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Collins Printed by University of Kansas Printing Service Lawrence, Kansas OCCASIONAL PAPERS MCZ LIBRARY SEP 2 9 1992 of the MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas number 151, pages 1-23 23 september 1992 The Status of the Hylid Frog Genus Ololygon and the Recognition of Scinax Wagler, 1830 William E. Duellman and John J. Wiens1 Division of Herpetology, Museum of Natural History, and Department of Systematica and Ecology, The University of Kansas, Lawrence, Kansas 66045-2454, USA ABSTRACT The hylid frog genus Scinax Wagler, 1 830 (type species: Hyla aurata Wied-Neuwied, 1821) has priority over Ololygon Fitzinger, 1843 (type species: Hyla strigilata Spix. 1824). A neotype is designated for Hyla aurata, which is compared with other species of Scinax. The genus is defined using external mor- phological, osteological, larval, and reproductive characters. Scinax is considered to be monophyletic and the sister group of Scarthyla. RESUMEN El genero hflido Scinax Wagler, 1830 (especie tipo Hyla aurata Wied-Neuwied, 1821) tiene prioridad sobre el genero Ololygon Fitzinger, 1843 (especie tipo Hyla strigilata Spix, 1824). Se designa un neotipo para Hyla aurata, el cual se compara con otras especies de Scinax. Se define el genero usando caracteres morfologicos, osteologicos, larvales, y reproductivos. Se considera a Scinax monofiletico y como el grupo hermano de Scarthyla. Key words: Hylidae, Generic nomenclature, Ololygon, Scinax. Prior to 1977, numerous species of neotropical Hyla were recognized as the Hyla rubra group by Leon (1969), Cochran and Goin (1970), and Duellman (1970a). Within this group, phenetic subgroups were recognized 'Present address: Department of Zoology, The University of Texas, Austin, Texas 78712-1064. USA. © Museum of Natural History. The University of Kansas. Lawrence. lSSN:0091-7958 2 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 151 by Duellman (1972) and Lutz (1973). Fouquette and Delahoussaye (1977) resurrected the generic name Ololygon Fitzinger, 1843, for frogs of the Hyla rubra group (sensu lato). They included 48 species in five species groups of Ololygon and referred 10 other species to the genus. Unfortu- nately, Fouquette and Delahoussaye (1977) overlooked the fact that among the species they included in Ololygon was the type species of Scinax Wagler, 1830, which, as noted by Pombal and Gordo (1991), has priority over Ololygon Fitzinger, 1843. Herein we provide (1) a discussion of the nomenclature relevant to the genus; (2) a definition of the genus Scinax, with a discussion of possible inter- and intrageneric relationships; and (3) a list of trivial names associ- ated with the genus. -v MATERIALS AND METHODS We examined cleared-and-stained skeletal preparations of 34 species, preserved adults of 42 species, and larvae of 20 species of Scinax, and comparative material representing most genera of hylids and centrolenids in the collection of The University of Kansas Museum of Natural History (KU) (Appendix I). Measurements were taken to the nearest 0.1 mm using needle-tipped calipers in the manner described by Duellman (1970a); snout-vent length is abbreviated as SVL. Formulae for toe webbing are based on Savage and Heyer (1967) as modified by Myers and Duellman (1982). Terminology of larval features follows Altig and Johnson (1989), and terminology for osteological features follows Duellman and Trueb ( 1 986) and Trueb ( 1 973 ). Acronyms for museum collections follow Leviton etal. (1985). NOMENCLATURE Scinax Wagler, 1830 Scinax Wagler, 1830:201. — Type species: Hyla aurata Wied-Neuwied, 1821, by subsequent designation (Stejneger, 1907:76). Ololygon Fitzinger. 1843:31. — Type species: Hyla strigilata Spix, 1824. by monotypy. Garbeana Miranda-Ribeiro, 1926:96. — Type species: Garbeana garbei Miranda- Ribeiro, 1926, by monotypy. Generic names. — In his classification of amphibians, Wagler (1830) proposed many new generic names with a list of included species. On page 201 he erected the genus Scinax and listed three species — S. aurata (Wied- Neuwied, 1821), 5. variolosa (Spix, 1824 [= Hyla punctata, fide Peters, 1872:214]), and 5. bipunctata (Spix, 1824). Listing several species under a new generic name without designating a type species was a common practice in the 19th Century. Leonhard Stejneger made valiant efforts to STATUS OF OLOLYGON AND SCINAX 3 tidy up earlier nomenclature, and in his treatise on the herpetology of Japan. Stejneger (1907:76) provided a generic synonymy of Hyla, in which he listed "1830. Scinax Wagler, Syst. Amph., p. 201 (type. H. awata)."' In the same synonymy Stejneger also listed type species for two other nomi- nal genera with lists of proposed included species. Schneider ( 1799) listed 11 species under Calamita; of these. Stejneger (1907) selected Rana arborea Linnaeus, 1758. as the type species. Wagler (1830) listed 11 species under Auletris; of these. Stejneger (1907) selected Hyla boans Linnaeus, 1758, as the type species. Similarly, of the seven species listed under Hylaplesia by Boie ( 1826), Stejneger ( 1937) chose Hyla punctata Daudin, 1802 (= Calamita punctatus Schneider. 1799) as the type species. All of these are valid subsequent designations of type species according to Article 69 of the International Code of Zoological Nomenclature (Anony- mous, 1985). In his list of genera of Hylidae, Fitzinger (1843:31) placed only one species in Ololygon — Hyla strigilata Spix, 1824. Miranda-Ribeiro (1926:96) described a new genus and species, Garbeana garbei. Hyla strigilata Spix, 1924. and Garbeana garbei are the type species of Ololygon Fitzinger. 1843. and Garbeana Miranda-Ribeiro. 1926. respectively, by monotypy. Thus, of the nominal species included in the Hyla rubra group (sensu lato, auctorum) and subsequently included in the genus Ololygon by Fouquette and Delahoussaye (1977) and Frost (1985), three of them are type species of genera. The earliest generic name is Scinax Wagler. 1830, with the type species Hyla awata Wied-Neuwied, 1821. Therefore, the correct generic name for this group of hylid frogs is Scinax Wagler, 1830; Ololygon Fitzinger, 1843, and Garbeana Miranda-Ribeiro, 1926, are sub- jective junior synonyms. The generic name. Scinax, is from the Greek skinos meaning quick or nimble, an appropriate name for these agile frogs. The gender is feminine. Type species. — In order to justify the use of Scinax and the placement of the generic synonyms, it is necessary to identify biologically the type species of the genera. Thus, we are concerned with the identities of Hyla aurata, Hyla strigilata, and Garbeana garbei. Garbeana garbei is readily identified. The holotype (MZUSP 277) was examined by Duellman, and many specimens are associated with this specific name (as Hyla garbei), a taxon widely distributed in the upper Amazon Basin (Duellman, 1970b; 1972). It is unfortunate that Hyla strigilata Spix, 1824, is the type species of Ololygon, because the holotype, ZSM 2369/0 from "Bahia," was destroyed in World War II ( Hoogmoed and Gruber, 1983). Peters ( 1 872) examined the type specimen ("Das einzige Original-exemplar") and noted discrepancies between it and Spix's illustration (1824:pl. X, fig. 3). Cochran (1955) examined the holotype of H. strigilata and associated many specimens 4 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 1 5 1 from eastern Brazil with the species, which she considered to be polytypic; she included Hyla brieni de Witte, 1930, and Hyla flavoguttata Lutz and Lutz, 1939, as subspecies. Cochran's concept of Hyla s. strigilata appar- ently included two or more species; she provided photographs ( 1955:pl. 14, figs. D, E) of a preserved specimen (ZMB 7504) from Caravellas, Bahia, Brazil. Lutz (1973) also was unsure of the identity of Hyla strigilata and tentatively referred only four specimens, all juveniles from Afonso Claudio, Espfrito Santo, to the species. Thus, the identity of Hyla strigilata remains questionable; presumably the name applies to a species in Bahia, Brazil, possibly one that is recognized by another name. However, based on the descriptions of the type by Spix (1824) and Peters (1872) and the illustra- tion in Spix (1824), Hyla strigilata is distinct from Hyla aurata. Wied-Neuwied (1821:249) described Hyla aurata from an unknown number of specimens taken in swamps at Santa Agnes, Bahia, Brazil. Wied-Neuwied (1824:531-533) provided a more detailed description. En- glish translations of both descriptions were given by Lutz (1973:167-168). Bokermann (1957) traced Wied-Neuwied's travels in Brazil and listed Wied-Neuwied's locality as "Santa Inez, fazenda (S. Agnes). Atualmente cidade de Santa Inez, BA, proxima ao rio Jequlrica." Bokermann (1966) gave the type locality of Hyla aurata as "Santa Inez (= 'S. Agnes'), Bahia." In 1965, the species was rediscovered at Maracas, Bahia, Brazil (Bokermann, 1969). Lutz ( 1973) did not mention Bokermann's locality, but noted that she had found Hyla aurata in 1941 at Salvador and at Cachoeira, near Sao Felix, on the Rio Paraguassii. Thus, the species has been reported from four localities from Salvador on the Atlantic coast to Maracas, about 160 km inland; all localities are between 12°30' and 13°26' S latitude. As was characteristic of the early 19th century, no type specimens were designated, and no types are known to exist. Some of Wied-Neuwied's types of frogs (e.g., syntype of Hyla crepitans) are in the American Mu- seum of Natural History, but there is no evidence for the existence of type specimens of Hyla aurata in that collection (Bokermann, 1969; D. R. Frost, pers. comm.). In the interests of nomenclatural stability, we designate a neotype of Hyla aurata Wied-Neuwied, 1821. Rules for the designation of neotypes are explicit in Article 75. d of the International Code of Zoological Nomenclature (Anonymous, 1985); these include ( 1 ) evidence that the neotype is consistent with what is known of the former name-bearing type; (2) evidence that the neotype came as nearly as practical from the original type locality; and (3) that the neotype is the property of a recognized scientific institution. Extant specimens of Hyla aurata are from three localities; of these, Maracas is the closest to the type locality. Maracas is about 70 km (airline) WSW of Santa Inez (= Santa Agnes, the stated type locality); both localities are inland. Maracas is at the headwaters of the Rio Jequirica, and Santa Inez is in the valley of the same STATUS OF OLOLYGON AND SCINAX 5 river. The other localities, Cachoeira and Salvador, are on the coastal plain, about 125 km NE and 150 km ENE (airline), respectively, from Santa Inez. The species was first found at Maracas in September 1965 by F. M. Oliveira, and additional specimens were collected there by F. M. Oliveira and W. C. A. Bokermann in November 1965. These specimens (WCAB 30726-36, 31857-906) were deposited in Bokermann's private collection in Sao Paulo, Brazil. One of these specimens, WCAB 31905, was donated to the Museum of Natural History, The University of Kansas, in 1969. This specimen satisfies all the requirements for designation as a neotype. Thus, KU 125383 hereby is designated as the neotype of Hyla aurata Wied- Neuwied, 1821. According to Article 75. f of the code, the place of origin of the neotype becomes the type locality. Therefore, the type locality of Hyla aurata becomes Maracas, Estado do Bahia, Brasil. Description of neotype. — Adult male; body moderately robust; head as wide as body, longer than wide; head length 36.4% of SVL; head width 32.5% of SVL; snout long, projecting well beyond margin of jaw, subacuminate in dorsal view and in profile: eye-nostril distance slightly greater than horizontal diameter of eye; canthus rostralis rounded; loreal region slightly concave; lips rounded. Top of head flat; interorbital distance about twice width of upper eyelid; nostrils barely protuberant dorsolateral^ at point above anterior margin of lower jaw: internarial area barely de- pressed. Supratympanic fold weak, barely obscuring upper edge of tym- panic annulus; tympanum round, posteroventral to eye; separated from eye by distance equal to 77% of diameter of tympanum. Forearm moderately robust; row of low ulnar tubercles; fingers short, unwebbed, bearing large, elliptical, terminal discs; width of disc on Finger III 85% diameter of tympanum; relative lengths of fingers I < II < IV < III; basal segment of Finger II greatly swollen. Palmar tubercle elevated, bifid; thenar tubercle low, elliptical; large, unpigmented nuptial excrescence on inner edge of thumb: subarticular tubercles conical; supernumerary tu- bercles subconical, present only on proximal segments of digits. Hind limb moderately robust; tibia length 57.0% of SVL; foot length 41.6% of SVL; numerous subconical tubercles on heel; rows of distinct tubercles on both inner and outer edges of tarsus; inner tarsal fold absent; inner metatarsal tubercle elliptical, elevated; outer metatarsal tubercle conical. Toes moder- ately long, bearing elliptical discs nearly as large as those on fingers; relative lengths of toes I < II < III = V < IV; toes about three-quarters webbed; webbing formula I 2— 2 II 1+— 2 III 1+— 2 IV 2— 1+ V; subarticular tubercles conical, smaller than those on fingers; supernumer- ary tubercles small, subconical, only on proximal segments of digits. Skin on dorsal surfaces of head and limbs smooth, on dorsal surfaces of body shagreened, on flanks weakly granular; numerous small tubercles on upper eyelids and dorsal surfaces of body and shanks; skin on belly and 6 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 151 ventral surfaces of thighs coarsely granular, on other ventral surfaces smooth. Cloacal opening directed posteroventrally at upper level of thighs; cloacal sheath short, unmodified. Vomerine odontophores slightly elevated, elliptical, slightly inclined posteromedially between ovoid choanae, nar- rowly separated medially, each bearing five teeth. Tongue broadly cordi- form, shallowly notched posteriorly, barely free behind. Vocal slit ellipti- cal, extending from midlateral base of tongue toward angle of jaws; vocal sac single, median, subgular, large, extending onto chest. Coloration in preservative: Dorsum tan with cream dorsolateral stripes extending from occiput immediately behind orbits to groin; stripe on left side interrupted and offset in scapular region; stripes bordered by series of small brown spots (Fig. 1 ). Canthal stripe brown, diffuse; interorbital bar brown, distinct, narrow, bordered anteriorly by much wider cream bar; upper lips and tympanum creamy tan. Three indistinct, transverse brown bars on each forearm; anterior and dorsal surfaces of thighs indistinctly mottled creamy tan and brown; posterior surfaces of thighs brown; dorsal shanks tan with indistinct brown spots and transverse bars; hands, feet, and venter creamy tan. Structurally, this specimen agrees with the original description by Wied- Neuwied (1921) and the description of the series from Maracas by Bokermann (1969). The color pattern of the neotype is less bold than that of the adult female illustrated by Bokermann (1969). Measurements (in mm): SVL 22.8, tibia length 13.0, foot length 9.5, head length 8.3, head width 7.4, internarial distance 1.6, interorbital dis- tance 3.2, upper eyelid 1.8, eye-nostril 2.7, eye diameter 2.6, eye-tympa- num 1.0, tympanum diameter 1.3. According to Bokermann (1969), the SVL of nine calling males is 23-25 mm (x = 23.7) and of seven gravid females, 23-24 (x = 23.7). Comparisons. — In comparison with other species of Scinax in north- eastern Brazil (Espirito Santo northward), S. aurata differs from S. nebulosa by lacking conical ulnar, tarsal, heel, and labial tubercles and bold black and off-white mottling on the posterior surfaces of the thighs. From S. eurydice, S. rubra, and S. x-signata, it differs by its smaller size (SVL of adult males < 25 mm vs. > 40 mm) and its lack of pale spots enclosed by dark brown or black on the posterior surfaces of the thighs. Scinax strigilata also is larger (holotype, 42 mm; Peters, 1872:216) and lacks dorsolateral stripes (Spix, 1824:pl. 10, fig. 3). Scinax argyreornata, S. catharinae, S. flavoguttata, and S. heyeri differ from S. aurata in having bold transverse bars across the hind limbs, shorter and higher rostra, and in lacking pale dorsolateral stripes. Scinax hoesemani has smooth skin and is uniform tan dorsally (except for a dark canthal-supratympanic stripe and a few small, brown flecks on the dorsum). Scinax crospedospila is larger (adult males > 28 STATUS OF OLOLYGON AND SCINAX Fig. 1. Neotypeof/Zv/aawrataWied-Neuwied, 1821; KU 125383, adult male, 22.8 mm SVL. mm SVL), lacks pale dorsolateral stripes and an interorbital bar, and has a dorsum with elongate, dark brown blotches. Lutz (1973) noted the similarity in coloration of S. cuspidata with Wied- Neuwied's (1821; 1824) descriptions of Hyla aurata. Scinax cuspidata is about the same size as S. aurata and has a similar dorsal coloration (Fig. 2), except that brown and cream interorbital bars are absent. Scinax aurata also differs from S. cuspidata in having distinct, cream tubercles on the dorsal surface of the shank. The coloration of S. aurata also resembles that of two other small species, S. exigua and S.fuscomarginata. Scinax exigua is smaller (adult males to 23 mm SVL) and has a smooth dorsum. Scinax fuscomarginata lacks a pale interorbital bar and tubercles on the dorsal surface of the shank. UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 151 v Fig. 2. Variation in dorsal color pattern in Scinax cuspidata. Left to right: KU 92008, 92006, 92007, 92005, all from Barro Sao Joao, Rio de Janeiro, Brazil. Remarks. — Bokermann (1969) provided some ecological and behav- ioral data for the species. Individuals were found in terrestrial and epi- phytic bromeliads. With the first rains of the season in November, males called from hidden perches on bushes and the stems of grasses within 1 m of temporary ponds. The call consists of a single note repeated at a rate of about 12 notes/s. Each note has a duration of about 0.2 s and consists of 9 pulses; the dominant frequency is at about 2000 Hz. A testis removed from the neotype contained no sperm (M. J. Fouquette, Jr., pers. comm.). GENUS SCINAX WAGLER, 1830 Diagnosis. — We tentatively consider the following three character states to be synapomorphies of the genus: ( 1 ) loreal region depressed; (2) discs on hand truncate, wider than long; (3) webbing between first and second toes reduced (absent or reduced to fringe along inner margin of second toe) relative to webbing on other toes (Fig. 3). Definition. — ( 1 ) Frontoparietal fontanelle variably exposed; (2) prootic and exoccipital fused in adults; (3) nasal slender, length 20-^4-0% of skull length, not in contact with maxilla or frontoparietal, well separated from pars facialis of maxilla in most species, articulating medially in a few species (e.g., S. acuminata , S. nasica); shape of maxillary process continu- ously variable from long and pointed (e.g., S. squalirostris) to short and rounded (e.g., S. cuspidata); (4) sphenethmoid well ossified; longer than wide in dorsal aspect; (5) alary process of premaxilla inclined anterodorsally; (6) quadratojugal long, slender, articulating with maxilla; (7) palatine process of maxilla usually present; (8) maxillary, premaxillary, and vomerine teeth small, blunt, pedicellate, always present; (9) vomerine odontophore transverse, not angular; (10) neopalatine (terminology of Trueb, 1992) usually present (absent in S. parkeri; variably present in S. staufferi), well developed, articulating with maxilla; (11) cultriform pro- cess of parasphenoid wide posteriorly, narrowing abruptly anterior to optic foramen, usually terminating posterior to level of neopalatine; parasphenoid STATUS OF OLOLYGON AND SCINAX Fig. 3. Feet of hylid frogs. A. Scinax rubra, KU 109471, showing characteris- tic reduction of webbing between Toes I and II. B. Hyla microcephala, KU 101606. showing primitive pattern of webbing between Toes I and II. Reproduced from Duellman. 1970a. alae short, narrow, oriented posterolaterally; (12) pterygoid well devel- oped, median ramus almost in contact with prootic; (13) ventral ramus of squamosal bowed anterodorsally in lateral aspect; otic ramus expanded into otic plate on crista parotica; zygomatic ramus acuminate, usually extending 50% of distance between ventral ramus and postorbital process of maxilla (estimated by projecting a straight line from the zygomatic ramus to the maxilla), but ranging from 20% (e.g., S. nebulosa) to 75% (e.g., 5. x-signata); (14) skull lacking dermal exostosis, co-ossification, casquing, and neomorphic dermal elements; (15) septomaxilla Type II (Trueb, 1970); (16) coronoid process of angulosplenial present, weakly developed; ( 17) posterolateral and anterolateral processes of hyoid always present; short anterior processes present on hyale; posteromedial processes long and ossified; commonly some calcification of hyoid plate posteriorly (Fig. 4); (18) plectral apparatus always present; pars externa plectri ori- 10 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 151 Fig. 4. Hyoids of hylid frogs. A. Smilisca phaeota, KU 169627, showing absence of anterior process of hyale. B. Scinax rubra, KU 123048, showing presence of anterior process of hyale. Scale = 5 mm. ented ventrally. elongate and not dilated distally in most species, entering tympanic ring dorsally; tympanic annulus large, occupying most of area between quadratojugal and zygomatic ramus of squamosal; (19) eight procoelous nonimbricate presacral vertebrae; atlas Type I (Lynch, 1973) (widely spaced cotyles); transverse processes of Presacral Vertebra IV not elongated posteriorly; processes of Presacral Vertebra VIII usually inclined anteriorly; sacral diapophyses weakly dilated; sacrococcygeal articulation bicondylar (Fig. 5); (20) omosternum and sternum always present, calcified in some specimens; suprascapula (cleithrum) bifurcate distally; (21) prepollex not enlarged or ossified; (22) intercalary elements between ulti- mate and penultimate phalanges cartilaginous, not mineralized in most species (mineralized in S. argyreornata, S. berthae, and 5. squalirostris); (23) adult size small to moderate, ranging from 18 mm in adult male S. exigua (Duellman. 1986) to 53 mm in adult female S. boulengeri (Duellman, 1970a); (24) coloration generally dull, with bold coloration on posterior surface of thighs in some species; (25) palpebral membrane clear; (26) skin smooth, shagreened, or tuberculate (with large tubercles on snout, mouth, and limbs in some species); otherwise lacking dermal folds, ap- pendages, membranes, or glands; (27) head longer than wide; (28) snout protruding, usually long, with depressed loreal region; (29) vocal sac single, median, subgular in most species, but paired, lateral, subgular in 5. jureia, S. ranki, and S. rizibilis (Andrade and Cardoso, 1987; Pombal and Gordo, 1991); (30) toes about one-half to three-quarters webbed; webbing reduced between first and second toes (relative to extent of webbing on other toes); fingers unwebbed or only webbed basally; (31 ) outer metatar- sal tubercle typically present in all species; (32) digital disks of hand dilated, truncate, wider than long; (33) nuptial excrescence inconspicuous STATUS OF OLOLYGON AND SCINAX 11 Fig. 5. Vertebral columns of hylid frogs. A. Phrynohyas venulosa, KU 92261. showing widely dilated sacral diapophyses. B. Scinax x-signata, KU 150871, showing narrow sacral diapophyses. Scale = 5 mm. (generalized type of Duellman, 1 970a) or absent: (34) depressor mandibulae with single slip; adductor m. posterior subexternus present (Starrett, 1968: data only for S. elaeochroa and S. rubra): (35) m. intermandibularis absent; submandibular bilobular extensions of the interhyoideus present in some species (Tyler, 1971); (36) sperm with two tail filaments (Fouquette and Delahoussaye, 1977); (37) diploid chromosome number 24 (Fouquette and Delahoussaye, 1977); (38) tadpoles with body relatively short posterior to orbit; eyes large, lateral, widely spaced in most species; tail usually deep and tapering to tip: dorsal fin extending onto body in most species; oral disc anteroventral; tooth row formula 2/3; marginal oral papillae with dorsal gap (except in members of the S. catharinae group); ventral papillae interrupted in S. crospedospila (Heyer et al., 1990) and interrupted by "labial arm" (muscular support for ventralmost tooth row) in most mem- bers of S. rostrata group; (39) mating call consisting of single pulsed note or series of short notes (Cardoso and Sazima, 1980; Duellman, 1970a, 1972; Peixoto and Weygoldt, 1987); (40) eggs deposited in standing water, streams (S. catharinae group), or in terrestrial bromeliads (5. perpusilla group; Peixoto, 1988a). Content. — Subsequent to Fouquette and Delahoussaye's (1977) recog- nition of Ololygon, new taxa have been named as Ololygon (O. trilineata 12 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 151 Hoogmoed and Gorzula, 1979; O. danae and O. exigua Duellman, 1986; O. atrata and O. melloi Peixoto, 1988a; O. chiquitana de la Riva, 1990; O. pedwmedinae Henle, 1991); some formerly nominal taxa have been resur- rected from synonymies and recognized as species of Ololygon (e.g., O. v- signata Peixoto, 1987; O. alcatraz Peixoto, 1988b). However, Almeida and Cardoso (1985) considered that variation in sperm morphology within Ololygon was too great to allow its use in defining the genus. Thus, other new species were named as Hyla (e.g., H. maracaya Cardoso and Sazima, 1980; H. canastrensis Cardoso and Haddad, 1982; H. agilis Cruz and Peixoto, 1983; H. ranki Andrade and Cardoso, 1987; H. jureia and H. littoralis Pombal and Gordo, 1991). Frost (1985) listed 54 species in Ololygon (= Scinax); two species listed by Frost (1985) in Hyla {Hyla albicans Bokermann, 1967, and Hyla ariadne Bokermann, 1967) obvi- ously belong in the genus. Hyla vigilans Solano, 1971, may belong to the genus (Frost, 1985:157). Subsequently, 12 new species have been de- scribed, three have been resurrected from synonymies, and one has been synonymized. Thus, the total number of recognized species in the genus now is 70. More than 100 trivial names apply to species of Scinax (Appen- dix II). Distribution. — The genus ranges from Mexico (Guerrero and Tamaulipas) through Central America to northern Uruguay and northern Argentina east of the Andes, to extreme northern Peru west of the Andes, and the Caribbean islands of Tobago, Trinidad, and St. Lucia. Most species occur in lowland rainforest, dry forest, or savanna at elevations of less than 500 m, but several species range to elevations of 1000 m in eastern Brazil, and an undescribed species ranges above 2000 m in the Andes of Peru (Duellman and Wiens, in prep.). The genus is most diverse in southeastern Brazil (Espfrito Santo to Minas Gerais and Sao Paulo). DISCUSSION Definition of Scinax. — Some discrepancies exist between our charac- terization of Scinax and other reports. According to our observations, the nasals are commonly less than 40% of the total skull length (vs. more than 40% in Leon, 1969, and Duellman, 1970a), and do not always have long, pointed maxillary processes (contra Leon, 1969, and Duellman, 1970a). Furthermore, the maxillary process of the nasal does not contact the max- illa in any of the species of Scinax we have examined (contra Duellman and de Sa, 1988). The presence of the palatine process of the maxilla varies both inter- and intraspecifically in Scinax (contra Leon, 1969, and Duellman, 1970a). According to Duellman and de Sa (1988), members of Scinax lack neopalatines (terminology of Trueb, 1992), but according to our observations, the neopalatines are present in all species except S. STATUS OF OLOLYGON AND SCINAX 1 3 parkeri and in some individuals of 5". staufferi. Duellman and de Sa ( 1988) indicated that all members of Scinax have cartilaginous intercalary ele- ments, however, we have observed mineralized intercalary elements in S. argyreornata, S. berthae, and S. squalirostris. As noted by Leon (1969), Duellman (1970a), and Duellman and de Sa (1988), the zygomatic ramus of the squamosal usually extends about 50% of the distance between the squamosal (dorsal head of ventral ramus) and maxilla, but this process can be considerably shorter (20% in S. nebulosa) or longer (75% in S. x- signata). Leon (1969) and Duellman ( 1970a) considered the sphenethmoid of Scinax to be wider than long, but our observations show that the sphenethmoid is actually longer than wide. The marginal oral papillae of the tadpole are complete in some species (in the S. catharinae group), rather than always having a dorsal gap as reported by Duellman (1970a). Also, a ventral gap in the marginal papillae may be present (e.g., S. crospedospila). We found nuptial excrescences to be present (although inconspicuous) on the prepollices of most male Scinax, whereas Duellman (1970a) stated that males lack nuptial excrescences. Intergeneric relationships and monophyly. — The genus Scinax (as Ololygon or Hyla) has been assigned to the subfamily Hylinae, a likely paraphyletic assemblage defined by the absence of the synapomorphies of the Hemiphractinae, Pelodryadinae, and Phyllomedusinae. Although rela- tionships within the Hylinae are unclear, we consider it heuristic to specu- late as to the relationships of Scinax. Our hypothesis is speculative because some of the character states that we postulate as derived occur in other hylid taxa and might prove to be plesiomorphic. We propose that Scinax belongs to a monophyletic group that includes the genera Scarthyla and Sphaenorhynchus (Fig. 6; numbered character states shown on tree corre- spond to numbers assigned to characters in following discussion). These three genera share numerous features, of which the following probably are derived (based on comparison with centrolenids and other hylids): (1) narrow (rather than strongly dilated) sacral diapophyses; (2) anteriorly inclined alary processes of the premaxillae; (3) tadpole with large, laterally placed eyes (polarity questionable; variable in Scinax). According to Fouquette and Delahoussaye (1977), the only genera in the Hylinae with double-filamented sperm are Scinax and Sphaenorhynchus (but single-filamented in Scarthyla: M. J. Fouquette. pers. comm.). Based on the survey of these authors, the presence of double-filamented sperm seems to be a plesiomorphic character state in hylids (present in centrolenids, bufonids, some leptodactylids, and pelodryadine hylids). The presence of single-filamented sperm might be a synapomorphy uniting all hylids exclusive of pelodryadines and the clade containing Scinax, Scarthyla (in which single-filamented sperm would be a reversal), and Sphaenorhynchus, or double-filamented sperm might indicate a relation- 14 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 151 itf .0 # tf ^ ff IS $*■ ■s <& 4-14 *» »♦ •9 $ * 20-22 15-16 Fig. 6. Hypothesized phylogenetic relationships of three genera of hylid frogs. Numbers by bars = character states numbered in text. ship between Scinax and Sphaenorhynchus (exclusive of Scarthyla) if these genera are in fact more deeply nested within the Hylinae. Among these three genera, Scinax and Scarthyla share (15) reduced webbing on the hand (relative to webbing on the feet) and (16) the presence of an anterior process on the hyale (see Fig. 3, and Duellman and de Sa, 1988:fig. 2). All three genera are demonstrably monophyletic. Members of Sphaenorhynchus examined share at least 1 1 synapomorphies (many of which are unique within the Hylidae): (4) poste- rior ramus of pterygoid absent; (5) zygomatic ramus of squamosal absent or reduced to small knob; (6) pars facialis of maxilla and alary process of premaxilla reduced; (7) postorbital process of maxilla reduced, not in contact with quadratojugal; (8) neopalatine reduced to sliver or absent (also in two species of Scinax); (9) pars externi plectri entering tympanic ring posteriorly (rather than dorsally); (10) pars externa plectri round; (11) hyale curved medially; (12) coracoids and clavicles elongate; (13) trans- STATUS OF OLOLYGON AND SCINAX 1 5 verse process of Presacral Vertebra IV elongate, oriented posteriorly; ( 14) prepollex ossified, bladelike. The monotypic Scarthyla is diagnosed by three apomorphies involving the larvae: (17) tadpole elongate; (18) tadpole with labial arm (also in Scinax rostrata group); (19) tadpoles capable of launching themselves out of water. The presence of a single-filamented sperm and mineralized intercalary elements (also mineralized in Scinax argyreornata, S. berthae, S. squalirostris, Sphaenorhynchus carneus and S. planicola) also might be apomorphies diagnosing Scarthyla. The monophyly of Scinax is corrobo- rated by at least three synapomorphies: (20) depressed loreal region; (21) webbing between first and second toes absent or reduced to fringe along margin of second toe; (22) discs on hand dilated, truncate, wider than long. McDiarmid and Altig (1990) suggested that the presence of a larval labial arm could be a synapomorphy allying the genus Scarthyla with some members of the Scinax rostrata group. If this were true, Scinax would be paraphyletic with respect to Scarthyla. Although the labial arm is unique to these species, we suggest that the synapomorphies uniting the species of Scinax (listed above), species of the Scinax rostrata group (see below), and some (but not all) species of the Scinax rostrata group with other species of Scinax (slender, longitudinally oriented nasals; elongate, rod-shaped pars externa plectri, black bars on posterior of thighs; tuberculate skin; tadpoles with bodies that are short and deep posterior to the orbit and deep caudal fins that extend onto the body) indicate that the labial arm evolved indepen- dently in these two lineages. Intrageneric relationships and taxonomy. — Seven species groups cur- rently are recognized within Scinax. Most of the species groups within Scinax were recognized by Fouquette and Delahoussaye (1977) on the basis of sperm head "types." In this section we review the evidence for the monophyly of the species groups and provide updated lists of their con- tents. The contents of the Scinax catharinae group were modified consider- ably by Peixoto and Weygoldt (1987). We follow their arrangement and also include Scinax littoralis, S. machadoi, and S. opalina. Thus, we consider the S. catharinae group to include the following species: S. albicans, S. ariadne, S. argyreornata, S. brieni. S. catharinae, S. flavoguttata, S. heyeri, S. humilis, S. littoralis, S. machadoi, S. obtriangulata, and S. opalina. The species for which reproductive and larval data are available share the derived features of oviposition in streams and tadpoles that lack a dorsal gap in the marginal labial papillae (S. ariadne, S. flavoguttata, S. heyeri, S. machadoi, S. opalina) or have the dorsal gap extremely reduced (S. obtriangulata, contra Heyer et al., 1990). The species of the Scinax catharinae and 5". perpusilla groups share short, truncate snouts (in dorsal view), a distinctive dorsal color pattern. 16 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 151 and known tadpoles having bluntly rounded snouts (features we presume to be derived). The S. perpusilla group was first recognized by Peixoto (1987) and includes the following species: S. alcatraz, S. atrata, S. littorea, S. melloi, S. perpusilla, S. v-signata. The species in this group share the synapomorphy of oviposition in terrestrial bromeliads. The Scina.x rizibilis group was recognized first by Andrade and Cardoso (1988) and contains three species (S. jureia, S. ranki, and S. rizibilis) sharing the synapomorphy of paired vocal sacs. The S. rostrata group was defined by Duellman (1972) and currently contains nine species: S. boulengeri, S. epacrorhina, S. garbei, S. kennedyi, S. nebulosa, S. pedromedinae, S. proboscidea, S. rostrata, and S. sugillata. All species in the group share an acuminate, depressed snout. Except for S. nebulosa, species for which tadpoles are known (S. boulengeri, S. garbei, S. pedromedinae, S. rostrata, S. sugillata) have fanglike teeth and a labial arm supporting the lowermost tooth row. All species except 5. kennedyi and 5. rostrata have conical ulnar, tarsal, and labial tubercles. Males of the S. rostrata group that we have observed (S. garbei, S. pedromedinae, S. sugillata), S. boulengeri (J. M. Ellingson, pers. comm.), and S. rostrata (J. D. Lynch, pers. comm.) often call in a head-down, vertical position; this unique behavior may be another synapomorphy of the group. We are unaware of any evidence to support the monophyly of the remaining three species groups. The Scina.x rubra group contains 10 spe- cies: S. blairi, S. chiquitana, S. cynocephala, S. elaeochroa, S.funerea, S. fuscovaria, S. hayii, S. quinquefasciata, S. rubra, and 5. similis. The S. staufferi group contains 13 species: S. agilis, S. baumgardeneri, S. berthae, S. danae, S. e.xigua, S.fuscomarginata, S. goinorum, S. nasica, S. parkeri, S. squalirostris, S. staufferi, S. trilineata (not listed in the group by Frost, 1985, but related to S. squalirostris, according to the original description), and S. wandae. The S. x-signata group contains six species: S. acuminata, S. boesemani, S. crospedospila, S. cruentomma, S. cuspidata, and S. x- signata. The remaining 1 1 species are unassigned to species groups; these are S. alleni, S. aurata, S. canastrensis, S. duartei, S. ehrhardti, S. eurydice, S. longilinea, S. maracaya, S. pachychrus, S. strigilata, and S. trachy thorax. Acknowledgments: This study was ancillary to a review of the Scina.x in Amazonian and Andean Ecuador and Peru, which is part of a project on patterns of speciation and biogeography of Andean anurans supported by a grant (BSR-8805920) from the National Science Foundation (William E. Duellman, principal investigator). We are grateful to Janna M. Ellingson, Darrel R. Frost, John D. Lynch, and Linda Trueb for helpful information; M. J. Fouquette, Jr. for information on sperm morphology; Amy Lathrop for the drawings; and John E. Simmons for the photographs. STATUS OF OLOLYGON AND SCINAX 1 7 LITERATURE CITED Altig, R., and G. F. Johnson. 1989. Guilds of anuran larvae: relationships among developmental modes, morphologies, and habitats. Herpetol. Monogr. 3:81-109. Almeida. C. G.. and A. J. Cardoso. 1985. Variabilidad em medidas dos espermatozoides de Hyla fuscovaria (Amphibia. Anura) e seu significado taxonomico. Rev. Brasil. Biol. 45:387-391. Anonymous. 1985. International Code of Zoological Nomenclature. Ed. 3. London: International Trust for Zoological Nomenclature. Andrade, G. V., and A. J. Cardoso. 1987. Reconhecimento do grupo rizibilis; descricao de uma especie de Hyla (Amphibia, Anura). Rev. Brasil. Zool. 3:433-440. Boie, H. 1826. Notice sur FErpetologie de File de Java. Ferussac Bull. Sci. Nat. 9:233-240. Bokermann, W. C. A. 1957. Atualizacao do itinerario da viagem do Principe de Wied ao Brasil (1815-1817). Arq. Zool. 10:209-251. Bokermann. W. C. A. 1966. Lista Anotada das Localidades Tipo de Anfibios Brasileiros. Servicio de Documentacao, Univ. Rural Sao Paulo. Bokermann, W. C. A. 1969. Notassobre 'Hylaaurata'Wied, 1824 (Anura, Hylidae). Rev. Brasileira Biol. 29:159-162. Cardoso, A. J., and C. F. B. Haddad. 1982. Nova especie de Hyla da Serra da Canastra (Amphibia. Anura, Hylidae). Rev. Brasil. Biol. 42:499-503. Cardoso. A. J., and I. Sazima. 1980. Nova especie de Hyla do sudeste Brasileiro (Amphibia. Anura. Hylidae). Rev. Brasil. Biol. 40:57-59. Cochran, D. M. 1955. Frogs of southeastern Brazil. Bull. U.S. Natl. Mus. 206:1-423 Cochran. D. M.. and C. J. Goin. 1970. Frogs of Colombia. Bull. U.S. Natl. Mus. 288:1-655. Cruz, C. A. G. de. and O. L. Peixoto. 1983. Uma nova especie de Hyla do Estado do Espirito Santo, Brasil (Amphibia. Anura, Hylidae). Rev. Brasil. Biol. 42:721-724. De la Riva, I. 1990. Una especie nueva de Ololygon (Anura: Hylidae) procedente de Bolivia. Rev. Espan. Herp. 4:81-86. Duellman, W. E. 1970a. The hylid frogs of Middle America. Monogr. Mus. Nat. Hist. Univ. Kansas 1:1-753. Duellman. W. E. 1970b. Identity of the South American hylid frog Garbeana garbei. Copeia 1970:534-538. Duellman, W. E. 1972. South American frogs of the Hyla rostrata group (Am- phibia, Anura, Hylidae). Zool. Mededel. Rijksmus. Nat. Hist. Leiden 47: 177-192. Duellman, W. E. 1986. Two new species of Ololygon (Anura: Hylidae) from the Venezuelan Guyana. Copeia 1986:864-870. Duellman, W. E., and R. O. de Sa. 1988. A new genus and species of South American hylid frog with a highly modified tadpole. Trop. Zool. 1:1 17-136. Duellman, W. E., and L. Trueb. 1986. Biology of Amphibians. New York: McGraw- Hill Book Co. 18 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 151 Fitzinger, L. J. F. J. 1843. Systema Reptilium. Vol. 1. Wien: Braunmuller and Seidel. Fouquette, M. J., Jr., and A. J. Delahoussaye. 1977. Sperm morphology in the Hyla rubra group (Amphibia: Anura: Hylidae), and its bearing on generic status. J. Herpetol. 11:387-396. Frost, D. R. (Editor). 1985. Amphibian Species of the World. Lawrence: Associa- tion of Systematic Collections. Henle, K. 1991. Ololygon pedromedinae sp. nov., ein neuer Knickzehen- laubfrosche (Hylidae) aus Peru. Salamandra 27:76-82. Heyer, W. R., A. S. Rand. C. A. G. da Cruz, O. L. Peixoto. and C. E. Nelson. 1990. Frogs of Boraceia. Arq. Zool. S. Paulo 31(4):23 1—410. Hoogmoed, M. S., and S. J. Gorzula. 1979. Checklist of the savanna inhabiting frogs of the El Manteco region with notes on their ecology and the description of a new species of treefrog (Hylidae, Anura). Zool. Mededel. Rijksmus. Nat. Hist. Leiden 54:183-216. Hoogmoed, M. S.. and U. Gruber. 1983. Spix and Wagler type specimens of reptiles and amphibians in the Natural History Musea in Munich (Germany) and Leiden (The Netherlands). Spixiana Suppl. 9:31 9^4 1 5 . Leon, J. R. 1969. The systematics of the frogs of the Hyla rubra group in Middle America. Univ. Kansas Mus. Nat. Hist. Pub. 18:505-545. Leviton, A. E., R. H. Gibbs. Jr., E. Heal, and C. E. Dawson. 1985. Standards in herpetology and ichthyology. Part. I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985:802-832. Lutz, B. 1973. Brazilian Species ^/'Hyla. Austin: Univ. Texas Press. Lynch, J. D. 1973. The transition from archaic to advanced frogs. Pp. 133-182 in J. L. Vial (ed.), Evolutionary Biology oftheAnurans. Columbia: Univ. Missouri Press. McDiarmid, R. W., and R. Altig. 1990 ("1989"). Description of a bufonid and two hylid tadpoles from western Ecuador. Alytes 8:51-60. Miranda-Ribeiro, A de. 1926. Notas para servirem ao estudio dos gymnobatrachios (Anura) Brasileiros. Arq. Mus. Nac. Rio de Janeiro 27:1-227. Myers, C. W., and W. E. Duellman. 1982. A new species of Hyla from Cerro Colorado, and other tree frog records and geographical notes from Western Panama. Amer. Mus. Novit. 2752:1-25. Peixoto, O. L. 1987. Caracterizacao do grupo "perpusilla" e reavaliacao da posi^ao taxonomica de Ololygon perpusilla perpusilla e Ololygon perpusilla v-signata (Amphibia, Anura, Hylidae). Arq. Univ. Fed. Rur. Rio de Janeiro, Itaguaf Jan./ Dez. 1987:37-49. Peixoto, O. L. 1988a. Duas novas especies de Ololygon do grupo "perpusilla" (Amphibia, Anura, Hylidae). Arq. Univ. Fed. Rur. Rio de Janeiro, Itaguaf Jan./ Dez. 1988:27-37. Peixoto, O. L. 1988b. Sobre o "status" taxonomico de Hyla catharinae alcatra: B. Lutz 1973, com a descricao de uma nova especie para o grupo "perpusilla" (Amphibia, Anura, Hylidae). Acta Biol. Leopoldensia 10:253-267. Peixoto, O. L.. and P. Weygoldt. 1987. Notes on Ololygon heyeri Weygoldt 1986 from Espfrito Santo, Brazil. Senckenbergiana Biol. 68:1-9. Peters, W. 1872. Uber die von Spix in Brasilien gesammelten Batrachier des Konigl. Naturalienkabinets zu Miinchen. Monatsber. Preuss Akad. Wiss. Berlin 1872:196-227. STATUS OF OLOLYGON AND SCINAX 1 9 Pombal. J. P., Jr., and M. Gordo. 1991. Duas novas especies de Hyla da floresta atlantia no Estado de Sao Paulo (Amphibia, Anura). Mem. Inst. Butantan 53:135-144. Savage, J. M., and W. R. Heyer. 1967. Variation and distribution in the treefrog genus Phyllomedusa in Costa Rica, Central America. Beitr. Neotrop Fauna 5:111-131. Schneider, J. G. 1799. Historiae Amphibiorum Naturalis etLiterariae. Vol. 1 . Jena: Friederici Frommanni. Spix, J. B. de. 1824. Animalia Nova sive Species Novae Testudinum Ranarum, quas in Itinere per Brasiliam Annis MDCCCXVII-MDCCCXX jussu et Auspiciis Maximiliani Josephi I. Bavariae Regis. Miinchen: F. S. Hubschmann. Starrett, P. 1968. The phylogenetic significance of the jaw musculature in anuran amphibians. PhD Dissertation. Ann Arbor: Univ. Michigan. Stejneger, L. 1907. Herpetology of Japan and adjacent territory. Bull. U.S. Natl. Mus. 58:1-577. Stejneger, L. 1937. Designation of genotype for Hylaplesia Boie. Copeia 1937: 139. Trueb, L. 1970. Evolutionary relationships of casque-headed tree frogs with co- ossified skulls (family Hylidae). Univ. Kansas Mus. Nat. Hist. Pub. 18:547-716. Trueb, L. 1973. Bones, frogs, and evolution. Pp. 65-132 in J. L. Vial (ed.), Evolutionary Biology of the Anurans . Columbia: Univ. Missouri Press. Trueb, L. 1992. Patterns of cranial diversity among the Lissamphibia. Pp. 000-000 in J. Hanken and B. K. Hall (eds.). The Vertebrate Skull. Vol. 2. Structural and Systematic Diversity. Chicago: Chicago Univ. Press. Tyler, M. J. 1971. The phylogenetic significance of vocal sac structure in hylid frogs. Univ. Kansas Mus. Nat. Hist. Pub. 19:319-360. Wagler, J. G. 1 830. Natiirliches System der Amphibien, mit Vorangehender Classi- fication der Sdugethiere und Vogel. Miinchen, Stuttgart and Tubingen: J. G. Cotta. Wied-Neuwied, M. A. P., Prinz von. 1821. Reise nach Brasilien in den Jahren 1815 bis 1817. Vol. 2. Frankfurt-am-Main: H. L. Bronner. Wied-Neuwied, M. A. P., Prinz von. 1824. Beitriige zur Naturgeschichte von Brasilien. Vol. 1. Verzeichniss de Amphibien. Weimar: Gr. H. S. priv. Landes- Industrie-Comptoir. APPENDIX I Specimens Examined for Definition of Scinax Unless otherwise noted, all specimens are in the collection of The University of Kansas Museum of Natural History (KU). Cleared-and-stained skeletal prepara- tions are noted by AA for Alizarin-red and Alcian-blue stained specimens and AR for specimens stained for Alizarin-red only. Acris crepitans 158146 (AA). Agalychnis callidryas 77495 (AA). Anotheca spinosa 71122, 86900 (AA). Aparasphenodon brunoi 152227, 92213 (AA). Aplastodiscus perviridis 92722, 92723 (AA). Argenteohyla siemseri 116935. 116937 (AA). Calyptahyla crucialis 192471. Centrolene geckoideum 164491. 178017 (AA). Centrolenella audax 178028 (AA). Centrolenella balionota 164713 (AA). Centrolenella buckleyi 178059. Centrolenella grandisonae 164685 (AA). Centrolenella griffithsi 118026. 118028 (AA). Corythomantis greeningi 125380. 20 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 151 Cryptobatrachus sp. 203998 (AA). Gastrotheca plumbea 202699 (AA). Gastrotheca rebeccae 196814 (AA). Hemiphractus probiscideus 123154 (AA). Hyla albopunctata 91933. Hyla andersoni 207334 (AA). Hyla aperomea 212086, 212087 (AA). Hyla arborea 148625. Hyla bistincta 140241. Hyla boons 166770, 150025 (AA). Hyla bogotensis 125363. Hyla bromeliacia 194224. Hyla carnifex 136210. Hyla circumdata 152233. Hyla ebraccata 77112. Hyla fasciata 205477 (AA). Hylo geographica 124171 (larvae). Hyla granosa 109496 (AA). Hyla gratiosa 116934 (larvae). Hyla hazelae 100969. Hyla lanciformis 104910 (AA), 152541 (larvae). Hyla marmorata 126433, 1 52455 (AA). Hyla melanomma 136795 (AA). Hyla microcephala 71242. Hyla miliara 101610. Hyla minuta 150308. Hyla miotympanum 200808. Hyla parviceps 205632 (AA). Hyla phyllognatha 180355 (larvae). Hyla picta 64873. Hyla pictipes 103691. Hyla pinorum 135225. Hyla pseudopuma 36599. Hyla pule he lla 182961 (AA). Hyla sumichrasti 57882. Hyla taeniopus 53827. Litoria caerulea 204667 (AA). Nyctimantis rugiceps 152476. Osteocephalus leprieurii 153390. Osteocephalus taurinus 175501 (AA), 124238 (larvae). Osteopilus dominicensis 84701 (AA). Osteopilus septentrionalis 87932, 55173 (larvae). Pachymedusa dacnicolor 78453 (AA). Phrynohyas venulosa 215379, 92261 (AA), 104189 (larvae). Phyllodytes luteolus 74250. Phyllomedusa tomopterna 200512 (AA). Plectrohyla glandulosa 209699 (AA). Plectrohyla quecchi 190244. Pseudacris clarkii 207238, 207419 (AA). Pseudacris triseriata 207482 (AA). Pternohyla fodiens 75395, 60385 (AA). Ptychohyla spinipollex 64128, 192890 (AA). Scarthyla ostinodactyla 215413, 205773-74 (AA), 205865 (larvae). Scinax acuminata 128916 (AR), 128917-21. Scinax argyreornata 91947, 148435-36, 148437 (AA). Scinax aurata 125383. Scinax baumgardneri 129753. Scinax berthae 91948 (AR), 91949-53. Scinax blairi 138215. Scinax boesemani 69745, 69764-69, 128357-58 (AA). Scinax boulengeri 64328 (AR), 87774 (AR), 102174-77, 103618-20 (AA), 104295 (larvae). Scinax catharinae 92184-86, 92188-89 (AR), 92306 (AA). Scinax chiquitana 217293-94, USNM 312659 (AA). Scinax crospedospila 91992-95, 91996 (AR). Scinax cruentomma 11930-53, 152980-82 (AR), 152984-85 (AR), 111928 (AA), 109491 (larvae). Scinax cuspidata 92001-03 (AR), 92005-08. Scinax dome 167088-91, 167769-70 (AR). Scinax duartei 92018 (AR), 92019-22. Scinax elaeochroa 68281-91 (AR), 68390 (larvae), 87772-73 (AR), 96009-12. Scinax epacrorhina 139242-45, 139247, 139249. Scinax exigua 167117-20, 167771-73 (AR). Scinax flavoguttata 92027, 92028 (AR). Scinax funerea 105168-79, 152979 (AR), 150148 (AA), 152559-60 (larvae). Scinax fuscomarginata 92029-3 1 , 92034 (AR). Scinax fuscovaria 160336, 182897-99. 182900-01 (AA), 185809. Scinaxgarbei 150176-28, 152986-91 (AR). 153257 (larvae). Scinax goinorum 179004. Scinax hayii 92049-51, 92052 (AA). Scinax humilis 112376. Scinax littorea 218436 (larvae). Scinax machadoi 152330-31, 152332 (larvae). Scinax madeirae 92066-67, 92068 (AA). Scinax melloi 218438 (larvae). Scinax nasica 92 107 ( AR), 160423. Scinax nebulosa 91917 (AR), 127988-89 (AR), 128365-66. 130154 (larvae). Scinax obtriangulata 152240,211301 (larvae). Scinax pachychrus 100344-45, 100346 (AA). Scinax parked 92117-18 (AR), 127845-48 (AA). 128552-54, 130152 (larvae). Scinax pedromedinae 205304-36, 205337-38 (AA), 205859 (larvae). Scinax perpusilla 92119-23, 92124 (AR), 218435 (larvae). Scinax proboscoidea 140900. Scinax quinquefasciata 218470-79, 166317 (AR), 202967 (larvae). Scinax rizibilis 92144^45. Scinax rostrata 77687 STATUS OF OLOLYGON AND SCINAX 2 1 (AR). 101586-90, 104244 (larvae). Scinaxrubra 109469-90, 123049 (AA), 74280 (AR), 152972-77 (AR), 109492 (larvae). Scinax squalirostris 92 172-75, 92 176-77 (AR). 197304 (larvae). Scinax staufferi 59924-27 (AR). 68614-15 (AR). 104162 (larvae), 136971-77. Scinax trachythorax 80862-64. Scinax v-signata 218437 (larvae). Scinax wandae 131717. Scinax x-signata 128517-20. 150871-72 (AA). Smilisca baudinii 60415 (AA). 71722 (larvae). Smilisca sila 91833 (larvae). Smilisca phaeota 133462, 169627 (AA). Sphaenorhynchus bromelicola 124668. Sphaenorhynchus carneus 146576-81. 183709 (AA). Sphaenorhynchus dorisae 178825 (AA). 180982 (larvae). Sphaenorhynchus lacteus 178826-37, 202769, 92288 (AA), 205458 (AA). Sphaenorhynchus orophilus 74308-74309, 92295-96. Sphaenorhynchus planicola 92297-99, 92300 (AA). Stefania evansi 181 125 (AA). Trachycephalus jordani 164484. Triprion petasatus 71736 (larvae). Triprion spatulatus 71744 (AA). 73841. APPENDIX II Species-group Names Associated with the Genus Scinax and Their Current Status Trivial name. Original combination. Author, and Date Current status acuminata (Hyla), Cope, 1862' 5. acuminata affinis (Hyla), Spix, 1824 S. .x-signata agilis (Hyla), Cruz and Peixoto, 1983 5. agilia alcatraz (Hyla catharinae), B. Lutz, 1973 S. alcatraz albicans (Hyla), Bokermann, 1967 S. albicans alleni (Scytopis), Cope. 1870 S. alleni altae (Hyla), Dunn. 1933 S. staufferi alterna (Hyla rubra), B. Lutz, 1973 S. rubra angrensis (Hyla catharinae), B. Lutz. 1973 S. catharinae argyreornata (Hylodes), Miranda-Ribeiro, 1926 S. argyreornata ariadne (Hyla), Bokermann, 1967 5. ariadne atrata (Ololygon), Peixoto, 1988 S. atrata aurata (Hyla), Wied-Neuwied, 1821 S. aurata baumgardneri (Hyla), Rivero, 1961 5. baumgardneri berthae (Hyla). Barrio, 1962 S. berthae blairi (Hyla), Fouquette and Pyburn. 1972 S. blairi bocainensis (Hyla catharinae). B. Lutz, 1968 5. catharinae boesemani (Hyla). Goin, 1966 S. boesemani boulengeri (Scytopis). Cope, 1887 5. boulengeri brieni (Hyla). DeWitte. 1930 5. catharinae caldarum (Hyla duartei), B. Lutz. 1968 S. duartei camposseabrai (Hyla). Bokermann. 1968 5. x-signata canastrensis (Hyla). Cardoso and Haddad, 1982 5. canastrensis catharinae (Hyla). Boulenger. 1888 S. catharinae 'Citations appearing only in this list are not given in the literature cited. 22 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 151 Trivial name. Original combination. Author, and Date Current status chiquitana (Ololygon), De la Riva, 1990 S. chiquitana coerulea (Hyla), Spix, 1824 5. x-signata coniwstris (Hyla), Peters, 1863 S. rubra crospedospila {Hyla), A. Lutz, 1925 S. crospedospila cruentomma (Hyla), Duellman, 1972 S. cruentomma crytanthus (Scytopis), Cope, 1874 S. rubra cule.x (Hyla), Dunn and Emlen, 1932 5. staufferi cuspidata (Hyla), A. Lutz, 1925 S. cuspidata cynocephala (Hyla), Dumeril and Bibron, 1841 5. cynocephala danae (Ololygon), Duellman, 1986 S. danae depressiceps (Hyla), Boulenger, 1882 S.funerea duartei (Hyla), B. Lutz, 1951 S. duartei dulcensis (Hyla). Taylor, 1958 S. elaeochroa egleri (Hyla), B. Lutz, 1968 S. nebulosa ehrhardi (Hyla), Muller, 1924 S. ehrhardti elaeochroa (Hyla), Cope, 1876 S. elaeochroa epacrorhina (Hyla), Duellman, 1972 S. epacrorhina eringiophila (Hyla strigilata), Gallardo, 1961 S. .x-signata eurydice (Hyla), Bokermann, 1968 5. eurydice evelynae (Hyla), Schmidt, 1944 S. sc/ualirostris e.xigua (Ololygon), Duellman, 1986 S. exigua fiebrigi (Hyla). Ahl. 1927 S. acuminata flavoguttata (Hyla). A. Lutz and B. Lutz, 1939 S. flavoguttata foliamorta (Hyla). Fouquette. 1958 S. rostrata funerea (Hyla), Cope, 1874 S.funerea fuscomarginata (Hyla). A. Lutz. 1925 S. fuscomarginata fuscovaria (Hyla), A. Lutz, 1925 S.fuscovaria garbei (Garbeana), Miranda-Ribeiro, 1926 S. garbei goinorum (Hyla), Bokermann, 1962 S. goinorum granulata (Hyla). Peters, 1871 S. .x-signata hayii (Hyla), Barbour, 1909 S. hayii hcyeri (Ololygon), Peixoto and Weygoldt. 1986 5. heyeri hiemalis (Hyla), Haddad and Pombal-Junior, 1987 S. hiemalis huebneri (Hyla rubra), Melin, 1941 S. rubra humilis (Hyla), B. Lutz, 1954 S. humilis inconspicua (Hyla rubra), Melin, 1941 S.funerea jureia (Hyla), Pombal and Gordo, 1991 S. jureia kennedyi (Hyla), Pyburn. 1972 S. kennedyi lateristriga (Hyla), Spix, 1824 S. rubra lindneri (Hyla), Muller and Hellmich, 1936 S. fuscomarginata lineomaculata (Hyla), Werner, 1899 S. rubra littoralis (Hyla), Pombal and Gordo, 1991 S. littoralis littorea (Hyla), Peixoto, 1988 S. littorea longilinea (Hyla), B. Lutz, 1968 S. longilinea lutzi (Hyla), Melin, 1941 S. garbei STATUS OF OLOLYGON AND SCINAX 23 Trivial name. Original combination. Author, and Date Current status machadoi (Hyla), Bokermann and Sazima. 1973 5. machadoi madeirae (Hyla). Bokermann. 1964 S.fuscomarginata maracaya (Hyla). Cardoso and Sazima, 1980 S. maracaya megapodia (Hyla). Miranda-Ribeiro. 1926. 1937 S.fuscovaria melloi (Ololygon). Peixoto, 1988 S. melloi mirim (Hyla). B. Lutz. 1973 5. rizibilis nasica (Hyla). Cope, 1862 5. nasica nebulosa (Hyla). Spix. 1824 S. nebulosa nigra (Hyla). Cope, 1887 S. x-signata obtriangulata (Hyla), B. Lutz, 1973 S. obtriangulata opalina (Hyla catharinae), B. Lutz, 1968 5. opalina orientalis (Hyla rubra). B. Lutz, 1968 S. rubra pachychrus (Hyla). Miranda-Ribeiro. 1937 S. pachychrus parkeri (Hyla). Gaige. 1929 S.parkeri pedromedinae(Ololygon). Henle, 1991 5. pedomedinae perpusilla (Hyla). A. Lutz and B. Lutz. 1939 S. perpusilla phrynoderma (Hyla), Boulenger, 1889 5. acuminata pickeli (Hyla), A. Lutz and B. Lutz, 1938 S. pachychrus proboscidea (Hyla), Brongersma, 1933 S. proboscidea queimadensis (Hyla p[erpusilla]). B. Lutz, 1973 S. v-signata quinquefasciata (Hyla), Fowler. 1913 S.quinquefasciata quinquevittata (Hyla), Cope, 1868 S. elaeochroa ranki (Hyla). Andrade and Cardoso, 1987 S.ranki rizibilis (Hyla). Bokermann, 1964 S. rizibilis robersimoni (Hyla). Donoso-Barros, 1965 S. rubra rostrata (Hyla), Peters, 1863 S. rostrata rubra (Hyla), Laurenti, 1768 5. rubra similis (Hyla), Cochran, 1952 5. similis simplex (Hyla catharinae). B. Lutz. 1968 S. obtriangulata squalirostris (Hyla). A. Lutz. 1925 5. squalirostris staufferi (Hyla). Cope. 1865 S. staufferi strigilata (Hyla). Spix, 1824 5. strigilata sugillata (Hyla), Duellman. 1973 S. sugillata trachythorax (Hyla), Miiller and Hellmich. 1936 S. trachythorax trapicheiroi (Hyla), B. Lutz, 1954 S. catharinae trilineata (Ololygon), Hoogmoed and Gorzula, 1979 5. trilineata v-signata (Hyla perpusilla), B. Lutz, 1968 5. v-signata wandae (Hyla). Pyburn and Fouquette, 1971 S. wandae x-signata (Hyla), Spix, 1824 S. x-signata 3 2044 093 361 715