RATTLESNAKE HARVEST IN FLORIDA 557

Histories,and Influenceon Mankind,2nd ed. Univ. TIMMERMAN, W. W. 1995. Home range, use, CaliforniaPress, Berkeley. and behavior of the eastern diamondbackrattle- NEILL, W. T. 1961. Giant rattlesnakes-past and pres- snake (Crotalus adamanteus) on the Ordway Pre- ent. FloridaWildl. 15:10-13. serve. Bull. FloridaMus. Nat. Hist. 38:127-158. SNYDER, B. 1949. Diamondbacksand dollarbills. Flor- TINKLE,D. W. 1962.Reproductive potential and cycles ida Wildl. 3:3-5, 16, 19. in female Crotalusatrox from northwesternTexas. TELFORD, S. R., JR. 1952. A herpetological survey in Copeia 1962:306-313. the vicinity of LakeShipp, Polk County,Florida. Q. J. FloridaAcad. Sci. 15:175-185. Accepted:5 August 1998.

Journalof Herpetology,Vol. 32, No. 4, pp. 557-565,1998 Copyright1998 Societyfor the Study of Amphibiansand Reptiles

Redescription of spiniger (Anura: ) and Description of Two New Reproductive Modes

CELIO E B. HADDAD13 AND JOSE P. POMBAL, JR.2

'Museum of VertebrateZoology, University of California,Berkeley, California 94720, USA, and 2Departamentode Vertebrados,Museu Nacional, Quinta da Boa Vista, 20940-040 Rio de Janeiro,Rio de Janeiro,Brasil

ABSTRACT.-We redescribe and describe its tadpole and its reproductive modes. This species has the following three alternative reproductive modes: (1) foam nest on pond and feeding tadpoles in pond (the typical mode for the Physalaemus); (2) foam nest on humid places on the forest floor near a pond, and feeding tadpoles in pond; (3) foam nest on water accumulated on the axils of terrestrial bromeliads and feeding tadpoles in pond. These last two modes were not included in the reviews of repro- ductive modes in anurans. The vocalizations of P?spiniger are described and compared with the vocalizations of P nanus, a sibling species.

Miranda-Ribeiro (1926) described Engysto- bal and Madureira, 1997): P bokermanniCardoso mops spinigera from Iguape, Sao Paulo State, and Haddad, P caete Pombal and Madureira, P southeastern Brazil, attributing the specific crombiei Heyer and Wolf, P maculiventris (A. name to Boulenger because the specimens were Lutz), P moreirae (Miranda-Ribeiro), P nanus so identified in the collection of the Museu Pau- (Boulenger), P obtectus Bokermann, P olfersii lista (currently Museu de Zoologia da Univer- (Lichtenstein and Martens), P signifer (Girard), sidade de Sao Paulo). However, Boulenger never and P spiniger (Miranda-Ribeiro). In recent years described (=Physalaemus) spinigera we collected and observed specimens of P spi- and accordingly the short account by Miranda- niger at several localities in southern and south- Ribeiro (1926) serves as the original description. eastern Brazil. Herein, we redescribe this species The specific name Engystomopsspinigera was ap- and its tadpole. The vocalizations are described parently forgotten and ignored by subsequent and compared with the vocalizations of P nanus, authors (e.g., Cochran, 1955; Bokermann, 1966a; a sibling species. Also, we provide information Lynch, 1970; Frost, 1985; Heyer and Wolf, 1989; on reproduction, including two new reproduc- Duellman, 1993). Lynch (1970) recognized four tive modes for anurans. in the Physalaemus:P bili- species groups genus MATERIALS AND METHODS gonigerus group, P cuvieri group, P pustulosus group, and P signifer group; P deimaticus and P Specimens used in the description or exam- rupestris are not presently allocated to any spe- ined for comparisons are deposited in AL-MN cies group (Sazima and Caramaschi, 1986; Car- (Adolpho Lutz collection, deposited in Museu amaschi et al., 1991). Ten species are presently Nacional, Rio de Janeiro, RJ,Brasil), CFBH (Celio allocated to the Physalaemussignifer group (Pom- F B. Haddad collection, deposited in Departa- mento de Zoologia, Universidade Estadual Pau- lista, Rio Claro, SP, Brasil), MNRJ (Museu Na- 3Present Address: Departamento de Zoologia, cional, Rio de Janeiro, RJ, Brasil), MZUSP (Mu- UniversidadeEstadual Paulista,13506-900 Rio Claro, seu de Zoologia da Universidade de Sao Paulo, Sao Paulo,Brasil, E-mail: [email protected] Sao Paulo, SP, Brasil), WCAB (Werner C.A. Bok- 558 C. F. B. HADDAD AND J. P. POMBAL, JR.

;, s. Wj *e- ,

0 VI

FIG. 1. Physalaemusspiniger, an adult male in life from Ilha do Cardoso,Canan6ia, SP, Brazil.

ermann collection, deposited in Museu de Zoo- RESULTS logia da Universidade de Sao Paulo, Sao Paulo, and with other ZUEC de Hist6ria Diagnosis Comparison Species.- SP, Brazil), (Museu Natural, A to the Physalaemus Universidade Estadual de species belonging signifer Campinas, Campinas, (sensu 1970), characterized (1) Additional examined are group Lynch, by SP, Brasil). specimens small size (males 17.1-21.3 mm SVL, females listed in 1. Appendix 20.6-22.0 mm SVL); (2) canthus rostralis dis- Abbreviations used in the measurements of tinct; (3) dorsal skin texture smooth to rugose; the adults are SVL HL (snout-vent length), (4) belly orange in life; (5) structure of the ad- (head length), HW (head width), ED (eye di- vertisement call. THL TBL ameter), (thigh length), (tibia length), Physalaemus spiniger is distinguished from P and FL All measurements are ex- (foot length). caete, P moreirae,P obtectus, and P olfersii by its in millimeters. The measurements of pressed smaller size (combined SVL of males ranging the adults follow Duellman (1970) and Cei 22.5-34.5 mm; Bokermann, 1966b; Heyer, 1985; (1980). Measured adult specimens were fixed in Heyer and Wolf, 1989; Heyer et al., 1990; Pombal 10% formalin and maintained in 70% ethyl al- and Madureira, 1997). By its larger size and dis- cohol. Eggs and tadpoles were fixed and main- tinct advertisement call, P spiniger differs from tained in 5% formalin. For measurements we P bokermanni(P bokermannimales with 15.3-17.0 used an ocular micrometer in a Zeiss stereo- mm SVL; see Cardoso and Haddad, 1985). Phys- microscope, except that SVL was measured with alaemus spiniger differs from P signifer by calipers. Drawings of the adult and tadpole smoother dorsal skin (more rugose in P signifer), were made using a Zeiss stereomicroscope with larger black spot on the inguinal glands, and a drawing tube. Vocalizations were recorded us- smaller mean length (P signifer males with SVL ing a Nagra E tape recorder with a Sennheiser = 20.1-22.4, x = 21.7, SD = 0.7, N = 20). Phys- ME 80 microphone at a tape speed of 19 cm/s. alaemus spiniger differs from P crombiei by the The sonagrams were produced by a Macintosh orange belly (pinkish in P crombiei;Heyer and computer coupled to the MacRecorded Sound Wolf, 1989) and distinct advertisement call (see System 2.0.5, using 8 bit resolution, 22 kHz sam- below and Heyer and Wolf, 1989). Physalaemus pling frequency, and FFT with 256 points. spiniger differs from P maculiventris by wider REDESCRIPTION AND REPRODUCTION OF PHYSALAEMUS SPINIGER 559

I \ci

B A,

t/

C

D

FIG.2. Physalaemusspiniger, CFBH 0307. (A) Dorsal and (B) lateral views of head; ventral views of (C) hand and (D) foot (scales = 3.0 mm). head and snout, and by posterior belly light col- chest; vocal slits present in males; choanae ored (boldly dark and light mottled in P macu- small, nearly round; tongue narrow, long; vo- liventris). Preserved specimens of P nanus and P merine teeth absent; maxillary teeth not visible, spiniger are indistinguishable; however, they but discernible by probe. Arms slender, fore- have different advertisement calls (see below). arms moderately robust in males (in females Description of Adults.-Body slender (Fig. 1); slender); fingers short; brown nuptial pad on head wider than long; snout rounded and pro- thumb in males; subarticular tubercles single, truding in dorsal and lateral views (Fig. 2A, B); protruding and rounded; outer metacarpal tu- nostril slightly protuberant, directed laterally; bercle large to small, rounded (81% of speci- canthus rostralis distinct; loreal region concave; mens) or ovoid (19% of specimens); inner met- eye slightly protuberant; tympanum indistinct acarpal tubercle large to medium, nearly ellip- to weakly distinct; distinct supratympanic fold tical, sometimes rounded; supernumerary tu- from tympanum to shoulder; narrow dorsolat- bercles small; finger tips not expanded; finger eral fold extending from posterior corer of eye lengths I = IV (or rarely IV < I) < II < III (Fig. to inguinal region; males with subgular vocal 2C). Legs moderately robust; tibia longer than sac, expanded externally, extending to border of thigh; foot with an inner metatarsal tubercle 560 C. F. B. HADDAD AND J. P. POMBAL, JR.

TABLE1. Measurements (in mm) of males and fe- 4- males of Physalaemusspiniger. -2 A N Males (N = 28) Females(N = 4) x SD Range x SD Range B 2 SVL 18.8 1.0 17.1-21.3 21.1 0.6 20.6-22.0 z e HL 5.2 0.4 4.0-5.8 5.9 0.3 5.7-6.5 HW 5.9 0.3 5.2-6.5 6.3 0.1 6.2-6.5 ED 2.1 0.2 1.8-2.6 2.3 0.3 2.0-2.7 THL 8.6 0.3 8.1-9.3 9.5 0.4 9.1-10.0 Lo2 TBL 9.3 0.4 8.5-10.5 10.2 0.3 9.9-10.7 FL 9.1 0.4 8.2-10.0 10.2 0.2 9.9-10.4 TIME 0.2 s

the arrow marks sometimes with a fine light ^^j^Uilgl,^ D border, in some specimens dorsal marks not ev- ident; a white line on the dorsolateral fold (sometimes absent); flanks below the dorsolat- eral fold, black; forearm light brown or gray, with a dark brown transverse bar; elbow dark IIIIII. . . E brown; thigh, tibia, and foot light brown, brown or gray, similar to the color of the dorsum; thigh and tibia with a dark brown transverse bar; two " MTT black bars on the upperparts of thigh and tibia; TIME , anal dark; cream or chest and 0.2 s region belly gray; throat gray or dark brown. Color in Life.-Dorsum yellowish brown to FIG. 3. Sonograms of the calls of Physalaemusspi- brown; dorsal marks (interorbital bar and ar- niger: (A) advertisement calls and (B) encounter calls row) dark brown; in some the dorsal = specimens of two interacting males; air temperature 15.5 C, marks are not evident; snout brown to cream; water temperature = 21 C; Ilha do Cardoso, Canan6ia, dark brown lateral stripe from snout, through Sao Paulo, Brazil. (C) Sonagrams of the calls of Phys- alaemus nanus: advertisement call and vocalization nostril, eye, tympanum, and arm insertion to with a black composed by two parts, respectively; air = 23.5, water groin; groin orange; inguinal gland = 22 C; Sao Joao do Rio Vermelho, Florian6polis, spot; forearm, thigh, tibia, and foot yellowish Santa Catarina, Brazil. (D) Waveform of the second brown to brown with a dark transverse bar; vocalization in Fig. 3A. (E) Waveform of the second chest and throat dark brown with white spots; vocalization in 3C. Fig. belly, axilla, and ventral surfaces of thigh, tibia, foot, and arm orange; anal region black. Vocalizations.-When an isolated male of Phys- slight protruding, elliptical to nearly rounded; alaemusspiniger is calling it emits pulsed adver- outer metatarsal tubercle small, slightly pro- tisement calls that are observed in the sonagram truding, rounded; well-developed subarticular as three sidebands between approximately 1.2 tubercles, single, protruding, round; supernu- and 2.1 kHz (Fig. 3A). However, the waveform merary tubercles small; toe tips slightly expand- shows a pulsed structure in the advertisement ed; toe lengths I < II < V

::.:

A

B

C

FIG.4. Tadpole of Physalaemusspiniger, Stage 36; (A) Dorsal and (B) lateral views; (C) oral disc (upper scale - 5.0 mm; bottom scale 0.5 mm). the last part of these vocalizationsthe sidebands range = 0.39-0.52, N = 8 vocalizations of one are replaced by a creaking sound as a result of male). This vocalization is composed of two dif- a decrease in pulse rate. The encounter calls ferent parts. In the first part there are six to 10 emitted by the first calling male do not show introductory notes (i = 8.13, SD = 1.36, N = 8 the strong initial descending and ascending fre- vocalizations of one male) similar to those ob- quenciesthat are observedin the encountercalls served in the advertisement calls; the second of the second calling male (Fig. 3B). part has a pulsed structure observed in the son- In Physalaemusnanus the mean durationof the agram as three to five sidebands very similar to advertisementcalls is 0.18 sec (SD = 0.01, range the advertisement and encounter calls of P spi- = 0.16-0.20, N = 10 vocalizationsof one male). niger (Fig. 3A-E). This vocalization has 7-9 notes (x = 7.9, SD = Tadpoles.-Larvae were obtained at the munic- 0.57, N = 10 vocalizations of one male) with a ipality of Sete Barras, State of Sao Paulo. The frequencybetween 1.0 and 2.3 kHz (Fig. 3C). A following description is based on a tadpole in second type of vocalization, emitted after the developmental Stage 36 (Gosner, 1960). Body emission of a variablenumber of advertisement ovoid in dorsal (Fig. 4A) and ventral views, de- calls, shows frequencies between 1.0 and 2.3 pressed/globular in lateral view (Fig. 4B); body kHz and mean duration of 0.46 sec (SD = 0.05, wider than high; snout rounded; eyes small, 562 C. F. B. HADDAD AND J. P. POMBAL, JR.

I20 200 km

FIG.5. Geographicdistribution of Physalaemusspiniger in southernand southeasternBrazil.

dorsolateral; nostrils dorsal, small and rounded; Iporanga; Saibadela, Sete Barras; Jacupiranga; nostrils about midway between the eyes and the Caraguatatuba) (Fig. 5). Although the P. signifer tip of snout; spiracle sinistral, its opening pos- group occurs from the State of Alagoas, in terior to the middle of body; cloacal tube me- northeastern Brazil, through southeastern to the dium sized, median; caudal musculature slen- State of Santa Catarina, in southern Brazil der; dorsal fin originating on body; dorsal fin (Frost, 1985; Pombal and Madureira, 1997), wider than ventral. Oral disc (Fig. 4C) directed most species of this group have restricted dis- ventrally and bordered by papillae, interrupted tributions (Heyer and Wolf, 1989), but P spiniger along a large area on the anterior labium; tooth has a moderately broad distribution. row formula 2 (2)/3 (1); jaw sheath strongly de- ReproductiveModes.-Physalaemus spiniger re- veloped and serrate; posterior jaw sheath v- produces during the rainy season (October-Feb- shaped. In preservative, body gray; caudal mus- ruary) near areas subject to flooding on the for- culature with scattered pigmentation; fins trans- est floor. Males call from the edges of forest lucent with scattered pigmentation. ponds, and the eggs are laid in a foam nest that Measurements of five tadpoles in develop- is deposited on the water, anchored on the mar- mental Stage 36: range (x, SD), total length 17.3- gins of ponds or on the humid leaf litter near 21.2 (19.56, 1.98); body length 5.8-7.5 (6.74, the ponds; embryos were observed developing 0.74); body height 3.5-4.1 (3.72, 0.27); body inside the foam mass, and feeding tadpoles width 4.3-5.7 (5.0, 0.62); interarial distance were observed developing in ponds. At Ilha do 0.7-1.0 (0.82, 0.13); interorbital distance 0.9-1.2 Cardoso, during the last part of the dry season (1.12, 0.13); eye-nostril distance 0.5-0.6 (0.56, (September), a pond where P spiniger repro- 0.05); eye diameter 0.8-1.0 (0.86, 0.09). duced dried up; after a light rain, a large chorus Distribution.-Physalaemus spiniger is known formed near the empty depression. Males were from the states of Parana (Guaraqueqaba) and observed calling during both day and night, Sao Paulo (Ilha do Cardoso, Cananeia; Estacao from terrestrial bromeliads that accumulate wa- Ecol6gica da Jureia, Iguape; Cavera do Diabo, ter near the depression. We counted 25 foam f

REDESCRIPTION AND REPRODUCTION OF PHYSALAEMUS SPINIGER 563

;? , .^' .

FIG.6. Foam nest (arrow)of Physalaemusspiniger on water accumulatedin the axils of a terrestrialbromeliad. nests: 21 on the water accumulatedin the axils among different species (see Heyer et al., 1996 of bromeliads (Fig. 6) and four on the leaf litter. and references therein). and The foam nests observed in bromeliadswere on P spiniger clearly have strong evolutionary affin- the leaves near the ground, between 2.6-22.0 cm ities observable by the indistinguishable mor- from the ground (x = 10.31, SD = 5.933, N = phology and by the resemblance of the pulsed 16). The mean number of eggs per nest, collect- structures present in the vocalizations of both ed from the bromeliads, was 29.7 (SD = 7.6, species. The study of vocalizations is a promis- range = 23-38, N = 3). We did not find tadpoles ing way to uncover other sibling species in the developing in the bromeliads,but embryoswere genus Physalaemus. developing inside the foam mass. During the We observed the following three alternative rainy season at liha do Cardoso we observed reproductive modes for Physalaemusspiniger: (1) foam nests on the water of temporary ponds foam nest on pond and feeding tadpoles in and on the leaf litter. Eggs are unpigmented, pond (Lynch, 1970; Mode 8 of Duellman and with an average diameter of 1.62 mm (SD = Trueb, 1986); (2) foam nest on humid places on 0.09, range = 1.4-1.8 mm, N = 15 eggs from the forest floor near a pond, and feeding tad- three dutches). poles in pond; (3) foam nest on water accumu- lated on the axils of terrestrial bromeliads and DISCUSSION feeding tadpoles in pond. These last two modes The Physalaemussignifer group contains spe- were not listed in the reviews of reproductive cies difficult to characterizeon the basis of mor- modes of anurans (Duellman, 1985, 1989; Duell- phological attributes of preserved specimens. man and Trueb, 1986). Preserved specimens of P nanus and P spiniger Foam nests in humid places of the forest floor, are morphologically indistinguishable.Howev- as observed for P spiniger, were observed for er, the differences between the advertisement three other species in the calls of P nanus and P spinigerare sufficient to group: P bokermanni,P. crombiei, and P signifer separateboth species. Advertisementcalls have (Haddad and Pombal, unpubl. data). However, been used to characterize sibling species of for P caete (Pombal and Madureira, 1997) and P because they are typically stereotyped,al- olfersii (Haddad and Pombal, unpubl. data) only lowing an unequivocal selection by the females, the generalized mode for the genus (foam nest- therefore acting as an isolating mechanism ing on pond and feeding tadpoles in pond) has 564 C. F. B. HADDAD AND J. P. POMBAL, JR. been observed. Cruz and Peixoto (1985) ob- todactylidae) from Minas Gerais, Southeastern served foam nests of an unidentified species, Brazil. Herpetologica 47:148-151. of on terrestrial bromeli- CARDOSO,A. J., AND C. F B. HADDAD. 1985. Nova es- probably Physalaemus, de do ads growing in a floor depression inside a forest pcie Physalaemus grupo signiferus (Amphib- ia, Anura, Leptodactylidae). Rev. Brasil. Biol. 45: at Santa Tereza, State of Espirito Santo, Brazil. 33-37. These authors concluded that the tadpoles CEI, J. M. 1980. of Argentina. Monit. would not meet suitable conditions to develop Zool. Ital. (N.S.). monogr. 2:1-609. in the bromeliads. The bromeliads could prob- COCHRAN,D. M. 1955. Frogs of Southeastern Brazil. ably provide the necessary humidity for keep- Bull U.S. Nat. Mus. 206:1-423. ing eggs and embryos alive until the rain water CRUZ,C. A. G., AND 0. L. PEIXOTO.1985. Sobre de- flooded the depression, allowing the embryos to sovas de Physalaemusem local inusitado. Arq. Univ. pass to the pond, enabling an early colonization Fed. Rur. Rio de Janeiro. 8:103-105. W. E. 1970. The of Middle of the pond by the embryos and tadpoles. These DUELLMAN, hylid frogs conclusions are valid for the of P America. Monog. Mus. Nat. Hist. Univ. Kansas 1: eggs spiniger 1-753. on the bromeliads. when deposited Probably, .1985. modes in anuran foam nests of are on the Reproductive amphib- Physalaemus deposited ians: phylogenetic significance of strate- forest floor or in the axils of terrestrial brome- adaptive gies. South Afr. J. Sci. 81:174-178. liads, the transportation of the embryos is made . 1989. Alternative life-history styles in anuran by the rain water that washes and carries the amphibians: evolutionary and ecological implica- foam nests and embryos until a nearby tempo- tions. In M. N. Bruton (ed.), Alternative Life-His- rary pond. tory Styles of , pp. 101-126. Kluwer Acad. The alternative options of reproductive Publ., Dordrecht. 1993. of the world: addi- modes described here for P. spiniger are appar- species related to rainfall and to tions and corrections. Univ. Kansas Publ., Mus. ently unpredictability Nat. Hist. Publ. 21:1-372. the of a foam nest the Spec. presence involving eggs. ,AND L. TRUEB.1986. of While or larvae are within a foam Biology Amphibians. eggs nest, McGraw-Hill, New York. most are from desiccation protected (Heyer, FROST,D. R. (ed.). 1985. Amphibian Species of the 1969). When the rains fill the pond, the larvae World. A Taxonomic and Geographical Reference. are released from the foam nest and enter the Allen Press and Assoc. Syst. Coll., Lawrence, Kan- pond with a temporal advantage over other an- sas. uran species reproducing at the pond. Different GOSNER,K. L. 1960. A simplified table for staging an- stages of a continuum from an aquatic to a ter- uran embryos and larvae with notes on identifi- cation. 16:183-190. restrial reproductive mode are known for spe- Herpetologica HEYER,W. R. 1969. The of the cies in the family Leptodactylidae (Heyer, 1969). adaptive ecology species is a groups of the genus (Amphibia, Lep- Physalaemus spiniger singular leptoactylid Evolution 23:421-428. because in one it is to todactylidae). population possible 1985. New species of frogs from Boraceia, Sao observe different reproductive modes as steps Paulo, Brazil. Proc. Biol. Soc. Wash. 98:657-671. a toward more terrestrial life. , AND A. J. WOLF. 1989. Physalaemuscrombiei thank U. (Amphibia: Leptodactylidae), a new frog species Acknowledgments.-We Caramaschi, from Santo, Brazil with comments on the W. E. Duellman, W R. R. A. and Espirito Heyer, Seigel, P. signiferus group. Proc. Biol. Soc. Wash. 102:500- D. B. Wake for helpful suggestions on the manu- 506. script; R. P. Bastos and M. Gordo for field assis- ,A. S. RAND,C. A. G. CRUZ,O. L. PEIXOTOAND tance; P. R. Nascimento for the drawings; L. M. C. E. NELSON.1990. Frogs of Boraceia. Arq. Zool, Castanho and I. Sazima for loaning specimens Sao Paulo, 31:235-410. of P spiniger; M. Martins for the helping with ,J. M. GARCiA-LOPEZ,AND A. J. CARDOSO.1996. the photos. Financial support was provided by Advertisement call variation in the Leptodactylus FAPESP (proc. 95/9269-2) and CNPq (research mystaceus (Amphibia: Leptodac- SEMA, Instituto Florestal de Sao tylidae) with a description of a new sibling species. fellowship). 17:7-31. Paulo, and Aracruz Celulose S. A. for Amphibia-Reptilia logistical LYNCH,J. D. 1970. status of the American at Jureia, Ilha do Cardoso, and Aracruz, Systematic support leptodactylid frog genera Engystomops,Eupemphix, respectively. and Physalaemus.Copeia 1970:488-496. MIRANDA-RIBEIRO,A. 1926. Notas servirem ao LITERATURECITED para estudo dos gymnobatrachios (Anura) brasileiros. BOKERMANN,W. C. A. 1966a. Lista Anotada das Lo- Arq. Mus. Nac., Rio de Janeiro, 27:1-227. calidades Tipo de Anfibios Brasileiros. RUSP, Sao POMBAL,JR., J. P., AND C. MADUREIRA.1997. A new Paulo. species of Physalaemus (Anura: Leptodactylidae) .1966b. Dos nuevas especies de Physalaemusde from the Atlantic Rain Forest of northeastern Bra- Espiritu Santo, Brasil. Physis 26:193-202. zil. Alytes 15:105-112. CARAMASCHI,U., L. C. CARCERELLI,AND R. N. FEIO. SAZIMA, I., AND U. CARAMASCHI. 1986. Descricao de 1991. A new speces of Physalaemus(Anura: Lep- Physalaemusdeimaticus, sp. n., e observac6es sobre REDESCRIPTION AND REPRODUCTION OF PHYSALAEMUS SPINIGER 565

comportamento deimatico em P nattereri 1797, 9975-10020 (Serra de Araraquara, PR). Physalae- (Steindn.)-Anura, Leptodactylidae. Revista de mus moreirae: MNRJ 464 (holotype, Paranapiacaba, Biologia 13:91-101. Santos, SP). Physalaemus nanus CFBH 081 (Sao Jose, SC); CFBH 2771, MNRJ 12827-32 (Florian6polis, SC). Accepted: 5 August 1998. Physalaemus obtectus: MNRJ 4025, 14206-07 (para- types, Linhares, ES). : MNRJ 2428 APPENDIX1 (Parati, RJ); MNRJ 0482, 5525-26, 12826 (Teres6polis, 1123, 6616-35 Additional Specimens Examined RJ). Physalaemussignifer: MNRJ (Duque de Caxias, RJ); MNRJ 2753, 12461-62 (Barro Branco, caete: 9712-17 Physalaemus MNRJ (Murici, AL); Duque de Caxias, RJ); MNRJ 2766, 12477-80 (Pendo- 9803 9848-50 MNRJ (holotype), 9801-02, 9804-05, tiba, Niteroi, RJ); MNRJ 12837-42 (topotypes, Jacare- (paratopotypes), 18280 (lot of tadpoles, Passo de Ca- pagua, Rio de Janeiro, RJ). Physalaemusspiniger: CFBH maragibe, AL). Physalaemusbokermanni: MZUSP 59551 312-17, 319-21, MNRJ 18474, ZUEC 6878, 6881-82 (holotype), MZUSP 59552, WCAB 48146, ZUEC 4520- (Canan6ia, SP); ZUEC 9333-43 (Caraguatatuba, SP); 21 (paratypes, Santo Andre, SP). Physalaemuscrombiei: CFBH 2479 (Eldorado, SP); MNRJ 18475-76 (Guara- MNRJ 17694-745 (Aracruz, ES). Physalaemus maculi- quecaba, PR); CFBH 307-10, 410, 835, MNRJ 18470- ventris AL-MN 684 (syntype, "Alto da Serra de Cu- 72, 18473, ZUEC 6876-77 (topotypes, Iguape, SP); batao"); MNRJ 4228-30 (Paranapiacaba, SP); MNRJ ZUEC 3250 (Jacupiranga, SP).