RATTLESNAKE HARVEST IN FLORIDA 557 Histories,and Influenceon Mankind,2nd ed. Univ. TIMMERMAN, W. W. 1995. Home range, habitat 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 Physalaemus spiniger (Anura: Leptodactylidae) 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 Physalaemus spiniger 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 genus 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 Engystomops (=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- ,<w I4 * a.. 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.