —Only known species of genus. Etymology.—Tantalos, Greek, mythological character symbolic of eternal torment. Referred Material—KU 18002, left mandible fragment with P,, M,, and that part of the mandible immediately below these two teeth. KU 18003, right mandible fragment with Mo, alveolus for Ms, and lacking horizontal ramus anterior to Ms, ascending ramus, and angle. KU 18004, isolated left M;. KU 18098, isolated right M?, heavily worn. KU 18342, edentulous left mandible fragment witn alveoli for M23, and lacking horizontal ramus anterior to M2 alveoli, ascending ramus, and angle. KU 18354, isolated right My). KU 18356, left mandible fragment with M, and alveoli for M>-;, lacking horizontal ramus anterior to M, and all of mandible behind M; alveolus. KU 18359, isolated right M!. KU 18404, isolated right M, trigonid. All the material referred to in this paragraph was collected at the same locality as the type specimen. NORTH AMERICAN ERINACEINE HEDGEHOGS 9 KU 18406, left mandible fragment with M,-3, lacking ascending ramus and horizontal ramus anterior to M,;. Found in the middle Cabbage Patch beds, Cabbage Patch locality 13 (KU-Mt-46), CSW 4, SE 4, sec. 1, T 10 N, R 12 W, Granite Co., Montana (= MV6547, Rasmussen, 1969: 132). TasLeE 1. Measurements (mm) of the Upper Dentition of Stenoechinus tantalus and Palaeoscaptor acridens S. tantalus S. tantalus P. acridens KU 18098 KU 18359 AMNH 22080 M' anterorposterior diameter 1 DD) M’, transverse diameter _........_.»___- 22 2.6 M’, anteroposterior diameter —__- L2 pe 1.6 M?, transverse diameter 1.9 aa 22 Lower Dentition.—( Figs. 2 and 3) Only indirect data are avail- able to give an indication of the dental condition anterior to Py. In transverse section, the horizontal ramus of KU 18001 has a medial and a lateral layer of compact bone with a space in between. On the internal side of the medial wall of this specimen, immediately anterior to the M,, are preserved traces of the partitions between the next four anterior alveoli (Fig. 2a). The most posterior of these alveoli were occupied by the roots of Py (Fig. 3a, c). Although the forward wall of the most anterior alveolus preserved is absent, enough of that alveolus is preserved to indicate that it probably had a significantly greater anteroposterior diameter than the one im- mediately behind. A similar size relationship exists between the same alveoli on a specimen of another small erinaceine, Palaeoscap- tor cf. acridens (AMNH 22082). On that mandible, the more pos- terior of these two alveoli was occupied by the rear root of P3 and the more anteroposteriorly elongated anterior alveolus was occupied by the forward root of P3 and the single root of Py (Fig. 5). If there was a bony partition between the forward root of P; and the root of Ps, it has been lost without a trace on both KU 18001 and AMNH 22082 and must have been much thinner than the other alveolar walls, for these two roots are quite close to one another on AMNH 22082. The most anterior alveolar wall preserved on the medial wal of the mandible of KU 18001 extends nearly to its ventral border. In erinaceids such as AmphechinusP rectus (AMNH 22084), which have an enlarged I, root alveolus extending posteriorly to the region immediately in front of the anterior root of Py, the alveolar borders of the C,-P; may be traced on the internal side of the medial wall of the ramus from the dorsal lip of their respective alveoli only halfway to the ventral border of the mandible. At that point, the alveolar walls of C,-P3 are cut off by the alveolar wall of I, because 10 oak 80 is ao — aa Ane om LO ~ pHs pruosiy “jy ae UT = oe ~--- = == Ee Qi narse ce Toyowelp Jol19}sodo19}ue “py oT ao aa ai ons c 3 ag | . GI ~ Wp pruoe, “jy VT ell os a a PR snapdoina snaopuiy OOO MINEUNIAN Bo VP. on oo Goa @ el snapdouna snaovumy COSC CINGEINING ga 8, Ge snapdoina snaovumug FOCI mong: © - eo gemetnag “eto iene sisuasuins snuiyooydmy ut OT “STA (comme Wo 1) oo eee ispupmpa snurysoydwy ub 2 Sta GiGIN com cg ayn he een ne 2 ispupmpa snuryoaydwy Ulf Si CORIO awe ek ee a ee ypnojouLoy snuryoaydwy 90.1n0s BXE aInjze1o}] 10 jequinu usuttoeds (wu) aze[Vg Jo UO}RSUOTY JO 90189q “¢ AIAV], 28 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY specimens available, the presence or nature of posterior palatine foramina in that crest cannot be determined. A slight groove is developed on either side of the midline at the anterior end of the palatine; it extends posteriorly to a small fora- men. Presumably this groove is homologous to the similar one found in galericines, in which the palatine nerve and artery lie. Midway between the aforementioned foramen and the transverse crest is a smaller palatine foramen. The bones of the palate are as thick and solid as those of the living galericines. No vacuities typical of the modern erinaceines are present in the palate. Although the snout is more elongate than in living erinaceines, the molars are more posterior relative to the zygomatic arch. The elongation is reflected in the high length-to-width ratio of the palate (Table 5). On KU 18163 this ratio is 2.41, on species of Amph- echinus edwardsi it is from 2.30 to 2.40, and in a sample of recent erinaceines it ranges from 1.84 to 1.99. The location of the base of the zygomatic arch, mainly opposite the M1}, is indicative of the degree of posterior displacement of the molars; however, a small segment of the base lies opposite the M?. In most modern erina- ceines the base of the arch is equally developed above both molars. Mandible.—( Fig. 9) The mental foramen lies below the anterior root of Py. A wide groove extends anterodorsally from the mental foramen and terminates below the partition between the alveoli for P, and C,;. In the region beneath P2-Ms, the mandible depth is nearly uniform. Moving away from this region in either direction, the depth gradually decreases. Fic. 11. Method of measuring the elevation of the anterior edge of the ascending ramus ( / a), elevation of the condyle ( / b), and inclination of the dorsal border of the midline symphysis ( / c) on the lower jaw. NORTH AMERICAN ERINACEINE HEDGEHOGS 29 Extending forward from beneath Ps, the unfused symphysis occupies the ventral three-quarters of the mandibular surface. The dorsal border of the symphysis is convex dorsally and roughly paral- lel to the ventral border, dipping posteriorly at an angle of 25° (Z cin Fig. 11)! The ascending ramus lies slightly labial to the midline of the mandible. Due to damage sustained by the only mandible avail- able, the anteroposterior length of the ascending ramus, precise position of the condyle, and extent of the angle cannot be deter- mined. The straight anterior border of the ascending ramus is posteriorly inclined at an angle of 76° ( 7a in Fig. 11). This angle is the same as Butler’s (1948:469-470) “elevation of anterior edge of coronoid process.” Contact between the dorsal and anterior edges of the ascending ramus is at a gently rounded corner. A strong lateral flange on the leading edge of the ascending ramus forms the anterior margin of the masseteric fossa. There is a ridge near the base and on the medial side of the ascending ramus. It extends posterordorsally from the anterior edge of the ascending ramus, passes immediately above the mandibular foramen (which is level with the tooth row), and continues a short distance further before being abruptly cut off by the damage described above. Because ridges similar to this are directed at the condyle on complete erinaceine mandibles, it appears that the condyle was quite low, with an angle of elevation (7 b in Fig. 11) of about 15°. This angle is the same as the “elevation of the condyle” as defined by Butler (loc. cit.). On the medial side of the angle can be seen the most anterior part of the inflected ridge which is directly below the mandibular foramen. Comparisons—Each of the three specimens from Montana identified as Amphechinus horncloudi are from different localities and stratigraphic positions in the Cabbage Patch beds. All are of the proper size to be parts of the unknown dentition, jaw, or skull of Amphechinus horncloudi originally described from South Dakota. However, with the exception of KU 18162, there is no reason other than geographic proximity to regard these specimens as conspecific with A. horncloudi rather than the slightly larger Amphechinus edwardsi of Europe or slightly smaller Amphechinus rusingensis of Africa. See Butler (1956: Tables 4 and 7) and Hiirzeler (1944:461) for measurements of the dentitions of these two species. KU 18097, a maxilla fragment with P*-M!, is similar to the species of Amphechinus in the proportions of the upper molars. Molars of modern erinaceines, Gymnurechinus, Mioechinus, and Postpalerinaceus are all more anteroposteriorly elongate. Palaeo- scaptor and Stenoechinus have much smaller molars. KU 18162, a mandible with I,, Ps, and Msg is similar to other species of Amphechinus and unlike other erinaceines except Palaeo- scaptor and Gymnurechinus. The P, is similar to that of the type of 30 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Fic. 12. Parvericius montanus. A. YPM 13956, type, right maxilla fragment, occlusal view, from Arikareean or Hemingfordian, Deep River beds, sec. 3, T 10 N, R 5 E, Meagher Co., Montana. x6. B. UMMP V61033, right P*, occlusal view, from late Barstovian, Egelhoff Quarry, from an unnamed lithic unit beneath the Valentine Formation and overlying the Rosebud Formation, SW corner of NE 4, SW 4, sec. 29, T 33 N, R 23 W, Keya Paha Co., Nebraska. «12. Cs UMP V61026, right M’, occlusal view, same locality as B. «12. Amphechinus horncloudi (SDSM 62113) and unlike any other known species of the genus in that the maximum transverse diam- eter is in the talonid region, not further forward across the trigonid. KU 18163, a partial skull with I'-P?, resembles Amphechinus edwardsi and Amphechinus rusingensis in the presence of an en- NORTH AMERICAN ERINACEINE HEDGEHOGS 3] Fic. 13. Parvericius montanus, UMMP V56569, left mandible. A. Lateral view. B. Occlusal view. C. Lingual view. From the late Barstovian, Egelhoff Quarry, from an unnamed lithic unit be- neath the Valentine Formation and overlying the Rosebud Forma- tion, SW corner of NE 4%, SW 4, sec. 29, T 33N, R 23 W, Keya Paha Co., Nebraska. 4.5. 32 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY larged I’. Among the other genera of Erinaceinae, only the much smaller Palaeoscaptor is known to possess as enlarged an [' (Tro- fimov, 1960:38 and Fig. 1). Parvericius Koerner 1940 Type Species.—Parvericius montanus Koerner 1940. Known Distribution —?32—?31 my (medial? Oligocene) Asia; medial Arikareean—late Barstovian (23—12.5 my, early to medial Miocene ), North America. Diagnosis.—Distinguished from Palaeoscaptor and Stenoechinus by the greater length of the M! relative to its width; from Palaeo- scaptor by absence of a metacone on M?; from all other erinaceines except Palaeoscaptor and Amphechinus by the presence of an en- larged I,; from Stenoechinus by a prominent paraconid taller than the metaconid on Py; from Amphechinus, Dimylechinus, Mioechinus, Palaeoscaptor, and Postpalerinaceus by the anteroposteriorly com- pressed M, trigonid; from Gymnurechinus, Palaeoscaptor, and Stenoechinus by absence of a talonid or postcingulum on M3; and from Dimylechinus by presence of M3 and M?*. Parvericius montanus Koerner 1940 Figures 7d, 12-15 Parvericius montanus Koerner 1940:841. Palaeoerinaceus minimus Bohlin 1942:23. Amphechinus ( Palaeoerinaceus ) cf. minimus Sulimski 1970:64. Holotype.—YPM 19356, right maxilla fragment with P?-M? com- plete except for lingual halves of P**. Type Locality and Stratigraphic Position—Collected from the Deep River beds in sec. 3, T 10 N, R 5 E, Meagher Co., Montana. Diagnosis.—Only known species of genus. Referred Material—MPUM 1551, right mandible with highly worn P,-Ms, tip of I, root, fragment of C;, P3 alveolus, horizontal ramus complete from C, to base of leading edge of the ascending ramus. Found in the middle Cabbage Patch beds, Bert Creek lo- cality 2 (MV6504-2), NW %, NE 4, SW %, SE 4%, sec. 1, T 10 N, R 12 W, Granite Co., Montana (Rasmussen, 1969:131-132). UCM 29507, isolated left M,. Found in the Split Rock Forma- tion, SW %, NW 4, sec. 36, T 29 N, R 90 W, Fremont Co., Wyoming. UCM 29853, isolated right M,. UCM 29955, left mandible with M,-3, posterior half of rear alveolus of P,, horizontal ramus com- plete from rear alveolus of P; to base of leading edge of ascending ramus. Found in the Split Rock Formation, NE 4%, NE 4%, NW 4, sec. 36, T 29 N, R 90 W, Fremont Co., Wyoming. F:AM 76704, complete left mandible, edentulous except for Py. UMMP V56569, left mandible lacking only P. and the angle. UMMP V57331, left mandible fragment lacking the region pos- NORTH AMERICAN ERINACEINE HEDGEHOGS 33 terior to the alveolus for the forward root of Mz with I,, C;, Py and alveoli for Iz, P3;, Mi and anterior alveolus for Mz. UMMP V61029, isolated left M, (this specimen appears to have been etched by acid). UMMP V61031, isolated right Ms. UMMP V61027, isolated right Mo. UMMP V61033, isolated left P*. UMMP V61034, isolated right Mt. UMMP V61026, isolated left M*. Found at Egelhoft Quarry in the SW corner of NE %, SW 4%, sec. 29, T 33 N, R 23 W, Keya Paha Co., Nebraska, in an unnamed lithic unit that uncon- formably overlies the Rosebud Formation and is unconformably overlain by the Valentine Formation. UMMP V61022, isolated left M;. Found at Norden Bridge Quarry in the SE %, SW 4%, sec. 33, T 33 N, R 23 W, Brown Co., Nebraska, in the same lithic unit as Egelhoff Quarry described in the previous paragraph. Upper Dentition—( Fig. 12) P’—Only the buccal half of this tooth is preserved on the single specimen available. The paracone is the largest and tallest cusp preserved on the tooth. The anterior side of the paracone is straight and the posterior side, concave. Wear has blunted the tip of the paracone, but, when unworn, the apical height of this cusp was probably no more than two-thirds the crown length. Directed posteriorly from the metacone, the metacrista abruptly turns in a buccal direction at the posterior border of the tooth, but no metastyle or other cuspules are devel- oped there. On the anterior side of the paracone base there is a narrow precingulum with a weak paracrista which extends ventrally Fic. 14. Parvericius montanus, MPUM 1551, right mandible. A. Lateral view. B. Occlusal view. From the medial Arikareean, middle Cabbage Patch beds, Bert Creek locality 2 (MV6504-2), NW %, NE #, SW #, SE 4, sec. 1, T 10 N, R 12 W, Granite Co., Montana (Rasmussen, 1969:131-132). x6. 34 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Fic. 15. Parvericius montanus, UCM 29955, left mandible. A. Lateral view. B. Occlusal view. From the Hemingfordian, Split Rock Formation, NW 4, sec. 36, T 29 N, R 90 W, Fremont Co., Wyoming. x6. from the cingulum toward, but not reaching, the apex of the cusp. Examination of the broken surface where the lingual half of the tooth broke away suggests that it was markedly shorter anteropos- teriorly than the buccal half; thus, the metastylar spur projected away from the body of the tooth as on the P*. On the buccal side of the tooth along the base of the paracone is a weakly developed cingulum. Two prominent roots are preserved, one above the parastylar region, the other over the metacrista. If a third root were present above the missing lingual region of the tooth, all trace of it has been lost. P',—This tooth is several times larger than the preceding P?. The height of the paracone, tallest cusp on the tooth, is about five- sixths of the tooth length. Two-fifths the height of the paracone, the protocone is lingual and slightly anterior to the cusp, as in living hedgehogs. Equally as large in basal dimensions, the hypocone is TABLE 6. Measurements (mm) of the Upper Dentition of Parvericius montanus YPM 13956 UMMP UMMP UMMP type V61033 V61026 V61034 P*, anteroposterior diameter _ 1.1 oe : P*, anteroposterior diameter _ 2.1 22 7 P*, transverse diameter _____. oe 22. me Be M’, anteroposterior diameter __ 2.1 a 2.0 2.2 M’, transverse diameter _______ 220 oes 2.4 2.6 M?’, anteroposterior diameter _ 1.6 : M?, transverse diameter _____. 1.8 { M®*, anteroposterior diameter __ 0.4 M®, transverse diameter ______ el NORTH AMERICAN ERINACEINE HEDGEHOGS 35 lower than the protocone and posterolingual to it. Although these two lingual cusps are linked together by a low crest, neither cusp is joined to the buccal structure of the tooth by a crest. Between the hypocone and the base of the metastylar spur is a planar surface that slopes posterodorsally. A small parastyle is present on the well-developed precingulum anterior to the base of the paracone. Although no paracrista is present, the prominent metacrista is di- rected posterobuccally away from the paracone and terminates posteriorly without a distinct metastyle. Likewise, the metastylar spur projects posterobuccally away from the body of the tooth so that its lingual half is narrower anteroposteriorly than the buccal half. Continuous with the precingulum is a weaker cingulum along the buccal side of the tooth. The three roots of the tooth lie above the protocone and hypocone, paracone, and metacrista, respectively; all are subequal in length and diameter. M'.—Largest tooth in the skull, the M! is somewhat broader but subequal in length to the P*. The height of the paracone, third tallest and third largest of the four principal cusps, is half the length of the tooth. The paracone is anterior to the metacone and their bases are confluent. Subequal in height and basal dimensions, the metacone and protocone are the tallest and largest of the four principal cusps. The anterior side of the metacone is steep; the posterior side is more gently dipping. The protocone is lingual and slightly anterior to the paracone. The hypocone is the lowest and smallest of the four principal cusps. The hypocone and metacone are as close to one another as the protocone and paracone. Linking the anterobuccal corner of the protocone and the anterolingual corner of the paracone are the well-developed preprotocrista and preprotoconule crista which grade imperceptibly into each other in the absence of a paraconule. In anterior view this loph has a broad, inverted V-shaped profile. The postprotocrista is directed posterobuccally from the protocone. This crista divides at a point lingual to the anterior edge of the metacone; one branch extends posterolingually to the hypocone whereas the other, the postmeta- conule crista, extends buccally to the metacone. No metaconule is developed on this tooth. A short paracrista crosses the precingulum and terminates against the base of the paracone at the anterobuccal corner of the tooth. No parastyle is present. The metacrista extends posterobuccally from the metacone for a distance equal to one-half the tooth length. A cingulum is developed along the entire margin of the tooth except at the posterior tip of the metastylar spur. In outline the tooth is approximately square, with prominent excava- tions on the posterior and lingual side. A well-developed metastylar spur projects posterobuccally away from the body of the tooth, and a much smaller parastylar spur projects anteriorly. Three roots are present—one above the paracone-paracrista region, a second OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 36 0 90 cs =: Oi ea ae a gs ae (PIM pruosyy “IN OT 10 -~ ae De ha ee ae yy sue] pruosny “WV We LT ---- sose os = a a a JayoUWIeIp IOLe}ysodo19}ue “jj ST CT Day CT Cle. a a> a mete qapias pruoyer “WW 9'I a 9'T eT Cuiny gaan Tae ee Ee ie TAPIA pruosiy “WW oT 16 | Gali Gali ile Ook eet. a A Yysue] pruos1y TIA CZ oS S'S VS Gil a ae a JayoUIeIp IoL1e}sodo19jue TPA ST = — ---- Gi0e ear as ee ee es JayoUIeIp ssi9AsuBI} Fg CT = ---- ness Clipe ke tage ee a JoyoWIeIp 1011e}sodo19jyue FF 69S9SAdNWNN S&66¢ WON €S86¢6 WON LOS6¢ WON TSst WOdWN SnuDJUOW SMILAAIDG JO UOIWUEP IOA\OT OY} JO (WUT) syUOWOINsBIIY *Y ATAV], 37 NORTH AMERICAN ERINACEINE HEDGEHOGS OT Sok aa GG ioe a as or Pes a es ae OT _: oi <. OT vel st Ih c ae ae JoyoweIp osioAsuely Fg Vilas: ey ae ees JoyeUIeIp IOL1eysodo10jue Fy GCOISAdWNWN FOLILWV'H LZEOISAdNWNN 6C0ISAdNNN ITSOT9AdNNN TeeLSA dNWN (penul}u0o) *) ATavy, 38 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY above the metacone-metacrista region, and a third above the proto- cone-hypocone region. M?.—Although smaller, the M? is basically similar to the M1}. The protocone and paracone are subequal in height and basal di- mensions, being the tallest and largest of the four principal cusps. The lingual region of the M? is the same as the M!; however, the buccal region, particularly the posterobuccal area, is different. The metacone has shifted position so that it is posterolingual to the paracone, not posterior. The metastylar spur projects posteriorly, not posterobuccally, and is reduced. The buccal border of the tooth is not subparallel to the midline of the skull, but is directed posteromedially. Similarly, the parastylar spur has shifted so that it projects anterobuccally rather than anteriorly. Narrow cingula are developed on all sides except the lingual. Only the two buccal roots of the tooth can be observed; relative to structures on the crown, their positions are the same as on the M!. M*.—This tooth is basically a mediolaterally elongated blade with a protocone at the lingual end and an equally large and tall paracone at the buccal end; the two cusps are joined together by a short, straight crest. A weak precingulum is developed along the anterior margin of this tooth. Two roots are developed on this tooth—one above the protocone and a second over the paracone. Lower Dentition.—(Figs. 13-15) I,—In morphology and size, this tooth is quite distinct from the incisor behind (I.). It is elon- gated, extends forward, and curves slightly upward from the an- terior end of the mandible for a distance about equal to the I,-P3 length, or 3.7 mm on the least worn specimen (UMMP V56569). Posteriorly, the root extends beneath and medial to the roots of I.-P. to terminate just in front of the anterior root of Py. In cross- section at the alveolar border, the tooth is a slightly distorted, dorsoventrally elongated oval with the blunt end dorsal and the medial side straight rather than curved. The tooth is mediolaterally compressed so that its dorsoventral diameter is about half again as great as its mediolateral diameter. At its alveolar border, the tooth is nearly as wide and deep as the mandible itself. Further posterior, beneath the canine, the dorsoventral diameter of the tooth is equal to about half the height of the mandible. I,.—This tooth and the somewhat larger canine behind are more similar in form to one another than either is to the I,. Presumably, the same could be said for the Ps, although the tooth is known only by its alveolus in this species. The single root of the I dips pos- teriorly at a shallow angle. Overlapping the I, ahead, the crown of the I, projects forward from the root. Along the midline is a low crest with two prominent swellings. The anterior swelling lies slightly forward of the midpoint of the tooth and is the protoconid; the second swelling lies at the posterior margin of the short talonid. The length of the tooth is about twice the width. NORTH AMERICAN ERINACEINE HEDGEHOGS 39 C,.—In addition to its somewhat larger size, the C, differs from the I; primarily in the relative sizes of protoconid and talonid cusp, which are slightly taller relative to the length and width of the tooth. P,.—This tooth is known in this species only by its single al- veolus: The P. was probably similar in size and morphology to the Iz, because the alveoli of the two teeth are of similar dimensions; in other erinaceines, they are similar in morphology. The posterior dip of the Py» alveolus is steeper than in I, or Cj. P,.—This tooth is markedly larger than the three immediately anterior to it. The protoconid is most prominent of the cusps; its height is nearly as great as the length of the tooth. Anteromedial to the protoconid is the lower paraconid, a well-developed cusp. In lateral view the paralophid linking the two cusps has a deep, acute V-shaped profile. The protolophid is directed medially from the protoconid and has a slight swelling on the lingual end, but no distinct metaconid. Along the buccal border of the tooth is a weak cingulum. This cingulum passes into the narrow talonid which is developed for the full width of the trigonid. On no specimen were the alveoli of this double-rooted tooth visible. M,—The length of the trigonid is four-fifths its width and approximately equal to half the length of the tooth. Lowest of the trigonid cusps, the paraconid is anterior to the metaconid; the bases of the two cusps are separated by a deep cleft. The proto- conid is posterobuccal to the paraconid; the two cusps are linked by a paralophid. In lateral view, the paralophid has a V-shaped profile that is acute but nearly a right angle, whereas in occlusal view, the paralophid is somewhat buccally convex. The protoconid is the tallest cusp on the tooth and has the greatest basal dimensions. On the least worn specimen (UCM 29853), the height of the proto- conid is equal to 85 percent of the tooth length. The protolophid joins the metaconid and protoconid; when viewed posteriorly, it has an acute, V-shaped profile. The metaconid is intermediate in height between the protoconid and paraconid. Ten percent wider than the trigonid, the talonid has only two cusps, one in each posterior corner. The entoconid is the taller of these two cusps and is intermediate in height between the meta- conid and paraconid. The entoconid is directly posterior to the metaconid and the hypoconid is slightly more buccally placed than the protoconid. In anterior view, the buccal margin of the proto- conid is nearly straight and that of the hypoconid, convex. Be- tween the entoconid and that part of the posterior wall of the tri- gonid beneath the apex of the metaconid is a low entocristid which encloses the talonid basin lingually. Directed anteriorly and slightly lingually from the hypoconid, the cristid obliqua abuts against the postvallid immediately behind the protoconid. A continuous, well- 40 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY developed, narrow cingulum extends along the entire buccal side of the tooth. At the base of the hypoconid, the buccal cingulum passes into the posterior cingulum which is directed dorsomedially so that it terminates lingually midway between the hypoconid and entoconid just below the crest of the postcristid. In the region of transition between the buccal and posterior cingula, these narrow cingula are even narrower. The partition between the alveoli of this double-rooted tooth has twinned vertical ridges characteristic of erinaceids. On the posterior wall of the rear alveolus is a single vertical ridge. The anterior alveolus is nearly circular with the posterior side slightly flattened. Its diameter is approximately 0.7 mm. The larger, pos- terior alveolus is roughly rectangular with an anteroposterior length of 0.8 mm and width of 0.9 mm. M,.—Although the Mb is only four-fifths as large as the M,, in other respects the teeth are quite similar. Therefore, only differ- ences between these teeth will be noted in the following two para- graphs. The trigonid length is only two-thirds as great as its width. The paraconid is not developed as a distinct cusp but rather is a small swelling at the anterolingual end of the paralophid. The antero- lingual quarter of the protolophid is at an abrupt angle with respect to the remainder of that crest, being oriented mediolaterally rather than anterolingually. The entoconid is more buccally placed on the tooth than the metaconid. When viewed anteriorly, the buccal margin of the protoconid is buccally convex. No buccal cingulum is developed on the most anterior quarter of the prevallid. M,.—This tooth differs from the two anterior molars in several ways: markedly smaller size; complete absence of a talonid; more anteroposteriorly expanded trigonid, its length being one-fourth greater than its width; and a single anteroposteriorly elongated root that may be weakly bifurcated. Only two distinct cusps are present on the trigonid—the protoconid and metaconid. The protoconid is taller and somewhat larger in basal dimensions than the metaconid; the bases of the two cusps are so close to one another that only a narrow notch separates them. No swelling is present at the lingual end of the paralophid to mark the presence of a distinct paraconid. Only the worn base of the paralophid remains on the best available specimen of this tooth. A well-developed, continuous, narrow basal cingulum is present along the buccal margin of the tooth, but no cingulum is present along the posterior side of the trigonid. Oval in outline, the single alveolus of this tooth departs slightly from the vertical so as to dip posteriorly. The length of the oval is 0.9 mm and the width, 0.5 mm. Mandible.—( Figs. 13-15) Circular in outline, the mental fora- men lies below the middle of Py, midway between the dorsal and NORTH AMERICAN ERINACEINE HEDGEHOGS 4] ventral borders of the ramus. Beneath Ps-Ms, the mandible is of a uniform depth; it gradually decreases in depth posteriorly and abruptly anteriorly. The angle of the jaw protrudes ventrally and posteriorly away from the body of the mandible and is slightly in- flected. The unfused midline symphysis of the mandible extends posteriorly to a point beneath the C,-P; juncture. The dorsal boundary of the symphysis dips posteriorly at an angle of 25° with respect to the dorsal edge of the mandible ( / c in Fig. 11). Be- neath I,, the symphysis covers the entire medial surface of the mandible; posteriorly it covers a progressively decreasing amount. Parallel and adjacent to the ventral margin of the midline symphysis is a faint depression which opens posteriorly and may have been the site for the insertion of the geniohyoideus muscle. The ascending ramus lies slightly labial to the midline of the mandible. Its anteroposterior width, measured from the condyle, is twice the maximum depth of the jaw. The anterior border of the ascending ramus departs slightly from the vertical and dips an- Fic. 16. Method of measuring distances between occluding surfaces. Cor- responding measurements on the upper and lower teeth are indicated by the same letter of the alphabet, capital letters for the upper teeth and small letters for similar measurements on the lower teeth. 42 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY teriorly; the angle of elevation ( Z a in Fig. 11) is 80°. Near its dorsal end, the anterior edge curves slightly posteriorly and dorsally and passes smoothly into the convex dorsal edge of the ascending ramus. On the leading edge of the ascending ramus, there is a sharp lateral flange that forms the anterior margin of the masseteric fossa. The prominence of this flange decreases dorsally. There is a ridge on the medial side of the ascending ramus. Beginning in the anteroventral corner of the ascending ramus immediately behind the tooth row, the ridge extends posterodorsally for a distance less than one-fourth the width of the ascending ramus. To the rear, the ridge terminates somewhat anterior to the mandibular foramen; between that point and the condyle, there is no ridge developed on the medial side of the mandible. The mandibular foramen is mid- way between the anterior and posterior borders of the ascending ramus and level with the tooth row. The condyle is well above the level of the tooth row, the angle of elevation ( / b in Fig. 11) being about 35°. In posterior view, the margin of the mandible has a knife-edge thinness except immediately below the condyle. The posterior border of the ascending ramus curves anterodorsally from the condyle and meets the dorsal edge of the ascending ramus at a sharp angle. Comvarisons.—In a living erinaceine such as Erinaceus euro- paeus, the distances between adjacent occlusal surfaces of the upper dentition are found to be close to those of the corresponding occlusal surfaces of the lower dentition. For example, the distance from the prevallum of M! to that of M? is nearly the same as that from the postvallid of M, to that of Ms. Because the prevallum of M! is adjacent to the postvallid of M, when the two teeth are in occlusion and likewise, the prevallum of M? is adjacent to the post- vallid of M» in the same circumstances; this result is hardly sur- vrising. See figure 16 for a diagramatic explanation and table 8 for a set of measurements in a sample of five recent specimens of E. europaeus. If the distances between the vrevalla of the upper molars of Parvericius montanus (YPM 13956, Fig. 12a) are compared to the TABLE 8. Measurements (mm) of the distances between occlusal planes in Erinaceus europaeus (See Fig. 16 and text for explanation ) AMNH(M) AMNH(M) AMNH(M) AMNH(M) AMNH(M) 34820 34821 149412 160470 164391 A 8.9 9.2 8.4 9.1 8.8 a 8.7 9.1 8.6 8.6 8.6 B 4.6 4.9 4.4 4.6 4.5 b 4.6 4.8 4.7 4.6 4.5 C 4.3 4.3 4.0 4.5 4.3 c 4.1 4.3 3.9 4.0 4.1 43 NORTH AMERICAN ERINACEINE HEDGEHOGS GT 6'T CT ea ia = = a. val al 6'1 LG 0% ei) 13 cs = ie Fil SG VS OF G¢ ie re — as ie as 6S TOOST AX “ed4} 69S9SA dNWWNNA ¢c66¢ WON 9S6ET Wd 4} 08066 HNINV STYDJUD} SNUDIJUOUL SNUDJUOU SNUDJUOUL SUA PIID SNULYIIAOUI4S SMIUIAD YT SMIUIAMW SMI1LIAID LOJADISOADIDT ( uolyeuR[dxe 10j 4x9} pue QT “SI 90S ) snuryoaouayg pue ‘snia1aaig “10jdvosoanjpg Ul sued [esn[o00 UsaA\joq s9oUR}SIP OY} JO (UU) S}USUTOINSeITY “6 ATAV I, =< @ Aa Oo td OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY distances between the postvallids of the mandible fragment pre- sumed to be that of P. montanus from the Split Rock Formation (UCM 29955, Fig. 5), the corresponding measurements are found to be similar to the degree found between individuals of Erinaceus europaeus of slightly different size (compare Tables 8 and 9). In addition, these upper and lower dentitions have been modified in the characteristic erinaceine pattern; M? reduced to a straight blade between the protocone and paracone; M; diminished to a trigonid lacking a talonid. These specializations of the most posterior upper and lower molars are functionally correlated with one another and are not known to occur singly. On the basis of the above evidence only, it could be postulated that the mandibles of Stenoechinus tantalus were the previously unknown lower jaws of Parvericius montanus. The distances be- tween the corresponding occlusal surfaces are nearly the same (Table 9); and, although the M; of S. tantalus does have a wide postcingulum, it conceivably did occlude with an M® similar to that on P. montanus. However, the M! and M®? referred to S. tantalus (found at the same locality as the majority of known lower denti- tions of that species) appears to be quite unlike those of P. mon- tanus; thus, the possibility of taxonomic synonymy seems improb- able. The material from Egelhoff Quarry further supports the assignment of the lower dentions, here inferred to belong to P. montanus, with the upper dentition on which the species is based. At that locality only one form of small erinaceine tooth is known for each position of the dental series represented, suggesting that all such teeth belong to a single snecies. The isolated P* and two M’’s from Egelhoff Quarry are quite similar to those of the type of P. montanus, whereas the lower teeth from there are 10 percent larger than those of the Split Rock mandible. Dentitionally, specimens of Parvericius montanus from Taben- buluk (originally described as Palaeoerinaceus minimus Bohlin, 1942), and from Nareen Bulak (originally described by Sulimski, 1970, as Amphechinus (Palaeoerinaceus) cf. minimus), are similar to specimens of P. montanus from North America. On the basis of dimensions of the teeth, this Asian material is most similar to older, smaller Arikareean and Hemingfordian North American specimens of P. montanus. There is a marked difference in the depth of the mandible; the North American specimens are about half again as deep as the Asiatic ones (Bohlin, 1942: Fig. 3d, d’, Plate 1, Fig. 4; Sulimski, 1970: Fig. 2d-f; Plate 19, Figs. 5-7). Besides P. minimus, Bohlin described another species in 1942 from Taben-buluk, Pa- laeoerinaceus kansuensis Bohlin (1942). This species is based on an incomplete edentulous mandible fragment. The mandibular depth is closer to the North American P. montanus than P. minimus, being only one-fifth larger. Furthermore, based on the alveoli, the Ce —E———— - m z = m 4 m aD o) JEEIEIS) INIT Eh IWOS snjojub, S RSS a SNHIALAOLNI | Pongoueie SAWAYO9IU9 —-$N/DIUD{ “S=om Ivh:1Z61 “YS9Y3SESN3Y 291:6961 ‘NSSSNWSVUY NOILO3S G3aYyNSVAW NOILOAS GSYNSVAW AVG NHOP HOLVd JSVEEVO 48 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY STRATIGRAPHIC RELATIONSHIPS Stratigraphic positions of the erinaceine insectivores discussed in this paper from Granite and Powell Counties, Montana, are illustrated in figure 18. Cabbage Patch beds shown in this figure range in age from medial to late Arikareean (late Oligocene and early Miocene), as based on correlations of the enclosed Cabbage Patch faunas to Arikareean faunas in the John Day of Oregon and Great Plains of South Dakota, Nebraska, and Wyoming. Cabbage Patch localities 2 (KU-Mt-9) and 4 (KU-Mt-12) and Bert Creek locality 2 (MV6504-2) occur within a single measured section. Cabbage Patch locality 13 (KU-Mt-46) occurs not far from this measured section and can be directly correlated into it. Because of poor exposures, erosional gaps, and distances up to several kilometers, correlation of Cabbage Patch locality 3 (KU-Mt-11) and Tavenner Ranch locality 2 (KU-Mt-21) to the measured section dis- cussed above is difficult, but each can be approximately correlated using the stratigraphic position of the range zone of the gopher Pleurolicus. Thus, each locality is correlated by recording whether it occurs above, within, or below the Pleurolicus range zone. Like- wise, an approximate correlation of this biostratigraphic zone can be made to the Pleurolicus range zone in the John Day Formation of Oregon (Rensberger, 1971:141, Fig. 69). Ten of the eleven specimens of Stenoechinus tantalus occur at a single horizon and locality (Tavenner Ranch locality 2) in the upper Cabbage Patch beds above the Pleurolicus range zone. AlI- though several rich fossil localities above and below this strati- graphic position of Tavenner Ranch locality 2 have been extensively searched and large bulk samples have been washed from some of them, only one other specimen of Stenoechinus has been found (at Cabbage Patch locality 13 which lies within the Pleurolicus range zone in the middle Cabbage Patch beds). Also occurring at Tay- enner Ranch 2 is a partial maxillary (KU 18097) and partial isolated tooth (KU 18405) of Amphechinus horncloudi. Other mammals found at this locality include Nanodelphys, Peratherium, Domnina, Proscalops, Meniscomys, Niglarodon, Paciculus, Leidymys, Plesios- minthus, Gregorymys, Mookomys, Paleocastor, Archaeolagus, Notho- cyon, Leptomeryx, and oreodonts, and will be reported on in a later paper by Rasmussen. Two additional specimens of Amphechinus horncloudi have been found within the Pleurolicus range zone in the middle Cabbage Patch beds (Fig. 18)—KU 18162 from Cabbage Patch locality 3, and KU 18163 from Cabbage Patch locality 2. Each was found associated with a fauna that is less varied but similar to that ac- companying A. horncloudi at Tavenner Ranch locality 2, except that Pleurolicus is present and Gregorymys is absent. An upper second molar of a small erinaceine (KU 18346) was NORTH AMERICAN ERINACEINE HEDGEHOGS 49 found at Cabbage Patch locality 4 within the Pleurolicus range zone in the middle Cabbage Patch beds. Although more than one ton of matrix has been washed from this locality and several hundred specimens have been recovered, this is the sole record of a hedge- hog in the fauna other than the genus Ocajila. Mammals from the locality include Peratherium, a mole, Ocajila, Domnina, Allomys, Niglarodon, Pleurolicus, Mookomys, a beaver, Eutypomys, Archae- olagus, an ochotonid, and Diceratherium. A lower jaw of a small hedgehog (MPUM 1551) referred to Parvericius montanus was found in the middle Cabbage Patch beds in association with Peratherium, a beaver, Pleurolicus, and Archaeo- lagus at Bert Creek locality 2. The Pleurolicus specimens collected at this locality are among the most primitive in the Cabbage Patch beds, indicating this locality is near the base of the Pleurolicus range zone. Several specimens of Stenoechinus have been found at Tavenner Ranch locality 2 in the Cabbage Patch beds and were not found at other localities stratigraphically above and below (except the single specimen from Cabbage Patch locality 13). Even at localities which were intensively sampled, the absence of Stenoechinus may be due to paleoecological differences among the various localities. The type locality of Sfenoechinus (Tavenner Ranch locality 2) is uniquely rich in terrestrial gastropods, whereas all other localities where hedgehogs were found (including Cabbage Patch locality 13) have very numerous freshwater gastropods. The scarcity of fresh- water gastropods and other aquatic organisms and the abundance of terrestrial animals at Tavenner Ranch locality 2 suggests that the area was relatively remote from aquatic environments during most of the episode of deposition of this particular bed. The sediment at Tavenner Ranch locality 2 is composed almost entirely of silt- sized glass shards with little clay and sand, and may have been partially wind-blown into the site of deposition. Root burrows are extremely common, indicating abundant vegetation. A grassy or wooded habitat in an area free from frequent flooding or standing water is indicated. Thus Stenoechinus may have lived in habitats which were drier than those indicated at the other localities where other types of hedgehogs have been found. Two isolated lower molars and a mandibular fragment of Parvericius montanus were found in Fremont Co., east central Wyoming, in the upper porous sandstone sequence of the Split Rock Formation in association with mammals of the Split Rock local fauna considered to be of Hemingfordian age, medial Miocene (Love, 1961:24; Munthe and Lander, 1973:497). All these speci- mens were found at or near locality 11-V of Love (1970). Two mandibles, three isolated lower teeth, and three isolated upper teeth of Parvericius montanus were found at Egelhoff Quarry in Keya Paha Co., north-central Nebraska, in an unnamed lithic 50 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY unit that unconformably overlies the Rosebud Formation and in turn is unconformably overlain by the Valentine Formation (R. H. Tedford, pers. com., 1972). In the same lithic unit and about 2 km southeast of Egelhoff Quarry, is the Norden Bridge Quarry in Brown Co., Nebraska, where an isolated M, of P. montanus was found. Both local faunas are regarded as Barstovian in age (C. W. Hibbard, pers. com., 1972). The type of Parvericius montanus Koerner (1940), a maxilla fragment, was found in Meagher Co., central Montana, in the Deep River beds. Unfortunately, Koerner never published a complete faunal list for the Deep River beds. However, the oreodont material he collected and described (Koerner, 1940), together with his ro- dent and lagomorph specimens described by Black (1961), indicate that some parts of the unit (as Koerner conceived it) are of Arik- areean age and other parts Hemingfordian. In other areas of North America, Promerycochoerus and Paciculus are known from Arik- areean deposits while Hypolagus and Monosaulax are known from Hemingfordian or later deposits (Schultz and Falkenbach, 1949; Black, 1961; Dawson, 1958; R. W. Wilson, 1960). CONCLUSIONS In the Arikareean (latest Oligocene and earliest Miocene), a general faunal interchange of mammals, particularly small forms, took place between the Old World and North America after an extended interval of almost no interchange during the medial and late Oligocene (R. W. Wilson, 1968). It is at that time, and pre- sumably as part of that episode of widespread interchange, that the North American history of the Erinaceinae appears to have begun with an invasion from Asia by at least two species. These are documented by the presence of two erinaceine species in Arik- areean deposits of North America—Amphechinus horncloudi and Parvericius montanus. Each species appears to be more closely related to known Oligocene species of Europe or Asia than to any other North American erinaceines. The Arikareean appearance of Stenoechinus tantalus n. gen. and n. sp. in North America may be the result of yet another invasion immediately prior to the Arik- areean. Or it is equally as likely that S. tantalus could represent the first record of a lineage endemic to North America through the Oligocene. Because no earlier species in either the Western or Eastern Hemisphere closely resembles S. tantalus, it is not possible at the present time to strongly support one alternative over the other. A poorly known species described here as “Erinaceinae, genus and species indet.” may be congeneric with one of the three other species. Largest of the North American erinaceine species, Amphechinus horncloudi can be readily allied with the early Oligocene to early NORTH AMERICAN ERINACEINE HEDGEHOGS 51 Miocene European group Amphechinus cayluxi, Amphechinus ar- vernensis, and Amphechinus edwardsi. The three European species and A. horncloudi are all quite similar to one another in size and morphology; apparently little change took place in this group during the time of its existence. Except for a somewhat deeper mandible, Parvericius montanus from North America is nearly identical to medial or late Oligocene east Asian specimens originally described by Bohlin (1942) as Palaeoerinaceus minimus. Parvericius would seem to be most simi- lar and closely allied to Amphechinus among the known erinaceine genera but more primitive in the shorter length of the M! and M? relative to their respective widths and in the more anteroposteriorly compressed M, trigonid. The M2? described as “Erinaceinae, genus and species indet.” appears to be yet another distinct species, because it is too large to be the M? of either Parvericius montanus or Stenoechinus tanta- lus and much too small to be the M? of Amphechinus horncloudi. In addition, it is distinguished from the M? of Paleoscaptor acridens in having a more prominent parastylar spur, and the postprotocrista and postmetaconule crista developed as two distinct ridges rather than as one continuous ridge. One of the three erinaceine genera described here has a post- Arikareean record in North America and a second one may. Par- vericius survived until the late Barstovian; this genus has been found in the samples of that age collected at Egelhoff and Norden Bridge Quarries in north-central Nebraska. Rich will describe these specimens in a future paper. John Storer is describing three molar talonids of a large erinaceid of the proper size to be Amvhechinus, collected at the medial Barstovian Kleinfelder Farm locality in southern Saskatchewan. Although none of the genera discussed here endured in North America after the Barstovian, the Erinaceinae were represented on this continent until at least Clarendonian time as attested by the presence of a single, isolated lower molar belong- ing to a member of this subfamily in the WaKeeney local fauna of Kansas (see R. L. Wilson, 1968: Fig. 12a-b). SUMMARY The history of the Erinaceinae in North America begins in the Arikareean (latest Oligocene and earliest Miocene) with the ap- pearance of four species: Amphechinus horncloudi, Parvericius montanus, Stenoechinus tantalus n. gen. and n. sp., and Erinaceinae, genus and species indet. The first two species listed are more closely related to various species known from Oligocene deposits of Europe and Asia than to any North American ones; hence, at least two separate erinaceine groups appear to have invaded North America. Presumably these invasions took place immediately prior to the 52 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY first record of these species in North America and were part of an episode of renewed, widespread interchange of mammals between North America and the Old World that occurred at that time (R. W. Wilson, 1968). Stenoechinus tantalus is not closely related to any known earlier forms; therefore, the Arikareean record of this species may be either the first discovery of a lineage whose unknown Oligo- cene history was in North America, or the descendant of yet another group that invaded North America from Asia in the Arikareean. The fourth, poorly known species may have been derived from one of the stocks that gave rise to one of the first three mentioned species, or it may represent yet another lineage with an independent history extending well back into the Oligocene. LITERATURE CITED AYMARD, AUGUSTE 1850. (No title.) Ann. Soc. Agric. Puy, 14:104-114. BERGGREN, W. A. 1971. Neogene chronostratigraphy, planktonic foraminiferal zonation and the radiometric time scale. Hungar. Geol. Soc. Bull., 101:162-169, 3 tables. Brack iC. G: 1961. Rodents and lagomorphs from the Miocene Fort Logan and Deep River Formations of Montana. Postilla, (48 ):1-20, 6 figs. DE BLAINVILLE, H. M. D. 1838. Recherches sur l’ancienneté des mammiféres insectivores a la surface de la terre; précédées de l’historie de la science a ce sujet, des principes de leur classification et de leur distribution géographique actuelle. C. R. Acad. Sci. Paris, 6:738-744. BoHLIn, B. 1942. The fossil mammals from the Tertiary deposit of Taben-buluk, western Kansu. Part I: Insectivora and Lagomorpha. In Hedin, Sven, Reports from the scientific expedition to the northwestern provinces of China. Stockholm. Vertebrate Paleontology. 6 (3): 1-113, 32 nes., Ipl. BuTLER, P. M. 1948. On the evolution of the skull and teeth in the Erinaceidae, with special reference to fossil material in the British Museum. Proc. Zool. Soc. London, 118:446-500, 28 figs. 1956. Erinaceidae from the Miocene of East Africa. Brit. Mus. Nat. Hist., Fossil Mamm. Afr., (11):1-75, 18 figs., 4 pls., 16 tables. 1969. Insectivores and bats from the Miocene of East Africa: New ma- terial. In Leakey, L. S. B., ed., Fossil Vertebrates of Africa. Aca- demic Press, New York and London. 1:1-37, 13 figs. Dawson, M. R. 1958. Later Tertiary Leporidae of North America. Univ. Kansas Paleont. Contrib., Vertebrata, (art. 6):1-75, 39 figs., 2 pls., 14 tables: FILHOL, HENRI 1879. Etude des mammiféres fossiles de Saint Gerand le Puy (Allier). Ann. Sci. géol. (Paris), 10:1-253, 30 pls. FRIANT, M. 1961. Les insectivores de la famille des Erinaceidae. L’evolution de leurs molaries au cours des temps géologiques. Ann. Soc. Géol. Nord. 81:17-30, 6 figs., 2 pls. NORTH AMERICAN ERINACEINE HEDGEHOGS 53 HURZELER, J. 1944, Uber einem dimyloiden Erinaceiden (Dimylechinus nov. gen.) aus dem Aquitanien der Limagne. Eclog. Geol. Helvetiae. 37 (2) :460- 467, 13 figs. Korrn_er, H. E. 1940. The geology and vertebrate paleontology of the Fort Logan and Deep River formations of Montana. Part I: New vertebrates. Amer. Jour. Sci. 238 (12) :837-862, 7 pls. Love, J. D. 1961. Split Rock Formation (Miocene) and Moonstone Formation ( Plio- cene) in central Wyoming. U. S. Geol. Surv. Bull. (1121-1) :1-39, 6 figs., 3 tables. 1970. Cenozoic geology of the Granite Mountains area, central Wyoming. U. S. Geol. Surv. Prof. Paper (495-C): viii-++154 pp., 61 figs., 13 tables. MACDONALD, J. R. 1970. Review of the Miocene Wounded Knee faunas of southwestern South Dakota. Bull. Los Angeles Co. Mus. Nat. Hist. (8):1-82, 32 figs., 53 tables, 2 maps. MACDONALD, L. J. 1972. Monroe Creek (early Miocene) microfossils from the Wounded Knee area, South Dakota. South Dakota Geol. Surv. Rept. Invests. 105: iii+43 pp., 14 figs. MatTrHew, W. D., and WALTER GRANGER 1924. New insectivores and ruminants from the Tertiary of Mongolia, with remarks on the correlation. Amer. Mus. Novitates (105) :1-7, 3 figs. Merrie, J. S. 1968. The Oligocene Hsanda Gol Formation, Mongolia: a revised faunal list. Amer. Mus. Novitates (2318):1-16, 4 figs., 2 tables. MunrTHE, JENS, JR. and FE. BrucE LANDER 1973. A reevaluation of the age of the Split Rock vertebrate fauna, Wyoming. Geol. Soc. Amer. Abs. with Programs 5(6):497. RaAsMussEn, D. L. 1969. Late Cenozoic geology of the Cabbage Patch area, Granite and Powell Counties, Montana. Univ. Montana unpubl. M.A. thesis: 1-188, 15 figs., 10 plates, 3 tables. RENSBEBGER, J. M. 1971. Entoptychine pocket gophers (Mammalia, Geomyidae) of the Early Miocene John Day Formation, Oregon. Univ. Calif. Publ. Geol. Sci. 90: 1-209, 76 figs., 22 pls., 15 tables. Ricu, T. H.V. and P. V. Ricu 1971. Brachyerix, a Miocene hedgehog from western North America with a description of the tympanic regions of Paraechinus and Podo- gymnura. Amer. Mus. Novitates (2477):1-58, 22 figs., 4 tables. ScHULTz, C. B. and C. H. FALKENBACH 1949. Promerycochoerinae, a new subfamily of oreodonts. Bull. Amer. Mus. Nat. Hist. 93 (art. 3):69-198, 26 figs., 8 tables, 6 charts. Srox., N. R., et al., eds. 1961. International Code of Zoological Nomenclature. International Trust for Zoological Nomenclature. London: xviii--176 pp. SULIMSKI, A. 1970. On some Oligocene insectivore remains from Mongolia. Palaeont. Polonica. (21):53-70, 2 figs., 2 pls., 6 tables. TRoFIMOV, B. 1960. [The insectivore genus Palaeoscaptor from the Oligocene of Asia.] Trudy Pal. Inst. Akad. Nauk SSSR 77:35-40, 3 figs., (Russian). 54 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY VAN CovuvERING, J. A. 1972. Radiometric calibration of the European Neogene. In W. W. Bishop and M. A. Miller, eds., Calibration of Hominoid Evolution. Edin- burgh, Scottish Academic Press:247-271, 2 figs., 2 tables. VAN VALEN, L. 1967. VIRET, J. 1929. 1938. Wi:son, R. 1968. WItson, R. 1960. 1968. New Paleocene insectivores and insectivore classification. Bull. Amer. Mus. Nat. Hist. 135(art. 5):217-284, 7 figs., 2 pls., 7 tables. Les faunes de mammiféres de |’Oligocéne supérieur de la Limagne Bourbonnaise. Ann. Univ. Lyon, (n.s. 1), (fase. 47): viii+328 pp., 32 figs., 32 pls. Etude sur quelques Erinacéidés fossiles spécialement sur le genre Palaerinaceus. Trav. Lab. Géol. Univ. Lyon, (fasc. 34, Mém. 28): 1232, 12 ies. 4 pl. 1 Systematics and faunal analysis of Lower Pliocene vertebrate as- semblage from Trego County, Kansas. Contrib. Mus. Paleo., Univ. Mich. 22(7 ):75-126, 17 figs., 20 tables. W. Early Miocene rodents and insectivores from Northeastern Colo- rado. Univ. Kansas Paleont. Contrib., Vertebrata, (art. 7):1-92, 131 figs. Insectivores, rodents, and intercontinental correlation of the Mio- cene. XXIII International Geological Congress 10:19-25, 1 table. Woop, A. E. and R. L. Konizesxt 1965. A new eutypomyid rodent from the Arikareean (Miocene) of Mon- tana. Jour. Paleontology. 39(3):492-496, 2 figs., 2 tables. UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly pub- lished in the above series are now published as Occasional Papers, Museum of Natural History. The Miscellaneous Publica- tions, Museum of Natural History, began with number 1 in 1946. Longer research papers are published in that series. Monographs of the Museum of Natural History were initiated in 1970. All manuscripts are subject.to critical review by intra- and extra- mural ‘specialists, final acceptance is at the discretion of the publications: committee. Institutional libraries interested in exchanging publications may obtain the Occasional Papers and Miscellaneous Publica- tions by addressing the Exchange Librarian, University of Kan- sas Library,- Lawrence, Kansas 66045. Individuals may pur- chase separate numbers of all series. Prices may be obtained upon request addressed -to Publications Secretary, Museum of Natural History, University of Kansas, Lawrence, Kansas 66045. Editor: LinpA TRUEB Managing Editor: WiLL1AM E. DUELLMAN PRINTED BY UNIVERSITY OF KANSAS PRINTING SERVICE LAWRENCE, KANSAS PN p27 ALIFOR ACADEMY O FEB - 6 1974 LIBRARY ) ole LLMe OCCASIONAL PAPERS of the MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas NUMBER 22, PAGES 1-27 JANUARY 25, 1974 A SYSTEMATIC REVIEW OF THE MARSUPIAL | FROGS (HYLIDAE: GASTROTHECA) OF THE ANDES OF ECUADOR By WILLIAM E.. DUELLMAN' Many species of marsupial frogs, genus Gastrotheca, occur in the Andes and associated cordilleras from western Venezuela south- ward to northern Argentina. Because of the paucity of specimens of many of the named taxa and confusing variation exhibited by some species, the taxonomy of the marsupial frogs has been chaotic. Duellman and Fritts (1972) reviewed the species occurring in the Andes to the south of the Huancabamba Depression in northern Pert and delimited seven species there as members of the Gastro- theca marsupiata group. These frogs differ in cranial characters from the species in the Huancabamba Depression and northward, all of which were referred to the Gastrotheca argenteovirens group by Duellman and Fritts (1972). The Ecuadorian species included in the latter group were: G. lojana Parker, 1932; G. monticola Barbour and Noble, 1920; G. plumbea (Boulenger, 1882); and G. riobambae (Fowler, 1913). Colombian and Venezuelan species assumed to belong to the same group were: G. argenteovirens (Boettger, 1892); G. aureomaculata Cochran and Goin, 1970; G. helenae Dunn, 1944; G. medemi Coch- ran and Goin, 1970; G. mertensi Cochran and Goin, 1970; and G. nicefori Gaige, 1933. It now seems apparent that these frogs are treated best as two distinct species groups. The Ecuadorian species listed above and two species named in this paper can be referred to as the Gastrotheca plumbea group, and the Colombian and Vene- zuelan species, as the Gastrotheca argenteovirens group. When bet- ter known, G. williamsoni Gaige, 1922, and an unnamed species re- lated to G. plumbea from Antioquia and Cundinamarca, Colombia, ‘Curator, Division of Herpetology, Museum of Natural History, University of Kansas. NI F SCIENCES 2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY may be referable to the Gastrotheca argenteovirens and plumbea groups, respectively. Thus, this paper is a review of the Gastrotheca plumbea group. The purpose of the present paper is to provide a synthesis of accu- mulated information and in so doing, 1) define the species, 2) as- sign names to taxonomically recognizable populations, and 3) sum- marize data on geographic variation, distribution, ecology, and life history. The results presented here, together with the synthesis of the southern Andean frogs by Duellman and Fritts (1972), will provide a basis for a future systematic assessment of the Gastro- theca argenteovirens group. ACKNOWLEDGMENTS For the loan of specimens or provisions of facilities in their re- spective institutions, I am grateful to F. Bernis, James E. Bohlke, F. W. Braestrup, Nelly Carrillo de Espinoza, Javier Castroviejo, Josef Eiselt, Alice G. C. Grandison, Jean Guibé, Konrad Klemmer, Alan E. Leviton, Robert F. Martin, Hymen Marx, Charles W. Myers, Giinther Peters, the late James A. Peters, Greta Vestergren, Charles F. Walker, and Ernest E. Williams. Much of the new material reported herein was collected by Thomas H. Fritts, John D. Lynch, and Linda Trueb; Arthur C. Echternacht, Bruce Mac- Bryde, Richard R. Montanucci, and John E. Simmons also aided in the field work. I appreciate their efforts in my behalf. Field work in Ecuador was supported in part by Watkins Mu- seum of Natural History Grants, The University of Kansas; travel to European museums was provided by a grant from the Penrose Fund (No. 5063) from the American Philosophical Society. I am indebted to Linda Trueb for critical review of my prose and to Thomas H. Fritts for his provocative comments during formulative phases of this work and critical review of the manuscript. My final acknowledgment is to Charles F. Walker, who has given freely his time and advice on marsupial frogs, and who offered valuable com- ments on the manuscript. MATERIALS AND METHODS I have examined 1125 preserved frogs (including type speci- mens of all taxa), 24 skeletons, and 50 lots of tadpoles referable to the species discussed in this paper, and I have studied each of the species in the field. Terminology follows that of Duellman (1970), except for two additional measurements: length of third finger is the distance from the proximal edge of the palmar tubercle to the tip of the finger; length of thumb is the distance from the proximal edge of the prepollical tubercle to the tip of the thumb. Through- out the text, specimens are listed by their catalogue numbers pre- ceded by the appropriate museum abbreviations, as follows: MARSUPIAL FROGS OF THE ANDES OF ECUADOR 3 AMNH American Museum of Natural History ANSP Academy of Natural Sciences of Philadelphia BMNH _ British Museum (Natural History ) CAS California Academy of Sciences CAS-SU | Stanford University Collection (in California Acadenty of Sciences ) FMNH _ Field Museum of Natural History FSM Florida State Museum KU University of Kansas Museum of Natural History MCZ Museum of Comparative Zoology, Harvard University MJP Museo Javier Prado, Lima, Pert MNCN_ Museo Nacional de Ciéncias Naturales, Madrid MNHN_- Museum National d’Histoire Naturelle, Paris NHMW_ Naturhistorisches Museum, Wien NHRM _~ Naturhistoriska Riksmuseet, Stockholm SMF Senckenbergische Museum, Frankfurt TNHC ~ Texas Natural History Collection UMMZ University of Michigan Museum of Zoology USNM United States National Museum UZM Universitets Zodlogiske Museum, Copenhagen ZMB Zoologisches Museum, Berlin THE GASTROTHECA PLUMBEA GROUP Species comprising this group have short to moderately long legs, narrow to moderately wide heads, relatively large hands, moderate to extensive exostosis of the dermal roofing bones, broad frontoparietals expanded into lateral flanges, the frontoparietal fontanelle covered by the frontoparietals, and a long cultriform process of the parasphenoid. All species have aquatic tadpoles. In contrast, members of the Gastrotheca argenteovirens group have long legs, broad heads, large hands, extensive exostosis of the dermal roofing bones and in some species co-ossification and cas- quing, and at least in some species a short cultriform process. Inso- far as known, all species in this group have aquatic tadpoles, but at least in G. argenteovirens, the tadpoles hatch at an advanced stage, and the larval period is short. Members of the Gastrotheca marsupiata group have short legs, narrow heads, small hands, no exostosis of the dermal roofing bones, small narrow frontoparietals not roofing the frontoparietal fontanelle, and a long cultriform process. Three species have aquatic tadpoles; in the other four development is completed in the maternal pouch. ACCOUNTS OF SPECIES In the following accounts, each taxon is diagnosed; the variation is discussed, and the distribution of each species is annotated. Per- tinent measurements and proportions are given in Table 1, and the frogs are illustrated in figures 1 and 2. Maximum snout-vent lengths are given in the diagnoses. OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY | 6¢1'0—6rS'0 TE0'0+ L990 VV 0—cES'0 6£0'0+229°0 60L°0—V0L'0 LOL'0 004'0—6L9'0 L€0'0-+0S9'0 1690 —899'0 ve9'0 L99°0—009'0 9¢0'0+0€9'0 6SL0—V99'0 c69'0 689 0—0FS 0 y6S'0 0&S'0—O8T'0 T0S'0 9¢9'0—L89'0 0c0'0+609'0 S69 0=5195'0 GVO 0+1¢9'0 6&9 0—T8¥'0 SS0'0+6ES'0 oA /vanueduid J, 896 0—81¢ 0 SL¢0—I¥G0 GLO‘ 0+S8¢'0 £10'0+808'0 6L60—€16 0 £86 0 _S¥G'0 910'0+6EC'0 010 0+6S¢'0 696 0 _VrE 0 c9t"0—8St'0 86 0 69E'0 GLG 0 9460 VLE'O—TVE'0 800'0+£96'0 TTO0'0+€SE'0 80¢°0—6L¢'0 COV 0—S0'0 9660 SPE'0 Tct'0 0860 LEC'O_ P60 VI0O'0+ T0€'0 €T0'O+ETE'O SCE 0816 0 ctv 0—L8t'0 GOE'0 8IV'0 086 0—LVG 0 v6G0—EL¢'0 L9G'0 S860 61096 0 90F'0—06E'0 L660 96£°0 STE 0 P66 0 LLE'O—EEC'0 O10'0+€0E'0 660'0+ TSE 18G'0—EGS'0 60¢'0—_S9¢ 0 €10'0+99¢'0 G10'0+98¢'0 8L¢'0—9FG 0 91E°0—9Le6 0 ST0°0+99¢'0 910'0+066'0 YyysueT peop] UIPIM pee /{EQSON-9AG /[eyig1019} UT 8LV'0—9IV'0 L10 0+GPFV'0 VEV 0—Z9E'0 Té0'0+9TF'0 cTe;0-—208'0 OTS'0 c6r 0—ZrV 0 6600+ L9V'0 OTS0 227 0 9LV'0 c8r 0—éSr'0 O10'0+0LF'0 GLV'0—Vetr'0 ccr'0 00S'0—CEF 0 8SPr'0 6LV'0—?r&Sr'0 9970 v9OV 0—LEV'0 TT0°0+0S7'0 609 0—OFF'0 €20'0+PLV'0 16V 0 LGV 0 96 0+C9P'0 "IAS /ysua'T 300, ver 0—ELE0 910°0+ZIF'0 OTF 0—LS¢'0 TZ0'0+€8E'0 STS'0—c0S'0 60¢'0 VES 0—GLV'0 0c0'0+C6r'0 60S°0— 6870 L6V'0 c6r O—LSV'0 G10'0+827'0 6¢S'0—897'0 csoc’0 L6G 0970 89F'0 66S 0 —L6r'0 ITs 0 00S'0—69F'0 G10'0+6LV'0 CSP 0 —C6E'0 L10';0+STV'0 ctv 0—T10r'0 €10'0+FI7'0 TAS /(psusT PIG, Cla ey 6GV'E+8' LV L8V—8 8E CVO E+V EV 0'T9—8'09 6°09 eto =097 69L°T+9'6P 0'89—L'6S v9 69 —_G GS OST e+7 Ls OLL—9T9 8°99 0'09—0°9F 0g V L9—L'0S css WSS = SO) O€SS+V'ES OLOm S24 OVE C+E'EO aS Oca Ouy Stvp+r 7s (TAS) Yisue'T }U9 A -NoUs (osuel “OUT, puodss ‘yxqS [ + UvoeU ‘OUTT 4si1,7 ) ‘DIAYJOLSDY) URIpUY fo suoT}IOodo1d pue s}UsWIOINSeOy “[ AIAV St me) aDquipqgoll **) ------ pyydouyotisd **) ~~ Daqund **) x9g pur sotoeds MARSUPIAL FROGS OF THE ANDES OF ECUADOR 5 Gastrotheca cavia new species Holotype —KU 148532, an adult female, 64.0 mm, from Isla Pequena, Laguna Cuicocha, Provincia Imbabura, Ecuador, 2890 m; one of a series collected on 31 October 1971 by William E. Duell- man and John E. Simmons. Paratopotypes—KU 138216-20, 24 July 1970, Thomas H. and Patricia R. Fritts; 139137-9, 30 January 1971, William E. Duellman; 143094, 148530-1, 148533-36, 31 October 1971, William E. Duell- man and John E. Simmons; 148537-40, 30 January 1971, William E. Duellman. Diagnosis.—1) Body robust, 58.5 mm in males, 67.2 in females; 2) snout round in dorsal view, rounded above and anteroventrally inclined in profile; 3) canthus rounded; 4) loreal region shallowly concave; 5) tympanum vertically elliptical; 6) supratympanic fold moderately heavy; 7) subarticular tubercles on hand large, round; 8) supernumerary tubercles on hand low, round; 9) palmar tubercle bifid; 10) fingers not webbed; 11) toes one-third webbed; 12) tar- sal fold low, extending one-third length of tarsus; 13) inner meta- tarsal tubercle low, ovoid, visible from above; 14) outer metatarsal tubercle low, flat; 15) subarticular tubercles on foot large, round; 16) supernumerary tubercles on foot small, low; 17) discs round; 18) dorsal skin shagreened, tubercular in tympanic region; 19) dor- sum green or tan with or without small irregular black spots; narrow bronze dorsolateral stripe usually present; 20) facial area uniform green or tan; canthal and labial stripes absent; 21) flanks tan with numerous small black spots; 22) dorsal surfaces of limbs plain or with small black spots; 23) posterior surfaces of thighs cream with black mottling; 24) venter cream with black mottling; 25) squa- mosal exostosed, in broad contact with maxillary; 26) temporal arcade complete in large individuals; 27) prevomers abutted medi- ally; 28) transverse processes on eighth presacral vertebra slightly inclined anteriorly. Gastrotheca cavia resembles G. riobambae in having short legs, a narrow interorbital distance, and a short snout. It differs from G. riobambae in having a distinct dorsolateral light stripe, small black flecks on the flanks, and dorsal markings comprised of small black spots. Gastrotheca riobambae lacks a dorsolateral light stripe and has large dark spots on the flanks and a dorsal pattern consist- ing of paired elongate dark marks; the dorsal pattern is shared with G. lojana and monticola. Gastrotheca plumbea and psychrophila have unmarked dorsal surfaces and uniformly dark flanks and ven- ters. The venter in G. cavia is cream with small dark spots. Variation—Whereas some individuals have only a few black flecks on the posterior part of the dorsum, others have more flecks over the entire dorsum; in more heavily flecked specimens, flecks on the shank tend to form transverse bars. In life, adults are green; 6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Fic. 1. A. Gastrotheca cavia, ?>SVL 60.0 mm, KU 139139. B. G. lojana, 6, SVL 56.5 mm, KU 148549. C. G. monticola, 2, 65.0 mm, KU 148568. MARSUPIAL FROGS OF THE ANDES OF ECUADOR i Fic. 2. A. Gastrotheca plumbea, 6, SVL 55.3 mm, KU 142614. B. G. psychrophila, 2, SVL 61.0 mm, KU 120650. C. G. riobambae, 6, 46.5 mm, KU 120725. 8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY the flanks are cream or bronze with black flecks, fused into reticu- lations in some individuals. The groin and posterior surfaces of the thighs are pale blue with black flecks. The dorsolateral stripe, ex- tending from the posterior edge of the eye to the groin, usually is bronze and distinct. In some individuals, the stripe is narrow and fragmented by black flecks; in others it is cream, and in one female it is absent. The throat is pale yellow; the rest of the venter is dull grayish white with small black spots. The iris is dull bronze, heavily marked with brown spots and black flecks. Juveniles are pale green or bronze-tan; the flanks and hidden surfaces of the limbs are pale greenish yellow with black spots, and the venter is pale yellow with or without black reticulations. Although most males and all females are green, three males are tan. Distribution—This species is known definitely from only two localities in the Cordillera Occidental in northwestern Ecuador (Fig. 3). The species is abundant on two rocky islands in a crater lake, Laguna Cuicocha, at an elevation of 2890 m on the south slope of Volcan Cotacachi. The other locality, Hacienda San Nicolas is at an elevation of 2000 m on the Pacific slope of the Cordillera Occidental. Two specimens from “Western Ecuador” (BMNH 1860.6.16.124-125) and one from Ibarra (?) (BMNH 1898.4.28.156 ) are referred to this species. Remarks.—Many adults were found in large terrestrial brome- liads in July, 1970, January and October, 1971. At the time of the last two visits, some adults were found beneath rocks. Brooding females were obtained in January and July, but not in October. Tadpoles were found in the lake surrounding the island in January, and at the same time two metamorphosing young were observed on reeds in the lake. Tadpoles have two upper and three lower rows of denticles. Superficially, Gastrotheca cavia resembles G. argenteovirens Boéttger; I have examined the type of the latter (SMF 2676) and have compared living and preserved specimens of argenteovirens with cavia. In comparison with G. cavia, G. argenteovirens has proportionately longer legs (ratio of tibia length to snout-vent length 0.488-0.514, x = 0.498 = 0.010, N = 5¢ 4; 0.486-0.514, x = 0.501 + 0.012, N = 4¢ @ ) and broader interorbital distance (ratio of interorbital distance to head width 0.335-0.371 + 0.014, x = 0.358, N = 5¢ 8; 0.369-0.395 x = 0.383 2 0.011, Nj==ieeeee The flanks and posterior surfaces of the thigh are mottled with dark blue, and the dorsum lacks black flecks. Furthermore, the cultri- form process of the parasphenoid is short, and in large individuals there is integumentary-cranial co-ossification. Etymology.—The specific name is the same as the generic name for the guinea pig (Caviidae), called cui in Quecchua the domi- MARSUPIAL FROGS OF THE ANDES OF ECUADOR 9 . CONTOUR 3000 M. 5000 M 100 KILOMETERS 1 Fic. 3. Distributions of Gastrotheca cavia (triangles), G. monticola (hexa- gons), G. plumbea (inverted triangles), and G. psychrophila (circles). nant Indian language in the Ecuadorian Andes. Cuicocha means lake of the guinea pig. Gastrotheca lojana Parker New combination Gastrotheca marsupiata lojana Parker, 1932:25 [Holotype—BMNH 1931.2.12.4 (RR 1947.2.31.13) from Loja, Provincia Loja, Ecuador; C. Carrion col- lector]. Diagnosis—1) Body depressed, 56.0 mm in males, 60.0 in fe- males; 2) snout round in dorsal aspect and in profile; 3) canthus rounded; 4) loreal region barely concave; 5) tympanum vertically elliptical; 6) supratympanic fold moderately heavy, angled postero- ventrally behind tympanum; 7) subarticular tubercles on hand 10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY large, round; 8) supernumerary tubercles on hand small, subconi- cal; 9) palmar tubercle bifid; 10) fingers not webbed; 11) toes one- half webbed; 12) tarsal fold low, extending one-half length of tar- sus; 13) inner metatarsal tubercle low, elliptical, visible from above; 14) outer metatarsal tubercle low, round; 15) subarticular tubercles on foot large, round; 16) supernumerary tubercles on foot low, round; 17) discs round; 18) dorsal skin shagreened; 19) dor- sum tan or green, with or without pair of elongate dark markings; narrow bronze or cream dorsolateral stripe present; 20) facial area green or tan; bronze canthal stripe and cream labial stripe present; 21) flanks dark brown with cream spots ventrally; 22) dorsal sur- faces of limbs green or tan, with or without narrow brown trans- verse bars; 23) posterior surfaces of thighs heavily mottled with bluish purple; 24) chin and chest gray; belly pinkish bronze; 25) squamosal exostosed, in narrow contact with maxillary; 26) tem- poral arcade complete; 27) prevomers narrowly separated medial- ly; 28) transverse processes on eighth presacral vertebra transverse. Gastrotheca lojana resembles G. monticola and riobambae in usually having a pair of elongate dorsal markings. It differs from G. riobambae in having fine pale reticulations laterally or dark flanks, nearly uniformly dark posterior surfaces of the thighs, and a dorsolateral light stripe; G. riobambae has large spots on the flanks, mottled posterior surfaces of the thighs, and lacks a dorsolateral light stripe. Gastrotheca monticola has mottled posterior surface of the thighs and further differs from G. lojana in having a wider dorsolateral stripe and a cream venter with dark spots, whereas the venter in G. lojana is dark brown with creamy-white spots. The other Andean Gastrotheca in Ecuador lack paired dorsal markings. Variation.—Variation in coloration can best be described by the following accounts of topotypic adult males (colors in life): KU 148549.—Tan above with greenish suffusion dorsolaterally; dorsolateral and labial stripes bronze; flanks and dorsal markings dark brown; upper surfaces of thighs bronze-tan; upper surfaces of shanks and posterior surfaces of thighs dull green; anterior surfaces of thighs dark brown; throat brown; belly brown with white spots; ventral surfaces of thighs pinkish brown; ventral surfaces of shanks bluish white. KU 148550.—Tan above with brown markings; face mask and anterior flanks dark brown; labial stripe creamy bronze; anterior and posterior surfaces of thighs mottled dark brown and _ blue; throat gray-brown; belly brown with white spots. KU 148551—Dorsum dull green with dark brown markings; flanks are dark brown; anterior and posterior surfaces of thighs and inner surfaces of shanks mottled dull blue and dark brown; throat dark bronze brown; belly brown with cream spots. MARSUPIAL FROGS OF THE ANDES OF ECUADOR 1] KU _ 126073.—Dorsum pale greenish brown with dark brown markings; labial and dorsolateral stripes cream; flanks gray-brown; groin and posterior surfaces of thighs creamy brown with green flecks. KU 142603.—Dorsum dull leaf green with no distinct markings; canthal, labial, and dorsolateral stripes bronze; flanks bronze- brown; groin and anterior and posterior surfaces of thighs purplish brown; throat pinkish bronze; belly same, becoming darker brown posteriorly, with white spots. In all specimens the iris is bronze. Four specimens from Cha- chapoyas, Departamento Amazonas, Peri (KU 138238-41) are colored somewhat differently: “Adults with leaf green dorsum; one with thin beige stripe from nostril through eye to inguinal re- gion and beige spots on flanks; all with white supralabial border; posterior thigh light leaf green; anterior thigh pale leaf green with few black flecks; venter yellow-beige; iris metallic orange. Juvenile with white patch at anus; dorsum tan with light brown blotches edged with dark brown; blotch between eyes and inverted U on body with few scattered small spots posteriorly; broad rich light brown stripe from eye to midflank; groin gray-white with small black blotches; posterior thigh flesh-pink, leaf green distally; supra- labial area beige-white” (T. H. Fritts, field notes, 1 May 1970). The dorsal markings are highly variable. Some individuals lack markings except for a few scattered spots. In most individuals a pair of broad longitudinal marks extend from the scapular region to the groin. In some individuals the marks are fragmented into anterior and posterior components, whereas in others the marks are confluent anteriorly. If an interorbital mark is present, it may or may not be connected to the body markings. Distribution—This species occurs at moderate elevations in the Huancabamba Depression and associated interandean valleys in northern Perti and southern Ecuador, where it has been found from elevations of 2100 to 2350 m in both Atlantic and Pacific drainages (Fig. 4). A record from Zamora, Ecuador, at an elevation of 1000 m on the Amazonian slopes is highly questionable; the specimen (BMNH 1933.6.24.45) is G. lojana, but the locality data probably are erroneous. Remarks.—In the Loja valley in southern Ecuador this species is found most frequently in large Agave, where the frogs seek shel- ter at the bases of the leaves by day and call by night. Individuals also have been found under rocks and in marshy meadows. At Chachapoas, Pert, they were beneath rocks and clods of earth in a cultivated field. Tadpoles were found in a grassy irrigation ditch. They are uniformly black and have two upper and three lower rows of denticles. 12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY is 81 j T l 0 | } - } 3} L 4 \ ~ = — z be Opa ne ~~ 600 M. CONTOUR ; coe 28 ae, ~ | OVER 3000M Dae Z a = (fe A Jo ae | J al 81 80 79 78 his 76 Fic. 4. Distributions of Gastrotheca lojana (triangles) and G. riobambae (circles ). One specimen (FSM 30080, an adult male having a snout-vent length of 48.5 mm) from 24 km WSW of Leimebamba, Departa- mento Amazonas, Pert, 3370 m tentatively is referred to G. lojana. This individual differs from all other G. lojana by having a dark brown dorsum with tan middorsal and dorsolateral stripes; the venter is mottled with dark gray. Futhermore, the locality is 1000 m higher than any other recorded for G. lojana. Parker (1932:25) named lojana as a subspecies of Gastrotheca marsupiata, a name which he applied to populations now known as G. riobambae. As noted in the foregoing diagnosis, G. lojana dif- fers from G. riobambae in a number of characters; there is no evi- dence for gene flow between the southern populations of G. riobam- bae and G. lojana. MARSUPIAL FROGS OF THE ANDES OF ECUADOR 13 Gastrotheca monticola Barbour and Noble Gastrotheca monticola Barbour and Noble, 1920:426 [Holotype—MCZ 5290 from Huancabamba, Departamenio Piura, Peri; G. K. Noble collector]. Gastrotheca marsupiata monticola—Parker, 1932:25. Gastrotheca monticola monticola—Vellard, 1957:39. Diagnosis—1) Body robust, 60.0 mm in males, 77.0 mm in fe- males; 2) snout round in dorsal view and profile; 3) canthus rounded; 4) loreal region barely concave; 5) tympanum nearly round, slightly higher than wide; 6) supratympanic fold moderately heavy, curved posteroventrally behind tympanum; 7) subarticular tubercles on hand large, round; 8) supernumerary tubercles on hand large, conical; 9) palmar tubercle bifid; 10) fingers not webbed; 11) toes one-half webbed; 12) tarsal fold tubercular, ex- tending full length of tarsus; 13) inner metatarsal tubercle elongate, visible from above; 14) outer metatarsal tubercle absent; 15) sub- articular tubercles on feet large, round; 16) supernumerary tuber- cles on hands small, round, flat; 17) discs round; 18) dorsal skin shagreened; 19) dorsum green or tan, usually with paired elongate dark markings; creamy bronze dorsolateral stripe present; 20) facial area green or tan; creamy bronze canthal stripe pesent; 21) flanks brown with cream and black flecks; groin blue; 22) dorsal surfaces of limbs green or tan with or without darker transverse bands; 23) posterior surfaces of thighs blue; 24) throat gray; chest and belly creamy gray with gray spots; 25) squamosal weakly exastosed, in moderately broad contact with maxillary; 26) temporal arcade complete in large individuals; 27) prevomers narrowly separated medially; 28) transverse processes on eighth presacral vertebra barely inclined anteriorly. Gastrotheca monticola differs from G. plumbea and psychro- phila in having a pale venter with black spots, instead of a uni- formly dark venter. Gastrotheca cavia resembles G. monticola in having a broad dorsolateral stripe and mottled posterior surfaces of the thighs, but G. cavia lacks paired longitudinal dark marks on the dorsum, characteristic of G. monticola, lojana, and riobambae. The latter species differs from G. monticola in having large dark spots, instead of fine reticulations, on the flanks, and in lacking a dorsolateral stripe. Gastrotheca monticola is most like G. lojana, from which it differs in having a pale venter with dark spots (dark with white spots in lojana), broader dorsolateral stripe, and more mottling on the flanks and thighs. Variation—The dorsum is green with paired longitudinal dark markings on the body; a large dark spot on the head, including the eyelids, is present in some individuals. The longitudinal marks extend from the eyelids or occipital region to the groin. In some individuals the marks are confluent anteriorly. The marks are green, usually darker than the dorsal ground color, and outlined 14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY with brown. The flanks are brown with or without bronze flecks. The dorsolateral and labial stripes are bronze-tan to metallic cream. The groin and hidden surfaces of the thighs vary from pale green to blue with cream and black mottling. The venter is creamy tan to creamy yellow with brown flecks; the vocal sac is gray. The iris is pale bronze. Comparison of specimens from Saraguro, Ecuador, with the type series from Huancabamba, Pert, revealed slight differences. The Peruvian specimens have less ventral spotting and more dark pigment on the posterior surfaces of the thighs. One Peruvian specimen (UMMZ 57747A), a male, has a highly fragmented dor- sal pattern. Distribution —Gastrotheca monticola occurs at elevations of 1600-2500 m on the Pacific slopes and associated intermontane val- leys in northern Pert and southern Ecuador (Fig. 3). The Cordil- lera Occidental in that region is dissected by many broad, dry val- leys, so it is unlikely that the distribution of G. monticola is con- tinuous. Remarks.—At Saraguro, Ecuador, adults were found beneath rocks in a pasture and in and along a vegetation-choked drainage ditch. At Girodn, Ecuador, adults were in Agave plants by day. Tadpoles were found in the ditch at Saraguro. The body is dull green above and greenish silver below; the tail is tan with green lichenous markings. The tadpoles have two upper and three lower rows of denticles. Parker (1932:25) considered G. monticola to be a subspecies of G. marsupiata (= riobambae). The differences noted in the diag- nosis obviate such an arrangement in the absence of evidence indi- cating genetic interchange. Cochran and Goin (1970:185) used the combination Gastrotheca monticola argenteovirens. Boettger named argenteovirens in 1892, whereas the name monticola dates from Barbour and Noble (1920). Moreover, the two taxa are con- siderably different and certainly not conspecific. Gastrotheca plumbea (Boulenger) Nototrema plumbeum Boulenger, 1882:417 [Holotype—BMNH _ 78.1.25.22 (RR 1947.2.31.19) from Intac, Provincia Pichincha, Ecuador; Mr. Buckley collector]. Gastrotheca plumbeum—Peters, 1955:346. Diagnosis —1) Body robust, 62.3 mm in males, 68.0 mm in fe- males; 2) snout round in dorsal view, angular above and inclined anteroventrally in profile; 3) canthus rounded; 4) loreal region bare- ly concave; 5) tympanum vertically elliptical; 6) supratympanic fold weak; 7) subarticular tubercles on hand moderate, round; 8) super- numerary tubercles on hand small, round; 9) palmar tubercle bifid; 10) fingers webbed basally; 11) toes one-half webbed; 12) tarsal MARSUPIAL FROGS OF THE ANDES OF ECUADOR 15 fold round, extending one-third length of tarsus; 13) inner meta- tarsal tubercle elliptical, visible from above; 14) outer metatarsal tubercle absent; 15) subarticular tubercles on foot moderate, round; 16) supernumerary tubercles on foot small, conical; 17) discs round; 18) dorsal skin shagreened; 19) dorsum green with narrow bronze dorsolateral stripe; 20) facial area green with bronze canthal and labial stripes; 21) flanks brown; 22) dorsal surfaces of limbs green, mottled or not with tan; 23) posterior surfaces of thighs bronze-tan; 24) venter greenish yellow; 25) squamosal not exostosed, in moderately broad contact with maxillary; 26) tem- poral arcade incomplete; 27) prevomers abutted medially; 28) transverse processes on eighth presacral vertebra strongly inclined anteriorly. Gastrotheca plumbea differs from all other Andean Gastrotheca, except G. psychrophila, in having a uniformly pigmented venter and a green iris. Like G. psychrophila, it also lacks dorsal mark- ings, but it differs from G. psychrophila in having paler venter, a dorsolateral light stripe, a green, instead of brown dorsum, and a green, instead of copper, iris. Variation—The dorsum invariably is unmarked bright green to tannish green. The canthal, labial, dorsolateral, and anal stripes are yellow. The loreal region, flanks, and posterior surfaces of the thighs are bronze, and the dorsal surfaces of the limbs are greenish bronze. The venter is yellowish tan with a greenish tint on the throat, and the ventral surfaces of the shanks are blue. A diffuse blue spot is present in the grain; the iris is green. Distribution —Gastrotheca plumbea is known from moderate elevations (1300-2350 m) on the Pacific slopes of the Cordillera Occidental in Ecuador (Fig. 3). Although the range of the species may extend northward into Colombia, it is doubtful if the species ranges into Peru; the dry valleys of the Huancabamba Depression probably are a barrier to southward dispersal. Remarks.—At Pilalé, Provincia Cotopaxi, Ecuador, where there are remnants of cloud forest, individuals were found in bromeliads in trees and on a cliff by day; males called from bromeliads at night. Gastrotheca psychrophila new species Holotype —KU 120760, an adult female, 61.0 mm, from the ridge between Loja and Zamora, 2850 m, 13-14 km E (by road) of Loja, Provincia Zamora-Chinchipe, Ecuador; obtained on 10 June 1968 by John D. Lynch. Paratopotypes—KU 120761, 10 June 1968, John D. Lynch; 141586, 21 May 1971, Richard R. Montanucci; 142631-7, 21-23 July 1971, William E. Duellman. Diagnosis —1) body depressed, 51.5 mm in males, 61.0 mm in 16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY females; 2) snout pointed with vertical rostral keel, in profile rounded above and anteroventrally inclined; 3) canthus angular; 4) loreal region flat; 5) tympanum slightly higher than wide; 6) supratympanic fold heavy, curved downward behind tympanum; 7) subarticular tubercles on hand large, round; 8) supernumerary tubercles on hand large, round; 9) palmar tubercle trifid; 10) fin- gers webbed basally; 11) toes slightly less than one-half webbed; 12) tarsal fold curved, two-thirds length of tarsus; 13) inner meta- tarsal tubercle elliptical, visible from above; 14) outer metatarsal tubercle low, round; 15) subarticular tubercles on foot large, round; 16) supernumerary tubercles on foot large, conical; 17) discs round; 18) dorsal skin shagreened; 19) dorsum uniform dark brown to grayish tan or dull green; 20) facial area colored like dorsum; bronze labial stripe in females; 21) flanks bluish black; 22) dorsal surfaces of limbs dark brown to grayish tan; 23) posterior surfaces of thighs bluish black; 24) venter grayish brown; 25) squamosal exostosed, in broad contact with maxillary; 26) temporal arcade incomplete; 27) prevomers abutted medially; 28) transverse proc- esses on eighth presacral vertebra strongly inclined anteriorly. Gastrotheca psychrophila differs from all other Andean Gastro- theca, except G. plumbea, in having a dark venter. It also is like G. plumbea in lacking dorsal blotches, but G. psychrophila differs from G. plumbea in lacking a dorsolateral light stripe and in having a darker venter, usually a primarily brown dorsum (green in G. plumbea) and a copper iris (green in G. plumbea). Variation.—Individuals of this species are capable of consider- able metachrosis. When frogs were found in bromeliads they were dark brownish black above and below; the flanks and posterior sur- faces of the thighs were dark bluish black. Later the dorsum changed to copper or bronze-tan with or without diffuse pale green blotches or streaks. The flanks are orange-brown or dark brown. The axilla, groin, and hidden surfaces of the limbs are bluish gray or bluish purple. The lips are dull bronze, and the iris is copper with black flecks. Distribution—This species is known only from the ridge be- tween Loja and Zamora, Ecuador (Fig. 3). Most individuals have been found on the upper eastern slope between 2750 m and the crest at 2850 m. Remarks.—The Loja-Zamora ridge is exceedingly wet; cold winds blow from the east. The vegetation near the crest consists of grasses and dense bushes. Large bromeliads are abundant on the ground and in the bushes. Adult G. psychrophila were found in the bromeliads and under rocks. Tadpoles were obtained from a grassy pond on the west side of the ridge. The tadpoles are black and have two upper and three lower rows of denticles. Etymology—tThe specific name is from the Greek psychros, MARSUPIAL FROGS OF THE ANDES OF ECUADOR i meaning cold, and philos, meaning having an affinity for; the name is used in allusion to the climate at the type locality. Gastrotheca riobambae (Fowler) Hyla riobambae Fowler, 1913:157 [Holotype—ANSP 16161 from Riobamba, Provincia Chimborazo, Ecuador; S. N. Rhoads collector]. Hyla quitoe Fowler, 1913:159 [Holotype ANSP 18238 from Quito, Provincia Pinchincha, Ecuador; S. N. Rhoads collector]. Chorophilus olivaceus Andersson, 1945:85 [Holotype —NHRM 1965 from “Rio Napo, 400 m.” (= ? Banos, Provincia Tungurahua), Ecuador; William- Clarke MacIntyre collector]. Gastrotheca m[arsupiata] ecuadoriensis Vellard, 1957:43 [Nomen nudum]. Gastrotheca riobambae—Duellman and Fritts, 1972:11. Diagnosis —1) Body robust, 48.7 mm in males, 51.2 in females; 2) snout rounded in dorsal view and in profile; 3) canthus rounded; 4) loreal region shallowly concave; 5) tympanum round; 6) supra- tympanic fold weak, curved posteroventrally behind tympanum; 7) subarticular tubercles on hand large, round; 8) supernumerary tubercles on hand small, subconical; 9) palmar tubercle bifid; 10) fingers not webbed; 11) toes one-fourth webbed; 12) tarsal fold curved, extending full length of tarsus; 13) inner metatarsal tuber- cle elliptical, visible from above; 14) outer metatarsal tubercle ab- sent; 15) subarticular tubercles on foot large, subconical; 16) super- numerary tubercles on foot small, round; 17) discs round; 18) dor- sal skin shagreened, with scattered pustules, tubercular in tympanic region; 19) dorsum green or tan, usually with pair of large elongate dark spots; 20) facial area green or brown; canthal and _ labial stripes absent; 21) flanks green, blue, or tan with dark brown or black spots; 22) dorsal surfaces of limbs green or tan, usually with elongate dark mark on thigh and irregular blotches or transverse bars on shank; 23) posterior surfaces of thighs dark brown with cream flecks; 24) venter cream with brown or gray mottling on chest and belly; 25) squamosal exostosed in large individuals, in moderately broad contact with maxillary; 26) temporal arcade in- complete; 27) prevomers abutted or narrowly separated medially; 28) transverse processes on eighth presacral vertebra transverse or slightly inclined anteriorly. Gastrotheca riobambae is like G. cavia in having short legs, a short snout, and a narrow interorbital distance, but it differs from G. cavia by having paired longitudinal dorsal marks, large spots on the flanks, and no dorsolateral light stripe. The dorsal color pat- terns of G. lojana and monticola are similar to that of G. riobambae, but both G. lojana and G. monticola have reticulated or plain flanks and dorsolateral light stripes. Gastrotheca plumbea and psy- chrophila have no dorsal markings and uniformly colored flanks; the former has a dorsolateral light stripe. Variation—The preceding diagnosis is based principally on 18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY topotypic material. Considerable variation, especially in coloration, obtains in this species. Five aspects of coloration are worthy of discussion (Table 2): 1. Dorsal ground color: The dorsum is either green or brown, varying to grayish tan in some individuals. The proportions of green to brown individuals is highly variable in local samples. At Papallacta on the high Amazonian slopes of the Cordillera Orien- tal, all individuals are green. The same is true at Guaranda on the Pacific slopes of the Cordillera Occidental. Approximately one-half of the frogs from the Cuenca Basin are green. At Riobamba, in the upper reaches of the Rio Pastaza drainage, 97 percent of the frogs are brown; at Banos, lower in the Rio Pastaza valley, 89 percent are brown. 2. Dorsal markings: Most specimens have dark dorsal blotches. In green frogs these are usually darker green, but in some the blotches are brown. The same variation occurs in brown frogs, with the addition of green blotches that are dark brown peripherally and bordered or not by cream. The blotches usually are in the form of a pair of broad, irregular marks extending from the eyelid or occiput to the rump. In a few individuals the blotches are frag- mented into a row of spots; in others they are expanded so as to cover most of the dorsum. With the exception of the series from Riobamba, all large samples contain some individuals lacking dor- sal blotches. Plain individuals make up less than one-third of each sample, except that from Guaranda, in which one-half of the specimens lack blotches. Most Gastrotheca riobambae lack definite dorsolateral light stripes, but these stripes are present in some speci- mens from Guaranda and the Cuenca Basin. 3. Thigh coloration: In most samples the posterior surfaces of the thighs are brown, gray, or tan (frequently with a green suffu- sion) with black flecks or small spots. In specimens from Riobamba and Guano the posterior surfaces of the thighs are tan or green with small cream flecks. The thighs are uniform dull blue in specimens from Papallacta and Biblian, bluish green from Cuenca, and blue with black flecks from Mulalé and Guaranda. 4. Flank coloration: The flanks are tan, green, gray, or blue, usually with black or dark brown spots. In specimens from Papal- lacta and Biblian the flanks are uniform blue. 5. Ventral coloration: Specimens from Papallacta are uniform- ly gray below. In all other samples the belly is cream; the belly is marked with black, gray, or dark brown flecks, spots, blotches, or reticulations in all other samples, except those from the Cuenca Basin, in which the belly is uniformly cream. In all specimens the iris is deep bronze to copper with black reticulations; males in all samples have dark brown to gray vocal sacs. 19 MARSUPIAL FROGS OF THE ANDES OF ECUADOR ggigg StF Lc‘0S Gc‘0g SS‘ LP 9S‘6Fr LECH €9°E¢ LG‘ZG 6F0S oa 2 (uau) TAS Xe wedi) wIeBaiy) syooH Yorig SUOT}BNI}E1 Youlg PoE ela sjods yorlg eee ies ABI sjods yorq leurs soyoro]q Prd ATOR u9918-9n[ gq aaa bel SyooH Youyq ‘onig sjods yoriq ‘useis-on[g Soo HE Yortq ‘useisg sjods yor[q ‘use13-Aeiyy SyooHE Yourq ‘ong ond ua018-on[g ete SyooR Yourq ‘onyg SyOoH WIeeIO SUs01S8-Ue ], usaI4y sjods yorlq ‘use1s-Aein SyooHf Youyq ‘onyig_ ond sjods yorlq ‘useis-on[g Urle[d 10 syoop yoR[q ‘useis-uUMOIg sjods yorlq ‘Avis on[g syur[ ‘aDquIDgOl DIAYIOJSDX) UL UOTyeLIVA oTydeIs09y °*% AIAV], SYOOHf YORlG ‘use1s-ue T, sysiy, 10119}s0g 8°Gz v' 2600 m) and Ama- zonian slopes of the Cordillera Oriental (> 1800 m), and in inter- andean valleys (> 2300 m) as far south as the slopes of Cerro Tinajillas in Provincia Azuay. The species occurs at elevations of 3860 m in the Paramo de Apagua, 3960 m at Paso de Guamani, and 4135 m on Volcan Antisana. The species occurs in extreme southern Colombia, but specimens resembling G. riobambae from Bogota, Departamento Cundinamarca, and San Pedro, Departamento An- tioquia, apparently are not conspecific. Remarks.—Gastrotheca riobambae occurs in a variety of habi- tats ranging from wet montane meadows to dry rocky hillsides. The species frequents ruderal situations—drainage and _ irrigation ditches, Agave, and corn fields. On cloudy or rainy days individuals are active, and males commonly call by day. Despite low tempera- tures (as low as 2-4° C), adults are most active at night. Tadpoles develop in still water. In large ponds the tadpoles aggregate in shallow water, but upon the slightest disturbance, they rapidly flee to deep water. As noted in the preceding discussion of variation, some popula- tions currently assigned to Gastrotheca riobambae may be spe- cifically distinct. I suggest that biochemical and karyological in- vestigations might be fruitful approaches to the taxonomy of this complex. DISCUSSION Apparent evolutionary trends in the Andean marsupial frogs are confusing. Members of the Gastrotheca marsupiata group are the most terrestrial and live in what seem to be suboptimal environ- mental conditions—dry interandean valleys. On the other hand, members of the Gastrotheca argenteovirens group and some mem- bers of the Gastrotheca plumbea group are arboreal and live in what seem to be more optimal anuran environments—cloud forest and wet paramo. Within the Gastrotheca plumbea group, two species (plumbea and psychrophila) inhabit cool moist environments. Gastrotheca plumbea lives in arboreal and terrestrial bromeliads in cloud forest, and G. psychrophila inhabits terrestrial bromeliads in wind-swept subparamo (Fig. 5). The other species in the Gastrotheca plumbea group principally inhabit drier interandean valleys and Pacific slopes of the Andes. In these areas the frogs live in paramo, Agave, and cultivated fields; G. cavia inhabits bromeliads in scrubby sub- paramo (Fig. 6). Among the members of the Gastrotheca plumbea group, G. riobambae is most like members of the Gastrotheca marsupiata 22 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Fic. 5. Type locality of Gastrotheca psychrophila, ridge east of Loja Ecua- dor. Note terrestrial bromeliads in foreground. group in having a relatively narrow head, least developed lateral flanges on the frontoparietals, and relatively small hands. The color pattern of G. riobambae is similar to that of G. peruana, the north- ernmost species in the Gastrotheca marsupiata group. I consider the Gastrotheca plumbea group to have been derived from the Gastrotheca marsupiata group and G. riobambae to be the most primitive member of the Gastrotheca plumbea group. All other members of the group have more extensive cranial ossification and exostosis. The major phyletic line in the Gastrotheca plumbea group has dorsolateral light stripes and moderately long snouts. Two members (G. lojana and monticola) of this line retain the paired dorsal blotches of G. riobambae, whereas the dorsum is plain in G. plumbea, a species most like members of the Colombian Gastro- theca argenteovirens group. The nearly unicolor G. psychrophila and the flecked G. cavia, each apparently represent independent derivatives from a G. riobambae-like stock. Vuilleumier (1971) documented Pleistocene changes in the flora and avifauna in the Andes. Her summary of geological, climatic, and biogeographic evidence demonstrates two glaciations in the Ecuadorian Andes. During glacial periods snow line was lowered as much as 700 m, and temperatures were depressed 4-11° C. The patterns of speciation and distribution of the Gastrotheca plumbea group are compatible with Vuilleumier’s paleobiogeographic hy- MARSUPIAL FROGS OF THE ANDES OF ECUADOR 23 Fic. 6. Type locality of Gastrotheca cavia, Isla Pequena, Laguna Cuicocha, Provincia Imbabura, Ecuador. Note bromeliads in scrubby trees in middle of picture. pothesis. Gastrotheca lojana and monticola are relictual popula- tions of warm-dry interglacial periods now isolated in lower and drier areas than other members of the group. During climatic de- pression in glacial periods, populations were isolated in interandean basins and the outer slopes of the Andes; thus, G. riobambae, plumbea, and psychrophila differentiated in the intermontane val- leys, Pacific slopes, and Amazonian slopes, respectively. A_ rio- bambae-like stock apparently was isolated from more southern populations by uninhabitable environments in the Nudo de Mo- janda during a glacial period and differentiated into G. cavia. Con- ceivably, the differentiation of the six species occurred at the time of the first glacial period. If so, the differentiation within G. rio- bambae and southward migration of G. lojana and monticola into northern Peru could be coincidental with the second glaciation. This proposed speciation model is similar to that suggested by Montanucci (1973) for the Andean microteiid genus Pholidobolus, a group of lizards inhabiting the same areas as Gastrotheca. SUMMARY The marsupial frogs of the genus Gastrotheca inhabiting the Andes and interandean valleys of Ecuador form a_ phylogenetic unit (Gastrotheca plumbea group) that is intermediate between the more southern Gastrotheca marsupiata group and the more 24 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY northern Gastrotheca argenteovirens group. The Gastrotheca plumbea group is characterized by a supraorbital flange on the frontoparietals and extensive exostosis, but no co-ossification, of the cranial roofing bones. Apparently all species in the group have free-swimming tadpoles. The Gastrotheca plumbea group contains six species: G. lojana Parker, G. monticola Barbour and Noble, G. plumbea (Boulenger ) and G. riobambae (Fowler). In addition, two new species are named herein: G. cavia from Laguna Cuicocha, Provincia Im- babura, Ecuador, and G. psychrophila from the ridge east of Loja, Ecuador. Gastrotheca riobambae is highly variable; some popula- tions may represent distinct species. The Gastrotheca plumbea group seems to have been derived from the Gastrotheca marsupiata group, and G. riobambae prob- ably is the most primitive member of the group. Speciation within the group evidently occurred through isolation of populations due to climatic fluctuation during the Pleistocene. RESUMEN Las ranas marsupiales del género Gastrotheca que habitan los Andes y valles interandinos del Ecuador forman una unidad filo- genetica (el grupo Gastrotheca plumbea) que es un grupo inter- mediano entre el grupo Gastrotheca marsupiata del sur y el grupo Gastrotheca argenteovirens del norte. El grupo Gastrotheca plumbea se caracteriza por tener una protuberencia supraorbital en los frontoparietales y prominencias extensas pero no co-ossifica- cién de los huesos que forman la cubierta craneal. Parece que todas las especies del grupo tienen renacuajos acuaticos. El grupo Gastrotheca plumbea tiene seis especies: G. lojana Parker, G. monticola Barbour and Noble, G. plumbea (Boulenger), y G. riobambae (Fowler). Ademas dos especies nuevas son nomi- nadas aqui: G. cavia de la Laguna Cuicocha, Provincia Imbabura, Ecuador, y G. psychrophila de la cordillera al este de Loja, Ecua- dor. Gastrotheca riobambae es muy variable; algunas poblaciones pueden representar especies distintas. El grupo Gastrotheca plumbea parece dirivarse del grupo Gas- trotheca marsupiata, y probablemente G. riobambae es la especie mas primitiva de este grupo. La diferenciacién en el grupo ocurre evidentemente como resultado del aislamiento de poblaciones de- bido las fluctuaciones climaticas durante el Pleistoceno. SPECIMENS EXAMINED Gastrotheca cavia “Western Ecuador,” BMNH 1860.6.16.124-125. Imbabura: Hacienda San Nicolas, 2000 m, UMMZ 92269, 92278-9, 92289-98; Ibarra, 2300 m, BMNH MARSUPIAL FROGS OF THE ANDES OF ECUADOR 25 (898.4.28.156; locality? ); Laguna Cuicocha, 2890 m, KU 138216-20, 139136- 9, 139439 (tadpoles), 139440 (young), 143094, 143537 (tadpoles), 148530- 42, 148543-4 (skeletons), 148545-7 (tadpoles), 148548 (young). Gastrotheca lojana Loja: Celica, 2130 m, BMNH 1931.11.3.3-4; Loja, 2150 m, BMNH 1931. 2.12.10-13, 1933.6.24.18-44, 1935.11.3.26-32, 1947.2.31.6-18, KU 120673-4; 2km N Loja, 2100 m, KU 142846 (tadpoles); 5 km N Loja, 2150 m, 138235- 6, 138237 (skeleton); 2 km E Loja, 220 m, KU 120675; 9 km E Loja, 2660 m, KU 121387 (tadpoles); 2 km S Loja, CAS 93898; 3 km W Loja, 2150 m, KU 138233; 5.5 km W Loja, 2330 m, KU 142603-8, 148549-51; 10 km W Loja, 2500 m, KU 138234. Zamora-Chinchipe: Zamora, 1000 m, BMNH 1933.624.45 (locality? ). PERU: Amazonas: Chachapoyas, 2340 m, KU 138238-41; 24 km WSW Leimebamba, 3370 m, FSM 47216 (ID?). Cajamarca: Cajamarca, MJP 204. Piura: Ayabaca, MJP 702 (2). Gastrotheca monticola Azuay: Girdn, 2240-2500 m, KU 138401-3. Loja: Saraguro, 2500 m, KU 138404-9, 138410 (skeleton), 138769 (tadpoles), 141565, 142609-13, 142847 (tadpoles), 148563-8, 148569-70 (skeletons), 148571 (tadpoles). PERU: Cajamarca: Bellavista, BMNH 1947.2.22.47-8, 1947.2.25.77-8; Querocotilla, MCZ 5328-20. Piura: Huancabamba, AMNH 7551, MCZ 5290- 3, 5296-7, 5299-300, 5302, 5304-7, 5309, 5312-15, 5317, 5319, 5328-30, SMF 2677, UMMZ 55747. Gastrotheca plumbea Azuay: Molleturo, 2350 m, ZMB 30057. Carchi: Atal, near San Gabriel, UMMZ 83655. Cotopaxi: Pilal6, 2320 m, KU 132413-22, 132423 (skeleton), 142614. Pichincha: Intac, 1200 m, BMNH 1947.2.31.19. Gastrotheca psychrophila Loja: 10 km E Loja, 2570 m, KU 142855 (tadpoles). Zamora-Chinchipe: 13-15 km E. Loja, KU 120760-2, 141585 (skeleton), 141586, 141595, 142631- To Gastrotheca riobambae Province Unknown: No specific locality, MNHN 965, 6227-9 (8), 9595; Andes, BMNH_ 58.7.25.21, 58.7.25.23, 58.7.25.25, 58.7.25.27-8, 58.7.25.31-3; Western Ecuador, BMNH 60.6.16.17, 60.6.16.127. Azuay: Bestion, AMNEI 13967; Cuenca, 2540 m, CAS 85172, KU 120676-723, 129797-8, SMF 2669- 79, USNM 61757-60, USNM-JAP 2345, 2347-8, 2350; 6. km N Cuenca, AMNH 71588-600; 9 km N Cuenca, CAS 85339-40, 93884-94; 18 km N Cuenca, CAS-SU 21851; 4 km E Cuenca, 2540 m, KU 138587-613, 138622-3 (skeletons), 138773 (tadpoles); 8 km SW Cuenca, AMNH 71601-2; 8.8 km NW Cuenca, 2620 m, KU 141583-4, 141594 (tadpoles); 9 km S Cumbe, 3300 m, KU 132536 (tadpoles); 10 km S Cumbe, 3350 m, KU 132392; 28.6 km S Cumbe, 3190 m, KU 142853 (tadpoles); 0.8 km S Cutchil, 2535 m, RU 14158252 1 km S: Cutehily 2720 m) KU 141572: 3.5 km_S Cutchil, 2785 m, KU 141579-81; 8.5 km S Cutchil, KU 141577-8; Lago de Sarogucho, 20 km W Cuenca, CAS 94114; Laguna de Zurucuchu, 3200 m, KU 121388 (tad- poles); Narihuina, MNHN 06-283; Rio Matadero, 8 km E. Cuenca, CAS-SU 21845-6; Rio Matadero, 9 km E Cuenca, CAS 94217 (tadpoles), 94218-22, CAS-SU 21847-8; Rio Matadero, 12 km E Cuenca, KU 129779-96; Sinicay, 2560 m, AMNH 17451-7, 17459-63, 17465-8, 17552, 17567. Bolivar: Guaran- da, 2640 m, KU 132403-12, 132531 (tadpoles), 132540 (young); 27.3 km 26 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY E Guaranda, 3800 m, KU 142850 (tadpoles); 2.5 km S Guaranda, 2650 m, KU 142616-9, 142851 (tadpoles), 142852, 148573; Guaranda-Riobamba road at Chimborazo border, 3700 m, KU 132541, 132542 (young). Canar: 3 km S Azogues, 2500 m, KU 138624-7, 138774 (tadpoles); Biblian, 2620 m, KU 141570-1, 141573, 142620-4, 147113; 8 km NW Biblian, 3420 m, KU 132537 (tadpoles); Camar, NHMW 6476, 6480, 2.8 km S Cafnar, 3150 m, KU 141574- 6; km 94, Guayaquil-Cuenca railroad, CAS 93899-900. Carchi: El] Carmelo (E] Pun), 2750 m, USNM-JAP 4946-7; Quebrada de Piedras, 20 km S Tulcan, 3400 m, KU 118120 (tadpoles); Tulcan, 3000 m, KU 117978-9, 118119 (tad- poles). Chimborazo: No specific locality, BMNH 1932.10.2.86; Guamote, USNM 33863; 1 km S Guano, 2500 m, KU 132354-89, 148581, 148582-4 (skeletons), 148592; Hacienda Alao, 15 km SE Pungala, 3100 m, KU 132543 (tadpoles); Laguna de Colta, 15 km SE Riobamba, 3400 m, NSNM-JAP 1728, 1730-3; Riobamba, 2780 m, ANSP 16161, KU 120732, MNHN 02-62(2), 02-350(2); 10 km N Riobamba, 2730 m, KU 138547-73, 138574-6 (skeletons ); 15 km E Riobamba, 2600 m, KU 120725-31, 120758-9, 121389 (tadpoles); Rosario, NHMW 6485; San Juan, 3160 m, KU 142615; 10 km W San Juan, 3160 m, KU 120724; 20 km SW Santa Rosa, 3700 m, KU 132348-9; Urbina Railroad Station, 3609 m, KU 132350-3; Volcan Chimborazo, USNM 103268- 74. Cotopaxi: El Porvenir, 2 km W Campamento Mariscal Sucre, 3620 m, KU 124167 (tadpoles); Guilo, 8 km E Pilalé, 3500 m, KU 132538 (tadpoles); Laguna de Limpios, N base Volcd4n Cotopaxi, 3890 m, KU 122593; 4 km S Latacunga, USNM 164337; 6 km S, 7 km E Latacunga, 2750 m, KU 127082-4; Mulalé, 2980 m, KU 141566, 146263-4, 146749-61, 146762-3 (skeletons); Paramo de Apagua, 3860 m, KU 132390-1, 132535 (tadpoles); 24.3 km E Pilalé, 3750 m, KU 142848 (tadpoles); 11.3 km W Pujili, 3500 m, KU 141567; 38.3 km W Pujili, 3350 m, KU 142849 (tadpoles); Rio Pita, N base Cerro Ingaloma, 3780 m, KU 122594-9. Imbabura: Ibarra, 2300 m, AMNH 10569-71, BMNH 98.4.28.152-4, NHMW 6482-6; N slope Nudo de Mojanda, 3650 m, KU 132393; Nudo de Mojanda, 4 km S San Pablo, 3050 m, KU 132394; Otovalo, 2550 m, KU 68708, 138587-613, MNCN 333; Quebrada San Miguel, 1 km N Otovalo, 2560 m, KU 117980; Quiroga, 2500 m, KU 1138577-86, 148585-6 (tadpoles). Napo: Laguna Papallacta, 3350 m, KU 109169 (tadpoles); Papallacta, 3130 m, KU 143095-102, 143538-40 (tad- poles), 143541, 148574-7; Rio Napo, 400 m, MNRM 1965 (locality?); Santa Barbara, 2625 m, USNM-JAP 4479, 4507; 1 km NW Santa Barbara, 2625 m, USNM-JAP 4487, 4491-3; 1 km SW Santa Barbara, 2625 m, USNM-JAP 4567; Volcan Antisana, 4135 m, AMNH 20127. Pastaza: Mera, 1140 m, AMNH 52852 (locality?). Pichincha: Intac, 1200 m, BMNH 78.1.25.20, FMNH 3607, NHMW 6481 (6) (locality?); Llave Pongo, AMNH 20140; Machachi, 2950 m, SMF 2667-8, UMMZ 47216; Paso de Guamani, 20 km E Pifo, 3960 m, KU 111626, 112316-7 (tadpoles), 127081, 127134 (tadpoles); W slope Paso de Guamani, 3940 m, KU 109170 (tadpoles), 109334-5; Quito, 2840 m, AMNH 20438-41, 20447-50, 20471-90, 60631; ANSP 18235, CAS- SU 2274, 11436-7, KU 94403, 111613-25, 112313-5 (tadpoles), 148416-28, 148578-80, 148587-91 (tadpoles), 148593-9, NMCN 156 (2), 158 (6), MNHN 34 (2), 1662 (2), 4878 (4), USNM 57804, USNM-JAP 1570-2, 1574, 1576-7, 1579, 1584, 1586-7, 1593, 1595, 1620-4, 1666, 1669, 1686, 2248-50, 2254, 2487, 2506-8; UZM 1474, 1477-8, 14424-93; between Rio Arturo and Taldadas, NE Cayambe, 3450 m, CAS-SU 8281; Rio Chiche, Valle de los Chillos, 2535 m, KU 152147-8; Santo Domingo de los Colorados, 500 m, AMNH 20147 (locality?). Tungurahua: Ambato, 2700 m, KU 120733-40, 121390 (tadpoles), USNM 164302; Bafos, 1800 m, CAS-SU 5082, FMNH 28091-2, 173661-80, KU 99123, 99124-9 (skeletons), 99130- 84, UIMNH 65539-675, USNM-JAP 5834-6, 6010-12, 6014, 6019-20; Chambo Grande, 7.6 km SE Pelileo, 2340 m, KU 141568-9, 142625-6, 146261-2; 10 km W Cotald, 3300 m, KU 132400-2; 1 km W Juan Benigno Vela, 3080 m, MARSUPIAL FROGS OF THE ANDES OF ECUADOR 27 KU 132395-6; Llanganati, near Rio Jorge, 3000 m, CAS-SU 17426-7; 4 km N Mocha, 3140 m, KU 120757; 10 km SW Mocha, 3700 m, KU 120741-56; Pelileo, 2600 m, MNHN 03/211; 3 km SSW San Miguelito, 2620 m, KU 132399; 12 km SW Santa Rosa, 3400 m, KU 132397-8. COLOMBIA: Cundinamarca: Bogota, BMNH_ 1919.3.6.37 (locality? ). Narino: Cuaspud, TNHC 40564-5. LITERATURE CITED ANDERSSON, L. C. 1945. Batrachians from east Ecuador collected 1937, 1938 by Wm. Clarke-MacIntyre and Rolf Blomberg. Arkiv Zool., 37A (2):1-88. Barsour, T. and G. K. Nose 1920. Some amphibians from northwestern Peru, with a revision of the genera Phyllobates and Telmatobius. Bull. Mus. Comp. Zool., 63 :395-427. BOULENGER, G. A. 1882. Catalogue of the Batrachia Salientia s. Ecuadata in the collection of the British Museum, ed. 2 London, xvi ++ 503 pp. Cocuran, D. M. and C. J. Goin 1970. Frogs of Colombia. Bull. U.S. Natl. Mus., 288: xii + 655 pp. DUELLMAN, W. E. 1970. The hylid frogs of Middle America. Monog. Mus. Nat. Hist. Univ. Kansas, 1:xi + 753 pp. DuUELLMAN, W. E. and T. H. Fritts 1792. A taxonomic review of the southern Andean marsupial frogs ( Hy- lidae: Gastrotheca). Occas. Papers Mus. Nat. Hist. Univ. Kansas, 9:1-37. Fow.Ler, H. W. 1913. Amphibians and reptiles from Ecuador, Venezuela, and Yucatan. Proc. Acad. Nat. Sci. Philadelphia, 55:153-176. JAMESON, D. L. and S. PEQUEGNAT 1971. Estimation of relative viability and fecundity of color polymor- phisms in anurans. Evolution, 25:180-194. Levins, R. 1968. Evolution in changing environments. Princeton Univ. Press, Princeton, ix + 120 pp. MonTANNUuCI, R. R. 1973. Systematics and evolution of the Andean lizard genus Pholidobolus (Sauria: Teiidae). Misc. Publ. Mus. Nat. Hist. Univ. Kansas, 59:1-52. Parker, H. W. 1932. Some new or rare reptiles and amphibians from southern Ecuador. Ann. Mag. Nat. Hist., (10) 9:21-26. PETERS, J. A. 1955. Herpetological type localities in Ecuador. Rev. Ecuatoriana Ent. Parasit., 2 (3-4) :335-352. VELLARD, J. 1957. Estudios sobre batracios andinos IV. El genero Gastrotheca. Mem. Mus. Hist. Nat. Javier Prado, 5:1-47. VUILLEUMIER, B. S. 1971. Pleistocene changes in the fauna and flora of South America. Sci- ence, 173 (3999 ):771-780. UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly pub- lished in the above series are now published as Occasional Papers, Museum of Natural History. The Miscellaneous Publica- tions, Museum of Natural History, began with number 1 in 1946. Longer research papers are published in that series. Monographs of the Museum of Natural History were initiated in 1970. All manuscripts are subject to critical review by intra- and extra- mural specialists; final acceptance is at the discretion of the publications committee. Institutional libraries interested in exchanging publications may obtain the Occasional Papers and Miscellaneous Publica- tions by addressing the Exchange Librarian, University of Kan- sas Library, Lawrence, Kansas 66045. Individuals may pur- chase separate numbers of all series. Prices may be obtained upon request addressed to Publications Secretary, Museum of Natural History, University of Kansas, Lawrence, Kansas 66045. Editor: LinpA TRUEB Managing Editor: W1Lu1AM E.. DUELLMAN PRINTED BY UNIVERSITY OF KANSAS PRINTING SERVICE LAWRENCE, KANSAS c CALIFORNIA ACADEMY OF SCIENCES FEB - 6 1974 LIBRARY OCCASIONAL PAPERS of the MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas NUMBER 23, PAGES 1-40 JANUARY 25, 1974 | SPECIATION IN FROGS OF THE HYLA PARVICEPS GROUP IN THE UPPER AMAZON BASIN By WILLIAM E.. DUELLMAN! and MArTHA L. CrumMpP?” As our knowledge of the amphibians in the upper Amazon Basin in South America increases, it becomes evident that the richness of the frog fauna is due to the occurrence there of representatives of many species groups and in some cases to the occurrence of several sympatric species belonging to one group. The latter phenomenon is especially noticeable in the Hyla leucophyllata and Hyla parvi- ceps groups with five and three species, respectively. The present paper deals only with the latter group. The primary purposes of this paper are to present analyses and interpretations of the morphological features of adults and_tad- poles, mating calls, breeding behavior, and ecological relationships of the three species at Santa Cecilia in Amazonian Ecuador. How- ever, as in most Amazonian frogs, some taxonomic problems must be dealt with before the biological problems can be discussed ef- fectively. Thus, we also present here a systematic review of the entire Hyla parviceps group, as presently understood. Acknowledgments We are indebted to the following persons for the loan of speci- mens or for the provision of working space in their respective insti- tutions: Werner C. A. Bokermann, Alice G. C. Grandison, Konrad Klemmer, Jean Lescure, Alan E. Leviton, Hymen Marx, Charles W. Myers, Giinther Peters, the late James A. Peters, Douglas A. Ross- 1 Curator, Division of Herpetology, Museum of Natural History, The Uni- versity of Kansas, Lawrence, Kansas 66045. * Research Assistant, Division of Herpetology, Museum of Natural History, The University of Kansas, Lawrence, Kansas 66045. 2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY man, Dorothy Smith, and Charles F. Walker. The field studies on these frogs were carried out at Santa Cecilia, Ecuador, where facili- ties were provided by Ing. Ildefonso Munoz B., to whom we extend our gratitude. Many of the specimens from Ecuador were collected by our field associates, who worked with us for varying periods of time from 1967 to 1972; we especially thank W. C. A. Bokermann, Stephen R. Edwards, Thomas H. Fritts, John D. Lynch, John E. Simmons, Linda Trueb, James W. Waddick, and Charles F. Walker. We are grateful to Jan Caldwell for analyzing the recordings of the mating calls, Albert E. Fisher and John E. Simmons for recording data, and Linda Trueb for executing the illustrations of the tad- poles. Field work in Ecuador was partly supported by Watkins Mu- seum of Natural History Grants and the F. William Saul Fund of the Museum of Natural History, and a grant from the Graduate School, The University of Kansas; further support was received from the National Science Foundation (GB 29557) and the Organi- zation for Tropical Studies. Duellman’s study of specimens in European museums was made possible by the American Philo- sophical Society (Penrose Fund No. 5063). The final work on this project is part of a study on the herpetofauna of Santa Cecilia, sup- ported by the National Science Foundation (GB 35483). Materials and Methods The systematic aspects of this study are based on the examina- tion of 720 preserved specimens, 26 cleared and stained specimens, and 19 lots of tadpoles. Recordings were made on a Uher-4000 Re- corder and analyzed on a Vibralyzer (Kay Electric Company). All measurements of morphological characters and calls were taken in the manner described by Duellman (1970). Webbing formulae were determined in the manner described by Savage and Heyer (1967). Tadpoles were staged according to Gosner’s (1960) system. All specimens are referred to by the following abbreviations: AMNH American Museum of Natural History BMNH British Museum (Natural History ) CAS California Academy of Sciences CM Carnegie Museum FMNH Field Museum of Natural History KU University of Kansas Museum of Natural History Le Lescure-Guiana (Jean Lescure, Paris ) LSU Louisiana State University Museum of Zoology NHMB Naturhistorisches Museum Basel SMF Senckenbergische Museum Frankfurt UIMNH University of Illinois Museum of Natural History UMMZ University of Michigan Museum of Zoology FROGS OF THE HYLA PARVICEPS GROUP 3 USNM-GOV United States National Museum (Gustavo Orcés V. collection ) WCAB Werner C. A. Bokermann, S4o Paulo, Brasil ZMB Zoologisches Museum Berlin SYSTEMATICS The Hyla parviceps Group Definition—1) Great sexual dimorphism in size; snout-vent lengths to 25 mm in males, 32 mm in females; 2) snout short, blunt; 3) tympanum visible, but tympanic ring indistinct or absent; 4) hands and feet moderately webbed; 5) axillary membrane abbrevi- ated; 6) thoracic glands absent; 7) calcars, ulnar and tarsal folds absent; 8) males having single, median, subgular vocal sac and no nuptial excrescences; 9) dorsum tan or brown with dark brown markings, with or without dorsolateral light stripes; 10) pale, vertical suborbital bars present; 11) thighs marked with cream, yellow, or orange spots; 12) anal, ulnar, and tarsal stripes absent; 13) iris pale gray with red ring around pupil; 14) nasals small, abutting or slightly overlapping sphenethmoid; 15) tectum nasi and septum nasi perichondrally ossified and synosteotically united with sphenethmoid; 16) frontoparictal fontanelle covered, or nearly so, by frontoparietals; 17) squamosals articulating with prootics; 18) quadratojugal reduced to small spur, or absent, not articulating with maxillary; 19) prevomerine teeth on posteromedially directed dentigerous processes between choanae; 20) presacral vertebrae nonimbricate, lacking neural crests, having transverse processes de- creasing in length posteriorly (3,6 perpendicularly transverse; 4,5 inclined posteriorly; 7,8 inclined anteriorly ); 21) sacral diapophyses expanded 53-77°, with convex edges; 22) tadpoles having ovoid bodies and xiphicercal tails with moderately deep fins not extend- ing onto body; 23) larval mouths anteroventral with one row of large labial papillae laterally and ventrally, robust serrate beaks, and no more than one row of denticles; 24) mating call consisting of short, high-pitched notes, followed or not by shorter secondary notes; 25) diploid chromosome number 30. Content.—Six species: Hyla bokermanni Goin, 1960; H. brevi- frons new species; H. luteocellata Roux, 1927; H. microps Peters, 1872; H. parviceps Boulenger, 1882; H. suwbocularis Dunn, 1934. Distribution—The Amazon Basin, coastal lowlands and slopes of southeastern Brasil, northern South America from French Guiana to Colombia, and eastern Panama. Comment.—The Hyla parviceps group is one of four groups of small Neotropical frogs that share many of the characters noted above. The other groups are—the Hyla leucophyllata group with about six species in South America and one in Central America, the Hyla microcephala group with perhaps a dozen species in South and Central America, and Hyla minuta, a widespread South Amer- 4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY ican “species” (see Cochran and Goin, 1970, for taxonomic com- ments). The Hyla leucophyllata and microcephala groups were de- fined by Duellman (1970). Twelve of the 25 species in these four groups have a diploid number of 30 chromosomes (Duellman, 1970; Bogart, 1973); the chromosome numbers in the others are unknown. All known tad- poles (18 species) have xiphicercal tails and reduced mouth parts. In addition, the species in the four groups are consistent in char- acters 4, 7, 8, 18, 19, 20, and 22. Members of the Hyla parviceps group differ from all of the other three groups by having (numbers correspond to statements in preceding definition): 1) more pro- nounced sexual dimorphism in size; 2) shorter snout; 3) tympanic ring indistinct or absent; 5) much less extensive axillary mem- brane; 9) sexual dimorphism in width of dorsolateral stripes; 10) suborbital bars; 11) thighs patterned; 13) iris color; 15) more perichondral ossification in the tectum nasi and solum nasi; 17) squamosals articulating with prootics. Although tadpoles of all of the groups have reduced mouth parts, there are consistent differences among the groups. Members of the Hyla parviceps and minuta groups have anteroventral mouths; there is one row of papillae in parviceps and two in minuta. Tadpoles of the Hyla parviceps group have no, or one, row of denticles below the beaks; minuta has one row. Tadpoles of the Hyla leucophyllata and microcephala groups have terminal mouths lacking rows of denticles; labial papillae are present in the former, absent in the latter. The mating calls of members of the Hyla leucophyllata and microcephala groups, and H. bokermanni in the parviceps group consist of primary and secondary notes; the calls of Hyla minuta and other members of the Hyla parviceps group consist of series of short notes. Cochran (1955) and Bokermann (1964) included Hyla microps in the Hyla marmorata group, although Bokermann (1964) defined the group in a more strict sense to include only four species (Hyla acreana, marmorata, melanargyrea, and senicula). These frogs dif- fer from the groups discussed above by having more extensive webbing, well-developed ulnar and tarsal folds, less cranial ossifi- cation, and different kinds of mating calls. They are like the other groups in general larval and karyological features. Other named taxa might be associated with the Hyla parviceps group or one of the other three groups; however, these species are too poorly known at the present time to be assigned to any group. These include: Hyla schubarti Bokermann, 1963.—Rondénia, Brasil, and Ama- zonian lowlands of Pert. FROGS OF THE HYLA PARVICEPS GROUP 5 Hyla leali Bokermann, 1964.—Rondonia, Brasil, and Amazonian lowlands of Ecuador and Pert. Hyla grandisonae Goin, 1966.—Guyana. Hyla oliveae Cochran and Goin, 1970.—Leticia, Colombia. Hyla riveroi Cochran and Goin, 1970.—Amazonian Colombia, Ecuador and Pert. Duellman (1969) named Hyla carnifex from the Pacific slopes of the Andes of Ecuador and placed the species in the Hyla parvi- ceps group. Although there are superficial similarities in adult structure and coloration, tadpoles, and mating calls, Hyla carnifex seems to be related to Hyla columbiana and variabilis in Colombia and is now excluded from the Hyla parviceps group. ANALYSIS OF CHARACTERS Three categories of characters were analyzed in adult frogs— morphometric, structural, and coloration. In addition, characters of the mating calls and tadpoles were utilized. Morphometric Characters Seven measurements were taken on representative samples of the six species. From these, five proportions were calculated (Tables 1 and 2). Analyses of variance showed highly significant differences (P = .001) in both sexes among species for the follow- ing: snout-vent length, tibia length/snout-vent length, and head width/snout-vent length. Equally significant differences exist for head length/snout-vent length in females and for tympanum/eye in males. Significant differences (P = .01) were found for foot length/snout-vent length in males. The differences among species in foot length/snout-vent length and tympanum/eye in females, and in head length/snout-vent length in males were not significant. Statistical comparisons of the sexes of three species revealed significant differences in snout-vent length between males and fe- males of all three and in different proportions in different species (Table 1). Adult females of all six species are much larger than adult males; this is especially noticeable in Hyla parviceps, in which the smallest gravid females are larger than the largest males. Structural Characters The species comprising the Hyla parviceps group are alike in most structural features; those features that are constant, or nearly so, are listed in the definition of the group. The tongue is cordiform in all species, deeply notched posteriorly in H. microps, and shal- lowly notched in the others. Ulnar and tarsal tubercles are absent in all species, except females of H. parviceps and both sexes of H. OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 6 SN 800°0 T00°0 ¢00°0 T00°0 600°0 SN ¥00'0 SN ¢00°0 100°0 O61 0 SN T10°0 SN 600'0 T00°0 £00°0 SN 900°0 SN S00°0 T00°0 0960 S00 OTO0'0 SN 000 SN f00°0 SN L00°0 SN 900°0 T00°0 staat dS } GhOO c6e'0 GI0'0 8860 GI00 66¢0 Tc0'0 = FaF'0 sc0';0 }3=6.: 9050 0L6°0 CV'es 990'0__ €IFr'0 cGI00 €6¢0 vIOO 60£°0 8c0O0 8 Ocr0 €c00 vVé6r0 00f'T 8h ES st00 61°0 e100 [0&0 8100 SsIe0 tc00 SIVO0 0c0'0 eLyV0 060'T 89°TS ds x So[RW9,J c6G'0 -VE'0 SOU Oia Groe Saece O aChOcoG0 - a..." -°. oA /ummuedus J, ISO -92'0 COO .Selo'O, 290e'0. -SC808G0-2 *S = TTAS/TaPI pee ZE'0 -8Z'0 SOOO ~ rie: ssa) AS04060 4s ~ = 2a TAS/tISUe] peo 9F'0 -8S°0 SOOM PZIOW sear: VEO=8e0 & ES TAS/tS8uUI] 300. cc’0 -9F'0 POU (Ce0'0y SPEC. “SS Os-pr0"* ss = TAS/Su9T| PIGLET, T'9Z- 9'1Z OF1O, “0650 S0PL OR 20S SE0cer =e (IAS) Wisuey yu9A-jnoug 56 GZ ‘PP Gs sdantaiwd DINAH 99'0 -€€'0 L000, SSE) Ses Ob “yi0 c60 2 = Se ee eA/ummue duré GEO -L3'0 C000 —110'0. 8600: “s2e0 90% 9 1. 6 = 80-90% 9 2. ¢ < 80% 9 B. Ulnar and tarsal tubercles: 0. Absent 1. Present FROGS OF THE HYLA PARVICEPS GROUP 27 C. Suborbital bars: 0. Absent 1. One 2. Two D. Canthal stripes: 0. Absent 1. Present E. Rostral stripes: 0. Absent 1. Present F. Dorsolateral stripes: 0. Absent in both sexes 1. Absent in males; present in females 2. Present in both sexes G. Anterior thigh: 0. Unicolor 1. Small yellow spots 2. Large yellow spot H. Dorsal thigh: 0. Unicolor 1. Small yellow spots I. Posterior thigh: 0. Unicolor 1. Large orange spot J. Venter: 0. Plain 1. Patterned K. Denticles (larvae): 0. One lower row 1. Absent The distribution of character states within the Hyla parviceps group reveals variation from 6 to 11 derived characters (Table 8). The greatest number of primitive states occurs in H. microps, fol- lowed by H. parviceps and brevifrons; H. bokermanni, luteocellata, and subocularis have the fewest primitive states. A phylogenetic arrangement (Fig. 7) was constructed by a method proposed by Camin and Sokal (1965). The branching se- quence defines one group having few derived characters (H. mi- crops and parviceps) and another with many derived characters (H. luteocellata, bokermanni, and subocularis). Hyla brevifrons is intermediate between these two groups. Although reproductive data are incomplete, the existing infor- mation supports the proposed phylogeny. Hyla microps and_par- viceps deposit eggs in water, whereas the other species (unknown for H. luteocellata) deposit eggs on vegetation above water. Quan- titative data on reproduction of three species in Ecuador reveal that in most features H. brevifrons is intermediate between H. boker- manni and parviceps. If the reproductive mode of H. luteocellata is the same as that for H. bokermanni and subocularis, a phylo- genetic construct based on reproductive characters would approxi- mate the arrangement based on morphological features. 28 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY microps (5) parviceps (6) brevifrons (6) luteocellata (7) < — subocularis (8) >< — bokermanni (8) G) ~ CO le ve: v1 nm G) 1) < A2BC!J C°E HK Me Fic. 7. Most parsimonious phylogenetic arrangement of species in Hyla parviceps group, based on 26 states of 11 characters. Letters indicate shifts from primitive to derived character states; superscripts indicate direction of change or degree of change in those characters represented by more than two states; X — change in dorsal pattern not accompanied by shifts in any one of the 11 characters; numbers in parentheses are numbers of evolutionary steps in each line. See Table 8 and text for character states and explanation. FROGS OF THE HYLA PARVICEPS GROUP 29 TABLE 8.—Character states of 11 characters in species in the Hyla parviceps group. (0 = primitive state; 1 and 2 = derived states; see text for list of characters and explanation. ) 2 S 2 : z Ss Z : s = 3 S S iF Character S 5 S 5 5 3 is) = = S 8 = As) ce) eS ~ Q DB on on cn en on on A 1 0 ] y) 2 ] B 0 0 0 i! G5" 0 C 2 2 2 il 1 0 D il 1 0 0 0 1 E 1 1 1 0 0 1 F 2, ! ny) 0 0 2 € yw 0 2 0 i 2 H 1 1 it 0 0 1 I 0 0 0 ] 0 0 J 0 0 0 ii ] 0 K 1 J ] 0 ] I Total gt 7 10 6 6.5 iG * Females only. The presence of a row of denticles in the tadpoles of Hyla microps is the only larval character that can be determined as primitive in the group. Otherwise, the known tadpoles differ from one another principally in coloration. We have no bases for deduc- ing evolutionary change in the coloration. We have incomplete data on mating calls, but the calls of H. parviceps and brevifrons consist of series of uniform notes, whereas H. bokermanni produces primary and secondary notes. Presumably the complex call of H. bokermanni is derived from the simple call of the others, a con- clusion compatible with the phylogenies deduced from morpho- logical and reproductive data. Zoogeography Recent works on distributional patterns in the tropical lowlands of South America emphasize Quarternary climatic fluctuations and the existence of forest refugia during times when climatic condi- tions were drier than present (Miiller, 1968; Haffer, 1969; Van- zolini and Williams, 1970). Miller and Schmithusen (1970) and Vuilleumier (1971) summarized Pleistocene climatic changes and interglacial sea transgressions in South America. Haffer (1969) provided evidence from avian distributions and differentiation for six forest refugia in Amazonian South America. Vanzolini and Wil- liams (1970) proposed four core areas for the differentiation of 30 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY populations of Anolis chrysolepis and hypothesized an evolutionary model of three expansions and two contractions of forest environ- ments resulting in isolation, differentiation, and introgression of Anolis chrysolepis. Accepting the preceding phylogenetic arrangement as a reason- able approximation of the actual evolutionary sequences of mor- phological characters in the Hyla parviceps group, we can super- impose the cladistics of the frogs on the biogeographical patterns in the lowland tropics. Our paleogeographic evolutionary model of the Hyla parviceps group is predicated on cyclic alternation of wetter and drier climates in the Pleistocene and Holocene, resulting in expansion and contraction of lowland tropical forests, and the existence of forest refugia during the drier periods (see Moreau, 1966, for paleogeographic evidence from Africa; Vanzolini and Williams, 1970, for a survey of the literature documenting changes in South America; and Van der Hammen and Gonzalez, 1960, for dating of palynological data). Duellman (1972) and Heyer (1973) suggested probable evolutionary histories of groups of South Amer- ican hylid and leptodactylid frogs with respect to vicissitudes of Quarternary climates. The evolutionary model is outlined, as follows: 1. The prototype of the Hyla parviceps group was widespread in Amazonia during a wetter period. This prototype had the primi- tive states of the characters listed in the preceding section, aquatic eggs, and a simple mating call. 2. Climatic desiccation resulted in contraction of forests and isolation of two populations: 1) the H. microps-parviceps stock in an Amazonian refugium; 2) the H. brevifrons-bokermanni-luteocel- lata-subocularis stock in an upper Amazonian refugium (Napo or Peruvian refugia of Haffer). The former stock developed more sexual dimorphism in size, ulnar and tarsal tubercles in females, one suborbital bar, and gray flecks on the venter. The second stock developed two suborbital bars, rostral stripes, yellow spots on the dorsal surfaces of the thighs; retained moderate sexual dimorphism in size; and lost the larval denticles. 3. During the subsequent pluvial period both stocks spread through the Amazon Basin; the H. microps-parviceps group ex- panded into eastern and southeastern Brasil, and the other stock invaded the coastal lowlands of northern South America and east- erm Panama. 4. Climatic desiccation resulted in the isolation of two popula- tions of the H. parviceps-microps group (H. microps in the Serra do Mar refugium of Miiller in southeastern Brasil and H. parviceps in an Amazonian refugium). The other stock was separated into four isolates. The prototypes of H. bokermanni and H. brevifrons were isolated in separate refugia (probably Haffers Napo and FROGS OF THE HYLA PARVICEPS GROUP 31 bokermann brevifrons /uteocellata microps parviceps subocularis bokermanni, brevifrons bokermanni, parviceps brevifrons, parviceps bokermanni, brevifrons, parviceps Fic. 8. Distribution of species in the Hyla parviceps group. Peruvian refugia); H. luteocellata was isolated in the Cordillera de la Costa in northern Venezuela (a core area proposed by Van- zolini and Williams, 1970), and H. subocularis was restricted to one or more of the small refugia in northwestern Colombia (Chocéd, Nechi, or Catatumbo refugia of Haffer, 1969). 5. The development of more equable climates resulted in the expansion of the Amazonian forests and of the ranges of the six populations of frogs. However, due to the dry caatinga and cerrado of eastern Brasil, H. microps remained restricted to southeastern Brasil. Hyla luteocellata expanded its range eastward into the Guianas, and H. subocularis moved into Panama. In the Amazon Basin, the ranges of three species (H. bokermanni, brevifrons, and parviceps) expanded from their respective refugia and became sympatric in the upper part of the basin (Fig. 8). EVOLUTION IN THE SYMPATRIC SPECIES If we accept the proposed phylogenetic arrangement and the suggested paleogeographic model, we are forced to hypothesize that the three species now occurring in sympatry in the upper Amazon Basin evolved through geographic isolation at different times (H. parviceps versus H. brevifrons and bokermanni) and in 32 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY different places (H. brevifrons and bokermanni). Presumably, genetic differences that evolved when the populations were isolated have been reinforced through selection in sympatric populations. Due to lack of data on any one of the species where it occurs in the absence of the other two, we are unable to determine if there is any character displacement. According to Van der Hammen and Gonzalez (1960), the cli- mate in northern South America has been becoming progressively wetter for the past 4000 years, thus, it is possible that the three species in the upper Amazon Basin have been sympatric for less than 4000 years, before which time they were isolated in forest refugia (Haffer, 1969). We are concerned here with those mechanisms which are operant in maintaining the specificity of the sympatric populations. Following Miiller’s (1942) classification of isolating mechanisms, as modified by Fouquette (1960), we recognize three major cate- gories of isolating mechanisms—anti-mating, courtship, and post- mating. The last category includes gametic incompatibility, hybrid inviability, and hybrid sterility; we have no information on these factors and no evidence of hybridization, so they are not considered here. Of the five kinds of anti-mating mechanisms, one (geographic isolation) has already been accounted for by the hypothesized differentiation through geographic isolation. Data on the others can be summarized, as follows: Habitat isolation—Relative to breeding activities, H. parviceps is a habitat generalist, usually present at sites where the other species breed. Hyla bokermanni and brevifrons usually breed at different sites. Seasonal isolation—Synchronous breeding activity is common, but at least H. bokermanni and H. parviceps have some different periods of intense breeding activity. Temporal isolation.—All three species are nocturnal breeders. Climatic isolation.—All three species have a positive response to periods of heavy rainfall. Fouquette (1960) recognized only two categories of courtship isolating mechanisms, whereas we recognize three; oviposition sites are considered: Auditory isolation—The mating calls of three species are dis- tinctly different in note repetition rate and duration of notes; fur- thermore, the call of H. brevifrons is distinctive in pulse rate, that of H. parviceps in dominant frequency, and that of H. bokermanni in the presence of secondary notes (Table 6). Mechanical isolation —Because the females of the three species have nearly identical snout-vent lengths, difference in size might be an effective mechanical isolating mechanism only in H. parvi- FROGS OF THE HYLA PARVICEPS GROUP 33 ceps, the males of which are much smaller than males of the other species. Ovipositional isolation—Hyla parviceps deposits its eggs in water and the other species place their eggs on vegetation over water. The apparent behavioral and probable physiological differ- ences of females and eggs, respectively, are potentially important factors in isolation. The three species broadly overlap in most categories of poten- tial isolating mechanisms, but there are distinct differences in two categories, ethological and ovipositional. Fouquette (1960) found that the mating call was the primary isolating mechanism operant in three sympatric species of Hyla in Panama. Duellman (1967) reached the same conclusion in a study of a breeding community of ten species of hylids in Costa Rica. Martof and Thompson (1958) and Littlejohn and Michaud (1959) provided experimental evidence demonstrating that females respond positively to the calls of the males and that females can discriminate between calls of males of their own and other species. Littlejohn (1964, 1965) pre- sented evidence on evolution of differences in mating calls through geographic isolation and reinforcement of these differences in sym- patric populations. We conclude that at least some of the differ- ences in mating calls in the three species of the Hyla parviceps group evolved at times when the populations were geographically isolated from one another; furthermore, these differences may have been reinforced through selection upon subsequent geographical sympatry. The differences in oviposition sites involve a modification in reproductive strategy from the presumed primitive strategy of placing eggs in water (H. parviceps) to a derived strategy of plac- ing eggs on vegetation over water (H. bokermanni and brevifrons). Salthe and Duellman (1973) showed that within a given anuran reproductive mode there exist positive correlations between ovum size, clutch size and female snout-vent length, and negative correla- tions between clutch size and ovum size and between ovum size and rate of development. Among the three species under con- sideration, Hyla parviceps with aquatic eggs has the largest clutch size, smallest ova, and shortest duration of development (Table 7). Females of the other species are no larger than H. parviceps; thus, as concluded by Salthe and Duellman (1973), the achievement of a larger ovum in H. bokermanni and brevifrons, without an in- crease in body size of the females, was by selection for a change in the site of intraoval development, in this case vegetation over water. Larger eggs produce larger hatchlings better adapted to make the transition from leaf to water. Because the reproductive modes are different in the two lineages in the group, it is most reasonable to assume that this differentiation took place only once and that by 34 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY the time the three species became sympatric in the upper Amazon Basin the differences in reproductive strategy were already in existence. However, the differences are important aspects of re- source partitioning in synchronously breeding frogs within one community. Salthe and Duellman (1973) noted that in small anurans: it seems to be more advantageous to maintain a certain minimal number of eggs in the clutch than it is to maintain the size of the hatchling. Presumably, clutch sizes are already at, or near, a lower limit allowable for the production of an adequate amount of genetic variability per unit time. Thus, in general, selection in small species seems to function by maintaining minimal clutch sizes and maximizing the sizes for relatively tiny hatchlings.” Our re- productive data indicate that despite differences in mode of re- production and ovum size, the hatchlings are about the same size in H. bokermanni and parviceps, but larger in H. brevifrons, which has the smallest clutches (Table 7). The problem of production of an adequate amount of genetic variability, as related to fecundity, per unit time perhaps is an important factor in the great amount of sexual dimorphism in size in the Hyla parviceps group. At Santa Cecilia, the three species in the group occur in swamps inhabited by many other species, of which 17 Hyla are known to breed in the same ponds or swamps as do members of the Hyla parviceps group. All of these are larger than the species in the Hyla parviceps group. Selection might favor reduction in size in response to competition for food, calling sites, and oviposition sites. However, reduction in size of the fe- males is more restricted than in males due to the limitations im- posed by egg-carrying capacities. Schoener (1967, 1969, 1970) documented evidence for sexual selection in size of West Indian Anolis, noting that extremes of sexual dimorphism in size are reached on islands inhabited by only one species of Anolis; on these islands the sexes exist ecologically as two species using different resource states, such as perches and food. However, in the Hyla parviceps group, the presence of many other species utilizing a variety of resource states negates the hypothesis of intraspecific se- lection in size solely for resource partitioning and suggests that the reproductive necessities of the females balance selection for small size, which is evident in males. SUMMARY Among the small Neotropical Hyla having a diploid number of 30 chromosomes and reduced mouth parts in the tadpoles, one group of six species is characterized by great sexual dimorphism in size, more cranial ossification, and aspects of coloration (patterned thighs, suborbital bars, and iris color). Six species are included in FROGS OF THE HYLA PARVICEPS GROUP 35 the group: H. bokermanni Goin, H. brevifrons new species, H. luteocellata Roux, H. microps Peters, H. parviceps Boulenger, and H. subocularis Dunn. Hyla rondoniae Bokermann, 1963, is placed in the synonymy of Hyla bokermanni Goin, 1960. The group is wide- spread in the Amazon Basin, southeastern Brasil, northern lowlands of South America and eastern Panama. Although interspecific differences exist in size, proportions, amount of webbing, and tubercles, the principal taxonomic char- acters are in the coloration. These include the number of pale suborbital bars, presence or absence of canthal and rostral stripes, presence and width of dorsolateral stripes, and pattern on the dor- sum, venter, and especially the thighs. All of the tadpoles have one row of labial papillae. Tadpoles of H. microps have one row of denticles; denticles are absent in the others. The known tadpoles are brightly colored and have different patterns of transverse bars or longitudinal stripes. The mating calls consist of series of short, high-pitched notes; H. bokermanni produces primary notes fol- lowed by shorter secondary notes. It is suggested that the species arose through isolation in forest refugia in the tropical lowlands during dry periods in the Pleisto- cene. The proposed phylogeny indicates that the two most primi- tive species now occupy different regions—H. microps in southeast- ern Brasil and H. parviceps in the upper Amazon Basin. The three most advanced species also have allopatric distributions—H. boker- manni in the upper Amazon Basin, H. luteocellata in coastal north- ern South America, and H. subocularis in eastern Panama and northwestern Colombia. Three species (H. bokermanni, brevifrons, and parviceps) occur at Santa Cecilia in Amazonian Ecuador. Intensive field studies there continuously for 13 months revealed slight differences in habitat and seasonal activity. One species (H. parviceps) lays eggs in the water, whereas the other two place eggs on vegetation over the water. Clutch size is larger and ovum size is smaller in H. parviceps than the others. It is suggested that differences in mating calls and oviposition sites are the primary isolating mechanisms operating in the sym- patric species. Furthermore, it is suggested that natural selection for small size in comparison with many sympatric congeners is balanced in females by the necessity for maintaining an adequate number of eggs to insure genetic variability in the population. In the absence of such pressure in the males, they have become much smaller than the females. RESUMEN Entre las pequenas ranas neotropicales del genero Hyla con un numero diploide de 30 cromosomas y renacuajos con bocas de 36 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY estructuras pequefias, un grupo de seis especies se caracteriza por el gran dimorfismo sexual de su tamafo, mayor osificacion craneal, y al- gunos aspectos de su coloracion (disenos en los muslos, barras sub- orbitales, y color del iris). Las seis especies incluidas en el grupo son: H. bokermanni Goin, H. brevifrons nueva especie, H. luteo- cellata Roux, H. microps Peters, H. parviceps Boulenger, y H. sub- ocularis Dunn. Hyla rondoniae Bokermann, 1963, se pone en el sinonomia de Hyla bokermanni Goin, 1960. Este grupo se encuen- tra difundido en Ila Cuenca Amazoénica, el sureste brasileno, las tierras bajas del norte de Surameérica, y el este panameno. Aun cuando existen diferencias interespecificas de tamano, pro- porciones, extensidn de las membranas interdigitales, y tubérculos, las principales caracteristicas taxondmicas giran alrededor de la coloracién. Esto incluye el numero de barras suborbitales claras, la presencia o auséncia de las rayas cantales y rostrales, la presencia y ancho de las rayas dorsolaterales, y el disefio dorsal, ventral, y especialmente de los muslos. Todos los renacuajos tienen una fila de papilas labiales. Los renacuajos de H. microps tienen una fila de denticulos que no existe en los otros. Los renacuajos que se conocen son de coloracién luminosa y tienen distintos disefios de barras transversales y rayas longitudinales. La llamada de reclamo consiste de series de notas cortas y altas; Hyla bokermanni produce notas primarias sequidas de notas secundarias mas cortas. Se sugiere que estas especies surgieron como resultado de su aislamiento en refugios en la selva de la tierra tropical baja durante los periodos secos del Pleistoceno. La filogenia propuesta sugiere que las dos especies mas primitivas ocupan ahora regiones dis- tintas—H. microps ocupa el sureste brasilefio y H. parviceps Ja parte superior de la Cuenca Amazoénica. Las tres especies mas evolucionadas tambien tienen distribuciones alopatricas—H. boker- manni en la parte superior de Ja Cuenca Amazonica, H. luteocel- lata en la costa norte de Suramérica, y H. subocularis en el este panameno y noroeste colombiano. Tres especies (H. bokermanni, brevifrons, y parviceps) se en- cuentran en Santa Cecilia en la Amazonia Ecuatoriana. Estudios continuos e intensivos de campo durante un periodo de 13 meses revelan pequenias diferencias en su “habitat” y en la actividad esta- cional. Una de las especies (H. parviceps) pone sus huevos en el agua, mientras que las otras dos los ponen en la vegetacioén sobre el agua. En H. parviceps el nimero de huevos es mayor y el tamano de los huevos es menor que en las otras especies. Se sugiere que diferencias en las Ilamadas de reclamo y los lugares de oviposicién son los mecanismos principales de aislami- ento en las especies simpatridas. Se sugiere ademas que la selec- cién natural a favor de un tamafio pequefio en estas especies en compariacion con muchos otros congéneres simpatridos esta con- FROGS OF THE HYLA PARVICEPS GROUP 37 trarestada en las hembras por la necessidad de mantener un nimero adecuado de huevos con lo que asegura la variabilidad genética de la poblacién. En Ja auséncia de esta presién, los machos se han mantenido mas pequenos. SPECIMENS EXAMINED Hyla bokermanni BRASIL: Acre: Cruzeiro do Sul, WCAB 12148. Rondénia: Rondonia, WCAB 7845. ECUADOR: Napo: Limon Cocha, UIMNH 64799-800, 90069; Santa Cecilia, KU 104432-3 (skeletons), 105118-9, 105121-2, 105125-6, 105131, 105189, 109355-7, 109452, 109468, 123051-8, 123060, 123062-71, 123089-91, 124193 (tadpoles), 126362, 126541-5, 126547, 126552-5, 143131, 143188, 146288-95, 146414, 146419, 146800-1 (tadpoles), 150026-51, 152290-1 (tad- poles), 152419-28, 152535, 152750-3 (skeletons), UMMZ 129325 (4). Pastaza: 3 km S Puyo, KU 127087. Hyla brevifrons BRASIL: Pard: IPEAN, 5 km E Belém, KU 127846. COLOMBIA: Putumayo: Santa Rosa de Sucumbios, AMNH 88068-80. ECUADOR: Napo: Lago Agrio, 126556-8; Puerto Libre, KU 123092-4; Santa Cecilia, KU 105063, 105107-11, 105120, 105127-30, 105132-4, 105190, 107003-4, 109449-51, 109453-67, 111786-802, 112341 (eggs), 123049-50, 123059, 123072-88, 125899-900 (tadpoles), 126342-61, 126363-71, 126546, 126548-51, 136298, 143189, 146281-7, 146413, 146415-8, 150052-74, 152429- 32, 152536-7 (tadpoles), 152754-8 (skeletons), UMMZ 129324 (2). Pastaza: Puyo, CAS 85142. FRENCH GUIANA: Inini: Crique Eleupocigne, Upper River Oyapak, LG 1119, 1308, 1311-12. PERU: Loreto: Pampa Hermosa, Rio Cushabatay, AMNH 42358; Rio Utoquinia-Rio Tapiche, AMNH 43576; Tacsha Huachiyacu, Rio Morona, AMNH 43045; Yurimaguas, Rio Huallaga, BMNH 84.2.18.53. Hyla luteocellata VENEZUELA: Aragua: Cumboto, UMMZ 113890 (2); Monte Oscuro, BMNH 1968.73-4, KU 125861-5, UMMZ 130085-6. Hyla microps BRASIL: Parand: Curitiba, UMMZ 104118 (2), 104129 (6), 104130, 104160. Rio de Janeiro: Barro Branco, SMF 41217; Novo Friburgo, ZMB 7472; Teresdpolis, BMNH 1947.2.14.10. Sdo Paulo: Campo Grande, Santo Andre, KU 74266-8, 74269 (skeleton), 92073-7, 92078 (skeleton); Campos de Jordao, KU 112398 (tadpoles); 10 km W Casa Grande, KU 129838. Santa Catarina: UMMZ 58515; Humboldt, AMNH 15573-82. Hyla parviceps COLOMBIA: Putumayo: Santa Rosa de Sucumbios, AMNH 88081-2. ECUADOR: Napo: Bermejo No. 4 (well site), Sierra Umbaqui, KU 122963; Cuyabeno, UIMNH 54134-6, 54172, 59625-8, 90108-9; Dureno, KU 105113-4; Lago Agrio, KU 126480; Limén Cocha, KU 99223-8, 99229 (skele- ton), UIMNH 64805, 90076; Santa Cecilia, KU 104428-31 (skeletons), 105046-62, 105064-106, 105112, 107048-9, 109433-46; 11764-84, 112339-40 (tadpoles), 122925-62, 123061, 126462-79, 143167-73, 146307-11, 146790-1 (tadpoles), 150309-56, 152303-4 (tadpoles), 152547 (tadpoles), 152759-63 (skeletons), UMMZ 129279 (9); Tena, UIMNH 59628, 90059-64; UMMZ 123903. Pastaza: Abitagua, UMMZ 90416-7; Chontoa, KU 120910-13, 38 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 121407; Mera, KU 120909; 3 km S Puyo, KU 127086, 127090; 25 km NNE Puyo, KU 146792 (tadpoles); Rio Conambo, mouth of Rio Shione, USNM- GOV 7143; Rio Oglan, upper Rio Curaray, USNM-GOV 9526; Sarayacu, BMNH 1947.2.13.51. Tungurahua: Llanganate area, FMNH 23522-3; Mirador, BMNH 99.10.30.49; Rio Negro, KU 120913-14, 121408. PERU: Cuzco: Pilcopata, KU 139228. Loreto: Balta, Rio Curanja, LSU 25719-57; Iquitos, AMNH 42086; headwaters Rio Utoquinia, AMNH 43314. Hyla subocularis COLOMBIA: Cordoba: Rio Manso, CM 50447-50. PANAMA: Darién: Laguna, KU 77348-59, 77620; Rio Chucunaque, AMNH 51777; Rio Chucunaque at first creek above Rio Tuquesa, AMNH 41117; Rio Ucurganti, 7 km above mouth, KU 116785, 116784 (tadpoles); Tacarcuna, KU 77360-403, 77621, 77688-90 (skeletons), UMMZ 124780. LITERATURE CITED BOKERMANN, W. C. A. 1963a. Duas novas espécies de Hyla de Rondénia, Brasil (Amphibia, Salientia). Rev. Brasil. Biol., 23:247-250. 1963b. Girinos de anfibios Brasileiros—1. (Amphibia,—Salientia). An. Acad. Brasil. Cien., 35:465-474. 1964. Notes on tree frogs of the Hyla marmorata group with description of a new species (Amphibia, Hylidae) Senckenberg Biol., 45:243- 254. 1966. Lista anotada das localidades tipo de anfibios Brasileiros. Impres. Serv. Documentacao, S40 Paulo, 183 pp. Boearr, J.P: 1973. Evolution of anuran karyotypes. in Vial, J. L. (ed.) Evolutionary biology of the anurans: contemporary research on major problems. Univ. Missouri Press, Columbia: 337-349. BOULENGER, G. A. 1882. Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British Museum, ed. 2, London, xvi+503 pp. 1920. Descriptions of two new frogs from Brasil. Ann. Mag. Nat. Hist., (9)5:122-124. Camin, J. H. and R. R. Soka 1965. A method for deducing branching sequences in phylogeny. Evo- lution, 19:311-326. Cocuran, D. M. 1955. Frogs of southeastern Brazil. Bull. U.S. Natl. Mus., 206:xvi+423 pp. Cocuran, D. M. and C. J. Goin 1970. Frogs of Colombia. Ibid., 288:xii+-655 pp. CoLwELL, R. K. and D. J. FuruyMa 1971. On the measurement of niche breadth and overlap. Ecology, 52: 567-586. DUELLMAN, W. E. 1967. Courtship isolating mechanisms in Costa Rican hylid frogs. Her- petologica, 23:169-183. 1969. A new species of frog in the Hyla parviceps group from Ecuador. Ibid., 25:241-247. 1970. The hylid frogs of Middle America. Monog. Mus. Nat. Hist. Univ. Kansas, 1:xi+753 pp. 1972. South American frogs of the Hyla rostrata group (Amphibia, Anura, Hylidae). Zool. Mededel., 47:177-192. FROGS OF THE HYLA PARVICEPS GROUP 39 Dunn, E. R. 1934. Two new frogs from Darien. Amer. Mus. Novitates, 747:1-2. FouQuETTE, M. J., JR. 1960. Isolating mechanisms in three sympatric treefrogs in the canal zone. Evolution, 14:484-497. Gorn, C. J. 1960. Description of a new frog of the genus Hyla from northwestern Brazil. Ann. Mag. Nat. Hist., 13(2):721-724. Gosner, K. L. 1960. A simplified table for staging anuran embryos and larvae with notes on identification. Herpetologica, 16:183-190. HAFFER, J. 1969. Speciation in Amazonian forest birds. Science, 165:131-137. Hever, W. R. 1973. Systematics of the marmoratus group of the frog genus Leptodac- tylus (Amphibia, Leptodactylidae ). Contri. Sci. Los Angeles Co. Mus., 251:1-50. KuucE, A. G. and J. S. Farris 1969. Quantitative phyletics and the evolution of anurans. Syst. Zool., 18:1-32. LITTLEJOHN, M. J. 1964. Geographic isolation and mating call differentiation in Crinia sig- nifera. Evolution, 18:262-266. 1965. Premating isolation in the Hyla ewingi complex (Anura: Hy- lidae). Ibid., 19:234-243. LITTLEJOHN, M. J. and T. C. MicHaup 1959. Mating call discrimination by females of Strecker’s chorus frog (Pseudacris streckeri). Texas Jour. Sci., 11:86-92. ez. 1B: 1973. Brazilian species of Hyla. Univ. Texas Press, Austin, xix-+260 pp. MartTor, B. S. and E. F. THompson 1958. Reproductive behavior of the chorus frog Pseudacris nigrita. Be- haviour, 13:243-258. ME In, D. 1941. Contribution to the knowledge of Amphibia of South America. Goteborgs Kungl. Vetensk.-och Vitterh.-Sam. Handl., Ser. B, 1(4): 1-71, MERTENS, R. 1950. Froschlurche aus Rio de Janeiro und seiner Umgebung. Wochen- schr. Aquar. Terrarien, 44:173-188. MULLER, H. J. 1942. Isolating mechanisms, evolution, and temperature. Biol. Symposia, 6:71-125. MULLER, P. 1968. Die herpetofauna der Insel von Sao Sebastiao (Brasilien). Saar- brucker Zeit., 68 pp. MUt.er, P. and J. SCHMITHUSEN 1970. Probleme der genese siidamerikanischer Biota. in Deutsche geog- raphische Forschung in der Welt von Heute. Kiel:109-122. Moreau, R. E. 1966. The bird faunas of Africa and its islands. London, 424 pp. PETERS, W. 1872. Uber eine Sammlung von Batrachiern aus Neu-Freiburg in Bra- zilien. Monatsb. Akad. Wiss. Berlin, 1872:680-684. RivERO, J. A. 1969. On the identity and relationships of Hyla luteocellata Roux (Am- phibia, Salientia). Herpetologica, 25:126-134. 40 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Roux, J. 1927. Contribution a l’érpetologie du Vénézuela. Verh. Naturf. Gesell. Basel, 38:252-261. SALTHE, S. N. and W. E. DUELLMAN 1973. Quantitative constraints associated with reproductive mode in anurans. in Vial, J. L. (ed.). Evolutionary biology of the anurans: contemporary research on major problems. Univ. Missouri Press, Columbia:229-249. SAVAGE, J. M. and W. R. HEYER 1967. Variation and distribution in the tree-frog genus Phyllomedusa, in Costa Rica, Central America. Beitr. Neotrop. Fauna, 5:111-131. SCHOENER, T. W. 1967. The ecological significance of sexual dimorphism in size in the lizard Anolis conspersus. Science, 155:474-477. 1969. Size patterns in West Indian Anolis lizards: I. Size and species diversity. Syst. Zool., 18:386-401. 1970. Size patterns in West Indian Anolis lizards. II. Correlations with the sizes of particular sympatric species—displacement and con- vergence. Amer. Nat., 104:155-174. TRUEB, L. 1973. Bones, frogs, and evolution. in Vial, J. L. (ed.). Evolutionary biology of the anurans: contemporary research on major problems. Univ. Missouri Press, Columbia:65-132. VAN DE HAMMEN, T. and E. GONZALEZ 1960. Holocene and late glacial climate and vegetation of Paramo de Palacio (Eastern Cordillera, Colombia, South America). Geol. en Mijnb., 39:737-746. VANZOLINI, P. E. and E. E. WiILLiamMs 1970. South American anoles: the geographic differentiation and evolu- tion of the Anolis chrysolepis species group (Sauria, Iguanidae). Arq. Zool., 19:1-298. VUILLEUMIER, B. S. 1971. Pleistocene changes in the fauna and flora of South America. Sci- ence, 173:771-780. Wacne_er, W. H., Jr. 1961. Problems in the classification of ferns. in Recent Advances in Botany. Univ. Toronto Press, Toronto:841-844. . //+Me 13300 20: Pe Ei OF Stences 1] ACADEMY OF SCIENCES APR 15 1974 ca! LIBRARY OCCASIONAL PAPERS of the MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas NUMBER 24, PAGES 1-60 MARCH 27, 1974 SYSTEMATICS AND EVOLUTION OF THE GREATER ANTILLEAN HYLID FROGS By LinpA TRUEB’ AND MICHAEL J. TYLER? The origins and number of ancestral stocks of the hylid fauna of the West Indies have been debated. The twelve species are now allocated to the genus Hyla. The Lesser Antillean treefrog fauna is depauperate, consisting only of a member of the Hyla rubra complex which occurs on the island of Saint Lucia. The occurrence of this small hylid on only one of the islands in the Lesser Antilles is puzzling, unless Hyla rubra is a relatively recent immigrant or adventive from the South American fauna, as seems probable. The species (or closely related members of the species group) is present on the continental islands of Trinidad and Tobago and is widespread on mainland South America. Hyla squirella has been introduced to Grand Bahama (Crombie, 1972), and Hyla cinerea is an apparent adventive to Puerto Rico (Albert Schwartz, pers. com.); both are common in the southeastern United States. Eight of the remaining species are endemic to either Jamaica or Hispaniola, and the ninth species is rather widespread; it occurs on Cuba and the Isle of Pines, the Bahamas, and southern peninsular Florida and recently has been introduced to Puerto Rico. Unlike Hyla rubra of the Lesser Antilles, H. cinerea of Puerto Rico, and H. squirella of Grand Ba- hama in the aforementioned nine species, there are no immediately obvious affinities between these Greater Antillean species and main- land representatives. The systematic relationships among the nine 1 Adjunct Curator, Division of Herpetology, Museum of Natural History, and Department of Systematics and Ecology, The University of Kansas. * Honorary Associate in Herpetology of the South Australian Museum, North Terrace, Adelaide, South Australia 5000. Associate in Herpetology, Museum of Natural History, The University of Kansas. OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 2 ‘SBO1J O[RUIOJ YNPe FO Sozis oBeIOAe o1e SIOqUINU [eOHeyWUOIYg “SuUTYO}eY-sso1o Aq PoJOIpPUT ST syDUOLUAq -das smdoajsEQ JO UONNAINSIG ‘sal[HUW JoyeoID Jo vuNe} pIAY UWEpIseI JO UONNGLYSIP SuIMOYs Sorpuy yap OY} FO dey “[ “OY eNSBEIPOI @sopeqieg (wwg7) Laplim ejAy eI to g eIon'y ‘1S (wwEe) aeuelew e|AH (wwTg) snauunig snjidoa}s¢ (WM HOT) Byeuayol| ejAyeldAled Oory O}Jeng be als) eynsurued (my) eyeaul|lyoind eA} (wmgy) luldiay e|Ay | (ww //) SisuadIUIWop snjidoa}s¢ a) (wu/eT) eySeA e|AH SiJeudLJUa}das snjidoa}sy V4 GREATER ANTILLEAN HYLID FROGS 3 species themselves remain obscure, although they have received considerable attention in the past from Dunn and Noble, among others. Our interest in these frogs is an outgrowth of several other morphological and systematic studies, which, in one way or another, have involved this isolated group of anurans. Because of the avail- ability of specimens, Hyla septentrionalis was used by Trueb (1966) to describe the histology and development of exostosis, casquing, and integumentary-cranial co-ossification. Although the results of this research subsequently proved to be extremely useful in sys- tematic considerations of other casque-headed, co-ossified hylids (Trueb, 1970), little was learned about the relationships of the casque-headed hylids of the Greater Antilles (Hyla septentrionalis, H, brunea, H. dominicensis, and H. lichenata). As a part of his larger study of the vocal sac structure in hylids, Tyler (1971) noted the occurrence of a unique type of throat musculature in three of the Greater Antillean frogs—septentrionalis, brunnea, and domini- censis. The accumulated morphological evidence suggested that the nine resident frogs were not so closely related as previously proposed by Dunn (1926), and prompted us to undertake this review of the taxa. The format of the following review has been dictated by the peculiarities of the problem and our own limitations. Regrettably, neither of us has been able to observe any of these species in their natural habitat. In order to provide a general understanding of the distributions of the Greater Antillean hylids and the kinds of habi- tat available to them, we have provided cursory statements of the major topographical and phytogeographical features of the Greater Antilles and Bahamas. Because few species are involved, we elected to compare and contrast each of the nine species with re- spect to various morphological features of adults and larvae, and characteristics of breeding. The analysis of characters includes in- formation from the literature and much new osteological and myo- logical data. Together, these data provide a base for the critical reinterpretation of the systematics, evolution, and zoogeographic history of the Greater Antillean hylid fauna. Acknowledgments For the loan of specimens or for the provision of working space in their respective institutions, we are indebted to Alice G. C. Grandison, Charles W. Myers, the late James A. Peters, Charles F. Walker, Ernest E. Williams, and Richard G. Zweifel. A grant from the Science and Industry Endowment Fund pro- vided by the Executive of the Commonwealth Scientific and Indus- trial Research Organisation of Australia enabled M. J. Tyler to visit 4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY The University of Kansas. We thank this organization for providing us the opportunity to collaborate on this research. Special thanks are extended to Mr. Ronald I. Crombie, who donated several specimens of Hyla marianae to the collections of The University of Kansas, and to Albert Schwartz and Ernest E. Williams for critical review of this manuscript. We are particularly grateful to Robert M. Mengel for his thorough editorial review, and to William E. Duellman who encouraged the preparation of this manuscript, critically reviewed it, and throughout its inception and writing served as a resourceful critic. Materials and Methods Measurements (Tables 1 and 2) and descriptions of external morphological features are based on examination of 140 preserved specimens. Myological observations were made on 27 preserved specimens, and osteological descriptions are based on 31 skeletal preparations. The majority of characters analyzed are used commonly and require no special explanation other than that provided in text, and in text references. In the section dealing with osteology, sev- eral characters are discussed which should be clarified. The first group of these involve the structure of the vertebral column (Fig. 2A and B). The widths of presacral vertebrae and the sacrum were determined by measuring the linear distance between the distal tips of the transverse processes to the nearest 0.1 mm with dial calipers (Fig. 2A). Such measurements are not an index to the actual length of the transverse processes; we have gauged the overall dimension of individual vertebrae without accounting for the orien- tation of the transverse processes. The resulting values are useful in generating a profile of vertebral shape and only of limited use in comparing individual vertebral elements (e. g., width of presacral III and sacrum). The anterior sacral angle (Fig. 2B) was deter- mined by measuring the angle between the longitudinal axis of the vertebral column and the leading edge of the sacral diapophysis. The posterior sacral angle is that angle between the longitudinal axis and the posterior edge of the diapophysis. Sacral diapophysis expansion can be calculated by subtracting the sum of anterior and posterior angles from 180°. Camera lucida drawings of the verte- bral columns were made, and the angles were measured with a protractor from these drawings. Two characters are of particular interest in the pelvis (Fig. 2C). The first of these, relative ilial length, was determined by measuring the greatest length of the ilium and the length of the acetabular portion of the pelvis—i.e., the distance from the preacetabular margin of the ilium to the posterior margin of the ischium. The pelvic angle is that angle, in lateral profile, between the acetabular GREATER ANTILLEAN HYLID FROGS 5 A B | presacral width > : anterior sacral angle sacral diapophysis expansion acetabular ilial length pelvic angle Fic. 2. Graphical explanation of postcranial osteological measurements. A. Ventral view of vertebral column of Hyla vasta. B. Ventral view of vertebral column of Osteopilus brunneus. C. Lateral aspect of pelvic girdle of Osteopilus septentrionalis. 6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY portion of the ilium and the horizontal plane of the ilial shaft. In all cases, the values assigned this character are visual estimates— acute, nearly 90°, or obtuse. Of the myological characters, the extrinsic musculature of the cloaca requires explanation here because these muscles previously have not been employed in anuran systematics. In fact, attention to the cloaca hitherto has been confined to descriptions of the musculature in such aberrant genera as Ascaphus (Du Toit, 1955) and Rhinophrynus (Noble, 1922). The nomenclature that we adopt (see Fig. 3) differs from that of previous authors. The most proximal muscle, for which we pro- pose the name Musculus ischeococcygeus is the M. compressor cloacae of Noble (1922) and some previous authors. We propose GE == eloacal=—= = == Ori ficou: = Grace min: == Dre, Clo: = ischial rim —— Se eee na Se ees SS ee ee Se ‘ ee ee See ee eee Fic. 3. Posterior aspect of thighs of Osteopilus septentrionalis. Superficial portion of M. gracilis minor removed from left side. Abbreviations: compr. clo., M. compressor cloacae; grac. min., M. gracilis minor; pro. clo., M. pro- tractor cloacae; pyr., M. pyriforms. GREATER ANTILLEAN HYLID FROGS 7 this new name because this muscle occupies a pre-cloacal position and does not in any way compress the cloaca. The adjacent muscle is a genuine cloacal compressor and our use of compressor cloacae for it is in agreement with the actions of Du Toit (1955). A pre- viously unreported muscle which we find occurs in many hylid species we name the M. protractor cloacae. Studies of musculature were assisted by the use of iodine/ potassium iodide staining technique described by Bock and Shear PON 2). We express the vocal sac aperture as a fraction of the mandible length. This was obtained from the maximal diameter of the vocal sac aperture (usually on a longitudinal axis), and the distance between the mandibular symphysis and the postarticular extremity of the mandible. DISTRIBUTION OF THE GREATER ANTILLEAN HYLID FAUNA At the risk of over-simplification, the Greater Antilles and Ba- hama Islands may be subdivided arbitrarily into five geographical subunits according to the distributions of the resident treefrogs (Fig. 1). These are: 1) Jamaica, 2) Hispaniola (Haiti and the Dominican Republic), 3) Cuba and the Isle of Pines, 4) the Bahama Islands, and 5) Puerto Rico. The following accounts are extremely brief and simplified, and intended only to introduce the reader to the habits, habitats, and distributions of the resident treefrogs. We also hope these accounts will serve as a modest, if superficial, sum- mary of the principal topographical and vegetational features of the geographical subunits. Information on the latter was derived from inspection of a variety of topographic and vegetation maps and the accounts of Verdoorn (1945), and Rabb and Hayden (1957). It is important to note that all the islands of the Greater Antilles are very diverse ecologically, each ranging from arid lowland deserts to high, upland deciduous or pine forests. The islands differ princi- pally in their physiographic complexity. Hispaniola is the most complex (see Schwartz, 1973), whereas Puerto Rico and Jamaica are smaller and more simple. Each of these islands consists mainly of a central mountain mass surrounded by lowlands. In contrast to Jamaica and Puerto Rico, the largest island, Cuba, is character- ized by three isolated mountain masses with surrounding lowlands. Jamaica.—The four endemic hylids which occur on this rela- tively small island (ca. 224 x 80 km) are Hyla lichenata, H. brun- nea, H. marianae, and H. wilderi. Topographically, the island is diverse. It is bisected longitudinally by a highland mass, the peaks of which lie between elevations of 1500 and 2100 m. The western and central three-fourths of the island comprise a plateau reaching elevations of 900 m. On the southern margin of Jamaica, there is 8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY an irregular coastal plain (maximally 24 km in width). A typically insular climate prevails, with the northeast sector receiving the highest annual precipitation, the central plateau somewhat less, and the southern coastal plain the least. The dry coastal plain is char- acterized by cactus, thorn bush, and dry scrub. In contrast, the northern aspect of the island is heavily wooded, with tropical rain- forest on the more remote slopes. The southern aspect of the island supports a drier, sclerophyllous vegetation consisting of park- savanna with scattered palms, pines, and scrub. The extensive cen- tral plateau is thickly wooded on its northern side and at higher elevations. Parkland and pasturage characterize lower elevations. The central and southern parts of the plateau are cultivated. Hyla brunnea is the most widespread of the four species; it occurs over almost the entire island at elevations below 1500 m (Schwartz and Fowler, 1973). This moderately large, casque- headed frog (adults attain 76 mm snout-vent length; Schwartz and Fowler, 1973) is found most often in arboreal and terrestrial brome- liads in open woods. Hyla wilderi seems to be widely distributed over the central plateau at elevations between 122 and 610 m (op. cit.). This species is small (adults reach about 30 mm snout- vent length) and is not casque-headed; it occurs sympatrically with brunnea and frequents arboreal and terrestrial bromeliads in open woods. Hyla lichenata is the largest (adults reaching 122 mm snout- vent length; Gosse, 1851) of the four species and is distinguished by its rugose patterned, casqued cranium. Although documentation is poor, the frog seems to be rather widely distributed throughout most of the island at low to moderately high elevations. In contrast to brunnea and wilderi, lichenata is reported to be an inhabitant of more densely wooded forests, where it has been observed to perch in hollows in branches 1.2 to 3.6 m above the ground (Dunn, 1926). Hyla marianae is moderately small (adults reaching 39 mm snout-vent length) and inhabits bromeliads in dense and scattered deciduous forests between elevations of 122 and 895 m (Schwartz and Fowler, 1973). It is at least partially sympatric with each of the three other species. All four species deposit their eggs exclu- sively in the water-filled leaf bases of bromeliads. Although they have been reported to inhabit both terrestrial and arboreal brome- liads, eggs have been found only in arboreal bromeliads. Appar- ently, none of the species occurs on the xeric, southern coastal plain. Hispaniola——Although Hispaniola is larger (ca. 640 x 288 km) and more diverse topographically and vegetationally than Jamaica, again, only four species of hylids occur on the island—Hyla do- minicensis, H. vasta, H. heilprini, and H. pulchrilineata; all are endemic. Hispaniola is the second largest Antillean island and probably the most complex physiographically. There is a central east-to-west mountain mass, the highest peak of which lies at an GREATER ANTILLEAN HYLID FROGS 9 elevation of 3175 m. The mountains are fragmented into a number of spurs and parallel chains. At higher elevations, the mountains support a mixed pine and broadleaf forest. Extensive areas of tropical rainforest occur at lower elevations in the central, north- eastern and eastern portions of the island, whereas the peripheral, low elevations and coastal plains are characterized by park-savanna with palms and scrub. Xeric lowland dry scrub associations exist along the northwestern coast and south-central coastal areas. Apparently, Hyla dominicensis is the most common and wide- spread of the resident hylids. The distribution of this relatively large frog (adults attain 99 mm snout-vent length) seems to be island-wide from the coastal lowlands to moderate elevations in the foothills. The species is principally an inhabitant of parkland- savanna and tropical rainforest environments. It is known to de- posit its eggs in stagnant pools of rainwater in the forest and along the edge of stream banks (Noble, 1927). The second large treefrog of Hispaniola is Hyla vasta (adults reach 137 mm in snout-vent length). This non-casque-headed species is considerably scarcer in collections than dominicensis and perhaps, therefore, less common. It occurs sympatrically with dominicensis at moderate to low eleva- tions and has been observed to lay its eggs in small gravel and stone depressions in quiet pools along streams (Noble, 1927). Schwartz (pers. com.) indicated that vasta is also partially sympatric with Hyla heilprini which breeds in torrenial streams. Hyla pulchriline- ata is the smallest member of the Hispaniolan hylid fauna. Adults attain a snout-vent length of 43 mm, and the cranium bears no dermal modifications. Published locality records for this species are scarce, but Schwartz (pers. com.) indicated that pulchrilineata is widespread on the island, occurring from the coastal lowlands to an elevation of about 636 m; thus, pulchrilineata seems to be sym- patric with dominicensis, vasta, and heilprini. Nothing has been reported concerning the larvae and reproductive behavior of this species. The fourth species, Hyla heilprini, is moderate in size (adults reaching 54 mm snout-vent length) and lacks cranial orna- mentation. The limited locality records for this frog indicate that it inhabits the central Hispaniolan highlands; however Schwartz (pers. com.) reported that the species is more widespread than suggested in the literature and implied that heilprini is not limited to the highlands. Noble (1927) reported that heilprini was found along streams at places where the water fell in cascades over rocks. Cuba and the Isle of Pines—Cuba, the largest of the Antillean islands, and its small insular associate, the Isle of Pines, have only one treefrog, the moderately large (adults up to about 75 mm snout-vent length), casque-headed Hyla septentrionalis. In con- trast to the other Greater Antillean treefrogs, this species is not endemic to a single island; it occurs in the Bahamas, southern OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 10 (S'FS-0'0¢ ) CGS (0'9S-F'9F ) cis (€'€S-S' LP) P'0S (9'°SS-T LP) Gos (9°9S-L'TS) PSs (¢°g9-G'°SG) 66S ($°G9-6'SF ) ELS (9'L9-8'FS ) 9°19 (%) oA /dwAy, (o'se-e'TE ) SCE ($°LE-L'8Z) OSE (O'TE-L'0E ) 8°0E (T7e-€'ee) g'se (§°8-6'9G) 1'8G (9'08-F'E3) 8G (L'1&-F' LZ) L'6Z (O'TE-8'LZ) £63 (%) MH/dOlI (9'SS-o'FE ) Tse (G'LE-T'PE ) g’°cs (7'8S-9'LE ) 09g (6'LE-S'FE) 0'8E (ZLE-S'FE ) ZOE (F'8E-0'FE ) 6'SE (O°ce->o IG) O'SE (ZS FE-E'0E ) O'SE (%) TAS/MH (L'¥E-T Se) PEE (0'9S-L'TS ) VPE (T'LE-E'Se ) SOE (6'8E-6'9E ) 0'8E (F'98-6'EE ) oss (9° LE-6'EE ) Lge (ESO 18) VEE (0'9€-3'ZE ) SVE (%) TAS/TH (0'FS-3'0S ) T'S ($'SS-3'6P) 9°GS ('ZS-T'0¢) 1g (8°ES-E'TS) 8°3S (L°SS-9'6F ) GIS (0°LS-T'6F ) TES (0'SS-F'SP) 80S (T'¥S-'9F) G'0S (%) GAS/ TEL (L'8S-F'S3) a CUZ (L=N) 6 (€°L3-8'€3) ed 9°67 (SI=N) ? wappn vjA (9°6E-L'L€ ) = 19¢ (Z=N) 6 (T'€E-F'ES) = 0°82 CSINOH apupiudu DIA (0°SS-3' 9F ) = © GF (V=N) 6 (€'PS-0' LP ) = ; GP (ZI=N) ? tuiidjay vii (8°GP-9' LE) = LO (9=N) 6 (S'6E-L'8Z) = OTe (OZ=N) ? Dwauyiuyond DIA ( urUr ) satoeds "IAS ‘IoyoureIp UNueduUtAy “GIN AL pue ‘yyue, vIqn “qL.L ‘ysue_ yUeA-ynous “TAS ‘e0URIsIP [eWqIOIOJUT “OT “Wp. peoy “MH ‘Wsue] peoy “TH :suonemaiqqy ‘euney pYAY uva[HUY JoyeaIg oY} JO Sioquiout so[[eUUs Jo suoniodoid pue sozis Apoq jo sesuvi pue suvouI Jo uOnRNGe], “[ ATAVy, 11 GREATER ANTILLEAN HYLID FROGS 8'F8 689 (0'8S-9'L¢ ) QL (O'TL-0'TS) PSs (F'S8L-8'SL) TLLs (0'0L-8'29 ) P99 « (T'E8-L'99 ) 9°€9 (0'°69-T'L¢) 9°E9 (9'08-L'GL) Gch (8°9L-9'€9 ) Says (%) oAq /dutA J, C'vé Tse (3 PE-6'SE ) L'¥E ($°L3-8'FG) S 9G (O'SP-F'FE ) L6ox (3 18-708 ) 808s (7'8P-S IP) CP (SPE-OFE) 6'6E (9°9F-S'0F ) eS 7 (0'SE-8'SE ) r'6E (%) MH/dOI ‘SUOQRIAZIQGe 10F [T 9[Ge 99S *(SC6T peseq ore suveul oy} Uasqe oe sosUI YOIYA\ UT Sooue}SUT 9SOY} UT SIF PSE 0'6E aS (9°GE-8'1e) (¥'08-S'6Z ) CGE 0'0€ (Ca Saks) (T'¥E-'08 ) ree ENG (6'SE-S'FE ) EEE oSos ($'FE-6'TE ) SI VES s ($°SS-8'TS ) (798-8 TE) ECS SFE (T'3E-0'6S ) ($°S8-6'08 ) 3°08 O'SE (TRe-Fes) (7'8E-0'SE ) 9°c¢ 9°c¢ (T'S8-F'Se ) (0'6E-Z'9E ) g'98 eye (%) (%) TAS/MH TAS/TH (Z'ZS-8 GP ) ors (7'9S-¢ 67 ) PSs 6 OF x SSP « (¢'9S-0'°6F ) GPs (€°SS-6 LP ) ETS (l'6P-L' PP ) L'OF ($°IS-1'SP) © LP (%) TAS/AEL 0'FOT 9°C6 (6° I¥1-8 T&T) 69ST (8'801-S'L8 ) L'96 GGL © os (L'86-Z'0L) SOL (9°€9-9'SS ) SL (0'0L-S'GS ) G09 (1'8h-0'1P7) G'GP a) TAS (I=N) 6 (I=N) ? pypuayoIYy pjiiyojidhjog (Z=N) 6 (ZI=N) ? pyspa oii (LI=N) 6 (OV=N) ? syouo.j4uajdas sm1do0ajsO (6I=N) 6 (O=N) ? SsisuadUiULop smjid0ajsO (9=N) 3 (9=N) ? snauunig smjidoa}sO satoeds ‘ZyIeMYOG pue ueueNC ) pepraoid jou o19A\ sosuer YOryM Url lode ainje1a}] B UO ‘OM} JO o[dures e& JO Suva }UBSoIdaI SYSTI9}se Aq poeyxleur soinsLy ‘euney pyAY uey[QUY IoyeorD oY} Jo Sioquiew 1081] Jo suoM1odoid pue sazis Apo fo sosuvr puv suvout Fo UOHLINGeL, “G ATV] 12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY peninsular Florida and Puerto Rico. Cuba differs from the smaller islands of Hispaniola and Jamaica in lacking a central highland mass; its mountainous areas are disjunct. A relatively low chain occurs on the western end of the island with peaks reaching eleva- tions of 728 m. South-centrally, the peaks of a small mountain mass reach 1156 m. The most extensive montane area lies along the southeastern coast and on the eastern end of the island where peaks reach elevations above 2000 m. There are restricted areas of mixed pine and broadleaf forests in the extreme eastern and western montane areas. Two low, isolated highland masses (ca. 310 m elevation) occur on the Isle of Pines. Tropical rainforest occurs principally in the south-central montane area of Cuba, whereas the southeastern coast supports xeric-adapted vegetation. Save for coastal mangrove and xeric scrub areas, most of Cuba is characterized by park-savanna vegetation of scattered palms and scrub. Hyla septentrionalis is widespread on Cuba and the Isle of Pines at low to moderate elevations, where it inhabits old cisterns, axils of palm leaves, banana plants, and drain pipes; eggs are laid in temporary drainage ditches, flooded basements, and old cisterns (Carr, 1940). Little is known concerning its habits in natural situations. The Bahamas.—The Bahama archipelago consists of some 29 major islands extending about 1128 km northwest to southeast. The relief of the islands is below about 67 m. Some of the larger islands are characterized by lengthwise ranges of hills, whereas others (e.g., the large island, Andros) have virtually no topographical re- lief. Running streams are rare and availability of fresh water is seasonal, depending upon rainfall. Vegetationally, the islands are characterized by coppice thorn bush and dry scrub. Hyla septen- trionalis is the only treefrog occurring naturally in the Bahamas; it has been reported from the Cayman Islands in addition to Little Bahama Bank, Grand Bahama Bank, San Salvador, Rum, Crooked and Acklin’s Islands. The species was introduced recently to Great Inagua Island (Schwartz, 1968). Hyla septentrionalis has become well established in southern peninsular Florida, where it occurs sympatrically with several North American hylids (Duellman and Schwartz, 1958). Puerto Rico.—The island of Puerto Rico has no native treefrogs. Hyla septentrionalis and H. cinerea are recent adventives (Duell- man and Crombie, 1970; Schwartz, pers. com.). The island, al- though small (ca. 160 x 48 km), is topographically diverse. The elevation of the central highlands reaches approximately 1220 m. A relatively narrow area of savanna-parkland occurs along the mar- gin of Puerto Rico. Mixed pine and broadleaf forests predominate in the central hills and tropical rainforest is restricted to the higher, central elevations. ee GREATER ANTILLEAN HYLID FROGS 13 ANALYSIS OF CHARACTERS External morphology.—Tables 1 and 2 summarize data on the sizes and proportions of nine species of the West Indian hylid fauna. Although the quality of these data vary with respect to sample size and uniformity, they are sufficient to substantiate several observa- tions on the physical features of these frogs. The species (wilderi, marianae, pulchrilineata, and heliprini) included in table 1 are small to moderate-sized frogs which range in mean size from 27.3 to 49.4 mm snout-vent length. The species (brunneus, dominicensis, septentrionalis, vasta, and lichenata) listed in table 2 are moderate to large frogs with snout-vent lengths between 60.5 to 136.9 mm. Ratios of tibia to snout-vent length are remarkably uniform among all species, with values lying between 46.9 and 54.5 percent; simi- larly the ratios of head length and width to snout-vent length vary little (30.0-39.0%). In contrast to the proportional uniformity of the foregoing measurements, the ratio of interorbital distance to head width parallels the pattern evident in overall size of frogs. The smaller species included in table 1 have relatively narrower inter- orbital areas (28.1-33.8%) than do the larger species (brunnea, dominicensis, septentrionalis, vasta, and lichenata) of table 2 (30.8- 44.5%). Likewise, relative to the diameters of the eyes, the tympana of the smaller species are smaller (50.4-61.1%) than those of the larger species (57.8-77.1%). Among the group of larger frogs, the tympanum-eye ratio of Hyla vasta is noticeably lower (ca. 58%) than the other species (63.6-77.1%). Visual inspection shows the tympanum of vasta to be small; the eye is not distinctly large. The dorsal skin of wilderi, pulchrilineata, marianae, and _ heil- prini is smooth. That of dominicensis and brunnea is smooth; how- ever the latter has scattered, small, round tubercles on the forearm. Hyla septentrionalis and vasta have similar tubercles scattered over the entire dorsum. In contrast to the other species, the dorsum of lichenata is strongly tubercular. The ventral skin of all species is granular to some degree. It is weakly granular on the abdomen and thighs of all the smaller species, except heilprini, in which it is coarsely granular; wilderi and marianae are distinguished by the presence of tubercles around the cloaca and heilprini by the pres- ence of a large, dermal submental gland. Granulation is moderately developed on the posterior abdomen and around the cloaca in brunnea. In septentrionalis it extends over the thighs and entire abdominal area, but is absent around the cloaca. The ventral re- gions of dominicensis, lichenata, and vasta are both coarsely granu- lar; lichenata is further distinguished by tubercular skin around the cloaca. Fringes are absent from the limbs of all species except vasta and lichenata. In the latter, a distinct dermal fringe is present along the outer edge of the forearm and tarsus. Hyla vasta bears more ex- 14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY tensive fringes along the forearm and the hind limb, extending the length of the tarsus and foot. In addition, it has a distinct series of tubercles on the mandible and a proliferation of tubercles at the tibia-tarsal joint. Dermal wrist folds are absent in all of the smaller species except septentrionalis. The tympanum is distinct in each species except wilderi, in which it is not obvious externally, and heilprini, in which it is “con- cealed.” Similarly, the supratympanic folds are scarcely evident in these two species, whereas they are distinct in the others. The nostrils are terminal in all species, but vary in their degree of pro- tuberance. In most the nostrils are not protuberant. They are slightly protuberant in dominicensis, lichenata, and brunnea, and noticeably protuberant in vasta. The cloacal opening lies posteriorly at the upper level of the thighs in all species. Dermal modifications in the cloacal region are absent in all species, except lichenata, vasta, and heilprini. A small horizontal supracloacal flap charac- terizes lichenata, whereas longitudinal folds of skin lie adjacent to the cloaca in heilprini. A profusion of tubercular skin growths distinguish vasta. Hands and feet——Hand morphology varies considerably among the West Indian hylids. All species have well-developed digital pads which are larger on the fingers than on the toes. Hyla heil- print is unique in having a prepollex which is evident externally, and a well-developed prepollical spine. Webbing is absent from the hand in wilderi and present basally between the fingers in marianae, pulchrilineata, and septentrionalis. Of the remaining species, the fingers of brunnea, dominicensis and lichenata are ap- proximately one-fourth webbed, whereas those of heilprini are one- half webbed and those of vasta three-fourths webbed. Subarticular tubercles are distinct and round on inner digits with a tendency toward becoming flattened and bifid on outer fingers in all species except wilderi, in which all subarticular tubercles are low, round, and barely evident. Supernumerary tubercles vary from total ab- sence in marianae and pulchrilineata, to presence of a few indistinct tubercles in vasta and wilderi, a moderate number of low, round tubercles in lichenata and septentrionalis, and presence of numerous tiny tubercles in brunnea, dominicensis, and heilprini. All species have inner palmer tubercles which are more or less elliptical in shape. Distinct outer palmar tubercles are absent in five species; whereas in the remaining four (wilderi, dominicensis, vasta, and pulchrilineata) two outer palmar tubercles are present. Males of at least seven of the species bear nuptial excrescences; these vary from flat horny pads in marianae, pulchrilineata, dominicensis, sep- tentrionalis and vasta, to a proliferation of small, keratinous gran- ules in brunnea, and the presence of large keratinous spines in GREATER ANTILLEAN HYLID FROGS 15 | re cm Fic. 4. Skull of Hyla vasta (University of Kansas, Museum of Natural History 84711). A. Dorsal view. B. Ventral view. 16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY wilderi. Nuptial excrescences are absent in heilprini; their presence or absence has not been noted in lichenata. Webbing is present between the toes and more extensively de- veloped in all species than finger webbing. The toes of wilderi are about one-fourth webbed, those of marianae one-third and pulchri- lineata and septentrionalis two-thirds webbed. The feet of brunnea, dominicensis, and lichenata are three-fourths webbed, whereas those of vasta and heilprini are fully webbed. Subarticular tubercles are round; they have a tendency to be flattened and bifid on the outer toes of dominicensis, heilprini, and lichenata. Plantar super- numerary tubercles are absent in vasta and heilprini, and few in number in all other species except brunnea and lichenata, in which their numbers are moderate. Inner metatarsal tubercles are present and vary between ovoid and flat, and elliptical and elevated in shape. Outer metatarsal tubercles are absent or scarcely evident in all species except pulchrilineata, marianae and lichenata; the tuber- cles are small and round in the latter species. Cranial Osteology—On the basis of cranial characteristics, the West Indian hylid fauna is easily subdivided into the small-sized and large-sized groups observed in tables 1 and 2. Although the observed classes correlate with body size, they are predicated on structural features of the skulls and will be discussed separately. Among the larger species there are three groups. The first is com- posed of septentrionalis, dominicensis, and brunnea, whereas the second and third groups each contain only one species, lichenata and vasta, respectively. Despite being the largest of the West Indian frogs (Table 2), vasta has the least well ossified skull (Fig. 4); it is only slightly wider than long and has no dermal modifications. The skulls of the moderate-sized septentrionalis-group also are slightly wider than long but, are distinguished by exostosis, casquing, and co-ossification (Fig. 5). A moderate development of dermal sculpturing resulting in patterns of pits, ridges, and bony tubercles is associated with this group. Hyla lichenata is distinctively different from both vasta and the septentrionalis group. The skull (Fig. 6) is distinctly wider than long and heavily exostosed, casqued, and co-ossified. Dermal sculp- turing consists of a distinctive pattern of peripheral, radial ridges which grade into a central pattern of pits and ridges. The skull of vasta is minimally roofed and delicate in structure (Fig. 4A). A dermal sphenethmoid is absent. Although nasals are moderately large, they are widely separated and barely overlap the anterior margin of the sphenethmoid. Similarly, the frontoparietals are minimally developed. They are not peripherally expanded and do not articulate with the maxillary. The palatines (Fig. 4B) are well developed, anteriorly convex, and articulate with the lateral edge of the sphenethmoid. The parasphenoid is large. The cultri- GREATER ANTILLEAN HYLID FROGS ly I cm Fic. 5. Skull of Osteopilus brunneus (University of Kansas, Mu- seum of Natural History 84693). A. Dorsal view. B. Ventral view. 18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY form process of this element is edentate and extends to the forepart of the orbit. The alae are posterolaterally inclined; their distal ends are widely separated from the medial rami of the pterygoid bones. The pterygoids are robust, triradiate and fully articulated; the anterior rami articulate with the maxillaries at the mid-level of the orbits. The prevomers are widely separated from the maxillary arch. Prevomerine teeth are situated on moderately broad, curved elevations and occur in a double row laterally. Cranially, dominicensis, septentrionalis and brunnea represent a group of broadly casqued, moderately exostosed, co-ossified hylids. Each species is characterized by a pattern of dermal sculpturing which consists of irregularly shaped pits and ridges, and prolifera- tions of small, bony tubercles (Fig. 5A). Along the margins of the dermal roofing bones and the maxillae, the tubercles tend to be aligned in series to produce patterns of low radial ridges. A dermal sphenethmoid is present in each species. The frontoparietals are extensively developed; a broad supraorbital flange is present and the paired elements articulate medially, completely obscuring the frontoparietal fontanelle. Posterolaterally, the frontoparietal com- pletely roofs the prootic region in mature females of each species. In septentrionalis and dominicensis the frontoparietal articulates with the otic arch of the squamosal, whereas no articulation is estab- lished between these elements in brunnea. The occipital margins of the frontoparietals are strongly convex in dominicensis and brunnea, but tend to be straight in septentrionalis. The nasals are extensive and with all adjacent elements articulated in all three species. The canthal ridges are straight in dominicensis, very slightly concave in septentrionalis, and markedly concave in brun- nea. Furthermore, brunnea has a much longer medial nasal articu- lation than the other two species. In dominicensis and septentrion- alis, the zygomatic arch of the squamosal articulates with the max- illary, whereas the arch is incomplete in brunnea. Palatines are well developed in brunnea and septentrionalis, but very poorly devel- oped in dominicensis. In contrast to the condition of vasta, the palatines of the septentrionalis group tend to be straight or to have their proximal ends turned anteriorly (Fig. 5B). The cultriform process of the parasphenoid is edentate and extends to the anterior level of the orbit in each species. The parasphenoid alae are per- pendicularly oriented in dominicensis and brunnea and slightly inclined posterolaterally in septentrionalis. Likewise, the distal ends of the alae are narrowly separated from the medial pterygoid ramus in dominicensis and brunnea and widely separated in septen- trionalis. The pterygoid is best developed in brunnea, in which the anterior arm articulates with the maxillary at the mid-level of the orbit and the medial arm articulates firmly with the otic capsule. The element is less robust and articulates with the maxillary at the ritish Museum of Natural z ee Ss 33 < 5 Se) oF Ss 5 jaa) is} Ss a Se 2. =s Siz Ss A < = a a Aa Lu aA oO Cam a Ss 2 a 20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY posterior level of the orbit in dominicensis and septentrionalis. In the latter species the medial pterygoid ramus is weakly articulated with the otic capsule, whereas in dominicensis the articulation is absent. The prevomers of the three species are similar in having single rows of teeth situated on broad, curved elevations. The pre- vomers are small and widely separated from the maxillary arch in dominicensis, whereas they are large and articulate with the maxil- lary arch in septentrionalis and brunnea. Hyla brunnea is further distinguished by the strong curvature of the prevomerine tooth row (Fig. 5B). The skull of lichenata is distinctly different from those of the other four species. Like the septentrionalis-group, it bears a dermal sphenethmoid and is broadly casqued and co-ossified. However, the skull is considerably broader than long and the pattern of exo- stosis (Fig. 6A) consists of rugose peripheral ridges which grade into a central pattern of bony pits and ridges. Although the skull is heavily casqued, it is less well developed than in the septen- trionalis group. The nasals are not completely articulated with the maxillaries. The frontoparietals extend posterolaterally over the otic region but are widely separated from the squamosal. The zygo- matic arch of the squamosal does not articulate with the maxillary. The palatines are robust and straight (Fig. 6B). The cultriform process of the parasphenoid is edentate and exceptionally short, extending only to the mid-level of the orbit. The parasphenoid alae are oriented perpendicularly and articulate distally with the medial rami of the pterygoids. The pterygoids are extremely robust and fully articulated; the medial ramus articulates with the maxillary at the mid-level of the orbit. The prevomers are moderate-sized bones separated from the maxillary arch. They bear single rows of teeth on elevations which are only slightly curved and oriented at a slight angle to one another. The cranial characteristics of the smaller members of the West- ern Indian hylid fauna (wilderi, marianae, heilprini, and pulchri- lineata) clearly represent the cumulative effect of different kinds of adaptive trends than those evident among the larger frogs. Corre- lated with their smaller body sizes (ca. 25-50 mm), the dimensions of the skulls are decreased, and ossification is reduced. The latter is especially evident in the absence of dermal modifications (includ- ing the absence of the dermal sphenethmoid), and the moderate to minimal development of dermal roofing bones; the frontoparietal foramen is evident in each species. The mandibles are incompletely articulated and, similarly, suspensory and bracing elements (squa- mosal, palatine, and pterygoid) tend to be reduced in size and incompletely articulated. Three different patterns of cranial archi- tecture are evident among these smaller hylids. The skulls of mari- anae (Fig. 8) and wilderi (Fig. 9) are characterized by the most GREATER ANTILLEAN HYLID FROGS 21 Fic. 7. Skull of Hyla pulchrilineata (University of Kansas, Museum of Natural History 84709). A. Dorsal view. B. Ventral view. marked reduction in ossification and by snouts which (in dorsal view) are broadly rounded. The skull of pulchrilineata (Fig. 7) is easily distinguished by its length and narrowness, and that of heilprini (Fig. 10) by its truncate snout. The narrow skull of pulchrilineata (Fig. 7A) is the most distinc- tive of these four species. The nasals are approximately rectangular in dorsal view and about twice as long as wide; their longitudinal axes nearly parallel those of the maxillary arch. About one-third of the length of the nasal projects anterior to the sphenethmoid. The premaxillary bears long, dorsolaterally divergent alary processes, but only a reduced palatine projection vetromedially. The prootic is narrow and laterally, abuts (but is not overlapped by) the otic ramus of the squamosal. The zygomatic ramus of this element extends approximately one-third the distance to the maxillary. The palatines are reduced, non-articulate and anteriorly concave (Fig. 7B). The parasphenoid bears a large cultriform process; the anterior end of the process is truncate and lies at the forelevel of the orbit. The pterygoid is small; the anterior ramus articulates with the maxillary at the posterior level of the orbit, whereas the medial arm does not bear a bony articulation with the otic capsule. The prevomers are small but bear moderately large dentigerous proc- esses. These processes are curved (anteriorly concave), and nar- rowly separated; each bears a single row of approximately nine teeth. The relatively greater breadth of the skulls of wilderi and mari- anae (Figs. 8 and 9) readily distinguishes them from pulchrilineata. The nasals are only slightly longer than wide, and their longitudinal axes parallel the maxillary arch. One-third (wilderi) to one-half 22 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Fic. 8. Skull of Hyla marianae (University of Kansas, Museum of Natural History 146838, female). A. Dorsal view. B. Ventral view. (marianae) the length of the nasal lies anterior to the spheneth- moid. The nasals of both species differ from those of pulchrilineata in having convex medial margins which broadly overlap the sphe- nethmoid. The alary processes of the premaxillary are moderately long in marianae as contrasted with wilderi; the processes are nearly vertical in both species. The palatine projection of the premaxillary is small in wilderi and minute in marianae. Like pulchrilineata, both marianae (Fig. 8A) and wilderi (Fig. 9A) have reduced prootics; but unlike the former, the otic rami of the squamosals in marianae and wilderi do not articulate with the prootics. Moreover, the zygomatic rami of the squamosals are longer, extending about one-half the distance to the maxillaries. The palatines of wilderi and marianae are anteriorly convex in contrast to those of pulchri- lineata. Those of wilderi (Fig. 9B) are reduced; they articulate with neither the maxillary nor the sphenethmoid, whereas the pala- tines of marianae (Fig. 8B) extend from the maxillary to articulate firmly with the sphenethmoid. The parasphenoids of both spe- cies are similar. The cultriform process of this element is edentate, acuminate anteriorly, and terminates at a level in the anterior third of the length of the orbit. The parasphenoid alae are widely sepa- rated from the medial rami of the pterygoid. The pterygoid of marianae is moderate in size and fully articulated; in contrast, that of wilderi is small and lacks a medial articulation with the otic capsule. The anterior rami of the pterygoids of both species articu- late with the maxillaries at the posterior levels of the orbits. The prevomers of marianae are relatively larger than those of wilderi. The bones are widely separated from the maxillary arches in both species and bear similar dentigerous processes which are straight, widely separated elevations. The processes bear single rows of teeth totaling 7-8 per row in marianae and 4-5 per row in wilderi. GREATER ANTILLEAN HYLID FROGS 23 Fic. 9. Skull of Hyla wilderi (University of Kansas, Museum of Natural History 84713, female). A. Dorsal view. B. Ventral view. Several features set apart the skull of heilprini (Fig. 10). Most apparent is its blunt, truncate snout. Unlike the other three species, the nasals are long and narrow (at least three times as long as the greatest width). Furthermore, the nasals are acuminate anteriorly with their anterior ends converging at the margin of the spheneth- moid. The longitudinal axes of the nasals are not parallel to the maxillary arch. The frontoparietals of heilprini are poorly devel- oped; thus a large part of the frontoparietal fontanelle is evident, particularly posteriorly. The premaxillary differs in several respects from those of the other small hylids. The alary processes are mod- erately small, dorsomedially convergent, and posteriorly inclined; the palatine processes are well developed. The prootics are wide in contrast to those of pulchrilineata, marianae, and wilderi. The squamosals overlap the prootics dorsolaterally and the zygomatic rami extend approximately one-half the distance to the maxillaries. The palatines are delicate, anteriorly convex, and articulate with the sphenethmoid medially. The parasphenoid is similar in structure to those of marianae and wilderi, differing from the latter and pulchrilineata in having alae which are only narrowly separated from the medial rami of the pterygoids. The pterygoids of heilprini are moderately large and fully articulated. The anterior pterygoid rami are much longer than those of the other species; they articulate with the maxillaries at approximately the midlevel of the orbit. The prevomers are moderately large and not closely associated with the maxillary arch. Each element bears a single row of teeth (5-6 per row) on a slightly curved elevation. Axial osteology—The vertebral columns of the West Indian hylids are uniformly characterized by eight procoelous, presacral vertebrae, none of which is fused. The cervical cotyles are widely separated, and there are no dermal modifications. The sacral verte- brae bear bicondylar articulations with unmodified coccyges. Within 24 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 10 mm Fic. 10. Skull of Hyla heilprini (University of Kansas, Museum of Natural History 90838, male). A. Dorsal view. B. Ventral view. these generalized limitations, there are morphological patterns (Fig. 2 A and B) which corroborate the species groups established on the basis of cranial structure. The presence of neural spines on the presacral vertebrae of the larger species distinguishes them from the smaller frogs. Among the larger members of the West Indian frogs, lichenata and vasta show the most specializations in the vertebral column associated with a trend towards advanced terrestrialism (i.e., ar- boreality ); in general, these modifications are most pronounced in lichenata. Although the neural spines are best developed on pre- sacrals I-V, they are present on all presacrals of lichenata. In vasta, the neural spines are obvious on presacrals I-IV and barely visible on the remaining presacrals. In both of these species (in contrast to the other frogs) the sacral diapophyses are wider than the widest transverse processes (presacral III); the diapophyses are propor- tionately widest in lichenata. The transverse processes are of nearly uniform width in lichenata; presacral III comprises 87 percent of the width of the sacral diapophyses, and each of presacrals IV-VIII, 75 percent of the sacral width. The pattern is less uniform in vasta (I1]—95% of the sacral width, IV—81%, and V-VIII—71%). In both species the transverse processes of presacral II and III are expanded distally. Hyla lichenata differs from vasta in bearing a small flange on the anterior margin of the transverse process of presacral II and a similar flange on the posterior margin of the transverse process of presacral III. The sacral diapophyses are only slightly dilated in vasta (ca. 20° and lichenata (ca. 31°). The differential expansion is the result of the orientation of the anterior margin of the sacral diapophyses. The orientation may be measured by calculating the angle between the anterior edge of the sacral diapophysis and the GREATER ANTILLEAN HYLID FROGS 25 longitudinal axis of the body. The angle approximates 80° in lichenata and 103° in vasta. The vertebral columns of the septentrionalis-group (septen- trionalis, dominicensis, and brunnea) show fewer advanced terres- trial specializations than those of vasta and lichenata (Fig. 2). Neural spines are best developed in dominicensis, in which they are evident on presacrals I-V; in brunnea, spines are present on presacrals I-IV, whereas in septentrionalis they are invariably pres- ent only on presacrals I-III. The sacral diapophyses of all three species are narrower than the widest transverse process of the pre- sacrals (III), in contrast to vasta and lichenata. The transverse processes of the presacrals tend to be subequal in width. In brun- nea, presacral III comprises 109 percent of the width of the sacral diapophyses, IV—89 percent, each of V and VI—79 percent, VII— 75 percent, and VIII—72 percent. The pattern is nearly the same in dominicensis except that presacral VIII is slightly wider than VII. Hyla septentrionalis demonstrates the strongest tendencies towards terrestrial modifications of the axial skeleton in this group; the transverse processes are more nearly equal in width (III—113% of sacral width, IV—100%, V—93% VI—87%, VII and VIII—85%). All three species are characterized by distal expansion of the transverse processes of presacral II and III. A minute anterior flange is present on the transverse process of presacral II in brunnea; a similar, but much larger flange is present in septentrionalis. The sacral diapo- physes are more broadly dilated in all three species than in vasta or lichenata. The angle of expansion is about 41° in both brunnea and dominicensis, and approximately 57° in septentrionalis. The corresponding anterior sacral angles are 70°, 74°, and 82° in sep- tentrionalis, dominicensis, and brunnea, respectively. The vertebral columns of the smaller hylids tend to show fewer specializations and greater uniformity of pattern than those of the larger frogs. In each of the four species, neural spines are absent and the width of the transverse processes of presacral III is greater than the width of the sacral diapophyses. The transverse processes lack flanges, and the sacral diapophyses are moderately dilated (43-48°). In pulchrilineata, which probably has the most general- ized axial structure of the groups, the width of presacral III com- prises 117 percent of the width of the sacral diapophyses. The widths of the transverse processes of the presacrals are subequal except those of VII and VIII (TMI—117%, IV—98%, V—85%, VI—80%, and VII and VIII—73%). The transverse processes of presacrals IT, III, and IV are slightly expanded distally. The angle between the anterior edge of the sacral diapophysis and the longitudinal axis of the body is 77°; the corresponding posterior angle is 55°. In the remaining three species (heilprini, wilderi, and marianae ), only the transverse processes of presacral IIT are expanded distally. 26 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Of the three, the axial morphology of heilprini is most similar to pulchrilineata, although the two are readily distinguishable. The presacrals of heilprini are clearly subequal in width (IIJ—111% of sacral width, [V—96%, V—87%, VI—8&6%, VII—80%, and VIII—75%). The sacral diapophysis expansion is nearly the same as pulchri- lineata (anterior angle, 77°; angle of expansion, 45°). Of the smaller species, marianae and wilderi show the highest degree of axial specialization. The relative widths of the transverse processes are slightly subequal in marianae (IIJ—110% of sacral width, IV— 97%, V—86%, VI—84%, VII—83%, and VIII—79%) and more nearly uniform in wilderi (III—105% of sacral width, IV—90%, and V-VIII —80%). The orientation and expansion of the sacral diapophyses is similar in both species. The anterior sacral angles are 75° and 80° in marianae and wilderi, respectively. Corresponding angles of sacral expansion are 43° and 48°. Pelvic osteology—tIn general, the pelvic structure of the West Indian hylids is quite generalized (Fig. 2C). The ilium and ischium are ossified and the pubis is cartilaginous or secondarily calcified in larger species. Dorsal acetabular expansion is minimal. Ilial protuberances are present, but moderately developed in all species. The ilium is devoid of dorsal crest and lateral expansions in all species except marianae in which a low crest is present. Values for relative ilial length (measured as a function of the length of the acetabular portion of the girdle) are pulchrilineata, 3.8; dominicensis, 3.7; septentrionalis and brunnea, 3.6; vasta, 3.5; lichenata, 3.3; wilderi, 2.9; and marianae, 2.1. One pattern emerges which apparently is independent of body size. Among the smaller frogs, the species with the least specialized vertebral column struc- ture (pulchrilineata) has the longest ilium. Similarly, the more generalized of the larger species (septentrionalis group) tend to have longer ilia than the more specialized vasta and lichenata. The septentrionalis group is characterized by obtuse angles between the ilial shaft and the preacetabular margin; that of vasta approaches 90°, whereas the angle is acute in lichenata. Hyla heilprini has an obtuse preacetabular angle; the angles of pulchrilineata and wilderi approximately 90° and that of marianae is acute. Throat musculature and vocal sac structure —All members of the West Indian hylid fauna have vocal sacs except marianae. The larger frogs are uniformly characterized by: 1) a posterolateral attachment of the fibers of the intermandibularis upon the ventral surface of the submentalis; 2) vocal sacs which are confined to a position above the interhyoideus muscle; and 3) presence of the adductor mandibulae externus superficialis. The septentrionalis- group is distinguished from vasta and lichenata by the differentia- tion of the intermandibularis to form a supplementary, apical ele- ment (Fig. 11). Most hylids, like vasta and lichenata, have an GREATER ANTILLEAN HYLID FROGS 27 Fic. 11. Superficial mandibular musculature of Osteopilus brunneus. Ab- breviations: imand. apic., supplementary apical element of M. intermandibu- laris. undifferentiated intermandibularis (Tyler, 1971). The interhyoid- eus muscle shows some variation in the development of lateral lobes. The lobes are absent in brunnea and lichenata; they are present and located posterior to the mandible in dominicensis and vasta. In septentrionalis, the lobes are present, small in size, and situated above the mandible. Similarly, the nature of the medial attachment of the intermandibularis muscle varies. In dominicensis and septentrionalis, the muscles meet at a tendinous raphe, whereas a slender aponeurosis is present in lichenata and brunnea, and a broad aponeurosis in vasta. In brunnea the vocal sac apertures are present as slits of mod- erate length, each being equivalent to approximately one-third of the length of the mandible. The remaining large species are char- acterized by vocal sac apertures which are small, circular orifices equivalent to one-eighth to one-tenth of the length of the mandible. This condition occurs infrequently in hylid frogs and represents a specialization over the moderate-sized slit. There are only two features common to all of the smaller species: (1) Absence of differentiation of the intermandibularis; (2) ab- sence of the adductor mandibulae externus superficialis. Postero- 28 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY lateral attachment of the fibers of the intermandibularis upon the ventral surface of the submentalis occurs in marianae, pulchri- lineata, and wilderi but not heilprini, where pulchrilineata is the only species lacking a median aponeurosis of the intermandibularis. The interhyoideus is least developed in marianae, which is unique in lacking a vocal sac, and most highly developed in pulchrilineata, in which there is a distinct, posterior, bilobular, supramandibular orientation. The remaining species have developed unilobular, postmandibular vocal sacs. In all species the vocal sac apertures are slit-like and approximately one-third of the length of the mandi- bles. Hyla heilprini is unique in possessing a dermal, submental gland comparable to that occurring in H. colymba, which is illustrated by Duellman (1970). Extrinsic musculature of the cloaca——The cloaca of all West Indian hylids is characterized by the presence of the protractor cloacae (Fig. 3). The muscle constitutes a bilateral extension of the ventral lip of the cloacal orifice, functionally permitting pro- traction of the orifice. The muscle occurs in all North American hylids examined, all casque-headed hylids, is present or absent in Hyla from Middle and South America, and does not occur in any from Australia and New Guinea. The only observed variation among the West Indian species is the inferior boundary of the protractor cloacae, and the nature of its attachments to the ischial ridge. In the larger species either a few muscle fibers or the median tendon extend to the ventral surface of the ischial ridge (Fig. 3). In the smaller species the muscle always terminates at the position posterior to the gracilis minor where the lateral halves of the protractor cloacae unite. From that site a very short tendon attaches to the posterior face of the ischial ridge. Breeding sites and larval morphology—The four Jamaican spe- cies (brunnea, lichenata, wilderi, and marianae) are known to breed only in bromeliads. None of the remaining species has been observed to utilize bromeliads for breeding. Hyla septentrionalis is apparently restricted to ponds and cisterns, whereas vasta and dominicensis utilize streams as well as pools. Hyla heilvrini breeds in torrential streams. No observations are recorded in the literature on either the breeding site or the larvae of pulchrilineata. The tadpoles of these frogs are diverse morphologically and easily distinguishable. The most generalized larvae are associated with the pond breeders. Noble (1927) reported that dominicensis laid its eggs in stagnant pools of rainwater in the forest or along the edge of a stream bank. The larva of dominicensis is character- ized by a short tail with well-developed fins, an interrupted labial disc and a 2/5 tooth row pattern. The tadpole of septentrionalis . ; 1 eo ny ha Se’ ae yh ee RP Ye Se Te ee GREATER ANTILLEAN HYLID FROGS 29 is similar, save for a 2/4 tooth row pattern. Hyla vasta lays its eggs in basins in gravel and stones on the edges of stream pools ( Noble, 1927); subsequent to hatching, the larvae move from the basins over wet stones into the stream. The larvae of vasta have a short, thick, muscular tail with deep fins, an uninterrupted labial disc, and minimally, a 4/6 tooth row pattern. The larvae of heilprini are the most obviously stream adapted with extremely thick, muscular tails, deep fins, uninterrupted labial disc and a 6/9 tooth row pat- tern. Relative to the non-bromeliad tapoles, the bromeliad larvae have longer tails, reduced fins, and reduced numbers of toothrows. The larvae of both brunnea and lichenata have incomplete labial discs and a tooth row pattern of 1/0; the tail of the lichenata larva is somewhat shorter than that of brunnea. The labial discs are complete in the larvae of marianae and wilderi; furthermore, tooth rows are absent and the lower beaks are unnotched in contrast to the remaining species. The tadpole of wilderi can be distinguished from that of marianae by the relatively shorter tail and presence of keratinous structures around the margin of the labial disc in the former. RELATIONSHIPS OF THE WEST INDIAN HYLIDS Our previous understanding of the West Indian hylids has been based principally on the work of Dunn (1926) and subsequent commentary by Noble (1927, 1931) and Williams et. al. (1963) who focused attention on the apparently complex and_ peculiar relationships of these frogs. Careful perusal of Dunn's (1926) discussion of the West Indian hylids yields several interesting ob- servations. Dunn considered brunnea, septentrionalis, and domini- censis to be closely related; he was the first to recognize them as the septentrionalis group. Similarly, he implied a relatively close relationship between wilderi and marianae. Dunn obviously was perplexed by the distributional symmetry of the small and large species (i.e., the occurrence of two large species in combination with two smaller species on both Jamaica and Hispaniola), in con- trast to the insular divergence of breeding habits (i.e., bromeliad breeders on Jamaica as opposed to the more generalized pond and stream breeders of Hispaniola). It is implicit, although not directly stated in his discussion, that Dunn was reluctant to assume more than one invasion of the West Indies by an ancestral hylid stock. Although the assumption is parsimonius in terms of historical zoo- geography, it severely restricted his corollary conclusions on the evolutionary relationships of the nine taxa. He implied that the septentrionalis group represented the most primitive derivative of an ancestral stock; thus, septentrionalis (sensu stricto) repre- sented a Cuban derivative, brunnea the Jamaican derivative, and dominicensis the Hispaniolan derivative. Dunn postulated that 30 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY lichenata, wilderi, and marianae represented derivative of brunnea, which diversified with respect to size while maintaining a special- ized arboreal mode of reproduction. He suggested that on His- paniola vasta, heilprini, and pulchrilineata evolved from a domini- censis-like ancestor. The Hispaniolan derivatives diversified with respect to size, but unlike their Jamaican counterparts, they also diversified with respect to breeding site (i.e., pond and stream) within the more generalized aquatic reproductive mode. Noble (1927) commented at some length on Dunn’s proposal in his discussion of the significance of life history data in the evolution of the Amphibia. He suggested that the tadpoles of the Hispaniolan hylids seemed to represent an orthogenetic series progressively modified to life in a mountain stream, thus strengthening Dunn’s argument in favor of a monophyletic origin for the group. Although, on the whole, Noble seemed to have agreed with Dunn's conclu- sions, he did question the high degree of parallelism involved in the evolution of the two insular faunas. On the basis of the comparative morphological evidence pre- sented herein, Dunn’s implied monophyletic origin of the West Indian hylids is an untenable hypothesis. The four smaller species— pulchrilineata, heilprini, marianae, and wilderi—are similarly dis- tinguished from the members of the septentrionalis-group and vasta and lichenata, by their size, generalized cranial structure, absence of dermal modification to the skull, and generalized pattern of throat musculature. Dunn (1926) suggested that marianae and wilderi might represent “neotenic” derivatives of a brunnea-like ancestor. Were this so, one would expect a striking resemblance in the pattern of osteological features between the smaller species and brunnea. Morphologically, brunnea unquestionably is distinct from marianae and wilderi; however, comparison of the two latter species suggests a close relationship between them. Myologically, marianae and wilderi differ in only three of the ten characters considered. Both are characterized by skulls with greatly reduced ossification and bluntly rounded snouts. The species share similar body pro- portions (Table 1), many features of soft morphology, and axial specializations such as short ilia, obtuse pelvic angles, expanded sacral diapophyses and transverse processes of nearly uniform width. Both are inhabitants of the central highlands of Jamaica, where they breed in bromeliads. Hyla marianae inhabits arboreal bromeliads in deciduous forest; its known altitudinal range is 122 to 895 m (Schwartz and Fowler, 1973). In contrast, the smaller species, Hyla wilderi, is known to inhabit both terrestrial and ar- boreal bromeliads in dense and scattered woodlands at elevations between 122 and 610 m (op. cit.). There seems to be no evidence linking either pulchrilineata or heilprini to marianae and wilderi. Osteologically, both Hispaniolan GREATER ANTILLEAN HYLID FROGS dl species are far more generalized than the Jamaican species; they are characterized by longer ilia, vertebral processes of subequal length, and more heavily ossified skulls. The resemblance of pulchri- lineata and heilprini is limited to an absence of arboreal speciali- zations in these species. Cranially, pulchrilineata is distinctly dif- ferent from any other West Indian hylid. Hylid heilprini is uniquely distinguished by its dermal submental gland and _ specializations adapting it to life along mountain streams of Hispaniola; included among these specializations are the presence of a propollical spine, well-developed palmar and plantar webbing, and extensive devel- opment of subarticular and supernumerary tubercles. Dunn was correct in his assessment of septentrionalis, brunnea and dominicensis. These three species comprise a closely related complex of moderately large frogs characterized by uniquely spe- cialized throat musculature (differentiated intermandibularis mus- cle). The skulls are heavily ossified (exostosed, casqued, and co- ossified ) and share many structural details in common, including presence of a dermal sphenethmoid and similar prevomerine mor- phology. In contrast to the specializations of the skull, the axial osteology of these three species is quite generalized. Sacral dia- pophysis expansion is moderate, and the transverse processes are distinctly subequal in length. The ilia of each of these species are long and the pelvic angle is obtuse. Although it is not at once obvious, lichenata and vasta are more similar to one another in many respects than either is to the sep- tentrionalis group. Both are extremely large frogs, sharing a gen- eralized pattern of throat musculature (i.e., differentiated inter- mandibularis muscle) and many arboreal specializations. The latter include dermal fringes along the outer margins of the fore- and hind limbs, and certain characteristics of the axial and pelvic skeleton. The ilia are somewhat shorter than those of the septen- trionalis group and the anterior pelvic angle is acute or perpen- dicular, instead of obtuse. These two species are unique among the West Indian hylids in having very slightly dilated sacral dia- pophyses, the total width of which is greater than that of the third presacral vertebrae. Furthermore, in both lichenata and vasta, the transverse processes of the vertebrae are long and nearly uniform in length. We are not implying that these two species are closely related; there is a host of morphological differences distinguishing them. On the other hand, the structural characteristics which they share make it seem highly unlikely that either species could have been derived from the septentrionalis group as proposed by Dunn. The most distinctive feature of lichenata is its heavily casqued, co- ossified skull. Although this would seem to indicate a relationship to the septentrionalis group, the basic cranial architecture is dis- tinctly different and certainly not derived from the pattern of these 32 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY frogs; furthermore, the undifferentiated intermandibularis sets it further apart from members of that group. Similarly, the vacuous, poorly ossified skull of vasta could only have arisen from the sep- tentrionalis group by heterochronous arrestment of bone deposition, which is not commensurate with the size attained by this species. On the basis of the accumulated evidence, there seem to be minimally six phylogenetic lines represented among the West Indian hylids—three having given rise to the four smaller species and three others ancestral to the five larger species. Of the smaller species, marianae and wilderi are the only species for which a common ancestor can be postulated. Unfortunately, at this time there is no clear evidence linking wilderi and marianae to any par- ticular group of hylids currently known from South or Central America. Although both species are highly specialized for an arboreal mode of existence, marianae is the more generalized. It is larger, and possesses fewer morphological specializations than wilderi. Presumably, the ancestral stock was similar to marianae— a moderate-sized bromeliad breeder inhabiting the central high- lands of Jamaica. Hyla marianae may represent a species closely related to the ancestral stock; the only obvious specializations of marianae are loss of the vocal sac and vocal slit aperture, and re- duction of ilial length. In contrast, wilderi is clearly a more highly derived species which has retained a vocal sac and become adapted to life in smaller bromeliads. The transverse processes of the verte- brae are more uniform in width than those of marianae, but the ilia are somewhat longer. The skulls of the two species are nearly identical, save for the reduced ossification of wilderi, which can be interpreted as developmental arrestment associated with overall reduction of size in this species. Palmar and plantar webbing is reduced in wilderi, but retained in marianae. The nuptial excres- cense of marianae is a flat horny pad, whereas that of wilderi is modified into a patch of keratinous spines. The larvae of both are highly specialized for their existence in bromeliads. The tadpole of marianae has a somewhat longer, slimmer tail with more reduced caudal fins than wilderi, but the larval mouth of the latter is much smaller and characterized by a distinctive ring of keratinous spines at its perimeter (Dunn, 1926). The presence of pulchrilineata and heilprini on Hispaniola must result from two separate invasions of that island by forms unrelated to one another and the marianae-wilderi complex. Although noth- ing has been reported concerning the breeding habits and larvae of pulchrilineata, the species seems to represent a generalized, wide- spread Hispaniolan hylid. Hyla heilprini, on the other hand, is a specialized stream-side inhabitant whose morphological characters are highly reminiscent of the wide-spread, South American Hyla GREATER ANTILLEAN HYLID FROGS 33 albomarginata group (see Duellman, 1970, for a résumé of this species group ). The septentrionalis group, vasta, and lichenata seem to us to represent three distinct phylogenetic lines. Morphologically, Hyla vasta seems to be related to the Hyla boans group of Central and South America. Members of the boans group are large species in- habiting lowland forests, and like vasta, they are characterized by extensive digital webbing. They have moderately or poorly ossified skulls which are very similar in structure to that of vasta (see Duellman, 1970, for a résumé of this species group). The vertebral columns in this group tend to have uniform transverse processes and narrowly dilated sacral diapophyses similar to those of vasta. The intermandibularis muscle always bears a vast, median aponeu- rosis which represents a derived condition as in vasta. Perhaps the most intriguing similarity involves breeding behavior. Males of some members of the boans group (boans, faber, and rosenbergi) build stream-side, water-filled nests of mud or gravel in which eggs are deposited and tadpoles develop. At localities where the sub- strate is not amenable to these nest building habits, boans has been observed (Duellman, pers. com.) to utilize water-filled depressions adjacent to streams exactly as reported for vasta by Noble (1927). Hyla brunnea, dominicensis, and septentrionalis form a very closely related species group characterized by their moderately large size and casqued, co-ossified crania. As a group, they are distinguished from vasta and lichenata by their differentiation of the intermandibularis to form an apical element, their obtuse pelvic angles, and generalized vertebral column structure in which the sacral diapophyses are narrower than the third presacral. These frogs seem to be highly adaptable, ecological generalists occurring over a wide elevational range (0-1500 m), in a variety of habitats. Although brunnea apparently breeds only in bromeliads, septen- trionalis and dominicensis breed wherever standing water is avail- able. Of the three species, septentrionalis and dominicensis are the most closely allied and probably more closely related to an ancestral type than brunnea. Externally, septentrionalis and dominicensis are almost indistinguishable, differing only slightly with regard to skin tuberculation, presence of a tarsal fold and less digital webbing in septentrionalis. The skulls of the two species differ in only minor details involving the shape of the occipital margin, orientation of the parasphenoid alae, and articulation of the medial rami of the pterygoids. The vertebral column of septentrionalis tends to have transverse processes of more uniform width than dominicensis. Hyla brunnea differs distinctly from both septentrionalis and do- minicensis in its cranial architecture and throat myology. Osteologically, the septentrionalis group seems to be most closely 34 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY allied with Osteocephalus, a widespread Amazonian genus recently reviewed by Trueb and Duellman (1971). Members of this genus tend to be moderately large, lowland generalists, which are pri- marily found at elevations below 500 meters, but do ascend the Amazonian slopes of the Andes to elevations of 1800 meters. Like the septentrionalis group, members of Osteocephalus are charac- terized by broad skulls which are exostosed with a tendency towards co-ossification. The basic cranial architecture of the two groups is the same; many details are similar, including the patterns of dermal sculpturing, the pterygoids, prevomers and dermal spheneth- moids. The body proportions are similar, as is the hand and foot morphology. Two principal differences prevail between these two groups. Osteocephalus, like the majority of Neotropical hylids, has an undifferentiated intermandibularis muscle. Unlike most other hylids, including the septentrionalis group, the vocal sacs of Osteocephalus are paired, posterior in position and when inflated, protrude posteroventral or posterolateral to the angles of the jaws. It has been suggested that both paired vocal sacs and the develop- ment of a supplementary apical slip to the intermandibularis repre- sent evolutionary specializations which are restricted in occurrence (Tyler, 1971). In view of the similarities of Osteocephalus and the septentrionalis group, it seems plausible to suggest that they may have had a common South American ancestor characterized by generalized throat musculature and moderately heavy cranial ossi- fication with a tendency toward exostosis and casquing. Such an ancestor may have invaded the West Indies and evolved the de- rived pattern of throat musculature characteristic of the septen- trionalis group. Subsequent isolation and evolution of an ancestral stock on Jamaica, Hispaniola, and Cuba may well have given rise to each of these three closely related species. The extra- and intra-Antillean relationships of lichenata are perplexing. The species shares a great many external features and characters of axial morphology as well as some features of the vocal apparatus with vasta, as pointed out above. However, the cranial architecture of lichenata is completely unlike that of vasta. Furthermore, these two species are widely divergent ecologically; lichenata is a tree cavity dweller which breeds exclusively in brome- liads. The skull of lichenata is most similar to those of the genus Trachycephalus, a South American group which apparently is allied closely with the generalized and widespread Osteocephalus (Trueb, 1970). The similarities of cranial structure of lichenata and Trachy- cephalus are striking. Both have broad skulls which are heavily casqued and co-ossified with the same pattern of dermal sculptur- ing. The prevomers are similar, bearing teeth on curved elevations. The palatines are well developed, and each bears a prominent transverse ridge. The anterior pterygoid ramus has a broad max- GREATER ANTILLEAN HYLID FROGS 35 illary articulation which tends to extend to the midlevel of the orbit, and the parasphenoid alae are perpendicularly oriented to the long axis of the bone. The principal area of difference involves cranial ossification. The dermal roofing bones of Trachycephalus are heavily ossified; consequently internal suspensory and bracing elements (pterygoid and palatine) tend to be reduced (e.g., no medial articulation of the pterygoid). In contrast, the dermal roof- ing elements of lichenata are less well developed and the palatine and pterygoid are robust, fully articulated bones. Furthermore, the cultriform process of the parasphenoid is exceedingly short in lichenata (see Trueb, 1970, for figure and osteological descriptions of Trachycephalus). The vertebral columns of Trachycephalus are characterized by subequal lengths of transverse processes and widely expanded sacral diapophyses in contrast to lichenata. Like the latter species, the intermandibularis muscle is undifferentiated in Trachycephalus, but basically similar to Osteocephalus and, un- like lichenata, the vocal sacs are paired and lateral. In view of these conflicting suites of characters, the following conclusions can be drawn concerning the relationships of lichenata. 1) Because of differences in throat myology and axial morphology (see Analysis of Characters), lichenata cannot be derived from the septentrionalis group. To assume that lichenata may have been derived from an ancestral stock of this group would require that either the differentiated intermandibularis was independently evolved three times or that lichenata developed an undifferentiated intermandibularis from the specialized, differentiated muscle. Neither assumption is especially parsimonius. 2) The structure of the vocal sac apparatus in lichenata precludes its being derived from either Osteocephalus or Trachycephalus; however, the struc- ture of the intermandibularis links lichenata more closely with these genera than to members of the septentrionalis group. Apparently, lichenata represents a separate invasion of the West Indies by a primitive stock related to the Osteocephalus-septen- trionalis complex. Although both the septentrionalis and lichenata ancestral lines retained the primitive subgular vocal sacs, the an- cestral septentrionalis line was distinguished by differentiation of the intermandibularis. The resemblances of the crania of Trachy- cephalus and lichenata may be accounted for by a common ancestral stock and parallel evolution of the frogs which have taken up a similar arboreal mode of existence. SYSTEMATIC ACCOUNTS The close affinities of the members of the septentrionalis group and their distinctiveness from all other Neotropical hylids suggest that these frogs have undergone considerable evolution in the iso- lation of the Greater Antilles and should be warranted generic 36 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY recognition apart from Hyla. Fitzinger (1843) applied the generic name Osteopilus to Trachycephalus marmoratus Dumeril and Bib- ron. Because Osteopilus is the earliest available name, the mem- bers of the septentrionalis group are herewith allocated to this genus. Osteopilus Fitzinger, 1843 Osteopilus Fitzinger, 1843:30 [Type species ——Trachycephalus marmoratus Bibron, 1842, in Ramon de la Sagra (=Hyla septentrionalis Duméril and Bibron), by monotypy; nec Hyla marmorata (Laurenti, 1768 )]. Diagnostic definition—1) Skull slightly longer than broad, or as long as broad; 2) dermal roofing bones of skull well ossified, exostosed, and co-ossified; 3) prenasal and internasal bones absent; 4) dermal sphenethmoid present; 5) dentigerous processes of pre- vomers large, curved, moderately to closely positioned, and bearing a single row of teeth; 6) cultriform process of parasphenoid of normal length, terminating anteriorly just posterior to the level of the palatines; 7) vocal sac single and subgular, confined to position above interhyoideus muscle; 8) submentalis muscle moderate in size and araphic; 9) intermandibularis muscle differentiated to form supplementary apical elements; 10) fibers of intermandibularis muscle attach to posterolateral portions of submentalis muscle; 11) protractor cloacae muscle fibers extend inferiorly to ventral surface of ischial ridge; 12) tympanum large, 63-77 percent of diameter of eye; 13) finger and toe discs large and round; 14) nup- tial excrescences present in breeding males; 15) inner metatarsal tubercle not modified for digging; 16) no fringes on hind or fore- limbs; 17) total width of sacrum less than total width of presacral vertebra III; 18) sacral diapophyses moderately dilated; 19) trans- verse processes of posterior presacral vertebrae subequal in width; 20) angle between ilial shaft and preacetabular margin obtuse. Content.—As defined here, the genus contains three known spe- cies—Osteopilus brunneus (Gosse), O. dominicensis (Tschudi), and O. septentrionalis (Duméril and Bibron). Description.—Refer to section on comparative morphology. Distribution—The Greater Antilles, including Cuba, Jamaica, Hispaniola, Puerto Rico, and Isle of Pines, southern penisular Florida, and the Bahama islands. The genus is most common at lowland localities, although it does occur at moderate elevations (1500 m) in the highlands of Jamaica and Hispaniola. Osteopilus brunneus (Gosse) new combination Hyla brunnea Gosse, 1851:361 [Type not designated; type locality Savanna-la- Mar, Parish of Westmoreland, Jamaica]. Trachycephalus scutigerus Cope, 1863:45 [Holotype—United States National Museum 6268 (now apparently lost) from “Jamaica”; Prof. C. B. Adams, collector]. Synonymy fide Dunn (1926:125). Hyla septentrionalis brunnea—Barbour, 1937:81. GREATER ANTILLEAN HYLID FROGS 37 Diagnosis——1) Size moderate, sexual dimorphism extreme; maximum observed snout-vent length in males 48.1 mm (mean 45.5 mm), in females, 70.0 mm (mean 60.5 mm); 2) skin on dorsum smooth except for scattered, small, round tubercles on forearm; 3) ventral skin finely granular on posterior half of abdomen; scat- tered, small tubercles around cloaca; 4) fingers one-fourth webbed; 5) numerous tiny supernumerary tubercles present on hand; 6) outer palmar tubercle absent; 7) nuptial excrescence of males a proliferation of small, keratinous granules; 8) tarsal fold absent; 9) toes three-quarters webbed; 10) moderate number of plantar supernumerary tubercles; 11) nostrils slightly protuberant; 12) skull longer than broad; 13) parasphenoid alae posterolaterally oriented to longitudinal axis of skull; 14) medial ramus of ptery- goid narrowly separated from parasphenoid ala and with firm articulation with otic capsule; 15) anterior ramus of pterygoid articulating with maxillary at mid-level of orbit; 16) prevomerine dentigerous processes strongly curved, narrowly separated medially; 17) sacral diapophyses moderately dilated (ca. 41°; anterior sacral angle ca. 82°; posterior sacral angle ca. 57°); 18) interhyoideus muscle without lobes; 19) intermandibularis muscle with slender, median aponeurosis; 20) vocal sac aperture a moderate slit; 21) anterior cornu of hyoid lacking anterolateral lobe; 22) larvae with long tail, reduced caudal fins, incomplete labial disc, notched lower beak, and toothrow pattern of 1/0. Osteopilus brunneus can be distinguished readily from the other two species in the genus by the shape of its snout, which is clearly truncate in lateral profile, terminal nostrils, smooth gular skin, finely granular abdominal skin, and absence of clearly defined transverse bars on the forearms. Distribution —This species is endemic to Jamaica where it is widespread at elevations below 1500 m. Remarks.—Although Osteopilus brunneus has been mentioned frequently in the literature, useful accounts are few and widely scattered. Gosse (1851:361) provided notes on the natural history of the species; these were summarized and augmented by Barbour (1910:288) and Dunn (1926:125). Dunn described the call, pre- ferred habits and sites of egg deposition, and figured the larvae and larval mouthparts (1926: Pl. 1, Fig. 2;,Pl) 2, Fig: 2). The best systematic accounts are those of Barbour (loc. cit.) and Lynn, in Lynn and Grant (1940:19-22). Included in the latter is information on colors in preservative and in life and their variation, comments on size, variation, and statements of habits, habitat, and distribution. Lynn, in Lynn and Grant (1940:Pl. 1) also provided semidiagram- matic drawings of brunneus. The species was illustrated also by Noble (1931:89, Fig. 28B). Discussion and comment on the rela- tionships of brunneus are available in Dunn (1926), Noble (1927), 38 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY and Duellman (1970). The most recent account of brunneus is provided by Schwartz and Fowler (1973). Although they acknow]- edged that specimens of this species are found most frequently in bromeliads, the authors also reported collecting individuals from leaves of shrubs and sides of trees (up to 2.1 m above the ground), and from the ground itself. The maximum snout-vent length they observed in male brunneus is 50.6 mm (mean, 45.6 mm; n=25), and in female brunneus, 78.5 mm (mean, 65.7 mm; n=25). Osteopilus dominicensis (Tschudi) new combination Hypsiboas dominicensis Tschudi, 1838:30 [Syntypes—Museum National d’His- toire Naturelle (Paris) 4614 (four specimens) from “Saint-Dominque’; presented by M. Alexandre Ricord]. Trachycephalus dominicensis—Duméril and Bibron, 1841:540. Trachycephalus ovatus Cope, 1863:44 [Holotype—Museum of Comparative Zoology (Harvard) 1518 from near Jérémie, Haiti; purchased from D. F. Weinland, 1859]. Synonymy fide Boulenger (1882:369). Hyla ovata—Boulenger, 1882:369 (part). Hyla dominicensis—Boulenger, 1882:370 (part). Hyla ( Trachycephalus ) ovata—Fischer, 1888:44. Hyla ( Trachycephalus ) dominicensis—Meerwarth, 1901:40. Hyla septentrionalis dominicensis—Barbour, 1937:81. Hyla dominicensis dominicensis—Mertens, 1938:332. Hyla dominicensis—Cochran, 1941:13. Diagnosis—1) Size moderately large, sexual dimorphism ex- treme; maximum observed snout-vent length in males 63.8 mm (mean 57.8 mm), in females 98.7 mm (mean 76.8 mm); 2) skin on dorsum smooth; 3) ventral skin of throat, abdomen, and postero- ventral surfaces of thighs coarsely granular; 4) fingers one-fourth webbed; 5) numerous, tiny supernumerary tubercles present on hand; 6) outer palmar tubercle present; 7) nuptial excrescence of males a flat, keratinous pad; 8) tarsal fold absent; 9) toes three- quarters webbed; 10) few plantar supernumerary tubercles; 11) nostrils slightly protuberant; 12) skull slightly longer than broad; 13) parasphenoid alae perpendicularly oriented to longitudinal axis of skull; 14) medial ramus of pterygoid narrowly separated from parasphenoid, but not articulating with otic capsule; 15) anterior ramus of pterygoid articulating with maxillary at posterior level of orbit; 16) prevomerine dentigerous processes moderately curved, narrowly separated medially; 17) sacral diapophyses moderately dilated (ca. 41°; anterior sacral angle ca. 74°; posterior sacral angle ca. 65°); 18) interhyoideus muscle with postmandibular lobes; 19) intermandibularis muscle without median aponeurosis; muscles meet at tendinous raphe; 20) vocal sac aperture a small orifice; 21) anterior cornu of hyoid with anterolateral lobe; 22) larvae with short tail, extensive caudal fins, incomplete labial disc, notched lower beak, and tooth-row pattern of 2/5. Osteopilus dominicensis can be distinguished from O. brunneus GREATER ANTILLEAN HYLID FROGS 39 by the shape of its snout, which is sloping in lateral profile, and by the coarse granularity of its gular and abdominal skin. It differs from septentrionalis in having more webbing between the fingers and toes, and from both septentrionalis and brunneus in the pres- ence of a distinct outer palmar tubercle. Distribution.—This species is endemic to Hispaniola where it is found from the coastal lowlands to moderate elevations (up to 1500 m) in the foothills and on the plateaus. Remarks.—Although dominicensis was named by Tschudi (1838), the first useful description to appear was that of Duméril and Bibron (1841:540-541). Save for an illustration of the adult (Noble, 1931:89, Fig. 28 D) and the larva and larval mouthparts (Noble, 1927, Pl. IX, Fig. A), no new information was published on dominicensis until Mertens (1939:34-35) reported on color variation, distribution, habitat, life history, and larval character- istics. A brief account of this species, including the description of an adult female, remarks on variation in adults and a diagnostic series of illustrations was provided by Cochran (1941:13-17, Fig. 3). Lynn (1958:154-155) commented on color variation in adults and the production of irritating skin secretions by dominicensis, and Williams, et. al. (1963:315) reported phragmotic behavior in the species. Egg deposition and tadpoles were described by Noble (1927:95). Discussion of the relationships of dominicensis are available in Dunn (1926), Noble (1927), Myers (1950), and Duell- man (1970). Osteopilus septentrionalis (Duméril and Bibron) new combination Trachycephalus marmoratus Duméril and Bibron, 1841:538. [Holotype.—— Museum National d'Histoire Naturelle (Paris) 4612, adult female from “Cuba”; donated by Ramon de Ja Sagra and Henri Delaroche]. Preoccu- pied by Hyla marmorata ( Laurenti, 1768:29). Hyla_ septentrionalis Duméril and Bibron, 1841:538. Substitute name _ for Trachycephalus marmoratus. Trachycephalus insulsus Cope, 1863:43 [Syntypes.—United States National Museum 12166 (adult male) and 167237 (adult female, formerly USNM 6266) from “Cuba”; collected by Felipe Poey]. Synonymy fide Boulenger (1882:368). Trachycephalus wrightii Cope, 1863:45 [Holotype——United States National Museum 5174 (adult female) from “District of Guantanamo, southeastern Cuba” (Oriente Province); collected by Charles Wright]. Synonymy fide Boulenger (1882:368 ). Hyla schebestana Werner, 1917:36 [Holotype.—presumably destroyed in World War II, formerly Zoologisches Museum (Hamburg), no number, an adult female from “Cuba”; donated by “Herrn Apotheker Sebesta”]. Synonymy fide Duellman and Cromble (1970:1). Hyla septentrionalis septentrionalis—Barbour, 1937:81. Hyla dominicensis septentrionalis——Mertens, 1938:333. Hyla dominicensis insulsa.—Mittleman, 1950:26. Hyla dominicensis.—Peterson, Garrett, and Lantz, 1952:63. Hyla insulsa.—Jaume, 1966:15. Hyla septentrionalis—Duellman and Crombie, 1970:92.1. 40 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Diagnosis —1) Size moderately large, sexual dimorphism ex- treme; average snout-vent length in males 55.3 mm, in females, 75.2 mm; 2) skin on dorsum with scattered tubercles; 3) ventral surfaces granular, tubercles absent around cloaca; 4) fingers basally webbed; 5) moderate number of low, round supernumerary tu- bercles present on hand; 6) outer palmar tubercle absent; 7) nup- tial excrescence of males a flat, horny pad; 8) weak tarsal fold present; 9) toes about two-thirds webbed; 10) few plantary super- numerary tubercles; 11) nostrils slightly protuberant; 12) skull slightly longer than broad; 13) parasphenoid alae posterolaterally oriented to longitudinal axis of skull; 14) medial ramus of pterygoid widely separated from parasphenoid and bearing only a weak articulation with the otic capsule; 15) anterior ramus of pterygoid articulating with maxillary at posterior level of orbit; 16) prevom- erine dentigerous processes moderately curved and separated me- dially; 17) sacral diapophyses dilated (ca. 57°; anterior sacral angle ca. 70°; posterior sacral angle ca. 53°); 18) interhyoideus muscle with small, supramandibular lobes; 19) intermandibularis muscle without median aponeurosis; muscles meet as tendinous raphe; 20) vocal sac aperature a small orifice; 21) anterior cornu of hyoid with anterolateral lobe; 22) larvae with moderately long tail, extensive caudal fins, incomplete labial disc, notched lower beak, and tooth-row pattern of 2/4. Osteopilus septentrionalis is distinguished from both domini- censis and brunneus by its tuberculate dorsal skin and basal web- bing between the fingers. The species differs from brunneus by the shape of the snout, which is sloping in lateral profile in septen- trionalis. It differs from brunneus by the shape of the snout, which is sloping in lateral profile in septentrionalis. It differs from do- minicensis by the absence of an outer palmar tubercle. Distribution.—The species is widespread at low and moderate elevations throughout Cuba and on the Isle of Pines. Osteopilus septentrionalis also occurs on the Cayman Island (Grand Cayman, Little Cayman, and Cayman Brac) and on some of the Bahama Islands northwest of the Mayaguana Passage (Little Bahama Bank, Grand Bahama Bank, San Salvador, Rum, Crooked, and Acklin’s islands). The species occurs on southern, peninsular Florida and has been introduced to Great Inagua Island and Puerto Rico. Remarks.—A complete review of the nomenclatural history, and pertinent literature including a description of the species, its fossil record and distribution was provided by Duellman and Crombie (1970:92.1-92.4). See Cochran (1961) for black and white and color photographs of septentrionalis. It is clear that lichenata is a highly specialized and distinctive frog which is neither closely allied to any member of the West GREATER ANTILLEAN HYLID FROGS 41 Indian hylid fauna nor any known representative from Central or South America. Because of its peculiar characteristics and endem- ism to Jamaica, we feel that this species should be accorded generic recognition. Calyptahyla new genus Type species —Trachycephalus lichenatus Gosse, 1851. Diagnostic definition—1) Skull broader than long; 2) dermal roofing bones of skull well ossified, exostosed, and co-ossified; 3) prenasal and internasal bones absent; 4) dermal sphenethmoid present; 5) dentigerous processes of prevomers large, straight, nar- rowly separated medially and bearing a single row of teeth; 6) cul- triform process of parasphenoid short, terminating anteriorly at mid-level of orbit; 7) vocal sac single and subgular; confined to position above interhyoideus muscle; 8) submentalis moderate in size and araphic; 9) intermandibularis muscle not differentiated; 10) fibers of intermandibularis muscle attach to posterolateral part of submentalis muscle; 11) protractor cloacae muscle fibers extend inferiorly to ventral surface of ischial ridge; 12) tympanum large, 68 percent or more of diameter of eye; 13) finger and toe discs large and round; 14) presence or absence of nuptial excrescences in males unknown; 15) inner metatarsal tubercle not modified for digging; 16) fringes present on hind and forelimbs; 17) total width of sacrum greater than total width of presacral vertebra III; 18) sacral diapophyses narrowly dilated; 19) transverse processes of posterior presacral vertebrae uniform in width; 20) angle between ilial shaft and preacetabular margin acute. Content.—One species, Trachycephalus lichenatus Gosse. Description.—Refer to section on comparative morphology. Distribution.—This genus is endemic to Jamaica. Etymology.—The generic name is derived from the Greek word kalypto, meaning to conceal or hide, and the Greek name Hylas from which the vocative Hyla is derived. Thus the name Calypta- hyla (feminine) alludes to the secretive nature of this frog which has been heard to call so often in the dense woodlands of Jamaica and has been collected so infrequently. Calyptahyla lichenata (Gosse) new combination Hyla crucialis Harlan, 1826:64 [Type not designated; type locality “Jamaica”’]. Nomen oblitum. Trachycephalus lichenatus Gosse, 1851:362 [Type not designated; type local- ity “the summit of Bluefields Mountain, Jamaica” ]. Trachycephalus anochlorus Gosse, 1851:364 [Type not designated; type lo- cality “Jamaica”]. Synonymy fide Boulenger (1882:370). Hyla lichenata.—Boulenger, 1882:370. Diagnosis —1) Size large, sexual dimorphism extreme; maxi- mum observed snout-vent length in males, 92.6 mm, in females, 122 mm; 2) skin on dorsum heavily tuberculate; 3) ventral skin 42 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY granular with paracloacal folds; 4) fingers one-fourth webbed; 5) moderate number of low, round supernumerary tubercles on hand; 6) outer palmar tubercle absent; 7) presence or absence of nuptial excrescence unknown; 8) tarsal fold absent; 9) toes three- fourths webbed; 10) plantar supernumerary tubercles few in num- ber; 11) nostrils slightly protuberant; 12) skull broader than long; 13) parasphenoid alae perpendicularly oriented to longitudinal axis of skull; 14) medial ramus of pterygoid articulating with otic capsule and distal edge of parasphenoid ala; 15) anterior ramus of pterygoid articulating with maxillary at mid-level of orbit; 16) prevomerine dentigerous processes straight, oriented at a slight angle to one another; and narrowly separated medially; 17) sacral diapophyses narrowly dilated (ca. 31°; anterior sacral angle ca. 81°; posterior sacral angle ca. 68°); 18) interhyoideus muscle without lobes; 19) intermandibularis muscle with slender, median aponeurosis; 20) vocal sac aperture a small orifice; 21) anterior cornu of hyoid without anterolateral lobe; 22) larva with relatively short tail, reduced caudal fins, incomplete labial disc, notched lower beak and tooth-row pattern of 1/0. Calyptahyla lichenata can be distinguished from any other known species of Neotropical hylid by the following combination of characters: 1) single subgular vocal sac; 2) rugosely sculptured, co-ossified cranium lacking labial flanges; and 3) heavily tuberculate dorsum. Distribution—This species is endemic to Jamaica and widely distributed in dense forest throughout the island at low and mod- erate elevations. Remarks.—Little is known about this immense casque-headed frog. The most complete account of the natural history and habits of lichenata was given by Dunn (1926:127-129). His descriptions were based largely on observations of Mr. E. Stuart Panton who described the call of lichenata and noted the preferred perch of this species in hollow trees or hollow branches of trees. The tad- pole and larval mouthparts were illustrated by Dunn (1926:PI. 1, Fig. 1; Pl. 2, Fig. 1). The adult was figured by Noble (1931:89; Fig. 28 A). The best illustrations and only complete systematic account available is that of Lynn, in Lynn and Grant (1940:22-24, Pl. II, Fig. 3). Included in the latter work are complete descrip- tions, comments on variation, and summaries of earlier literature. Discussion of the relationships of Calyptahyla lichenata may be found in Dunn (1926), Noble (1927), Myers (1950), and Duell- man (1970). It should be noted that an application has been sub- mitted to the International Commission on Zoological Nomenclature (Trueb, 1972) requesting suppression of the specific name crucialis, a nomen oblitum of 145 years. For the most recent account of this species refer to Schwartz and Fowler (1973). GREATER ANTILLEAN HYLID FROGS 43 The remaining five species of Greater Antillean hylids seem to be allied closely with the mainland Hyla. This suggests that either these frogs are more recent derivatives of mainland ancestors, or that in isolation, evolutionary change has had less dramatic effects on these phylogenetic lines than on Osteopilus and Calyptahyla. We think that until new evidence suggests otherwise, vasta, mari- anae, wilderi, heilprini, and pulchrilineata should be retained in the genus Hyla. A diagnostic definition of the Greater Antillean Hyla is provided below to distinguish members of this group from Osteopilus and Calyptahyla. For a synonymy of the genus Hyla, refer to Duellman (1970:173), whose arrangement has been modi- fied only by the resurrection of Litoria Tschudi (Tyler, 1971). Hyla Laurenti, 1768 Diagnostic definition——1) Skull longer than broad, as long as broad, or broader than long; 2) dermal roofing bones poorly to moderately well ossified; not exostosed, casqued or co-ossified; 3) prenasal and internasal bones absent; 4) dermal sphenethmoid absent; 5) dentigerous processes of prevomers widely separated or not, large or small, curved or straight, usually bearing a single row of teeth; 5) cultriform process of parasphenoid of normal length, terminating anteriorly between the mid-level of the orbit and the level of the palatines; 7) if present, vocal sac single and subgular; 8) submentalis moderate in size and araphic; 9) intermandibularis not differentiated; 10) fibers of intermandibularis with or without attachment to posterolateral part of submentalis muscle; 11) pro- tractor cloacae muscles extending inferiorly, or not, to ventral surface of ischial ridge; 12) tympanum moderately large, 50.5-61.0 percent of diameter of eye; 13) finger and toe discs large and round; 14) nuptial excrescences usually present in breeding males; 15) inner metatarsal tubercle not modified for digging; 16) no fringes on hind or forelimbs; 17) total width of sacrum more or less than total width of presacral vertebra III; 18) sacral diapophyses slightly or moderately dilated; 19) transverse processes of posterior presacral vertebrae uniform or subequal in width; 20) angle be- tween ilial shaft and preacetabular margin varying from obtuse to acute. Content.—As defined here, Hyla includes five Greater Antillean species—Hyla vasta Cope, H. pulchrilineata Cope, H. heilprini Noble, H. marianae Dunn, and H. wilderi Dunn. Description.—Refer to section on comparative morphology. Distribution.—Within the Greater Antilles, members of this genus are known only from Jamaica (marianae and wilderi) and Hispaniola (vasta, pulchrilineata, and heilprini). 44 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Hyla vasta Cope Hyla vasta Cope, 1871:219 [Holotype-——Academy of Natural Sciences (Phila- delphia) 2097 from “Haiti, San Domingo”; collected by Dr. William M. Gabb]. Diagnosis—1) Size large, sexual dimorphism extreme; maxi- mum observed snout-vent length in males 108.8 mm (mean 96.7 mm), in females 141.9 mm (mean 136.9 mm); 2) skin on dorsum strongly tuberculate; 3) ventral skin coarsely granular; tubercles around cloaca; 4) fingers three-fourths webbed; 5) very few, in- distinct supernumerary tubercles on hand; 6) outer palmar tubercle present; 7) nuptial excrescence in breeding males a flat, keratinous pad; 8) tarsal fold absent; 9) toes fully webbed; 10) plantar super- numerary tubercles absent; 11) nostrils strongly protuberant; 12) skull broader than long; 13) parasphenoid alae posterolaterally oriented to longitudinal axis of skull; 14) medial ramus of pterygoid articulating with otic capsule, widely separated from parasphenoid ala; 15) anterior ramus of pterygoid articulating with maxillary near mid-level of orbit; 16) prevomerine dentigerous processes large, slightly curved, having a moderate medial separation and a single row of teeth medially and a double row laterally; 17) sacral diapophyses narrowly dilated (ca. 20°; anterior sacral angle ca. 103°; posterior sacral angle ca. 57°); 18) interhyoideus muscle with postmandibular lobes; 19) intermandibularis muscle with broad, median aponeurosis; 20) vocal sac aperture a small orifice; 21) anterior cornu of hyoid without anterolateral lobe; 22) larva with short, thick tail, extensive caudal fins, complete labial disc, notched lower beak, and tooth-row pattern of 2/5. Hyla vasta is distinguished easily from all other West Indian hylids by its size, tuberosity, and lack of a casqued, co-ossified cranium. It is unique among the Greater Antillean Hyla in having a sacrum which is wider than the third presacral vertebra. Distribution —This species is endemic to Hispaniola where it occurs from sea level to an elevation of approximately 1728 m (Schwartz, pers. com. ). Remarks.—Although Noble (1923a; 1923b) wrote two popular accounts describing his quest for the giant tree frog, Hyla vasta, the first noteworthy information published about this frog, its habits, habitats and life history is that of Mertens (1939:35-36). A standardized systematic account and illustrations of the species ap- peared in Cochran (1941:19-22, Fig. 5). Photographs of this re- markable frog in life were provided by Noble (1923a; 1923b). A drawing of the species appeared in Noble (1931:89, Fig. 28C), and the tadpole and larval mouthparts were illustrated by Noble (1927: Pl. IX, Fig. B). Discussion of the relationships of Hyla vasta may be found in Dunn (1926), Noble (1927), Myers (1950), and Duell- man (1970). GREATER ANTILLEAN HYLID FROGS 45 Hyla pulchrilineata Cope Hyla pulchrilineata Cope, 1869:163 [Holotype-——Academy of Natural Sciences (Philadelphia) 14495 from “east St. Domingo”; collected by Dr. William M. Gabb]. Diagnosis——1) Size moderately small, some sexual dimorphism; maximum observed snout-vent length in males 39.5 mm (mean 31.6 mm), in females 42.8 mm (mean 40.1 mm); 2) skin on dor- sum smooth; 3) ventral skin slightly granular on belly and thighs; no tubercles around cloaca; 4) fingers webbed basally; 5) palmar supernumerary tubercles absent; 6) two round, outer palmar tu- bercles present; 7) nuptial excrescence in breeding males a flat, horny pad; 8) tarsal fold absent; 9) toes two-thirds webbed; 10) few plantar supernumerary tubercles; 11) nostrils not protuberant; 12) skull longer than broad; 13) parasphenoid alae slightly postero- laterally oriented to longitudinal axis of skull; 14) medial ramus of pterygoid not articulating with otic capsule, and widely separated from parasphenoid ala; 15) anterior ramus of pterygoid articulating with maxillary at posterior level of orbit; 16) prevomerine dentig- erous processes moderate sized, straight, and having a narrow medial separation and a single row of teeth; 17) sacral diapophyses dilated (ca. 48°; anterior sacral angle ca. 77°; posterior sacral angle ca. 55°); 18) interhyoideus muscle with small, supramandibular lobes; 19) intermandibularis muscles lacking aponeurosis, muscles meet at tendinous raphe; 20) vocal sac aperture a moderate slit; 21) anterior cornu of hyoid with anterolateral lobe; 22) larva un- known. The absence of a dermal submental gland and prepollical spine, and the smaller size of pulchrilineata readily distinguish this spe- cies from heilprini. The characteristic dorsal pattern of stripes and the narrow head of pulchrilineata differentiate it from marianae and wilderi. Distribution—Hyla_ pulchrilineata is endemic to Hispaniola where it is widespread occurring from the coastal lowlands to an elevation of approximately 636 m on the island (Schwartz, pers. com. ). Remarks.—There is a paucity of published information on pulchrilineata. Although the calling sites and voice of this species are known (Schwartz, pers. com.), no tadpoles have been asso- ciated with the species. The best available systematic account of pulchrilineata is that provided by Cochran (1941:17-19 and Fig. 4). She included a description, comments on variation and an illustra- tion. A photograph appeared in Noble (1923a:115). Hyla heilprini Noble Hyla heilprini Noble, 1923c:1 [Holotype—American Museum of Natural His- tory 11401, adult male, from Lo Bracita, Prov. Pacificador, Dominican Republic; collected on August 20, 1922, by G. K. Noble]. 46 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Diagnosis—1) Size moderate, no sexual dimorphism in size noted; maximum observed snout-vent length in males 54.3 mm (mean 49.4 mm), in females 52.0 mm (mean 49.3 mm); 2) skin on dorsum smooth; 3) ventral skin coarsely granular on belly and thighs; no tubercles around cloaca; 4) fingers one-half webbed; 5) numerous small, palmar supernumerary tubercles; 6) outer palmar tubercles absent; 7) breeding males lacking a nuptial excres- cence; but having an extruding prepollex and propollical spine; 8) tarsal fold absent; 9) toes fully webbed; 10) plantar supernumer- ary tubercles absent; 11) nostrils not protuberant; 12) skull broader than long; 13) parasphenoid alae posterolaterally oriented to longi- tudinal axis of skull; 14) medial ramus of pterygoid articulating with otic capsule and separated from parasphenoid; 15) anterior ramus of pterygoid articulating with maxillary at mid-level of orbit; 16) prevomerine dentigerous processes moderate sized, slightly curved, and having a wide medial separation and a single row of teeth on each; 17) sacral diapophyses dilated (ca. 45°; an- terior sacral angle ca. 77°; posterior sacral angle ca. 58°); 18) inter- hyoideus muscle without separate lobes; 19) intermandibularis muscles with broad median aponeurosis; 20) vocal sac aperture an elongate slit; 21) anterior cornu of hyoid without anterolateral lobe; 22) larva with thick, short, muscular tail, extensive caudal fins, complete labial disc, notched lower beak and tooth-row pattern of 6/9. Hyla heilprini is distinguished uniquely from all other West Indian hylids by either one of two characters—presence of a dermal submental gland and/or presence of a prepollical spine. Moreover, it is the only species having an elongate vocal slit aperture and cloacal musculature characterized by a short protractor muscle which terminates above the proximal end of the gracilis minor. Distribution.—The species is endemic to Hispaniola, where it is widespread (occurring near torrential streams) between elevations of sea level and approximately 1728 m (Schwartz, pers. com. ). Remarks.—Noble’s (1923c) original description of Hyla heil- print is probably the most comprehensive of the type descriptions provided for any of the Greater Antillean species. In her account of heilprini, Cochran (1941:22-24) offered only brief remarks on vari- ation in addition to Noble’s description, which is quoted in a large part. She also illustrated the adult (Cochran, 1941:23; Fig. 6). The larva and larval mouthparts are figured by Noble (1927:Pl. IX, Fig. C), and a photograph of the adult appeared in Noble (1923b: 119). Discussion of the relationships of heilprint may be found in Dunn (1926) and Noble (1927). GREATER ANTILLEAN HYLID FROGS 47 Hyla marianae Dunn Hyla marianae Dunn, 1926:129 [Holotype-—Museum of Comparative Zoology (Harvard) 11122 from Spauldings, Clarendon Parish (altitude 2900 feet); collected in August, 1925, by E. R. Dunn]. Diagnosis—1) Size moderately small, sexual dimorphism marked; maximum observed snout-vent length in males 33.1 mm (mean 28.0 mm), in females 39.6 mm (38.7 mm); 2) skin on dor- sum smooth; 3) ventral skin weakly granular; tubercles present around cloaca; 4) fingers basally webbed; 5) palmar supernumerary tubercles absent; 6) outer palmar tubercle absent; 7) nuptial ex- crescence in breeding males a flat, horny pad; 8) tarsal fold absent; 9) toes one-third webbed; 10) few, round, plantar supernumerary tubercles present; 11) nostrils not protuberant; 12) skull as broad as long; 13) parasphenoid alae posterolaterally oriented to longi- tudinal axis of skull; 14) medial ramus of pterygoid articulating with otic capsule, and widely separated from parasphenoid; 15) anterior ramus of pterygoid articulating with maxillary at level of posterior half of orbit; 16) prevomerine dentigerous processes moderately small, straight, and having a wide medial separation and a single row of teeth on each; 17) sacral diapophyses mod- erately dilated (ca. 43°; anterior sacral angle ca. 75°; posterior sacral angle ca. 62°); 18) interhyoideus muscle without postman- dibular lobes; 19) intermandibularis muscles with slender, median aponeurosis; 20) vocal sac aperture absent; 21) anterior cornu of hyoid with anterolateral lobe; 22) larva with long tail, reduced caudal fins, complete labial disc, unnotched lower beak and tooth- row pattern of 0/0. Hyla marianae most closely resembles H. wilderi from which it can be distinguished by its larger size, distinct tympanum, and presence of basal webbing between the fingers and a distinct outer metatarsal tubercle on the foot. The absence of a dermal submental gland and prepollical spine differentiate this species from heilprini. Superficially, marianae might be most easily confused with pulchri- lineata which, in contrast to marianae, is characterized by a pattern of narrow dorsal stripes and a head which is distinctly longer than broad. Distribution—This species is endemic to Jamaica where it is found in dense and scattered deciduous forest between elevations of 122 and 895 m (Schwartz and Fowler, 1973). Remarks.—Apart from Dunn’s (1926:129) brief species descrip- tion and the systematic account of Lynn, in Lynn and Grant (1940: 27-28), there is virtually no published information on marianae. The larva and larval mouthparts were illustrated by Dunn (1926: Pl. 1, Fig. 3; Pl. 2, Fig. 3) and semidiagrammatic drawings of the adult were provided by Lynn and Grant (1940:PI. II, Fig. 5). Dis- 48 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY cussion of the relationships of marianae may be found in Dunn (1926) and Noble (1927). More recently, Schwartz and Fowler (1973) provided a systematic account of this species. Hyla wilderi Dunn Hyla wilderi Dunn, 1925:161 [Holotype—Museum of Comparative Zoology (Harvard) 10500, an adult male from Moneague, Saint Ann Parish, Jamaica (altitude 1200 feet); collected in March, 1925, by Dr. and Mrs. H. H. Wilder]. Hyla shrevei Taylor, 1952:1 [Holotype—Museum of Comparative Zoology (Harvard) 26769, male, from “La Loma, Chiriquicito, Republica de Panama” (apparently in error); collected by E. R. Dunn and C. Duryea]. Synonymy fide Goin (1959:340). Diagnosis.—1) Size small, little sexual dimorphism; maximum observed snout-vent length in males 27.3 mm (mean 25.8 mm), in females 28.7 mm (mean 27.3 mm); 2) skin on dorsum smooth; 3) ventral skin weakly granular; tubercules present around cloaca; 4) fingers not webbed; 5) few palmar supernumerary tubercles; 6) two outer palmar tubercles present; 7) nuptial excrescence in breeding males a group of keratinous spines; 8) tarsal fold absent; 9) toes one-fourth webbed; 10) few, low, rounded plantar super- numerary tubercles present; 11) nostrils not protuberant; 12) skull broader than long; 13) parasphenoid alae posterolaterally oriented to longitudinal axis of skull; 14) medial ramus of pterygoid not articulating with otic capsule, and widely separated from para- sphenoid; 15) anterior ramus of pterygoid articulating with maxil- lary at posterior level of orbit; 16) prevomerine dentigerous proc- esses small, patch-like, and having a wide medial separation and single row of teeth on each; 17) sacral diapophyses dilated (ca. 48°; anterior sacral angle ca. 80°; posterior sacral angle ca. 52°); 18) interhyoideus muscle without postmandibular lobes; 19) inter- mandibularis muscles with slender, median aponeurosis; 20) vocal sac aperture a moderate slit; 21) anterior cornu of hyoid without anterolateral lobe; 22) larva with relatively short tail, reduced caudal fins, complete labial disc with keratinous “spines” arranged around perimeter, unnotched lower beak, tooth-row pattern of 0/0. Hyla wilderi can be distinguished from marianae by the smaller size, indistinct tympanum, absence of webbing between the fingers, and absence of an outer metatarsal tubercle in the former. Absence of a chin gland and prepollical spine differentiates this species from heilprini. The broader head and absence of dorsal stripes dis- tinguishes wilderi from pulchrilineata. Distribution.—This species is endemic to Jamaica where it is widely distributed in open woodlands between elevations of 122 and 610 m (Schwartz and Fowler, 1973). Remarks.—A complete systematic account of Hyla wilderi is provided by Lynn, in Lynn and Grant (1940:25-26); he also illus- GREATER ANTILLEAN HYLID FROGS 49 trated the species (Pl. III, Fig. 4). The larva and larval mouth- parts are figured by Dunn (1926:Pl. 1, Fig. 4; Pl. 2, Fig. 4). Rela- tionships of this species are discussed by Dunn (1926) and Noble (1927). The most recent account of wilderi is that of Schwartz and Fowler (1973). They reported finding individuals in both arboreal and terrestrial broemliads and observed a maximum snout- vent length in males of 27.9 mm (mean, 25.5 mm; n=25) and 29.2 mm in females (mean, 26.4 mm; n=25), ORIGIN OF THE GREATER ANTILLEAN HYLID FAUNA Graham (1972) summarized the paleophysiography of the Caribbean Basin and noted that there is no evidence that the Greater or Lesser Antilles either were rafted to their present posi- tions or connected to continents in the past. In the absence of contrary evidence, he assumed that the islands have maintained their characteristic arc-configuration (extending from the Yucatan Peninsula to Venezuela) throughout the Tertiary, although the spatial relationships between individual islands may have varied considerably during middle and late Tertiary times. The variation in spatial relationships supports Williams's (1969) statement that the lowest islands of the Lesser Antilles may have been submerged several times during the Pleistocene. Graham (1972) proposed that the flora of the Antilles is derived from 1) long distance dis- persal of floral elements from Yucatan-northern Central America, and 2) from northern South America. In the probable absence of any inter-island land connections among the Greater Antilles, and the absence of sustained connections among the islands of the Lesser Antilles, inter-island dispersal of most forms would then depend, at least in part, upon waif dispersal by wind and water. The relatively high frequency with which rafts of sufficient size to transport plants and small animals occur in the Caribbean has been documented by King (1962). Among the reptiles and amphibians of the Antilles, three groups have been studied extensively; these are the hylid and leptodac- tylid frogs, and the anoline lizards. The extensive publications of Schwartz on leptodactylid frogs are taxonomic and are not con- cerned with the distributional patterns, ecology, and zoogeographic history of the group. In contrast, the studies of Williams (e.g., 1969, 1972) delve into the phylogenetic relationships and zoogeo- graphic history (particularly as it relates to inter-island dispersal, colonization, competition, coexistence, and speciation) of the nu- merous anoline lizards of the Antilles. In view of Williams’ work especially, we are tempted to compare our zoogeographic conclu- sions with his. As these conclusions relate to our hypothesis, Wil- liams states (pers. com.) that with respect to anoline lizards he 50 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY believes, “. . . that the Greater Antilles have received relatively few invaders, usually one per major group [see Williams, 1969] and that complex radiations involving considerable island inter- change have occurred.” He further postulates that there have been only two invasions of Anolis—“From two colonists—one to Puerto Rico or Hispaniola—I would derive a fauna of ca. 70 species. I would indeed recognize two other anoline invasions, one to provide Cuba with the two species of Chamaeleolis and another to provide Hispaniola with Chamaelinorops.” In considering Williams’ state- ments with respect to our conclusions, there are several basic dif- ferences in the faunas under study that should be noted. First, we are dealing with amphibians, not reptiles. Although both groups can, and undeniably have, dispersed over water, the chances for successful dispersal and subsequent colonization in anurans are less than those for lizards for obvious structural and physiological reasons. Furthermore, among anurans of the Antilles (bufonids, leptodactylids, and hylids) the hylids are the least likely to be able to disperse successfully over water for reasons discussed be- yond. Finally, we are dealing with a much smaller extant fauna, and have no way of estimating possible or probable extinctions if, in fact, they occurred. Given these limitations, we have attempted to construct a zoogeographic model compatible with the available paleophysiographic evidence, morphological evidence, and _ syste- matic conclusions resulting from this study. The Greater Antillean hylids may have been derived reasonably from three sources—North America, northern Central America, and South America. As a possible source, North America merits the least consideration. Other than the occurrence of the Puerto Rican adventive, Hyla cinerea, no North American hylids occur in the Greater Antilles. Doubtless, this is a result of the fact that during much of the Cenozoic, peninsular Florida was submerged, thus greatly increasing the distance between the North American land mass and the Greater Antilles. Osteopilus septentrionalis apparently is a recent immigrant to southern peninsular Florida. The species may have rafted to Florida from nearby Cuba and/or the Bahamas, or it may have been introduced by man; both alternatives are feasi- ble, and perhaps both occurred. As Graham (1972) pointed out, the historical association of the Greater Antilles with Yucatan-Central America is enigmatic. Be- cause of certain floral and faunal similarities, it is thought currently (Khudoley and Meyerhoff, 1971) that there was a connection be- tween Cuba and northern Central America and that this connection may have been in the form of closely spaced island “stepping stones.” If this is so, then frogs, like plants, may have dispersed by means of rafts to Cuba from Central America. Of those species for which related, mainland forms may be postulated, only Hyla GREATER ANTILLEAN HYLID FROGS 51 vasta and Hyla heilprini have related taxa occurring in Central America. We have proposed that vasta is related to the Hyla boans group. The greatest diversity of this group is in South America, although three species (boans, crepitans, and rosenbergi) range into Central America as far as eastern Panama and southeastern Costa Rica; there is an isolated population of Hyla crepitans at Tela, Honduras (Duellman, 1970). Similarly, a representative of the Hyla albormarginata group, to which heilprini may be related, occurs in Central America. Of the nine species comprising this group, one species, Hyla rufitela, occurs in the Caribbean lowlands from east-central Nicaragua to Panama; the remaining species are South American. Thus, it is possible to assume the existence of lines ancestral to heilprini and vasta in Central America. If these ancestral anurans did disperse from Central America into the Greater Antilles, then inter-island dispersal progressed from the Yucatan-northern Central America area to Cuba or Jamaica, and thence eastward to Hispaniola. Given this dispersal route, we might anticipate finding one or more of these or related species on Jamaica or Cuba. Although such forms are unknown, it is possible that they existed from Puerto Rico, which is inhabited by only two recently introduced hylids, Osteopilus septentrionalis and Hyla cinerea. Probably the strongest argument against this pro- posed route of dispersal is found in the prevailing wind and ocean currents which are generally east-west rather than west-east. Under these conditions, it is highly improbable that the frogs could have successfully rafted from Central America to the Greater Antilles. However, the possibility exists that dispersal took place during the separation of North and South America at the Atrato (Bolivar) Trough at a time when ocean currents may have been substantially different. Thus ancestral heilprini and vasta may have successfully colonized Hispaniola and been excluded from Puerto Rico either by competition or inter-island ocean currents which precluded their rafting westward from Hispaniola to Puerto Rico. A third available alternative is that some, or all, of the Greater Antillean hylids originated in South America and rafted by way of the Lesser Antilles into the Greater Antilles. At least two facts favor this hypothesis. Of all the species for which related mainland forms can be postulated (i.e., all except marianae, wilderi, and pulchrilineata), the related mainland groups are most diverse in tropical South America; this includes the Hyla boans group, Hyla albomarginata group, Osteocephalus and Trachycephalus. Further- more, the existence of substantial rafts which originated from the Orinoco and reached Barbados and St. Vincent has been docu- mented (King, 1962). Ocean currents are such that they would favor waif dispersal in a northward or northwestward direction throughout the West Indies and Bahama Islands. This seems to be 52 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY the simpler of the two possible hypotheses, save for the absence of any of the Greater Antillean frogs or related forms on the Lesser Antilles and Puerto Rico (except for Osteopilus septentrionalis, a recent adventive). Because of the small sizes of the Lesser Antillean islands and the presence of leptodactylids on these islands, we must assume that the hylids have been, for the most part, unsuccessful colonists (except for the member of the Hyla rubra complex as noted in the Introduction). For the present we must assume that competi- tion has excluded this diverse group of species from Puerto Rico, although it seems unlikely. Despite the difficulties with this alterna- tive, we favor South American origins of the Greater Antillean hylid fauna because the hypothesized dispersal routes are amenable with ocean and wind currents as we know them today, and because of the richness of the tropical South American frog fauna as contrasted with that of the Yucatan-northern Central American area. Although not the largest island in the Greater Antilles, His- paniola has extremely diverse habitats. If we assume a South American origin of the Greater Antillean hylid fauna, then His- paniola is also nearer the mainland source of frogs than Jamaica, Cuba, or the Bahamas. Because of its size, relative distance from the mainland, and variety of habitats, we should expect a more diverse anuran fauna on Hispaniola than on the remaining islands. This, in fact, turns out to be the case. Hispaniola support 2 species of Bufo (Bufonidae), 1 species of Leptodactylus (Leptodactylidae), at least 42 species of Eleutherodactylus (Leptodactylidae), and 4 hylid species. Because of the degree of their morphological dis- tinctiveness (see Analysis of Characters), the hylid species which occur on Hispaniola seem to represent four distinct phylogenetic lines, and therefore, four separate, successful invasions. Although there is distributional sympatry, the species differ in size and/or habitat preference so that interspecific competition would appear to be minimal or non-existent. There seems to be a co-adjustment of species pairs by size (Fig. 1) reminiscent of Schoener’s Second Rule as stated by Williams (1972:57-58) in his discussion of the anoles of the West Indies. Both Osteopilus dominicensis and Hyla pulchrilineata appear to be generalists most common at lower ele- vations; however, dominicensis is about half again as large as pulchrilineata. Hyla vasta is between two and three times as large as Hyla heilprini; both use streams as breeding sites, although heilprini breed in swifter parts of the streams than vasta. It would seem that competition might occur between vasta and dominicensis; however, vasta is about one-third larger than dominicensis, and dominicensis breeds in quiet, rather than moving water. Jamaica is much smaller than Hispaniola and, based on the suggested dispersal scheme, more distant from the mainland; thus, 2 We owes. es a el ea a> Ss GREATER ANTILLEAN HYLID FROGS 53 we should expect it to have a lower immigration rate, and therefore, fewer species (MacArthur and Wilson, 1967:22-23). Only one spe- cies of Bufo (Bufonidae) occurs on Jamaica, and it is an adventive (Schwartz and Fowler, 1973). In addition to the four hylids, there are 15 species of Eleutherodactylus (Leptodactylidae), three of which have been introduced (op. cit.). Although Jamaica supports the same number of hylid species as Hispaniola, the species repre- sent three distinct phylogenetic lines and therefore, separate in- vasions; one of the species (brunneus) is closely related to Oste- opilus dominicensis of Hispaniola. Perhaps an ancestral stock of Osteopilus successfully colonized Hispaniola and subsequently Jamaica, evolving into dominicensis and brunneus, respectively. Because of the distinctiveness of Calyptahyla lichenata from all other Antillean anurans, we think it represents a separate invasion, as does the ancestral stock of marianae and wilderi. The hylids of Jamaica are much more restricted ecologically than those of His- paniola, although the same kind of size differential prevails (Fig. 1). Williams et al. (1963) suggested that at some time in the past absence of standing water on Jamaica forced the resident treefrogs to breed in bromeliads. Regardless of the operant selective factor, these four species have adapted successfully to this limited resource. The two larger species, brunneus and lichenata, use larger brome- liads for breeding. Calyptahyla lichenata is about one-third larger than Osteopilus brunneus and frequents tree-hole cavities, a habit not observed in brunneus. Hyla marianae and H. wilderi are one- half to one-third the size of the larger hylids and therefore use smaller bromeliads for breeding than does lichenata or brunneus. Of the two smaller hylids, wilderi is smaller than marianae. We suggest that wilderi evolved from a marianae-like ancestor in the process of adapting to life in smaller bromeliads. Cuba is much larger than Hispaniola and Jamaica, and has far more extensive arid to semi-arid lowlands than either. Although Cuba is somewhat farther from South America than Jamaica, it is, as pointed out, much larger and has an interesting anuran fauna relative to Jamaica and Hispaniola. Five species of Bufo ( Bufoni- dae), about 24 species of Eleutherodactylus, and one species of Sminthillus (Leptodactylidae) occur on Cuba in addition to the single hylid representative. In view of its position, extensive low- land habitat, and anuran fauna, it is not surprising that Cuba (and the Isle of Pines) has only one hylid, Osteopilus septentrionalis. Osteopilus septentrionalis is very similar in habits and morphology to O. dominicensis of Hispaniola; this suggests that their common ancestor successfully invaded Cuba from Hispaniola. The ability of this group to colonize successfully is substantiated by the oc- currence of O. septentrionalis in southern Florida and the Bahamas, 54 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY and by its apparent success on Great Inagua Island and Puerto Rico where it has been introduced recently. Parenthetically we note that, except for the absence of naturally occurring hylids, Puerto Rico supports an anuran fauna about as diverse as that of Jamaica: two species of Bufo (Bufonidae), one Leptodactylus, and about 12 species of Eleutherodactylus (Lepto- dactylidae). Thus, of the anurans inhabiting the Greater Antilles, the leptodactylids seem to have been the most successful in inter- island colonization and subsequent adaptive radiation and specia- tion. This success may be related to the relative independence from fresh water these taxa have achieved by the evolution of specialized breeding habits—namely, the utilization of foam nests and more commonly, the direct development of young from eggs deposited in terrestrial situations. Bufonids, on the other hand, seem to have been the least successful colonists. They are more diverse on Cuba with five species, whereas only two species each occur on Hispaniola and Puerto Rico, and none occurs naturally on Jamaica (a distribu- tional disparity which may relate, in part, to the relative abundance of lowland habitats on the various islands). At first thought, it would seem that a typical bufonid is hardier and less susceptible to the possible inequitable conditions encountered in waif dispersal than the smaller eleutherodactyline. However, larger size may inhibit bufonid dispersal; the smaller frog is more apt to find suit- able protective cover (eé.g., bromeliads) in which to secret itself than the larger frog. Furthermore, bufonids have not developed the breeding specializations typical of their leptodactylid relatives. As a rule, bufonids (and, in particular, Bufo) deposit their small, pigmented eggs in rosary-like strings in ponds or other temporary fresh water sources. Thus, whereas a founding leptodactylid popu- lation might be established by overwater dispersal of terrestrial eggs, the possibility of a similar mode of dispersal in Bufo is negli- gible. The diversity of hylids in the Greater Antilles closely paral- lels that of the bufonids, except that the hylids have achieved their greatest diversity (at both generic and specific levels) on Jamaica and Hispaniola instead of Cuba—a fact which we believe is ex- plained most logically by two facts: 1) Ecologically, the greatest area of Cuba is more suitable to habitation by bufonids than by hylids. 2) There have been multiple invasions of the Greater An- tilles by hylids from South America and subsequent competition with the more numerous and adaptively leptodactylids. The fact that we postulate more numerous invasions and far less speciation and inter-island dispersal than does Williams with respect to anoline lizards necessarily relates to several significant factors. Systematic study has shown the West Indian anoline lizards to be a closely related group, whereas our results reflect a great amount of morphological diversity, indicating that the Antillean hylids ————————— GREATER ANTILLEAN HYLID FROGS 55 represent several diverse phylogenetic stocks—a fact which is in- controvertible in our minds. Given that frogs probably are less successful in overwater dispersal than lizards, how do we account for the disparity in the number of postulated invasions? One possi- ble explanation centers on extinction. The anoles have been extra- ordinarily successful in radiating into available niches (Williams, 1969, 1972) on the various islands. The question arises then, how many invasions of lizards, other than anolines, may have resulted in unsuccessful colonization because of competition with resident anoles? The apparent minimal amount of speciation of the various lines of hylids is probably related to 1) the limited number of niches available to them, and 2) competition with the more numerous and adaptively labile leptodactylids. SUMMARY A comparative study is undertaken of nine species of hylids inhabiting the Greater Antilles and Bahamas. Although information from the literature concerning the habits, habitats, breeding sites and larvae is included, we concentrate on morphological compari- son of these frogs. Included in the morphological analysis of char- acters are sizes and proportions, characteristics of the skin, and features of the hands and feet. Osteological data on the cranium, vertebral column, and pelvic girdle are provided for each species, and in addition, data on throat and cloacal myology are presented. On the basis of our findings, we conclude that the nine species, all of which were allocated heretofore to the genus Hyla, are best placed in more than one genus. Hyla lichenata has been assigned to a new genus, Calyptahyla, which is similar to Trachycephalus of South America. Hyla septentrionalis, H. dominicensis, and H. brun- nea have been allocated to Osteopilus Fitzinger, 1843. These frogs differ in various myological and osteological features from Hyla and probably are related most closely to Osteocephalus of South America. The remaining five taxa are retained in the genus Hyla, although, with the exception of H. marianae and H. wilderi, they do not appear to be related closely to one another. Hyla vasta may be related to the Hyla boans group of South America, and Hyla heilprini to the Hyla albomarginata group, also of South America. We are unable to postulate any close mainland relative for Hyla pulchrilineata, or H. marianae and H. wilderi at this time. We propose that the present hylid fauna of the Greater Antilles has been established by at least five separate invasions from the mainland. Although other possibilities are discussed, we favor a hypothetical colonization of the Greater Antilles by frogs which have dispersed northward through the Lesser Antilles from the South American mainland. In this way we account for the greater diversity of hylids of the nearest island, Hispaniola, as compared 56 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY to Jamaica and Cuba. Jamaica is far more diverse with respect to its resident hylids than Cuba which is farther from the South Ameri- can mainland. By analyzing the sites and habitat preferences of the hylids residing on any given island we find an interesting con- formity to the model of co-adjustment of species pairs by size applied recently to the Antillean anoles. RESUMEN Hemos emprendido un estudio comparativo de nueve especies de los hilidos que habitan las Bahamas, Cuba, Hispaniola, Puerto Rico y Jamaica. Aunque incluimos informacion sobre la literatura tocante a los habitos, las habitaciones, los lugares de criar, y las larvas, concentramos en una comparacién morfoldgica de estas ranas. Incluimos en el analisis morfolégico de caracteres las di- mensiones y proporciones, las caracteristicas de la piel, y los sem- blantes de las manos y los pies. Proveemos datos osteolégicos sobre el craneo, la columna vertebral, y la pelvis para cada especie, y ademas presentamos datos de musculos de la garganta y de la cloaca. A base de nuestras observaciones, concluimos que las nueve especies, todas las cuales se han atribuido previamente al género Hyla, representan mas de un género. Nosotros asignamos Hyla lichenata a un género nuevo, Calyptahyla, que es similar a Trachy- cephalus de Sudamérica Hyla septentrionalis, H. dominicensis y H. brunnea han atribuido a Osteopilus Fitzinger, 1843. Estas ranas difieren substancialmente en algunos semblantes morfolégicos y osteologicos de Hyla, y estas estan emparentados probablemente mas estrechamente a Osteocephalus de Sudamérica. Retenemos los cinco especies restantes en el género Hyla, aunque, con la excepcion de H. marianae y H. wilderi, que no parecen estar emparentados unos con otros. Posiblemente, Hyla vasta este emparentada con el grupo Hyla boans de Sudamérica, y Hyla heilprini con el grupo Hyla albomarginata de Sudamérica también. No podemos postular ningtn pariente cercano del continente para Hyla pulchrilineata, H. marianae, y H. wilderi. Proponemos que la fauna corriente de los hilidos de las islas mayores de las Antillas se ha establecido por cinco invasiones del continente por lo menos. Aunque discutimos otra posibilidades, preferimos una colonizacién hipotética de las islas majores de las Antillas con ranas que hayan dispersado hacia el norte por las islas menores de las Antillas del continente de Sudamérica. De este modo, explicamos la diversidad mayor de los hilidos de la isla mas cercano Hispaniola, comparado con Jamaica y Cuba. Jamaica es mas diversa con respecto a los hilidos residentes que Cuba, que queda mas lejos del continente de Sudamérica. Mediante andalisis de los ta- manos y las preferencias en habitaciones de los hilidos que viven GREATER ANTILLEAN HYLID FROGS 57 en alguna isla particular, hallamos una conformidad notable al modelo ( Williams, 1972:57-58) que ha aplicado a los lagartillos del genero Anolis de las Antillas. LITERATURE CITED Barsour, T. 1910. Notes on the herpetology of Jamaica. Bull. Mus. Comp. Zool., 52(15):273-301, pls. 1-2. 1937. Third list of Antillean reptiles and amphibians. Bull. Mus. Comp. Zool., 82(2):77-166. BipRoN, G. 1842. In Ramon de la Sagra. Historia fisica, politica y natural de la Isla de Cuba. Part 2, Historia natural [not seen]. Bock, W. J. and C. R. SHEAR 1972. A staining method for gross dissection of vertebrate muscles. Anat. Anz. 130:222-227. BOULENGER, G. A. 1882. Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British Museum, ed. 2, London, xvi + 655 pp. Carr, A. F., JR. 1940. A contribution to the herpetology of Florida. Univ. Florida Publ., Biol. Sci. Ser., 3(1):1-118. CocurAan, D. M. 1941. The herpetology of Hispaniola. Bull. U.S. Natl. Mus., viii + 398 pp. 1961. Living amphibians of the world. Doubleday & Co. Inc., New York. 199 pp. Coreg, E. D. 1863. On Trachycephalus, Scaphiopus, and other Batrachia. Proc. Acad. Nat. Sci. Philadelphia. 15:43-54. 1869. Seventh contribution to the herpetology of tropical America. Proc. Acad. Nat. Sci. Philadelphia, 11: 147-169. 1871. Ninth contribution to the herpetology of tropical America. Proc. Acad. Nat. Sci. Philadelphia, 23 (part 2) :200-224. DUELLMAN, W. E. 1970. The hylid frogs of Middle America. Monog. Mus. Nat. Hist., Univ. Kansas, 1:xi + 753 pp. DvuELLMAN, W. E. and R. I. CromBIr 1970. Hyla septentrionalis. Cat. Amer. Amph. Rept., 92:1-4. DUELLMAN, W. E. and A. SCHWARTZ 1958. Amphibians and reptiles of southern Florida. Bull. Florida State Mus., Biol. Sci., 3(5) :181-324. Dumeri, A. M. C. and G. BiBRON 1841. Erpetologie generale ou histoire naturelle complete des reptiles. vol. 8. Paris, vi + 792 pp. Dunn, E. R. 1925. A new tree-toad from Jamaica. Occas. Papers Boston Soc. Nat. Hist., 5:161-162. 1926. The frogs of Jamaica. Proc. Boston Soc. Nat. Hist., 38(4):111- 130, pls. 1-2. Du forr, Cy A: 1955. The “tail” of Ascaphus: A historical resume and new histological- anatomical details. Ann. Univ. Stellenbosch. 31: A (1):1-71. 58 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY FIscHER, J. G. 1888. Herpetologische Mitteilungen IV. Uber eine Kollektion Reptilien und Amphibien von Hayti. Jahrb. hamb. wiss. Anstalt. 5, S. 23-45. FirzinceEr, L. 1843. Systema reptilium. Vienna, ix + 106 pp. Gorn, C. J. 1959. A synonym and a homonym in the frog genus Hyla. Copeia, 1959 (4) :340-341. Gossg, P. H. 1851. A naturalist’s sojourn in Jamaica. London [not seen in entirety]. GRAHAM, A. 1972. Some aspects of Tertiary vegetational history about the Caribbean Basin. Mem. I Congr. Latinoamericano Bot.:97-117. HaRLAN, R. 1826. Descriptions of several new species of Batrachian Reptiles, with observations on the larvae of frogs. Amer. Jour. Sci. and Arts, 10 (Arti"7): 53-65. JauMgE, M. L. 1966. Catalogo de los anfibios de Cuba. Mus. “Felipe Poey” Acad. Cien. Cuba, Trab. Divulg., 35:1-21. Kuubo.ey, K. M. and A. A. MEYERHOFF 1971. Paleogeography and geological history of Greater Antilles. Geol. Soc. Amer. Mem., 129:199 pp. [not seen]. Kinc, W. 1962. The occurrence of rafts for dispersal of land animals into the West Indies. Quart. Jour. Florida Acad. Sci., 25(1):45-52. LAURENTI, J. N. 1768. Specimen medicum, exhibens synopsin Reptilium emendatum cum experimentis circa venena et antidota reptilium Austriacorum. Vienna, 214 pp., pls. 1-5. Lynn, W. G. 1958. Some amphibians from Haiti and a new subspecies of Eleuthero- dactylus schmidti. Herpetologica, 14:153-157. Lynn, W. G. and C. Granr 1940. The herpetology of Jamaica. Bull. Inst. Jamaica, Sci. Ser., 1:1-132. MacArtuur, R. H. and E. O. Witson 1967. The theory of island biogeography. Princeton University Press, Princeton, New Jersey, xi + 203 pp. MEERWARTH, H. 1901. Die westindischen Reptilien und Batrachier des naturhistorischen Museums in Hamburg. Mitth. Naturh. Mus. Hamburg, 18:1-41 [not seen]. MERTENS, R. 1938. Amphibien und Reptilium aus Santo Domingo, gesammelt von Prof. Dr. H. Boker. Senckenbergiana, 20(5):332-342. 1939. Herpetologische Ergebnisse einer Reise nach der insel Hispaniola, Wesindien. Abhandl. Senckenberg. Naturforsch. Gesellsch., 449: 1-84, 10 pls. MirTLEMAN, M. B. 1950. Miscellaneous notes on some amphibians and reptiles from the southeastern United States. Herpetologica, 6(1):20-24. GREATER ANTILLEAN HYLID FROGS 59 Myers, G. S. 1950. The systematic status of Hyla septentrionalis, the large tree frog of the Florida Keys, the Bahamas and Cuba. Copeia, 1950(3):203- 214. NoBLE, G. K. 1922. The phylogeny of the Salientia. I. The osteology and the thigh musculature; their bearing on classification and phylogeny. Bull. Amer. Mus. Nat. Hist., 46( 1): 1-87. 1923a. In pursuit of the giant tree frog. Nat. Hist., 23(2):105-116. 1923b. Field studies of Dominican tree frogs and their haunts. Nat. Hist., 2002) 117-121" 1923c. Six new batrachians from the Dominican Republic. Amer. Mus. Novitates, 61:1-6. 1927. The value of life history data in the study of the evolution of the Amphibia. Ann. New York Acad. Sci., 30:31-128, pl. 9. 1931. The biology of the Amphibia. McGraw-Hill Book Co., Inc., New York, xiii + 577 pp. PETERSON, H. W., R. GARRETT, and J. P. LANtTz 1952. The mating period of the giant tree frog Hyla dominicensis. Herpe- tologica, 8(3):63. Rass, G. B. and E. B. HaybEn, Jr. 1957. The Van Voast-American Museum of Natural History Bahama Islands Expedition Record of the expedition and general features of the islands. Amer. Mus. Novitates, 1836:1-53. SCHWARTZ, A. 1968. The geckos (Sphaerodactylus) of the southern Bahama Islands. Ann. Carnegie Mus., 39( 17) :227-271. 1973. Six new species of Eleutherodactylus (Anura, Leptodactylidae ) from Hispaniola. Jour. Herpetology, 7(3) :249-273. ScHwartTz, A. and D. C. FowLer. 1973. The Anura of Jamaica: a progress report. Studies on the Fauna of Curacao and other Caribbean Islands, 43( 142) :50-142. TAyYLor, E. H. 1952. A new Panamanian treefrog. Breviora, (1):1-3. TRUEB, L. 1966. Morphology and development of the skull in the frog Hyla sep- tentrionalis. Copeia, 1966( 3) :562-573. 1970. The evolutionary relationships of casque-headed treefrogs with co- ossified skulls (family Hylidae). Univ. Kansas Publ., Mus. Nat. Hist., 18:547-716, pls. 1-12. 1972. Hyla crucialis Harlan, 1826 (Amphibia): Proposed suppression under plenary powers. A.N.(S). 1892. Bull. Zool. Nomencl., 29 (1):39-40. Trues, L. and W. E. DUELLMAN 1971. A synopsis of Neotropical hylid frogs, genus Osteocephalus. Occas. Papers Mus. Nat. Hist., Univ. Kansas, 1: 1-47. TscuHupDt, J. J. 1838. Classification der Batrachier mit Berucksichtigung der fossilien Thiere dieser Abtheilung. Memb. Soc. Sci. Nat. Neuchatel, 2:1-100. Lyre, MJ. 1971. The phylogenetic significance of vocal sac structure in hylid frogs. Univ. Kansas Publ., Mus. Nat. Hist., 19:319-360. 60 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY VERDOORN, F. 1945. Plants and plant science in Latin America. Frans Verdoorn, editor. Chronica Botanica Co., Waltham, Massachusetts, vol. 16: xxxvii + 381 pp. WERNER, F. 1917. Uber einige neue Reptilien und einin neuen Frosch des Zoologischen Museums in Hamburg. Mitt. Zool. Mus. Hamburg, 34:31-36. WILuuaMs, E. E. 1969. The ecology of colonization as seen in the zoogeography of anoline lizards on small islands. Quart. Rev. Biol., 44(4):345-389. 1972. The origin of faunas. Evolution of lizard congeners in a complex island fauna: A trial analysis. In Evolutionary Biology. Th. Dob- zhansky, M. K. Hecht, and W. E. Steere Eds., Appleton-Century- Crofts, Vol. 6:47-89. WiuuraMs, E. E., B. Sureve, and P. S. HuMpHREY 1963. The herpetology of the Port-au-Prince region and Gonave Island, Haiti. Parts I-II. Bull. Mus. Comp. Zool., 129(5):293-342, pls. 1-5. 133 Ul) CALFOR oo i} ACADEMY OF SCIENCES | | JUN 2 1 1974 OCCASIONAL PAPERS of the MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas NUMBER 25, PAGES 1-39 JUNE 17, 1974 THE MAMMALS OF CARTER COUNTY, SOUTHEASTERN MONTANA By Ricuarp P. Lampe,’ J. KNox Jones Jr.,” RosBert S. HOFFMANN,* AND ELMER C. BirNEy* This report treats the more than 50 species of mammals occurring in a segment of the Northern Great Plains defined by the political boundaries of Carter County, Montana, the southeasternmost county in the state. The mammalian fauna of this area has, until now, been relatively poorly documented. Carter County (Fig. 1), organized in 1917 and named after Thomas Henry Carter, Montana’s first Congressional representative, has an area of approximately 3300 square miles and slopes north- eastward on the average at 10 feet to the mile (Bauer, 1924:233). The county is characterized by gently rolling hills, extensive flats, steep ridges, and occasional “badlands.” The Boxelder Creek Valley drains northeastward through the center of the county. This valley varies in elevation from 3200 to 3500 feet above sea level and gener- ally is featureless except for the intermittent tributaries of Boxelder Creek that flow either northwestward or southeastward. The north- ern portion of the valley, referred to as Alkali Flats, is bordered by ‘Bell Museum of Natural History, University of Minnesota, Minneapolis, 55455. This paper is based in part on research accomplished while the senior author was a graduate student at the Museum of Natural History and Depart- ment of Systematics and Ecology, The University of Kansas. 2 Associate Vice President for Research and Dean of the Graduate School, Texas Tech University, Lubbock, 79409 * Curator of Mammals, and Professor of Systematics and Ecology, Museum of Natural History, The University of Kansas, Lawrence, 66045 “Curator of Mammals, Bell Museum of Natural History, University of Minnesota, Minneapolis, 55455 2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 104°30' MILL IRON 5 EKALAKA HILLS CHALK BUTTES LONG PINE HILLS CAPITOE® PINIBEE @ 104°30' Fic. 1. Map of Carter County, Montana, showing location of place-names mentioned in text. Camp Crook, Harding Co., South Dakota, is located at the black dot on the right-hand margin of the map. THE MAMMALS OF CARTER COUNTY, MONTANA 3 three disjunct pine-clad ridges, Chalk Buttes to the west (maximum elevation, 4215), Ekalaka Hills to the north (maximum elevation, 4115), and Long Pine Hills to the east (maximum elevation, 4130). The Long Pine Hills and Ekalaka Hills are separated by Boxelder Creek, whereas the latter and Chalk Buttes are separated by a small valley drained by the intermittent Little Beaver Creek. Little Beaver and Boxelder creeks are tributaries of the Little Missouri River, which traverses the southeastern corner of the county. The extreme western part of the county lies in the Powder River drainage. Largest of the three pine-clad ridges are the L-shaped Long Pine Hills, which extend north and south for approximately 18 miles with an east-west base of 11 miles. Lillegraven (1970:832) de- scribed the Long Pine Hills as part of a system of “. . . Tertiary erosional remnants standing above the Late Cretaceous rocks of northwestern South Dakota, southwestern North Dakota, and south- eastern Montana.” The southern and western sides of these ridges tend to have more cliffs and support less vegetation than do the eastern and northern slopes, which slope gradually onto the plains (Bauer, 1924:233; Lillegraven, 1970:832). Landslides of Tertiary rocks in the Long Pine Hills and Finger Buttes have been part of the erosional sequence of the area (Gill, 1962:731). South of Boxelder Creek Valley, the rolling landscape is broken by numerous, sparsely vegetated buttes. Formations such as Chim- ney Butte, Sheep Mountains, and Finger Buttes extend as a chain southwestward from the Long Pine Hills, forming the divide be- tween the drainages of Boxelder Creek and the Little Missouri River. Soils of Carter County are of four types (Kellogg, 1938). Chest- nut soils of the Williams-Morton-Bainville series have developed over glacial tills, sandstone, and shale and are found in the northern and southwestern parts of the county. In the extreme northwestern corner, brown soils have developed over shale and sandstone. Litho- sols of the Pierre type, with parent material of Cretaceous shale, occur in southeastern Carter County. The rough, eroded areas or “badlands” of the north-central part of the county have developed over clay and shale of Tertiary formation. Banks and washes of the many dry stream beds found throughout the county are sandy. Underlying or partially exposed sedimentary rocks of Cretaceous deposits include Benton (shale), Niobrara (shale), Pierre (shale), and Fox Hill (sandstone). Tertiary deposits include the Ludlow and Hell Creek members of the Lance formation, the Tongue River member of the Fort Union formation, the White River formation, and the Arikaree formation (Bauer, 1924; for additional information on geologic formations see also Alden, 1932, and Perry, 1962). Much of Carter County is covered by mixed-grass prairie, com- plemented by pine-clad ridges and sagebrush flats. Part of the Sioux Division of Custer National Forest, the Chalk Buttes, Ekalaka Hills 4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY (known also as the Ekalaka Forest), and Long Pine Hills support predominantly ponderosa pine (Pinus ponderosa). Mature stands of — this pine are relatively open; they usually are restricted to ridges — and slopes, only seldom extending onto the grassy lower valleys that _ dissect the ridge formations. Northern and eastern slopes generally — support denser stands than do southern and western slopes, which — tend to be steeper and have more cliffs. Sawlog and pole stands — generally accommodate an understory of mixed grasses, forbs, and — woody shrubs, whereas stands of saplings have a poorly developed — understory in response to the greater density of young trees and — consequent minimal sunlight penetration. Park-like stands of pine commonly occur on lower slopes and subordinate ridges along with snowberry (Symphoricarpos racemosus), various grasses, shrubs, and forbs. Juniper (Juniperus scopulorum) occurs irregularly and small stands of aspen (Populus tremuloides) occupy some slopes. Thickets of common snowberry, fleshy hawthorne (Crataegus sp.), wild plum (Prunus sp.), chokecherry (P. melanocarpa), buf- falo-berry (Shepherdia sp.), gooseberry and currant (Ribes sp.), serviceberry (Amelanchier alnifolia), and rose (Rosa sp.) occupy some of the wetter ravines. Boxelder (Acer negundo), green ash (Fraxinus pennsylvanica), and some ponderosa pine are found in the drainageways leading from the ridges. The grassland drainages sup- port discontinuous stands of deciduous trees—mainly cottonwood (Populus sargentii), willow (Salix sp.), boxelder, and green ash-— characterized as the Northern Floodplain Forest (Kichler, 1964); the Little Missouri River is lined with a heavier growth of riparian forest than that found along smaller streams throughout the county. Grasslands of the county are of the wheatgrass-needle grass com- munity (Kiichler, 1964) with dominants of western wheatgrass (Agropyron smithii), blue grama (Bouteloua gracilis), needle-and- thread grass (Stipa comata), and green needle-grass (Stipa viridula). Grasslands occupying the valleys and meadows of the Long Pine Hills are characterized as a sedge-blue grama-bluestem-wheatgrass community (Jonas, 1966:7) but “. . . under more favorable condi- tions needle-and-thread grass would possibly replace wheatgrass in importance.” Flats in the northern and central areas of the county are commonly vegetated by fringed sagebrush (Artemisia frigida), big sagebrush (A. tridentata), and silver sagebrush (A. cana), with an understory of short grasses and prickly pear (Opuntia humifusa). Carter County has a semiarid continental climate (Table 1) that is shielded from the northward flow of precipitation-laden air by the Black Hills. Slightly more than half of the mean annual precipi- tation falls in May, June, and July. Seasonal fluctuation in temper- ature is great and fluctuations are common from day to day. The contrast between summer and winter temperatures is exemplified by an average of 34 days with a maximum temperature above 90°F and THE MAMMALS OF CARTER COUNTY, MONTANA 5 TaBLE 1. Climatological data from Ekalaka, Montana (1931-1952), and Camp Crook, South Dakota (1896-1967). Temperature is recorded in degrees Fahr- enheit and precipitation is measured in inches. Ekalaka, Montana Camp Crook, South Dakota Temperature Mean annual 44.1 43.8 Mean January 176 17.3 Mean July 71.8 112 Record high 108 114 Record. low —43 —57 Precipitation Mean annual ile peAT/ ees 7a Record maximum 21.97 (1963) 24.07 (1927) Record minimum 6.35 (1936) 4.73 (1936) Mean annual snowfall O23 So an average of 33 days with a minimum temperature below zero. During winter, prevailing northwest winds often accompany snow storms, creating drifts in protected areas and leaving open fields bare. It is noteworthy that weather conditions in July, 1972, were unseasonably cold and wet, including temperatures below 32°F and freezing rains. Our interest in the mammals of Carter County stemmed from studies by one of us (Jones) conducted in Harding County, South Dakota, during the 1960’s as well as previous work in Montana by Hoffmann. Kenneth Andersen visited the Long Pine Hills briefly in May of 1968 and obtained a small collection of mammals. Subse- quently, in the period 29 June to 24 July 1970, a field party from The University of Kansas made extensive collections from the Long Pine Hills and other localities in northern and eastern Carter County, and Jaime Pefaur, a graduate student at The University of Kansas, studied populations of small mammals on a north-facing slope at the eastern edge of the Long Pine Hills until mid-August of the same year. Birney and Lampe collected mammals for the Uni- versity of Minnesota in the Long Pine Hills, Ekalaka Hills, and Chalk Buttes in July 1971. Merlin Tuttle and Larry Heaney made additional collections for Minnesota in Carter County in July 1972. Early reference to the mammals of Carter County was made by Captain W. F. Reynolds (1868) during his brief passage through the area in 1859. Subsequent military explorations in this area, such as General George Custer’s expedition to the Black Hills in 1874, were concerned primarily with matters other than faunal observa- tions (see also Tumer, 1974). E. A. Preble, in an undated manu- script, probably written about 1910, listed mammals from eastern Montana and briefly commented on the habitat preference and abundance of 57 species. Other references in the literature to mam- mals of Carter County are those of Visher (1914), Couey (1946), 6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Tryon (1947), Hoffmann et al. (1969a,b), Andersen and Jones | (1971), Pefauer and Hoffmann (1971), and Jones et al. (1973). | Several specimens from Carter County have been incorporated in | revisionary studies such as those by Nelson (1909), Bailey (1915), Hollister (1916), Jackson (1928), Howell (1929, 1938), and Gold- | man (1944). Acknowledgments We acknowledge the field assistance of the students who were — enrolled in the Field Course in Vertebrate Zoology at The University of Kansas in the summer of 1970 as well as the members of several other field parties from Kansas who visited Carter County periodi- cally. Merlin D. Tuttle and Lawrence R. Heaney collected mam- mals in Carter County for the University of Minnesota for 10 days in July of 1972. The summer field course was financed in part by grant GZ-1512 from the National Science Foundation. Minnesota field parties in both 1971 and 1972 were supported by grants to Birney from the Graduate School, University of Minnesota and from the Grassland Biome Section of the International Biological Program (NSF Grant GB-13096). We especially thank District Rangers T. S. Burns and Richard McElfresh and their staff for assistance and sincere interest in our study. We also wish to acknowledge the cooperation and hospitality shown our field parties by various U.S. Forest Service personnel (Sioux Division, Custer National Forest) and numerous residents of Carter County, Montana, and Harding County, South Dakota. Identifications of ectoparasites were provided by Drs. Eleanor K. Jones (ticks), J. M. Kinsella (fleas), Richard B. Loomis (chiggers), and Nixon Wilson (mites). ACCOUNTS OF SPECIES The following accounts treat the 53 species of mammals pres- ently on record from Carter County, Montana. Additionally, com- ments on 15 species of unverified occurrence are appended. Speci- mens examined (a total of 488) are listed in telegraphic style and are ordered first from north to south and, secondarily, from west to east. All specimens examined are deposited in the Museum of Natural History at The University of Kansas or in the Bell Museum of Natural History at the University of Minnesota (noted as MMNH). All measurements used are in millimeters (lengths of embryos are for crown-rump) and weights are expressed in grams. In certain cases, measurements are given as means, with extremes in parentheses. THE MAMMALS OF CARTER COUNTY, MONTANA | ORDER INSECTIVORA Sorex cinereus haydeni Baird, 1858 Masked Shrew Specimens examined (26).—3 mi S, 1.75 mi E Ekalaka, 6 (MMNH); 5.5 mi N, 10 mi W Camp Crook, 3500 ft, 20 (17 MMNH). The masked shrew evidently is a common inhabitant of pro- tected, mesic habitats in the pine-covered portions of the study area. Only three specimens, all adults, were captured during approxi- mately four weeks of intensive trapping in summer of 1970. Most trapping efforts were with break-back traps, although buried can traps were tried unsuccessfully in several areas in the Long Pine Hills that seemed suitable for shrews. Three additional specimens, two adult females and a shrew of unknown sex, were captured in 1971 in break-back traps in the mesic draw at Lantis Spring and in lush vegetation around a nearby beaver pond fed by the spring, the site of capture of two males in 1970. The third specimen from 1970, a female, was taken above the draw on snowbrush-covered ground near a clearing. On 10 July 1972, five can-traps were set near water at Lantis Spring. Four of these contained masked shrews the following morn- ing. Eleven more can-traps and 75 Museum Specials then were set in mesic habitat along the spring and near the beaver pond. No shrews were captured in the Museum Specials, but six more were taken from the cans. Even after an additional 50 Museum Specials, baited with a mixture of rolled oats and peanut butter, were set near the spring, all four shrews subsequently captured were taken from the 16 buried cans. A few days later, two Sorex were among the animals taken from 202 Museum Specials set in tall grass and reeds near a marsh in the Ekalaka Hills. Seventeen can-traps set near the marsh yielded four additional masked shrews. Brown (1967) and Spencer and Pettus (1966) demonstrated the efficiency of can-trapping for shrews. Our studies indicate that the masked shrew is not uncommon in Carter County. Instead, it seems to be common in isolated or semi-isolated populations associated with permanent sources of water and relatively undisturbed habi- tats. Genoways and Jones (1972:5) trapped Sorex cinereus at seven localities in southwestern North Dakota, but Andersen and Jones (1971:388) did not report shrews from adjacent Harding County, South Dakota, although can-traps were not employed there. Three specimens from 5 mi SE Ekalaka were recorded by Jackson (1928: 53), and were probably those mentioned by Preble (MS) from the Sioux National Forest, near Ekalaka. Testes of 12 males taken in mid-July 1972 were 4 mm in mean length. One of eight females collected at that time carried four embryos, each 12 mm in length; another had placental scars but apparently had completed lactation. 8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Sorex merriami leucogenys Osgood, 1909 Merriam Shrew Specimen examined (1).—5 mi N, 3.5 mi W Camp Crook, 3400 ft, 1. Our specimen of Merriam’s shrew, the fifth to be recorded from Montana (Pefaur and Hoffmann, 1971:247), was trapped on 22 July on a north-facing grassy slope at the eastern edge of the Long Pines. An adult male with enlarged flank glands and testes 5 mm in length, it had the following selected external and cranial measurements: total length, 101 mm; tail length, 39 mm; hind foot length, 11 mm; weight, 4.8 gms; condylobasal length, 16.5 mm; palatal length, 7.2 mm; cranial breadth, 8.3 mm; least interorbital breadth, 3.7 mm; maxillary breadth, 5.2 mm; maxillary toothrow length, 6.2 mm. Owing to the large size of this shrew, especially apparent in the cranial dimensions, we assign it to the subspecies leucogenys (see Long, 1965, and Hooper, 1944, for representative measurements of S. m. leucogenys and S. m. merriami), even though the type locality of S. m. merriami lies only approximately 150 miles to the west. ORDER CHIROPTERA Seven species of bats were collected in Carter County in 1970. Collections were made principally in the Long Pine Hills, but a few bats were obtained in adjacent areas and at Medicine Rocks. Speci- mens obtained during that summer were reported by Jones e¢ al. (1973) together with comments on habitat, reproduction, molt, ac- tivity, dietary habits, and ectoparasites of these seven bats. Included in the present accounts are specimens obtained in the Long Pine Hills, Ekalaka Hills, and Chalk Buttes in 1971 and 1972. One spe- cies, Plecotus townsendii, was not captured in Carter County until July 1972 and thus was not treated by Jones ef al. (1973). Myotis evotis evotis (H. Allen, 1864) Long-eared Myotis Specimens examined (7).—4 mi S, 1 mi E Ekalaka, 4 (MMNH); 5.5 mi S, 1 mi E Ekalaka, 1 (MMNH); 7 mi N, 10 mi W Camp Crook, 3800 ft, 2 (MMNH). All specimens are adults, and were netted over reservoirs in the Long Pine Hills and the Chalk Buttes and in a campground clearing in the Ekalaka Hills. At Stagville Spring, 4 mi S and 1 mi E Eka- laka, 11 long-eared myotis were collected over a spring-fed watering tank located in a brush-lined ravine. Seven of these were banded and released. Three of four banded females were lactating; the three males gave no external evidence of reproductive activity. Four adult males taken in mid-July had testes that were 3, 3, 4, and 5 mm in length, and three were in the process of seasonal molt. THE MAMMALS OF CARTER COUNTY, MONTANA 9 Myotis leibii ciliolabrum (Merriam, 1886) Small-footed Myotis Specimen examined (1).—4 mi S, 1 mi E Ekalaka, 1 (MMNH). A lactating female in the process of seasonal molt was netted 18 July above a livestock tank in the Ekalaka Hills. This specimen is the only small-footed myotis known from the county outside of the Long Pine Hills, where six were obtained in 1970 (Jones et al., 1973). Myotis lucifugus carissima Thomas, 1904 Little Brown Myotis No specimens of the little brown myotis were captured in 1971 or 1972. Jones et al. (1973) reported 12 individuals that had been netted or shot over water in the Long Pines in 1970. Myotis volans interior Miller, 1914 Long-legged Myotis Specimens examined (7).—4 mi S, 1 mi E Ekalaka, 2 (MMNH); 5.5 mi S, 1 mi E Ekalaka, 3 (MMNH); 7 mi N, 10 mi W Camp Crook, 2 (MMNH). We found the long-legged myotis to be common in most areas of buttes and ridges in Carter County although none was taken in the Chalk Buttes. All specimens collected in mid-July are adults. One female was lactating and two males had testes that were 3 and 4 mm in length. Seven adults were netted over a livestock tank near Stag- ville Spring in the Ekalaka Hills on 17 July 1971; of five subsequently banded and released, two were reproductively active females, one lactating and the other pregnant. Lasionycteris noctivagans (Le Conte, 1831) Silver-haired Bat Although found to be common in the Long Pine Hills in 1970 (Jones et al., 1973), only a single silver-haired bat was captured in the subsequent two summers. This specimen, an adult male netted over a beaver pond at Slick Creek Spring, 7 mi N, 10 mi W Camp Crook, was banded and released. Eptesicus fuscus pallidus Young, 1908 Big Brown Bat Specimens examined (4).—11 mi S, 7.5 mi W Ekalaka, 1 (MMNH); 7 mi N, 10 mi W Camp Crook, 3800 ft, 3 (MMNH). Two adult male big brown bats were banded and released at the locality (Slick Creek Spring) listed above, and an additional two adult females and one adult male were preserved. An adult male (testes 8 mm in length) was netted over a small reservoir in the Chalk Buttes in 1971. We failed to capture this species in south- eastern Montana in 1972. 10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Lasiurus cinereus cinereus (Palisot de Beavois, 1796) Hoary Bat Specimens examined (6).—4 mi S, 1 mi E Ekalaka, 1 (MMNH); 11 mi S, 7.5 mi W Ekalaka, 2 (MMNH); 7 mi N, 10 mi W Camp Crook, 3800 ft, 3 (MMNH#). Volant juvenile hoary bats were netted in mid-July in both 1971 and 1972. Two adults, a lactating female and a male, also were cap- tured. Testes of the male were 8 mm in length on 13 July. One juvenile female was banded and released in the Long Pines in 1971. Plecotus townsendii pallescens (Miller, 1897) Townsend’s Big-eared Bat Specimen examined (1).—7 mi N, 10 mi W Camp Crook, 1 (MMNH). A reproductively inactive, adult female Plecotus was captured in a mist net stretched over a beaver pond in the Long Pine Hills on 12 July 1972. Preble (MS) reported the species from adjacent Powder River County and Hoffmann et al. (1969b:741) considered it wide- spread in Montana. Andersen and Jones (1971:372) considered big- eared bats uncommon in Harding County, South Dakota. The ab- sence of this bat in our 1970 collections (Lampe, 1971; Jones et al., 1973) and in 1971 and the paucity of specimens from adjacent areas attest to the apparent rarity of the species in this portion of its range. OrpDER LAGOMORPHA Lepus townsendii campanius Hollister, 1915 White-tailed Jackrabbit Specimens examined (9).—12 mi E Ekalaka, 3200 ft, 1; 6.5 mi N, 5.5 mi W Camp Crook, 3400 ft, 1; 5.5 mi N, 3 mi W Camp Crook, 1; 2 mi N, 14 mi W Camp Crook, 1; 2 mi N, 4.5 mi W Camp Crook, 3700 ft, 1; 2.5 mi S, 3 mi W Camp Crook, 3200 ft, 1; 5.5 mi S, 3.5 mi W Camp Crook, 1; 6 mi S, 3 mi W Camp Crook, 3200 ft, 1; 12 mi S, 4 mi W Camp Crook, 1. White-tailed jackrabbits are common in the grasslands of Carter County, particularly in the meadows and broad valleys within the Long Pine Hills. All of our specimens were collected in 1970 from along roads or roadside ditches. An adult female taken on 3 July carried three embryos, each of which was 48 mm in length. Another examined on 7 July was lac- tating; both females were actively molting. Three subadult males collected in early and mid-July had testes that were 24, 18, and 15 mm in length, whereas an adult male obtained on 5 July had testes 559 mm long. One jackrabbit was parasitized by ticks, Dermacentor andersoni. Sylvilagus audubonii baileyi (Merriam, 1897) Desert Cottontail Specimens examined (2).—17 mi E Ekalaka, 3200 ft, 2. This cottontail was seen on sagebrush flats and in grasslands as THE MAMMALS OF CARTER COUNTY, MONTANA 11 well as on the pine-covered ridges of the Long Pine Hills. Two S. audubonii were obtained from a small formation in the “badlands” north of the Long Pines. Preble (MS) listed specimens from the “head of Boxelder Creek,” no doubt the same individuals reported by Nelson (1909:234) as simply from “Boxelder Creek.” Addition- ally, Andersen and Jones (1971:373) reported a specimen obtained on a sparsely covered pine ridge at the extreme eastern edge of the Long Pines in Harding County, South Dakota. A subadult male examined on 8 July had testes 14 mm in length. An adult female shot on 9 July was actively molting and carried six embryos (three in each uterine horn) in addition to a seventh in the right uterine horn that, in light of its smaller size (9 mm in length as compared to 14 mm for the others), was apparently being re- sorbed. Sylvilagus floridanus similis Nelson, 1907 Eastern Cottontail Specimens examined (4).—5.5 mi N, 10 mi W Camp Crook, 1 (MMNH); 2 a 4.5 mi W Camp Crook, 3700 ft, 1; 8 mi S, 3 mi W Camp Crook, 3200 be The eastern cottontail occurs in Montana only in Carter County, where it is restricted to riparian habitats. One specimen was shot along a small spring-fed stream at the southeastern edge of the Long Pine Hills and a juvenile was trapped in a thicket on high ground at Lantis Spring; the other two were collected in dense, woody vege- tation along the Little Missouri River. Hall and Kelson (1951:52) reported specimens of S. f. similis from Boxelder Creek (25 mi SW Sykes—Hall, 1951:159), Capitol, and the Little Missouri River (7 mi NE Albion—Hall, loc. cit.) in southern Carter County. It is of interest that our four specimens and those cited by Hall and Kelson (loc. cit.) all were collected in the eastern third of Carter County. The apparent absence of S. floridanus in western Carter County may be of importance with respect to the distribution of S. nuttallii, which evidently has been displaced by S. floridanus in ad- jacent states (Genoways and Jones, 1972:11; Turner, 1974:61). A female taken on 30 June carried six embryos. Two males shot in early July had testes 40 and 55 mm in length. These three speci- mens, all adults, were actively molting from winter to summer pelage. ORDER RODENTIA Eutamias minimus pallidus (J. A. Allen, 1874) Least Chipmunk Specimens examined (15)—17 mi E Ekalaka, 3200 ft, 1; 4 mi S, 1 mi E Ekalaka, 2 (MMNH); 4.5 mi S, 1 mi E Ekalaka, 2 (MMNH); 4 mi N, 8 mi W Camp Crook, 3650 ft, 4; 4 mi N, 7.5 mi W Camp Crook, 3800 ft, 1; 2 mi N, 4.5 mi W Camp Crook, 3700 ft, 3; 1.5 mi N, 5 mi W Camp Crook, 4100 ft, 2. 12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Chipmunks are common in the pine-covered hills of southeastern Montana. Specimens were obtained from ridges and roadsides where fallen logs and rocks provided suitable cover. Two adult males were collected at Capitol Rock, a large butte along the south- ern edge of the Long Pine Hills, 1.5 mi N and 5 mi W Camp Crook, that is characterized by gullies, steep and eroded slopes, and pale- colored soils. An adult male obtained from a similar, but smaller area of “badlands” isolated in the grasslands of northeastern Carter County, 17 mi E Ekalaka, was markedly grayer than specimens col- lected in the Long Pines and Ekalaka Hills. This animal was actively molting on 8 July. A chipmunk was observed running over a bare, vertical cliff-facing at the crest of Trenk Pass, 11 mi S, 6.5 mi W Ekalaka, in July 1971, but no specimens were collected in the Chalk Buttes. An adult female obtained in late June and one taken in July were lactating and still retained winter pelage. Eleven subadults taken in July were molting. A young individual, one of four obtained on 25 July from 4 mi N and 8 mi W Camp Crook, had a scar, no doubt caused by a cuterebrid larva, on the right shoulder. Preble (MS) mentioned specimens from the Sioux National For- est and Alzada and commented that chipmunks inhabited “indiffer- ently the evergreen forests, bushy canyons, and even sections almost without forest cover... .” Howell (1929:44) reported specimens from Alzada, Ekalaka, 8 mi E Sykes, and 5 mi E Sykes. Spermophilus tridecemlineatus pallidus J. A. Allen, 1874 Thirteen-lined Ground Squirrel Specimens examined (33).—11.5 mi N, 3 mi E Ekalaka, 5; 16 mi E Eka- laka, 2; 1 mi S, 1 mi W Ekalaka, 1 (MMNH); 3.5 mi S, .25 mi W Ekalaka, 2 (MMNH); 3.75 mi S, 2.25 mi W Ekalaka, 3 (MMNH); 4 mi S, 1.5 mi W Ekalaka, 1 (MMNH); 5 mi N, 3.5 mi W Camp Crook, 3400 ft, 18; 6 mi S, 4.5 mi W Camp Crook, 3200 ft, 1. This ground squirrel is common in short grass and sagebrush throughout the area of study. Juveniles represented more than half of the total number of specimens obtained. Eight of eleven speci- mens collected in an upland meadow along the eastern edge of the Long Pine Hills during the period 8 to 13 July were juveniles, and only one of seven individuals from southwest of Ekalaka was adult. Preble (MS) commented on the destruction of crops caused by thirteen-lined ground squirrels, “especially in the southeastern part of the state.” Howell (1938:114) reported specimens from Albion, Capitol, and Medicine Rocks. An adult female taken on 14 July had seven placental scars and eight corpora albicantia. The testes of eight adult males obtained in mid-July had a mean length of 5.8 mm; those of eight juveniles collected on comparable dates averaged 4.9. THE MAMMALS OF CARTER COUNTY, MONTANA 13 Fleas, Thrassis sp., and ticks, Ixodes sculptus, were found on these thirteen-lined ground squirrels. Cynomys ludovicianus ludovicianus (Ord, 1815) Black-tailed Prairie Dog Specimens examined (4).—5 mi N, 6 mi E Ekalaka, 3400 ft, 3; 1 mi S, 13 mi E Ekalaka, 3200 ft, 1. Once a common inhabitant of grasslands throughout Carter County, the prairie dog was found in areas overgrazed by herds of bison and other ungulates. With the decline of native herbivore populations, the prairie dog became associated with domestic live- stock, often becoming a “pest,” particularly in areas overgrazed by cattle and sheep. Visher (1914:89) reported the existence of large “towns in Harding County, South Dakota, in the early 1900's, one of which covered “several sections . . . west of the Little Missouri River.” Such colonies most likely occurred also in Carter County. Rather than directly treating the cause of poor range condition, rodent control campaigns tended to treat only symptoms—that is, large populations of prairie dogs. The efficiency of programs de- signed to eradicate this sciurid is attested to by the fact that Cyno- mys now occurs only in relatively small, disjunct populations in Carter County and adjacent regions. Three prairie dogs were shot in mid-day from a colony of approx- imately 200 individuals northeast of Ekalaka. The colony was lo- cated on flat ground covered with sagebrush (Artemesia tridentata), below the dam of a farm pond. Another specimen was obtained from a half-grown badger that was observed crossing a road with a prairie dog in its mouth. As members of our field party approached the badger, it dropped the prey and took shelter in a nearby burrow. Hollister (1916:14) reported a specimen from Boxelder Creek and Andersen and Jones (1971:375) noted two specimens from the grasslands of northwestern Harding County, South Dakota, border- ing the eastern Long Pine Hills. An adult female taken on 14 July contained eight placental scars and eight corpora albicantia. Two males, an adult and subadult, collected on 14 July both had testes 8 mm in length. Fleas, Opiso- crostis hirsutus, and ticks, Ixodes kingi, were found on one of three individuals collected northeast of Ekalaka. Sciurus niger rufiventer E. Geoffroy St.-Hilaire, 1803 Fox Squirrel Specimens examined (2).—Sec. 16, T 5S, R 62 E, 2. Although fox squirrels are known to occur in the riparian cotton- wood forests of the Yellowstone and Big Horn rivers in central Mon- tana (Hoffmann et al., 1969a:589), their presence in southeastern Montana has not been documented previously. Hibbard (1956:525) 14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY discussed the westward spread of fox squirrels in North Dakota along the Missouri and James rivers as well as their wooded tribu- taries in the south-central part of that state. Our specimens verify the occurrence of fox squirrels in the Little Missouri drainage. Be- cause we know of no introductions of this species in this area of Montana, our records probably indicate a southward migration from the valley of the Missouri River. Both specimens are adult males in winter pelage, collected along the Little Missouri River in east-central Carter County on 14 Jan- uary 1971 by John Teigen, Jr. In this area, the banks and flood plain of the Little Missouri support riparian stands of boxelder, green ash, plains cottonwood, and willows. Selected external and cranial measurements of our specimens are: total length, 490, 488 mm; tail length, 212, 208 mm; hind foot length, 68, 68 mm; ear length, 28, 28 mm; weight, 645, 623 gms; condylobasal length, 56.8, 58.8 mm; palatal length, 32.1, 32.5 mm; zygomatic breadth, 35.2, 35.2 mm; mastoid breadth, 27.6, 27.2 mm; postorbital breadth, 19.2, 19.1 mm; maxillary toothrow length, 11.4, 11.5 mm; testes length, 23, 25 mm. Tamiasciurus hudsonicus dakotensis (J. A. Allen, 1894) Red Squirrel Specimens examined (14).—5.5 mi S, 1 mi E Ekalaka, 2 (MMNH); 7 mi N, 10 mi W Camp Crook, 3800 ft, 1; 6 mi N, 10 mi W Camp Crook, 1 (MMNH); 5.5 mi N, 10 mi W Camp Crook, 2 (MMNH); 4 mi N, 9 mi W Camp Crook, 3950 ft, 5; 3.75 mi N, 7.75 mi W Camp Crook, 2 (MMNH); sec. 1, Ty0.5,iR 6) ee, Red squirrels were collected in moderately dense stands of pon- derosa pine. Two were taken in the Ekalaka Hills in 1972 and others were collected in the central part of the Long Pine Hills in all three years of field work. Red squirrel “sign” was noted by Lampe and Birney in the Chalk Buttes, but no individuals were seen. Five squirrels were obtained in 1970 from an immature pine stand of 10 to 15 acres located on a gentle westward-facing slope marked by small ravines. Additional areas were hunted thoroughly, but only two other specimens were taken; a third was sighted 6 mi N and 10 mi W Camp Crook. Three individuals were feeding on ponderosa pine cones when shot, but no indication of regular feeding stations was observed in 1970. Three red squirrels were shot and numerous others were seen during a two-day visit to the central Long Pine Hills in 1971. A midden with a surface area of approximately 60 square feet was found on a small north-facing slope at a place 8.5 mi N and 11 mi W Camp Crook. The midden was extremely moist as a result of its proximity to a nearby spring. Smaller caches and middens were seen elsewhere in the Long Pines around fallen and hollow trees. THE MAMMALS OF CARTER COUNTY, MONTANA 15 The scarcity of caches on the forest floor in stands of ponderosa pine was discussed by Finley (1969:238). The dryness of sites oc- cupied by this pine, the increased circulation of air and penetration of sunlight caused by the lack of lower limbs, and the open nature of stands create conditions unsuitable for the storage of cones. Caches seen in the Long Pines were associated with mesic or shel- tered conditions, whereas those observed in the Chalk Buttes were small, shallow subterranean chambers. The marked variability in cone production of ponderosa pine (Finley, 1969:261) may account for much of the variation in red squirrel population density noted in the three years of our study. Preble (MS) mentioned that red squirrels were fairly common near Ekalaka and east of Sykes and that specimens from these places appeared to be typical T. h. dakotensis. Visher (1914:88) reported a specimen shot in the Long Pines in July of 1910, and Miller and Kellogg (1955:263) included southeastern Montana in the range of dakotensis. Andersen and Jones (1971:375) commented that speci- mens from the Long Pine Hills were clearly assignable to dakotensis on the basis of color (see also Hoffmann and Jones, 1970:374, fig. 7; and Turner, 1974:80). Average (and extreme) external and cranial measurements of 12 adults (five females, seven males) are: total length, 343.4 (322-365) mm; tail length, 133.9 (115-144) mm; hind foot length, 53.9 (50-58) mm; ear length, 28.0 (25-30) mm; condylobasal length (four fe- males, seven males), 47.6 (46.7-49.1) mm; palatal length, 26.7 (26.0-27.3) mm; zygomatic breadth (four females, five males), 29.2 (28.9-30.4) mm; mastoid breadth (three females, six males), 24.1 (23.3-24.5) mm; postorbital breadth (four females, seven males) 14.2 (13.5-14.7) mm; maxillary toothrow length, 9.0 (8.8-9.2) mm. Five of seven females collected from late May through July were lactating; two contained three placental scars each. Two females obtained on 18 and 20 July were young of the year. Seven adult males taken during July had testes that had an average length of 14.2 (10-20) mm. Thomomys talpoides bullatus Bailey, 1914 Northern Pocket Gopher Specimens examined (34).—16 mi E Ekalaka, 1; .5 mi S, 12 mi E Ekalaka, Il; 3 mi S, 1 mi E Ekalaka, 1 (MMNH): 3 mi S, 1.75 mi E Ekalaka, 3 (MMNH); 12 mi S, 16 mi E Ekalaka, 3600 ft, 3; 8 mi N, 8 mi W Camp Crook, 2; 7 mi N, 3 mi W Camp Crook, 3400 ft, 1; 6 mi N, 10 mi W Camp Crook, 1; 6 mi N, 9 mi W Camp Crook, 4000 ft, 1; 5.5 mi N, 10 mi W Camp Crook, 1 (MMNH); 5.5 mi N, 9.5 mi W Camp Crook, 1; 5 mi N, 6 mi W Camp Crook, 3500 ft, 2; 5 mi N, 3.5 mi W Camp Crook, 3400 ft, 12; 2 mi N, 4.5 mi W Camp Crook, 3700 ft, 1; 6 mi S, 4.5 mi W Camp Crook, 3200 ft, 3. The northern pocket gopher is common in Carter County. Our specimens were collected primarily in the Long Pine Hills from 16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY roadside ditches, sparsely pine-clad ridges, and meadows. Four in- dividuals were trapped in similar habitats in the Ekalaka Hills and adjacent areas. Three were taken along a sandy arroyo located 6 mi S and 4% mi W Camp Crook. Bailey (1915:102) referred a specimen from Alzada to Thomo- mys talpoides bullatus, the type locality of which is just to the west of Carter County at Powderville, Powder River County. Swenk (1941:3) later included this specimen in his description of T. t. pierreicolus, a subspecies supposedly delimited by the distribution of soils of the Pierre series. Andersen and Jones (1971:376) exam- ined a specimen from Harding County, also referred by Swenk (1941:3) to pierreicolus, and found it clearly referable to bullatus. Five adults collected in early and mid-July and a subadult taken in early July were molting. Two adult males obtained on 8 July had testes that were 10 and 13 mm in length, whereas those of a subadult taken the same day were 4 mm. An adult and a subadult collected on 18 July had testes that were 6 and 5 mm, respectively. An adult female taken on 8 July and two obtained 12 July contained, re- spectively, 13, 10, and six placental scars. The following ectoparasites were found on our specimens of Thomomys talpoides: a flea, Foxella ignota albertensis; four mites, Androlaelaps geomys, Androlaelaps fahrenholzi, Haemogamasus sp., and Hirstionyssus sp.; two ticks, Ixodes kingi and I. sculptus; and two lice, Geomydoecus thomomus and G. wardi. Fleas, mites, ticks and lice were found together on one specimen from the Long Pine Hills. Perognathus fasciatus olivaceogriseus Swenk, 1940 Olive-backed Pocket Mouse Specimens examined (30).—11.5 mi N, 3 mi E Ekalaka, 1; 17 mi E Eka- laka, 3200 ft, 1; 4.5 mi S, 1 mi E Ekalaka, 1 (MMNH); 5.5 mi N, 3.5 mi W Camp Crook, 3400 ft, 22; 6 mi S, 4.5 mi W Camp Crook, 3200 ft, 5. The olive-backed pocket mouse was found to be common in grazed and ungrazed meadows on sandy soils. A series of 11 adults and 11 young was collected from a north-facing slope at the eastern edge of the Long Pine Hills. Five individuals were obtained in an area of short grass and sage 6 mi S and 4.5 mi W Camp Crook, along a sandy arroyo. Specimens from 8 mi NE Albion were mentioned by Preble (MS). Adult females taken on 7 and 8 July carried embryos (five meas- uring 11 mm and nine measuring 13 mm, respectively). A female bearing three placental scars was obtained on 9 July and one of two adult females taken on 10 July carried three embryos and had seven placental scars, whereas one of two taken on 11 and 12 July con- tained five embryos in addition to six placental scars. Two females trapped on 16 and 17 July carried seven and six embryos, respec- THE MAMMALS OF CARTER COUNTY, MONTANA i tively, the latter also containing five placental scars. One taken on 19 July had four corpora lutea and one obtained on 25 July was nonparous. An adult female examined on 29 July contained 11 pla- cental scars of at least two ages. One obtained on 1 August was lactating and had nine placental scars, and another trapped on 5 August carried five embryos and had six placental scars. Two adult males taken on 8 and 11 July had testes that were 11 and 6 mm in length, respectively. Average and extreme external and cranial measurements for 10 adults (two males, eight females) are: total length, 128.9 (125-136) mm; tail length, 59.7 (55-65) mm; hind foot length, 16.9 (16-18) mm; ear length, 6.6 (6-8) mm; basal length, 17.6 (17.2-18.3) mm; occipitonasal length (one male, six females), 11.9 (11.3-12.2) mm; interorbital breadth, 4.7 (4.5-5.0) mm; depth of cranium, 8.5 (8.3- 8.8) mm. Jones (1953:522) assigned an olive-backed pocket mouse from 8 mi NE Albion to P. f. olivaceogriseus. Andersen and Jones (1971: 376) and Genoways and Jones (1972:16) considered specimens from northwestern South Dakota and southwestern North Dakota, respectively, to be intergrades between the subspecies fasciatus and olivaceogriseus, but in both cases assigned specimens to fasciatus, mostly on the basis of color. Specimens from Carter County average slightly smaller and paler in color than do mice from those two ad- jacent areas. Therefore, we use the name olivaceogriseus to repre- sent olive-backed pocket mice from southeastern Montana, although they undoubtedly are intergrades between the two races. Our specimens of P. fasciatus were parasitized by a flea, Me- ringis sp., and three species of mites, Androlaelaps fahrenholzi, Haemogamasus sp., and Hirstionyssus sp. Perognathus hispidus paradoxus Merriam, 1889 Hispid Pocket Mouse Specimen examined (1).—5 mi N, 3.5 mi W Camp Crook, 1. A nulliparous, subadult female, the first record of this species from Montana (Pefaur and Hoffmann, 1971:247), was trapped on a grassy slope at the eastern edge of the Long Pines. Mammals trapped in the same or adjacent trap lines included Sorex merriami, Spermophilus tridecemlineatus, Thomomys talpoides, Reithrodon- tomys megalotis, Peromyscus maniculatus, Microtus ochrogaster, Microtus pennsylvanicus, Mus musculus, and Zapus hudsonius. The hispid pocket mouse has also been recorded from adjacent Harding County, South Dakota (Andersen and Jones, 1971:377; Birney and Lampe, 1972:466), and northwestern Crook County, Wyoming (Long, 1965:618). 18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Dipodomys ordii terrosus Hoffmeister, 1942 Ord’s Kangaroo Rat The kangaroo rat evidently is uncommon in southeastern Mon- tana, being found in sandy areas along dry streams as well as on flats. We obtained no D. ordii, but its presence was reported to us, and Preble (MS) mentioned specimens from the Sioux National Forest near Ekalaka. Additionally, this rat is known from adjacent Harding County, South Dakota (Andersen and Jones, 1971:377), and Bowman County, North Dakota (Genoways and Jones, 1972: 18), and from but a few miles west of the Carter County boundary at Powderville, Powder River County, Montana (KU 14885-86 and 14912-13). Castor canadensis missouriensis Bailey, 1919 Beaver Beaver dams and freshly cut trees were noted along many of the streams and springs in the Long Pine Hills. An adult was observed at dusk on a large pond at Slick Creek Spring, 7 mi N and 10 mi W Camp Crook, 3800 ft, a place where numerous dams (the largest 35 feet long and 10 feet high) had been built across steep ravines, the sides of which were clad with mature stands of ponderosa pine. The bark of some pines in an area up to 150 feet away from the ponds had been girdled to a height of about two feet. Pines nearest the ponds bore old scars and many were dead, whereas those on the periphery of the girdled area were freshly cut. No lodges were observed, although a cache of deciduous limbs and saplings was noted on a pond below Slick Creek Spring. Active dams also were found at other localities in the Long Pine Hills, as follows: Wick- ham Gulch, 2 mi N and 4.5 mi W Camp Crook, 3700 ft; Lantis Springs, 5.5 mi N and 10 mi W Camp Crook, 3500 ft; 4 mi N and 7.5 mi W Camp Crook, 3800 ft. Visher (1914:89) and others have reported beaver activity in Harding County, South Dakota—on Boxelder Creek and along the Little Missouri River. Beaver probably occur in these drainages in Carter County as well. Reithrodontomys megalotis dychei J. A. Allen, 1895 Western Harvest Mouse Specimens examined (2).—5 mi N, 5.5 mi W Camp Crook, 3500 ft, 1; 5 mi N, 3.5 mi W Camp Crook, 3400 ft, 1. This harvest mouse evidently is uncommon in southeastern Mon- tana. Our only two specimens, a subadult male and an adult male (testes 8 and 9 mm in length, respectively) were trapped in July from dense grassy areas in the eastern Long Pine Hills. THE MAMMALS OF CARTER COUNTY, MONTANA 19 Peromyscus maniculatus nebrascensis (Coues, 1877) Deer Mouse Specimens examined (152).—11.5 mi N, 3 mi E Ekalaka, 23; 16 mi E Ekalaka, 3150 ft, 3; 17 mi E Ekalaka, 3200 ft, 15; 2 mi S, 1.5 mi E Ekalaka, 1 (MMNH); 3 mi S, .75 mi E Ekalaka, 7 (MMNH); 3 mi S, 1.75 mi E Eka- laka, 2 (MMNH); 3.75 mi S, 2.25 mi W Ekalaka, 4 (MMNH); 4.5 mi S, 1 mi E Ekalaka, 2 (MMNH); 11 mi S, 7.5 mi W Ekalaka, 2 (MMNH); 11 mi S, 7 mi W Ekalaka, 2; 11 mi S, 6.5 mi W Ekalaka, 3 (MMNH); 10.5 mi N, 5 mi W Camp Crook, 3400 ft, 1; 9 mi N, 11 mi W Camp Crook, 3600 ft, 1; 9 mi N, 3 mi W Camp Crook, 3400 ft, 2; 8.5 mi N, 8 mi W Camp Crook, 1; 7 mi N, 3 mi W Camp Crook, 3300 ft, 1; 5.5 mi N, 10 mi W Camp Crook, 3950 ft, 8 (6 MMNH); 5 mi N, 6 mi W Camp Crook, 3500 ft, 14; 5 mi N, 3.5 mi W Camp Crook, 3400 ft, 31; 4 mi N, 8 mi W Camp Crook, 3650 ft, 7; 3.75 mi N, 7.75 mi W Camp Crook, 1 (MMNH); 2 mi N, 4.5 mi W Camp Crook, 3700 ft, 7; 8 mi S, 3 mi W Camp Crook, 3200 ft, 14. The deer mouse was by far the most common and widely dis- tributed mammal in Carter County. We obtained specimens from all terrestrial habitats except dense stands of ponderosa pine having a sparse understory. Adults, subadults, and young (in juvenile pelage) were obtained in various stages of seasonal and maturational molt throughout July and in early August. A pregnant female was still in worn winter pelage on 25 July and an adult male and a pregnant female were molting to summer pelage as late as 22 July. Adults of both sexes were taken in fresh summer pelage as early as 1 July, but most indi- viduals obtained early in July still were in winter pelage. In the first half of July, 12 females carried an average of 5.2 (3-7) embryos, the crown-rump length of which ranged from 3 to 14 mm. Eight individuals collected at this same time had an average of 6.1 (4-9) recent placental scars and two individuals were lactating. One captured during this period in 1972 had 11 placental scars of at least two age categories. In the second half of July, nine females carried an average of 5.0 (3-6) embryos that varied from 3 to 18 mm in length. An average of 4.5 placental scars was noted for four indi- viduals. Thirty-five adult males collected in the first half of July had testes with a mean length of 9.6 (8-11) mm, and 15 taken in the last half of July had testes whose average length was 9.9 (8-13) mm. Additionally, we obtained two females on 30 June that had four and five recent placental scars. Five males obtained on that date had testes that were 10.2 (10-11) mm in mean length. A subadult female taken on 1 August carried three 2 mm embryos. Peromyscus maniculatus were parasitized by the following ecto- parasites: a mite, Androlaelaps fahrenholzi; ticks, Dermacentor andersoni and Ixodes sculptus; a louse, Hoplopleura hesperomydis; and fleas, Callistopsyllus terinus, Monopsyllus eumolpi eumolpi, Monopsyllus wagneri, and Monopsyllus sp. Three specimens were parasitized by cuterebrid larvae. 20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Onychomys leucogaster missouriensis (Aubudon and Bachman, 1851) Northern Grasshopper Mouse Specimen examined (1).—11.5 mi N, 3 mi E Ekalaka, 1. A subadult male was trapped in a sandy draw bordered by short grass in Medicine Rocks State Park, an area of massive rock out- croppings in northern Carter County from which Preble (MS) also listed specimens. Although 240 traps were set on three consecutive nights in this area in mid-July, no additional Onychomys were ob- tained; however, Peromyscus maniculatus, Spermophilus tridecem- lineatus, and Perognathus fasciatus were obtained from the same or adjacent trap lines. The rarity of grasshopper mice in eastern Mon- tana was earlier mentioned by Preble (MS). Our male, taken on 15 July, was molting actively as evinced by patches of new pelage on both shoulders and rump. The testes were 12 mm in length. Neotoma cinerea rupicola J. A. Allen, 1894 Bushy-tailed Woodrat Specimens examined (3).—11 mi S, 6.5 mi W Ekalaka, 1 (MMNH); 7 mi N, 3 mi W Camp Crook, 3400 ft, 1; 3.75 mi N, 7.75 mi W Camp @rooko4 (MMNH). Our three bushy-tailed woodrats were nulliparous females, ap- parently young of the year. One, trapped on 5 July under a rock ledge, was molting on the sides and rump. The other two, taken on 12 and 17 July, were completing molt into adult pelage. One was trapped under a large rock near the summit of Trenk Pass in the Chalk Buttes. The other was removed from an abandoned building. Nests and old fecal pellets of woodrats were found frequently in the Long Pine Hills; Merlin Tuttle examined contents of owl pellets (thought to be those of a great horned owl) from an abandoned farm in the Ekalaka Hills that contained remains of these woodrats. Andersen and Jones (1971:380) recorded a specimen from 7 mi N and 2.5 mi W Camp Crook, South Dakota, at the extreme eastern edge of the Long Pine Hills. Although none of our specimens was fully adult, we assign them to the subspecies rupicola rather than orolestes. They are indistin- guishable from a series assigned to rupicola by Andersen and Jones (1971), which includes several individuals of comparable age from Harding County, South Dakota. We are convinced that woodrats living in the several hills and buttes of this tri-state area are of a single taxon. THE MAMMALS OF CARTER COUNTY, MONTANA 2] Microtus ochrogaster haydenii (Baird, 1858) Prairie Vole Specimens examined (6).—9 mi N, 3 mi W Camp Crook, 3300 ft, 3; 5 mi N, 3.5 mi W Camp Crook, 3400 ft, 3. Three specimens were trapped in an ungrazed upland meadow, 5 mi N and 3.5 mi W Camp Crook, and the remaining three were collected from the semi-marshy banks of a small stream in a lightly grazed pasture of sweet clover. M. pennsylvanicus and M. ochro- gaster were taken together at the former locality; however, 600 trap nights in the vicinity of the stream at the latter locality produced no specimens of pennsylvanicus. Jones (1964:228), Turner (1974:116), and others have noted that the prairie vole occupies riparian habi- tats in the absence of pennsylvanicus but is excluded from such habitats when both species are present. One praire vole harbored a flea, Orchopeas leucopus, and a chig- ger, Euschoengastia setosa. Microtus pennsylvanicus insperatus (J. A. Allen, 1894) Meadow Vole Specimens examined (71).—1.75 mi S, 1 mi E Ekalaka, 1 (MMNH); 2 mi S, 1.5 mi E Ekalaka, 2 (MMNH); 3 mi S, 1.75 mi E Ekalaka, 27 (MMNH); 8.5 mi N, 8 mi W Camp Crook, 3400 ft, 1; 5.5 mi N, 10 mi W Camp Crook, 10 (MMNH); 5 mi N, 10 mi W Camp Crook, 2; 5 mi N, 6 mi W Camp Crook, 3500 ft, 1; 5 mi N, 3.5 mi W Camp Crook, 3400 ft, 5; 3.75 mi N, 7.75 mi W Camp Crook, 18 (MMNH); 2 mi N, 4.5 mi W Camp Crook, 3700 ft, 1; 8 mi S, 3 mi W Camp Crook, 3200 ft, 3. The meadow vole was the most abundant microtine in Carter County. It was found in grasslands and lush riparian habitats in the Long Pine Hills as well as along the Little Missouri River. Although we have no specimens from within the Ekalaka Hills, many of these voles were trapped along streams that flow northward from there. The number of specimens collected was small in comparison to the density of runways observed in many areas that were trapped. Preble (MS) earlier mentioned specimens from Medicine Rocks, Ekalaka, Capitol, and Albion. Of 26 females obtained in July, 10 carried an average of 6.7 embryos (5-9) that ranged in length from 4 to 29 mm. Four females were recorded as lactating in early and mid-July; three other indi- viduals had recent placental scars. Testes of 34 adult and subadult males taken in mid-July were 14.6 (9-20) mm in average length and those of an adult taken in early August were 18 mm. Molt was observed in adult, subadult, and juvenile individuals collected in July and early August. Our specimens of meadow voles were parasitized by a mite, Laelaps kochi, and a louse, Hoplopleura acanthopus. 22 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Lagurus curtatus pallidus (Merriam, 1888) Sagebrush Vole Specimen examined (1).—16 mi E Ekalaka, 3150 ft, 1. An adult male, trapped on 8 July on a sagebrush flat in the north- eastern part of the county, was the only sagebrush vole obtained. Fecal pellets and runways occasionally were found under large clumps of sagebrush in this area, but extensive trapping, employing both break-back and can traps, produced no additional individuals. Peromyscus maniculatus and Spermophilus tridecemlineatus were obtained in this habitat. Bailey (1927:101) earlier commented on the difficulty of collecting sagebrush voles in North Dakota. The species evidently is relatively rare and of localized occurrence along the eastern edge of its range. Preble (MS) mentioned specimens from Medicine Rocks, 15 mi N Ekalaka [probably 11.5 mi N and 3 mi E Ekalaka], and from 5 mi SE Ekalaka. Birney and Lampe (1972:466) reported the only specimens of sagebrush voles known from South Dakota. The spe- cies also is known adjacent to Carter County in southwestern Pow- der River County (Hoffmann et al., 1969a:592) and southwestern Bowman County, North Dakota (Genoways and Jones, 1972:24). Our specimen was molting as evinced by small irregular patches of new pelage on head, back, and sides, and had testes that were 11 mm in length. External and cranial measurements are: total length, 139 mm; tail length, 19 mm; hind foot length, 16 mm; ear length, 12 mm; weight, 38.2 gms; condylobasal length, 25.9 mm; palatal length, 14.8 mm; interorbital breadth, 3.1 mm; zygomatic breadth, 15.5 mm; maxillary toothrow length, 7.0 mm. Mus musculus Linnaeus, 1758 House Mouse Specimens examined (3).—3 mi S, 1.75 mi E Ekalaka, 1 (MMNH); 5 mi N, 3.5 mi W Camp Crook, 3400 ft, 2. Two adult house mice, a male and a female, were collected on a north-facing slope of a grassy meadow at the eastern edge of the Long Pine Hills. Both were taken on a grid that previously had been live-trapped for 10 days in a trap-release program, then snap- trapped. No house mice were obtained in live-traps on this grid nor were any taken in either live-traps or snap-traps on two adjacent grids. Our third specimen, an adult male, was trapped in a marsh along with Zapus hudsonius, Microtus pennsylvanicus, and Pero- myscus maniculatus. Preble (MS) commented that house mice generally were dis- tributed in eastern Montana in the proximity of railroads. Andersen and Jones (1971:381) recorded a single individual, trapped along a fencerow, from adjacent Harding County, South Dakota. THE MAMMALS OF CARTER COUNTY, MONTANA 23 Zapus hudsonius campestris Preble, 1899 Meadow Jumping Mouse Specimens examined (33).—8 mi S, 1.75 mi E Ekalaka, 21 (MMNH); 5.5 mi N, 10 mi W Camp Crook, 3 (MMNH); 5 mi N, 5.5 mi W Camp Crook, 3500 ft; 2; 5 mi N, 3.5 mi W Camp Crook, 3400 ft, 2; 4 mi N, 8 mi W Camp Crook, 3650 ft, 2; 3.75 mi N, 7.75 mi W Camp Crook, 1 (MMNH); 2 mi N, 4.5 mi W Camp Crook, 3700 ft, 1; 8 mi S, 3 mi W Camp Crook, 3200 ft, 1. Twenty-one of our specimens were trapped from a large marsh at the north edge of the Ekalaka Hills. The remainder were ob- tained from riparian and marshy habitats in the Long Pine Hills, along the Little Missouri River, and on a grassy upland slope at the eastern edge of the Long Pines. Of our 33 specimens, all caught in July, 25 were adult, and only eight were juvenile or subadult (see Krutzsch, 1954:377, for criteria of age classes). Testes of 13 adult males were 6.1 mm in mean length whereas those of six young males were 4.7 mm. Of the 14 females in our sample, six were carrying embryos that ranged in length from 6 to 19 mm. Mean and modal number of embryos per pregnant female was 6.0 and 7, respectively. Three adult and five young females were neither pregnant nor lac- tating. Preble (MS) recorded specimens from Ekalaka and the Sioux National Forest. Intergradation between the subspecies campestris and_ inter- medius has been suggested in Harding County, South Dakota (An- dersen and Jones, 1971:382), and in Dawson County of east-central Montana (Krutzsch, 1954:448). Average and extreme cranial meas- urements of 19 adults (11 males, 8 females) are: condylobasal length, 21.2 (20.1-22.2) mm; occipitonasal length, 23.3 (22.3-24.0) mm; palatal length, 10.1 (9.7-10.5) mm; mastoidal breadth, 10.3 (10.0-10.7) mm; maxillary toothrow length, 3.7 (3.5-4.1) mm. In their more ochraceous color, our specimens more closely resemble campestris from Harding County, South Dakota and the Black Hills, than intermedius from North Dakota. Erethizon dorsatum bruneri Swenk, 1916 Porcupine Specimen examined (1).—7 mi N, 3 mi W Camp Crook, 3400 ft, 1. An adult male weighing 27 pounds was shot in the late afternoon of 4 July in a den located under a rock ledge on a pine-clad ridge along the eastern edge of the Lone Pine Hills. In addition, we ob- served an individual in a meadow, 5 mi N and 3.5 mi W Camp Crook, and two others on a road near Lantis Spring, 5.5 mi N and 10 mi W Camp Crook. Fresh sign of porcupines was seen elsewhere in the Long Pines. The species evidently is fairly common there and, presumably, in adjacent pine-covered areas. 24 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY ORDER CARNIVORA Canis latrans latrans Say, 1823 Coyote The coyote population of Carter County was apparently ex- tremely low at the time of our studies, perhaps as a result of active predator control practices, including 1080 poison. This compound is currently banned, and coyotes are reported increasing in many places. Our field parties neither saw nor heard coyotes. Visher (1914:90) reported that during the winter of 1910-11 one trapper collected 102 coyotes in the Ekalaka Forest. During the period July 1970 to June 1971, bounties were paid on four coyotes purportedly taken in Carter County. Andersen and Jones (1971:382) listed two specimens obtained by a federal trapper during the winter of 1961- 62 along the eastern edge of the Long Pine Hills, 6 mi N and 4 mi W Camp Crook, Harding County, South Dakota. Canis lupus nubilus Say, 1823 Wolf The wolf, although now extirpated (Goldman, 1944:442), once ranged throughout Carter County. Bailey (1907:10) reported that in 1893 “wolves were common at Alzada and Powderville, and were very destructive to stock, especially calves and colts.” Visher (1914: 90) noted that during the winter of 1909-10 a trapper took four wolves in the Ekalaka Forest, and Goldman (1944:444) listed a specimen from Ekalaka. Extirpation of the wolf in Carter County most likely paralleled in time its disappearance from Harding County, South Dakota (see Andersen and Jones, 1971:383). Populations may have remained slightly longer than elsewhere in the Long Pine Hills and forested areas near Ekalaka, although such areas were exploited heavily by commercial trappers. Vulpes vulpes regalis Merriam, 1900 Red Fox We took no specimens of this fox in Carter County, but an imma- ture female, captured approximately 5 mi W Camp Crook, was held as a pet by a local family. An adult was sighted on the night of 26 July 1970 on a sagebrush flat just north of the Long Pines and an- other was seen on the night of 17 July 1971 on a U.S. Forest Service road in the Ekalaka Hills. Visher (1914:90) reported the red fox as “not rare about the Long Pines” in the early part of the century. During the period 1 July 1970 to 16 July 1971, Carter County paid bounties on 958 red foxes. The extent to which predator con- trol programs have affected red fox populations is unknown, but Andersen and Jones (1971:383) reported that in adjacent Harding THE MAMMALS OF CARTER COUNTY, MONTANA 25 County, South Dakota, this fox “seems to maintain considerably higher populations than does the coyote,” in spite of control pressure. Ursus arctos horriblilis Ord, 1815 Grizzly Bear The skull of a grizzly bear displayed in the Carter County Mu- seum in Ekalaka, was found in 1920 on Cottonwood Creek, T 5 S and R 61 E, in southern Carter County. The specimen evidently had been shot inasmuch as there appears to be a bullet hole above the orbits. Visher (1914:91) noted that during the early 1890's a grizzly bear was killed a short distance southwest of Camp Crook. Additionally, a cave wall in the Cottonwood Creek area carries the inscription “Killed Bear in nest” (M. E. Lambert, pers. com.), which may refer to this or the following species. Ursus americanus americanus Pallas, 1780 Black Bear Visher (1914:91) reported that “Bears have been recently killed in the Long Pine and Ekalaka forests, but their day of extermination is here near at hand.” Preble (MS) indicated that black bears for- merly were widely distributed in eastern Montana. Procyon lotor hirtus Nelson and Goldman, 1930 Raccoon Our only records of the raccoon in Carter County are of tracks found in the Long Pines along the edge of a beaver pond, 4 mi N and 8 mi W Camp Crook, and a specimen from 3 mi W Alzada (Hoffmann et al., 1969a:595). An adult was observed on several nights in July, however, on a ridge at the extreme eastern edge of the Long Pine Hills just inside Harding County, South Dakota, and Andersen and Jones (1971:384) reported a litter of five young found in May in this same area. They noted that this species “evidently is one of the commonest carnivores” in northwestern South Dakota and raccoons likely occur along the Little Missouri River and in other suitable riparian habitats in Carter County. Mustela frenata longicauda Bonaparte, 1838 Long-tailed Weasel Specimens examined (2).—3 mi S, 1.75 mi E Ekalaka, 1 (MMNH); 7 mi N, 4 mi W Camp Crook, 1. The mummified body of a long-tailed weasel in winter pelage was found on a sparsely forested ridge at the eastern edge of the Long Pine Hills. From size and cranial characters, we judge this specimen to be a female. A second weasel, a young male, was cap- tured in a hand-held rat trap by Larry Heaney at the entrance to a 26 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY pocket gopher burrow on 17 July. A partially eaten gopher had been removed from the burrow minutes before following a minor tug-of-war with the weasel. The rat trap was then baited with meat of the dead gopher. Preble (MS) noted that this species “. . . ap- pears generally distributed but nowhere abundant” in southeastern Montana. Mandibular formation is anomalous in the mummy; the length of the right mandible is 25.8 mm whereas the length of the left man- dible is 28.1. Extension of the left mandible caused a slight labial curve in the distal portion of the right mandible, nonalignment of the left upper and lower canines, and malocclusion of upper and lower incisors. The lower left canine does not occlude with the upper, but is directed more anteriorly and occludes with the left i2 and i3, which in turn have been pushed posteriorly toward the upper left canine. Both second lower incisors are situated posterior to 11 andii3. Mustela nigripes (Aububon and Bachman, 1851) Black-footed Ferret Specimen examined (1).—Piniele, 1. Cahalane (1954:419) reported that an adult male black-footed ferret was killed on 3 November 1953 on a highway north of Alzada, in southern Carter County. Additionally, Henderson et al. (1969) listed seven localities of record in Harding County, South Dakota, two of which are in townships adjoining Carter County (T 19 N, R 1 E, and T 15 N, R1E), and we have examined museum specimens from Custer County, Montana, adjacent to Carter County to the northwest. A skin and partial skeleton (KU 14411) of an adult mentioned by Hoffmann et al. (1969a:597) was obtained in October 1944 at Piniele in southwestern Carter County. In view of its small size, this speci- men most likely represents a female. Selected cranial measurements are: condylobasal length, 66.8 mm; palatal length, 30.8 mm; mastoid breadth, 33.8 mm; least interorbital breadth, 16.7 mm; postorbital breadth, 10.7 mm; maxillary toothrow length, 19.1 mm. Close dependence of the black-footed ferret on prairie dogs as a major food source no doubt has had a serious impact on the current status of this mustelid, although it may never have been abundant. Efforts to reduce or extirpate prairie dogs continue in Carter County and elsewhere in southeastern Montana. Taxidea taxus taxus (Schreber, 1778) Badger Specimen examined (1).—8 mi S, 3 mi W Camp Crook, 3700 ft, 1. Badgers probably occur throughout Carter County. A subadult male with testes that were 49 mm in length was trapped on 8 July THE MAMMALS OF CARTER COUNTY, MONTANA 27 at a den located in the north face of a gulley along the Little Mis- souri River. The stomach of this specimen contained the remains of three Thomomys talpoides, a small bird, five bees, and unidentified plant material. A subadult observed crossing a road in the sage- brush flats to the north of the Long Pine Hills, 1 mi S and 13 mi E Ekalaka, 3200 ft, in the late afternoon of 14 July escaped to a nearby burrow after it dropped an immature prairie dog that it carried in its mouth. Our one specimen harbored a tick, Ixodes kingi. Mephitis mephitis hudsonica Richardson, 1829 Striped Skunk Specimens examined (4).—8.5 mi N, 8.5 mi W Camp Crook, 3400 ft, 2; 6 mi N, 9 mi W Camp Crook, 2. Striped skunks evidently are common in the Long Pine Hills and surrounding areas. Two of our four specimens were collected from a densely wooded slope in the Long Pines—one trapped along a game trail and the other picked up in the field as a skeleton. The other two specimens, a young male (testes 2 mm in length) and an adult female (six placental scars), were shot on the night of 16 July along a road in rolling grasslands just to the north of the eastern part of the Long Pines. Striped skunks were observed frequently in the Long Pine Hills. Our only sight record other than from that area was of a young animal killed on a road in the vicinity of Sheep Mountain, 4 mi S and 11 mi W Camp Crook. Andersen and Jones (1971:385) re- ported an observation from just east of the Long Pines at a place 6 mi N and 2.5 mi W Camp Crook, Harding County, South Dakota. The young male shot on 16 July was parasitized by a tick, Ixodes kingi. Felis concolor hippolestes Merriam, 1897 Mountain Lion Mountain lions, “formerly of quite general distribution over the eastern part of Montana” (Preble, MS), have been extirpated in the southeastern part of the state. Visher (1914:91) noted that the mountain lion that visited Harding County, South Dakota, during the winter of 1910-11 “was doubtless a roamer from the Long Pines or Ekalaka Forests.” The presence of this cat in Carter County prob- ably now is limited to an occasional transient. ORDER ARTIODACTYLA Cervus elaphus canadensis Erxleben, 1777 Wapiti or Elk Prior to the appearance of homesteaders and hunters, elk were 28 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY common on the plains of eastern Montana. In a summary of his- torical accounts of big game in Montana, Koch (1941:368) noted “that they [elk and bison] occurred in vastly greater numbers in the plains country, with some timber or mountain shelter, than they did in the higher mountain country. ...” Preble (MS) mentioned that elk formerly were distributed over southeastern Montana and Visher (1914:87) reported that the “last [elk] were killed in 1879 when a large crew cut trees for the N.P.R.R. in the Long Pines.” A fragment of an elk antler was found (M. E. Lambert, pers. com.) in “a timbered area of the Chalk Buttes about twenty miles south of Ekalaka.” Odocoileus hemionus hemionus (Rafinesque, 1817) Mule Deer Mule deer commonly were seen in meadows and occasionally along ridges in the Lone Pine Hills and Chalk Buttes. Preble (MS) reported this deer to be generally distributed throughout eastern Montana in the early part of this century. Mule deer may be more numerous in Carter County than are white-tailed deer; in the Long Pines, however, we found the latter to be the more numerous. Sight records other than from the Long Pine Hills are from 1.5 mi N and 5.9 mi E Ekalaka (six bucks) and 4 mi SE Ekalaka (four does and one fawn). Odocoileus virginianus dacotensis Goldman and Kellogg, 1940 White-tailed Deer Specimens examined (6).—8.5 mi N, 13 mi W Camp Crook, 2 (MMNH); 5.5 mi N, 10 mi W Camp Crook, 3400 ft, 1; 5 mi N, 3.5 mi W Camp Crook, 3400 ft, 1; 4 mi N, 9 mi W Camp Crook, 3950 ft, 2. We frequently observed white-tailed deer in the Long Pine Hills, mainly along the pine-covered ridges and in meadows adjacent to such ridges. Our only observation of white-tails outside the Long Pines was of a doe, accompanied by a fawn, resting among decid- uous brush along the Little Missouri River at a place 8 mi S and 3 mi W Camp Crook. Preble (MS) reported white-tailed deer “. . . to be more commonly met with in the southeastern part of the State than elsewhere [in eastern Montana].” No group of more than five deer was observed. Several does with fawns repeatedly were seen grazing in grassy draws in early and mid-July. All of our specimens are skulls picked up in the Long Pines. Antilocapra americana americana (Ord, 1815) Pronghorn Pronghorns regularly were seen in the grasslands, and occasion- ally in meadows, along the margin of the Long Pines in the summer THE MAMMALS OF CARTER COUNTY, MONTANA 29 of 1970. Beer (1944:45) reported that in 1941 Carter County had the largest number of pronghorns, an estimated 2500, of any county in Montana. He also noted that in severe winters pronghorns mi- grated into the area from South Dakota, seeking shelter among the breaks and ridges along the Little Missouri River. Andersen and Jones (1971:387) reported sighting herds of up to 50 individuals in adjacent Harding County, South Dakota; all of our observations, however, were of groups of five or less. While conducting an aerial census of pronghorns in Carter County on 28 January 1948, Thompson (1949:313) observed an adult golden eagle attacking a single adult male pronghorn near the rear of a herd of 64 individuals. In three consecutive attacks, the eagle grasped the back of the animal and rode with outspread wings as the pronghorn fled with the herd across open grassland. We have no such observations of predation, but golden eagles were sighted in the Long Pines as well as over adjacent grasslands. An analysis of pronghorn stomach contents, conducted in Carter County from September through December of 1944 and 1945 (Couey, 1946:367), revealed that sagebrush (Artemisia tridentata and A. cana), snowberry (Symphoricarpos sp.), and snakeweed (Guterriezia sp.) composed 75 per cent of the diet by volume. Bison bison bison (Linnaeus, 1758) Bison Vast herds of bison are known to have occurred along the Mis- souri and Yellowstone rivers as well as across the prairies of what is now Carter County until the middle of the 19th century. During the exploration of the Yellowstone River and its tributaries in 1859, Captain W. F. Raynolds and his party crossed the extreme south- western corner of Carter County. While camped near the present border between Carter County, Montana, and Crook County, Wy- oming, on 21 June, he noted (1868:33): “We are now in the buffalo region, and small herds are to be seen in all directions. Their pres- ence may explain some of the prevalent barrenness, as they consume all the grass in their paths.” During an expedition to the Black Hills in July 1874, Custer traveled down the Little Missouri River and entered southeastern Carter County, then part of the Montana Ter- ritory. Custer found (1875:1) “this valley [of the Little Missouri River] almost destitute of grazing .. .” a condition perhaps attribu- table to bison. J. A. Allen (1877:545) noted that by 1877, between the Platte and Missouri rivers, bison were confined to the area of the Bighorn, Tongue, and Powder rivers. Indian hostilities, which prevented ex- ploitation of the eastern Montana herds, were curtailed in the mid- 1870’s, opening the territory to “civilization” and the extirpation of the remaining bison. Visher (1914:88) reported “a small bunch” in 30 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Harding County, South Dakota, in 1886 and commented on the occurrence of numerous bullet-riddled skulls testifying to the activ- ities of “market shooters.” Ovis canadensis auduboni Merriam, 1901 Bighorn Sheep Bighorn sheep once were abundant in eastern Montana (Preble, MS) and specimens were collected in the late 1800's in the badlands and high plateaus along the Little Missouri River in North Dakota (Buechner, 1960:20). The mounted head of a bighorn displayed in the Carter County Museum, Ekalaka, Montana, was obtained by Dan Bowman in 1893 from the Powder River near Locate, Custer County, Montana. The only record of bighorn in Carter County consists of two pairs of horn cores, one “dug out of a spring twenty miles west of Ekalaka on Spring Creek,” and the other “found on Tie Creek south of the Long Pines” (M. E. Lambert, pers. com.). The specimens are tenta- tively assigned to O. c. auduboni, a subspecies now considered ex- tinct (Cowan, 1940:542). OTHER SPECIES Three additional species, the muskrat (Ondatra zibethicus), the mink (Mustela vison), and the bobcat (Lynx rufus) most assuredly are found in Carter County as indicated by the availability of suit- able habitats and the presence of recent records (Andersen and Jones, 1971) from nearby areas. Although we obtained neither speci- mens nor sight records of these species, all three were reported by local residents. Additionally, during the fiscal year 1970-71, bounties were paid in Carter County for 130 bobcat and lynx. Individuals were not identified to species, but we believe that most, if not all, were Lynx rufus. Species of Unverified Occurrence Twelve species listed below may occur (or in the recent past probably occurred) in Carter County. The possible presence of 10 species is inferred by their distribution and habitat preference in areas adjacent to Carter County. A published account of migration of the Norway rat (Tryon, 1947) and the mention of lynx in an unpublished manuscript by Preble also are noted. Preble’s mention of Lepus americanus from the forested areas near Ekalaka almost certainly was in error as this hare now occurs no nearer Carter County than in the Killdeer Mountains of North Dakota (Genoways and Jones, 1972:9), Richland County, Montana (Hoffmann e¢ al., 1969a:584), and the Bighorn Mountains of Wyoming (Long, 1967: 548). Sylvilagus nuttallii grangeri (J. A. Allen, 1895).—Although this cottontail is not presently known from Carter County, it may be THE MAMMALS OF CARTER COUNTY, MONTANA 31 found along the wooded and brushy streams in the hills along the western and southern edge of the county. Specimens have been taken a short distance to the south in Crook County, Wyoming (Long, 1965:544). Marmota flaviventris dacota (Merriam).—The yellow-bellied marmot is known to occur in the Black Hills (Turner, 1974:68) and adjacent regions in Wyoming (Long, 1965:570). Hoffmann et al. (1969a:568) reported marmots from Rosebud County, Montana, which is west of Carter County. The presence of yellow-bellied marmots among rocky outcrops in Carter County is possible, al- though we observed no sign of the species. Reithrodontomys montanus albescens Cary, 1903.—The plains harvest mouse has been reported (Andersen and Jones, 1971:378) from adjacent Harding County, South Dakota, where it was rare in upland grassy habitats. In northeastern Wyoming, it is known from northern Campbell County (Long, 1965:626). The species has not been reported from Montana. Peromyscus leucopus aridulus Osgood, 1909.—The white-footed mouse may be present in the county in small discontinuous popula- tions, similar to the isolated population reported by Andersen and Jones (1971:378) in adjacent Harding County, South Dakota. The species is known from Crook County, Wyoming (Long, 1965:634), and elsewhere in Montana (Hoffmann et al., 1969a:590). However, we trapped unsuccessfully for this species in a variety of habitats. Rattus norvegicus (Berkenhout, 1769).—Tryon (1947:188) in- cluded northeastern Carter County within the range of migration of rats that had reached eastern Montana in the late 1940’s from North Dakota. Migrational routes in eastern Montana followed highways and railroads, and therefore, were primarily in valleys. We know of no specimens of the Norway rat from Carter County. Vulpes velox (Say, 1823).—The swift fox no doubt occurred in Carter County within historic times, but there are no actual records of its presence. Hoffmann et al. (1969a:594) regarded the species as “now probably extinct in the state.” However, it is noteworthy that a specimen was taken in nearby Slope County, North Dakota, on 20 February 1970 (Pfeifer and Hibbard, 1970:835). Urocyon cinereoargenteus ocythous Bangs, 1899.—Although no records of the gray fox are known from Montana, an adult male has been recorded (Jones and Henderson, 1963:283) from 43 miles east of Carter County at Deer Ear Buttes, Butte Co., South Dakota. This record constituted a considerable westward extension of the then known range and possibly is indicative of a trend in dispersal of this species. Mustela erminea muricus (Bangs, 1899).—No specimens of this weasel have been reported from eastern Montana. However, the 32 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY proximity of a recent record (Long, 1965:692) from Crook County, Wyoming, at a place 40 miles south of Carter County, suggests the possible occurrence of the ermine in the southeastern part of the state. Gulo gulo luscus (Linnaeus, 1758).—The wolverine, although probably never abundant, likely occurred in southeastern Montana within historic time. Spilogale putorius interrupta (Rafinesque, 1820).—Southeastern Montana was included in the distribution of the spotted skunk by Hoffmann and Pattie (1968:117), although no specimens have been reported from the area. Visher (1914:91) commented that the spotted skunk was more abundant than the “large skunk” in north- western South Dakota, but Andersen and Jones (1971:385) more recently reported the species as rare there. Lontra canadensis pacifica (Schreber, 1776).—Preble (MS) com- mented that the otter once occurred in small numbers in eastern Montana, and Visher (1914:91) reported a specimen taken on the Little Missouri River in Harding County, South Dakota. We follow van Zyll de Jong (1972) in the use of generic and subspecies names of the otter. Lynx canadensis canadensis Kerr, 1792.—Lynx populations in Montana have recently shown a marked increase, but this cat is un- common in the southeastern part of the state (Hoffmann et al., 1969a:598). No records of the species are known from Carter County, where it no doubt once occurred and may again be found. Although it has been suggested that L. canadensis is conspecific with L. lynx of the Old World, we follow Kurtén and Rausch (1959: 41) in regarding them as specifically distinct. ZOOGEOGRAPHIC COMMENTS Faunal Components The 50 native North American species known as recent inhabi- tants of southeastern Montana exhibit five distributional patterns, the implications of which have been discussed by Hoffmann and Jones (1970). The largest faunal grouping consists of species having widespread distributions. These species either occupy much of tem- perate North America, or have distributions that do not coincide with any major faunal unit. The 23 species so categorized are: Myotis leibii, Myotis lucifugus, Lasionycteris noctivagans, Eptesicus fuscus, Lasiurus cinereus, Castor canadensis, Peromyscus manicu- latus, Erethizon dorsatum, Canis latrans, Canis lupus, Vulpes vulpes, Ursus arctos, Ursus americanus, Procyon lotor, Mustela frenata, Taxi- dea taxus, Mephitis mephitis, Felis concolor, Cervus elaphus, Odo- coileus hemionus, Odocoileus virginianus, Antilocapra americana, and Bison bison. Many of these mammals are vagile and eurytopic species, or species with specializations that are met in more than a ~— THE MAMMALS OF CARTER COUNTY, MONTANA 33 single biotope (for example, the beaver). The wolf, grizzly bear, black bear, mountain lion, elk, bison, and bighorn sheep have been extirpated in southeastern Montana directly or indirectly by man. The reduction of coyote populations elsewhere in Montana ( Hoff- mann et al., 1969a:596) and in Wyoming (Robinson, 1961, and Lin- hart and Robinson, 1972) had been followed by increased popu- lations of other carnivores, such as skunk, badger, lynx, and red fox; the last-mentioned species recently has extended its range westward in Montana (Hoffmann et al., 1969a:594). Although these wide-ranging carnivores occurred in grasslands, the mountain lion and black bear were more likely to be found among rocky buttes and ridges and the raccoon is more typical of the floodplain forests. The porcupine, although associated princi- pally with coniferous forests, herein is considered a “widespread species” because of its extensive distribution. The mule deer and pronghorn are essentially western species, whereas the white-tailed deer is a wide-ranging species. Although a common plains mammal, the bison had an extensive northern and eastern distribution. Three of the five bats are apparently permanent residents of the area, Lasiurus and Lasionycteris being migrants. Boreomontane species—Three bats, Myotis evotis, Myotis vo- lans, Plecotus townsendii, as well as Thomomys talpoides, Neotoma cinerea, and Ovis canadensis are primarily montane species, al- though the first, second, third, and sixth species also have extensive southwestern distributions. Four species (Sorex cinereus, Eutamius minimus, Tamiasciurus hudsonicus, and Microtus pennsylvanicus ) occur in boreal areas to the north as well as montane areas to the west. Zapus hudsonius, basically a boreal species, also inhabits much of the eastern deciduous forest. All of these species, except the three bats, have developed distinctive subspecies on the Northern Great Plains. Steppe species.—Steppe species have evolved in close association with the grassland environment; they are characteristic of, and more or less restricted to, such areas. Steppe species occurring in south- eastern Montana are Lepus townsendii, Cynomys ludovicianus, Spermophilus tridecemlineatus, Perognathus fasciatus, Perognathus hispidus, Microtus ochrogaster, and Mustela nigripes. Indicative of such autochthonous influence is the close predator-prey relationship of the prairie dog and the black-footed ferret. Sonoran species.—Species that have invaded the area from the Southwest are Sylvilagus audubonii, Dipodomys ordii, Reithrodon- tomys megalotis, and Onychomys leucogaster. The last two species now inhabit much of the Northern Great Plains as well as areas of the Great Basin. Great Basin species——Two species have invaded the plains from the Great Basin. The sagebrush vole (Lagurus curtatus) most likely 34 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY entered the Northern Great Plains through the Wyoming Basin (Hoffmann and Jones, 1970:370). Knowledge of the distribution of Sorex merriami, although this shrew is widespread in the Great Basin, is limited by a lack of specimens. Eastern species——The eastern cottontail (Sylvilagus floridanus) reaches its northwesternmost known distribution in Carter County, where it is closely associated with riparian habitats. Found pri- marily east of the Rocky Mountains, one subspecies of this rabbit, S. f. similis, has developed on portions of the Northern and Central Great Plains. Another eastern species, Sciurus niger, is a recent in- vader, and most likely entered the area from the northeast along the Little Missouri River, where it is restricted to the deciduous flood- plain forest. Unverified species—Eleven of the 12 species of unverified oc- currence in Carter County are native North American mammals. Of these 11, five (Sylvilagus nuttallii, Marmota flaviventris, Mustela erminea, Gulo gulo, and Lynx canadensis have boreal or montane affinities, two (Peromyscus leucopus and Urocyon cinereoargenteus ) are associated with the easern forest, three (Reithrodontomys mon- tanus, Vulpes velox, and Spilogale putorius) are steppe species, and one (Lontra canadensis) is a widespread species. Faunal Comparisons In order to add greater perspective to the zoogeographic rela- tionships of the mammalian fauna of southeastern Montana, we have compared it with the faunas from two other sites on the Northern Great Plains. Southern Saskatchewan, a Transition and Upper So- noran zone of parkland and prairie bordered on the east, south, and west by the Saskatchewan and Qu’Appelle rivers, is representative of the northern extent of the Great Plains. Northwestern Nebraska (Sioux, Dawes, and Sheridan counties) comprises a southern site that includes the Pine Ridge district of the High Plains and the Pierre Plains of the Missouri Plateau. Carter County, Montana, and ad- jacent Harding County, South Dakota, form a unit representing a central location on the Northern Great Plains. The mammals of southern Saskatchewan have recently been examined by Beck (1958) and Soper (1961). The distributional records of Jones (1964) and McDaniel (1967) were used to compile a list of mammals of northwestern Nebraska. The foregoing ac- counts and those by Andersen and Jones (1971) for adjacent Hard- ing County were combined for the mammalian fauna of the central site. The past and present distribution of many carnivores and artiodactyls is uncertain and members of these two orders thus are excluded from the following discussion. Of a total of 51 species of small mammals found at the three sites, 24 species are common to all. Although the number of THE MAMMALS OF CARTER COUNTY, MONTANA 35 TABLE 2. Composition of small mammal faunas of three regions on the Northern Great Plains; see text for discussion and sources of figures. Northwestern Northwestern South Dakota Southern Taxa Nebraska Southeastern Montana Saskatchewan Insectivora 3 2 4 Chiroptera ite 8 6 Lagomorpha 4 3 4 Rodentia 3 29, pa) species present at each site is similar (Table 2), the species compo- sition varies considerably. Of the 11 species restricted to southern Saskatchewan, six (Sorex vagrans, Microsorex hoyi, Sylvilagus nut- talii, Lepus americanus, Glaucomys sabrinus, and Clethrionomys gapperi) are of boreal or montane distribution and three (Blarina brevicauda, Lasiurus borealis, and Marmota monax) have affinities with the eastern deciduous forest. Two sciurid rodents, Spermophi- lus richardsonii and Spermophilus franklinii, are restricted to the northern site and are, respectively, of Great Basin and (eastern) steppe affinities. A montane bat species, Plecotus townsendii, is, among the three faunas, the only species found only in the south- eastern Montana-northwestern South Dakota area. The five species found only in northwestern Nebraska consist of two steppe species (Geomys bursarius and Perognathus flavescens), two Sonoran spe- cies (Lepus californicus and Spermophilus spilosoma), and one eastern species (Scalopus aquaticus). It is apparent, although not surprising, that the northern and southern sites support faunistic elements of somewhat different distributional affinities. It is inter- esting to note that but a single species is found solely at the central site. The two species common only to the northern and central sam- ples are the boreal Tamiasciurus hudsonicus and the invader from the Great Basin, Lagurus curtatus. Two steppe species (Pero- gnathus hispidus and Reithrodontomys montanus), two Sonoran species (Sylvilagus audubonii and Reithrondontomys megalotis), two montane species (Myotis evotis and Myotis volans), one eastern deciduous woodland species (Sciurus niger), and one Great Basin species (Sorex merriami) are common to the central and southern sites. There are no species shared by the northern and southern sites to the exclusion of the Montana-South Dakota region. As its geographic location suggests, the Montana-South Dakota sample represents an “intermediate” fauna inasmuch as all but one species is shared with either the northern or southern samples. The presence of eight species shared by the central and southern sam- ples in comparison to two species common to the northern and cen- tral units is noteworthy. The boreomontane element, prevalent in 36 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY southern Saskatchewan, is limited at the central site and to an even greater degree at the southern site, probably owing to such features as temperature, amount of precipitation, and nature and extent of forest habitats. Grassland invaders have been aided in their post- Wisconsin dispersal by the continuity of the grassland environment. Several of the steppe and grassland-associated species present at the southern site have reached the Montana-South Dakota area, but are absent in Saskatchewan. SUMMARY Fifty-three species of mammals are reported to have occurred in Carter County, Montana, within historical times. In addition, 12 species are noted as having a possible but unverified occurrence in the county. Annotated accounts of these 65 species provide com- ments on their distributional status and natural history in southeast- ern Montana. Zoogeographically, this part of Montana supports a mammalian fauna composed of widespread, boreomontane, steppe, sonoran, Great Basin, and eastern species. Species with widespread distributions comprise the largest single group. LITERATURE CITED ALDEN, W. C. 1932. Physiography and glacial geology of eastern Montana and adjacent areas. U.S. Geol. Surv., Prof. Paper, 174: viii + 133. ALLEN, J. A. 1877. History of the American bison, Bison americanus. Ann. Rept. U.S. Geol. and Geogr. Surv. Territories, 9:443-587. ANDERSEN, K. W., and J. K. JONEs, Jr. 1971. Mammals of northwestern South Dakota. Univ. Kansas Publ., Mus. Nat. Hist., 19:361-393. BaILeEy, V. 1907. Wolves in relation to stock, game and the national forest reserves. Bull. U.S. Forest Service, 72:1-31. 1915. Revision of the pocket gophers of the genus Thomomys. N. Amer. Fauna, 39:1-136. 1927. A biological survey of North Dakota. Ibid., 49:vi + 1-226. BAuveEr, C. M. 1924. The Ekalaka lignite field, southeastern Montana. Bull. U.S. Geol. SUEV Dl 21-268. Beck, W. H. 1958. A guide to Saskatchewan mammals. Spec. Publ. Saskatchewan Nat. Hist. Soc., 1:1-52. BEER, J. 1944. Distribution and status of the pronghorn antelope in Montana. Jour. Mamm., 25:43-46. Birney, E. C., and R. P. LAMPE 1972. Sagebrush vole (Lagurus curtatus) in South Dakota. Amer. Midl. Nat., 88:466. Brown, L. N. 1967. Ecological distribution of six species of shrews and comparison of sampling methods in the central Rocky Mountains. Jour. Mamm., 48 :617-623. THE MAMMALS OF CARTER COUNTY, MONTANA 37 BuECHNER, H. K. 1960. The bighorn sheep in the United States, its past, present, and future. Wildlife Monogr., 4:1-174. CAHALANE, V. H. 1954. Status of the black-footed ferret. Jour. Mamm., 35:418-424. Couey, F. M. 1946. Antelope foods in southeastern Montana. Jour. Wildlife Megt., 10:367. Cowan, I. McT. 1940. Distribution and variation in the native sheep of North America. Amer. Midl. Nat., 24:505-580. CustTER, G. A. 1875. A report of the expedition to the Black Hills. Exec. Doc., Senate, 2nd Sess., 43rd Congress, 32:1-9. FINLEY, R. B., Jr. 1969. Cone caches and middens of Tamiasciurus in the Rocky Mountain region. Pp. 233-273, in Contributions in mammalogy (J. K. Jones, Jr., ed.), Misc. Publ. Mus. Nat. Hist., Univ. Kansas, 51:1-428. Grenoways, H. H., and J. K. Jones, Jr. 1972. Mammals from southwestern North Dakota. Occas. Papers Mus., Texas Tech Univ., 6:1-36. Cir, }._ Ri: 1962. Tertiary landslides, northwestern South Dakota and southeastern Montana. Bull. Geol. Soc. Amer., 73:724-736. GoLpMAN, E. A. 1944. Classification of wolves. Pp. 387-507, in The wolves of North Amer- ica (S. P. Young and E. A. Goldman), Amer. Wildlife Inst., Wash- ington, D.C., xvi + 636 pp. Harn. E.R. 1951. A synopsis of the North American Lagomorpha. Univ. Kansas Publ., Mus. Nat. Hist., 5:119-202. Hatt, E. R., and K. R. KELson 1951. Comments on the taxonomy and geographic distribution of some North American rabbits. Ibid., 5:49-58. HENDERSON, F’. R., P. F. SPRINGER, and R. ADRIAN 1969. The black-footed ferret in South Dakota. South Dakota Dept. Game, Fish and Parks, Pierre, 37 pp. HIBBARD, E.. A. 1956. Range and spread of the grey and fox squirrel in North Dakota. Jour. Mamm., 37:525-531. HorrMAanwn, R. S., and J. K. Jonss, Jr. 1970. Influence of late-glacial and post-glacial events on the distribution of Recent mammals on the Northern Great Plains. Pp. 355-394, in Pleistocene and Recent environments of the central Great Plains (W. Dort, Jr., and J. K. Jones, Jr., eds.), Spec. Publ. Dept. Geol., Univ. Kansas, 3:1-433. HoFFMann, R. S., and D. L. PATTIE 1968. A guide to Montana mammals. ... Univ. Montana, Missoula, x + 133 pp. HorrManwn, R. S., P. L. Wricut, and F. E. Newsy 1969a. The distribution of some mammals in Montana. I. Mammals other than bats. Jour. Mamm., 50:579-604. HoFrrMann, R. S., D. L. Pattie, and J. F. BELL 1969b. The distribution of some mammals in Montana. II. Bats. Ibid., 50: 737-7Al1. 38 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY HO.uisTeEr, N. 1916. A systematic account of the prairie-dogs. N. Amer. Fauna, 40:1-37. Hooper, E. T. 1944. Additional records of the Merriam shrew in Montana. Jour. Mamm., 25:92. Howe tu, A. H. 1929. Revision of the American chipmunks (genera Tamias and Eutamias). N. Amer. Fauna, 52:1-157. 1938. Revision of the North American ground squirrels, with a classifica- tion of the North American Sciuridae. Ibid., 56:1-256. Jackson, H. H..T. 1928. A taxonomic review of the American long-tailed shrews (genera Sorex and Microsorex). N. Amer. Fauna, 51:vi + 1-238. Jonas, R. J. 1966. Merriam’s turkey in southeastern Montana. Tech. Bull., Montana Fish and Game Dept., 3:1-36. Jones, J. K., Jr. 1953. Geographic distribution of the pocket mouse, Perognathus fasciatus. Univ. Kansas Publ., Mus. Nat. Hist., 5:515-526. 1964. Distribution and taxonomy of mammals of Nebraska. Ibid., 16:1- O50, Jones, J. K., Jn., and F. R. HENDERSON 1963. Noteworthy records of foxes from South Dakota. Jour. Mamm., 44:283. Jones, J. K., Jn., R. P. Lamps, C. A. SPENRATH, and T. H. Kunz 1973. Notes on the distribution and natural history of bats in southeastern Montana. Occas. Papers Mus., Texas Tech Univ., 15:1-12. KELLoGG, C. E. (ed. ) 1938. Soils of the United States. Pp. 1019-1161, in Soils and Men, Year- book of Agriculture, U.S.D.A., xvi + 1232 pp. Kocu, E. 1941. Big game in Montana from early historical records. Jour. Wildlife Mgt., 5:357-370. Krutzscu, P. H. 1954. North American jumping mice (genus Zapus). Univ. Kansas Publ., Mus. Nat. Hist., 7:349-472. KUCHLER, A. W. 1964. Potential natural vegetation of the conterminous United States. Spec. Publ., Amer. Geogr. Soc., 36:v + 1-39 + map 4+ 116 plates. KurTEN, B., and R. Rauscu 1959. Biometric comparisons between North American and European mammals. Acta Arctica, Fasc. XI. E. Munksgaard, Copenhagen, 44 pp. LAMPE, R. P. 1971. The mammals of southeastern Montana. Unpubl. M.A. thesis, Univ. Kansas, Lawrence, ii + 68 pp. LILLEGRAVEN, J. A. 1970. Stratigraphy, structure, and vertebrate fossils of the Oligocene Brule Formation, Slim Buttes, northwestern South Dakota. Bull. Amer. Geol. Soc., 81:831-850. LinHART, S. B., and W. B. Rosinson 1972. Some relative carnivore densities in areas under sustained coyote control. Jour. Mamm., 53:880-884. Lone G: A. 1965. The mammals of Wyoming. Univ. Kansas Publ., Mus. Nat. Hist., 14:493-758. THE MAMMALS OF CARTER COUNTY, MONTANA 39 McDAanlrceL, L. L. 1967. Merriam’s shrew in Nebraska. Jour. Mamm., 48:489-493. MILLER, G. S., and R. KELLocG 1955. List of North American Recent mammals. Bull. U.S. Natl. Mus., 205:xii + 954. NELSON, E. W. 1909. The rabbits of North America. N. Amer. Fauna, 29:1-314. PeFaur, J. E., and R. S. HOFFMANN 1971. Merriam’s shrew and hispid pocket mouse in Montana. Amer. Mid. Nat., 86:247-248. Perry, E. S. 1962. Montana in the geologic past. Bull. Montana Bur. Mines and Geol., 26:vi + 1-78. PFEIFER, W. K., and E. A. Hisparp 1970. A recent record of the swift fox (Vulpes velox) in North Dakota. Jour. Mamm., 51:835. PREBLE, E. A. MS. List of the mammals of eastern Montana. On file, Dept. Zoology, Univ. Montana, Missoula, 12 pp. RAYNOLDs, W. F. 1868. Journal of Captain W. F. Raynolds. Pp. 18-127, in Report on the exploration of the Yellowstone River . .., Exec. Doc., Senate, 2nd Sess., 40th Congress, 77:1-174. [erroneously printed as Ist Sess.] Rosinson, W. B. 1961. Population changes of carnivores in some coyote-control areas. Jour. Mamm., 42:510-515. SopPER, J. D. 1961. Field data on the mammals of southern Saskatchewan. Canadian Field-Nat., 75:23-40. SPENCER, A. W., and D. Petrus 1966. Habitat preferences of five sympatric species of long-tailed shrews. Ecology, 47:677-683. SwENK, M. H. 1941. A study of subspecific variation in the Richardson pocket gopher (Thomomys talpoides) in Nebraska, with description of two new subspecies. Missouri Valley Fauna, 4:1-8. THOMPSON, W. K. 1949. Predation on antelope. Jour. Wildlife Megt., 13:313-314. Tryon, C. A, Jr. 1947. Entrance and migration of the Norway rat into Montana. Jour. Mamm., 28:188-189. TURNER, R. W. 1974. Mammals of the Black Hills of South Dakota and Wyoming. Misc. Publ. Mus. Nat. Hist., Univ. Kansas, 60:1-178. VAN ZYLL DE JONG, C. G. 1972. A systematic review of the Nearctic and Neotropical river otters (genus Lutra, Mustelidae, Carnivora). Life Sci. Contrib., Royal Ontario Mus., 80:1-104. VISHER, S. S. 1914. Report on the biology of Harding County, northwestern South Da- kota. Bull. South Dakota Geol. Surv., 6:1-103. UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly pub- lished in the above series are now published as Occasional Papers, Museum of Natural History. The Miscellaneous Publica- tions, Museum of Natural History, began with number 1 in 1946. Longer research papers are published in that series. Monographs of the Museum of Natural History were initiated in 1970. All manuscripts are subject to critical review by intra- and extra- mural specialists; final acceptance is at the discretion of the publications committee. Institutional libraries interested in exchanging publications may obtain the Occasional Papers and Miscellaneous Publica- tions by addressing the Exchange Librarian, University of Kan- sas Library, Lawrence, Kansas 66045. Individuals may pur- chase separate numbers of all series. Prices may be obtained upon request addressed to Publications Secretary, Museum of Natural History, University of Kansas, Lawrence, Kansas 66045. Editor: RicHarp F.. 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