Comparative Cytogenetic Studies of Eleven Species of the Group

Heredifas 125: 37-46 (1996)

Comparative cytogenetic studies of eleven species of the Tru~idurustur~~~~~~ group (Sauria, Tropiduridae), with banding patterns

SANAE KASAHARA’, KATIA CRISTINA MACHADO PELLEGRINO*, MIGUEL TREFAUT RODRIGUES3 and YATIYO YONENAGA-YASSUDA2

’ Departamento de Biologia, Instituto de Biocigncias, Universidade Estadual Paulista, UNESP, Rio Claro, SP, Brasil Departamento de Biologia, Instituto de Biocigncias, USP, Slio Paulo, SP, Brasil ’ Departamento de Zoologia, Instituto de Biocigncias, USP, Sclo Paulo, SP, Brasil

&SAHARA, S., PELLEGRINO, K. C. M., RODRIGUES,M. T. and YONENAGA-YASSUDA, Y. 1996. Compar- ative cytogenetic studies of eleven species of the Tropidurus forquatus group (Sauria, Tropiduridae), with banding patterns. - Heredifas 125 37-46. Lnnd, Sweden. ISSN 0018-0661. Received May 24, 1996. Accepted October 10, 1996

The chromosomes of 173 specimens representing eleven species of the Tropidurus forquatus group, from 33 localities in Brazil, were analysed after Giemsa staining, C-banding, NORs, and replication banding techniques. A karyotype with 2n = 36, including 12 macrochromosomes and 24 microchromosomes (12 M + 24 m), and sex determination of the XY:XX type were found in Tropidurus cocorobensis, T. erythrocephalus, T. etheridget, T. hispidus, T. hygomi, T. monfanus, T. mucujensis, T. oreadicus, and T. forquatus. The two other species, T. ifambere and T. psammonasfes, presented 2n = 36 (12 M + 24 m) karyotype only in females while males had 2n = 35 (12 M + 23 m). due to the sex determination of the X, X2Y:XlXI X2X2 type. Other interspecific differences as well as some intraspecific variation regarding the NORs and C-banding patterns have been observed, mainly in the microchromosome set. On the contrary, the macrochromosomes were highly conservative. Although consistent karyotypic diversity occurred in the torquarus group, the cytogenetic data obtained up to now did not allow us to clarify the phylogenetic relationships of the species. Nevertheless, the geographical distribution of the distinct cytotypes in T. hispidus and T. rorquafus suggested that more than one species might be involved in each case.

Sanae Kasahara, Departamento de Biologia, Instifuto de BiociZncias, Universidade Esfadual Paulista, UNESP, CP 199, CEP 13506-900, Rio Claro, SP, Brad

Until recently, the former genus Tropidurus was this analysis, the former western Tropidurus of considered a natural assemblage of open forma- the occipitalis and peruvianus groups were consid- tion lizards widely distributed throughout the ered as Microlophus and Plesiomicrolophus, while South American continent and the Galapagos the eastern species, previously included in the Islands. Several taxonomic revisions of Tropidurus melanopleurus, nanuzae, and torquatus groups, the species have been made over the last years ( DIXON genus Tapinurus, and the species of the mainly and WRIGHT 1975; ORTIZ1980; CEI 1982; GUDY- forest genera Plica, Strobilurus, and Uracentron NAS and SKUK 1983; RODRIGUES1981, 1984a, were grouped in a redefined genus Tropidurus, 1984b, 1986, 1987), and five relatively homoge- rendering it monophyletic. One of the points that neous species groups were accepted to belong to remains obscure in the cladogram concerns the the genus: occipitalis, melaizopleurus, peruuianus, position and relationship of the former torquatus torquatus, and nanuzae ( RODRKXJES1988). Al- group. Once considered a monophyletic complex though these studies improved the species level of species, based on the absence of a dorsal crest taxonomy of this complex and it remains largely of scales, the group does not appear so in the accepted, the generic concept of Tropidurus has cladogram. Although we think that this charac- changed very recently. teristic is synapomorphic for the group (includ- FROST(1992) carried out an extensive phyloge- ing Tapinurus), much more data are necessary netic analysis of Tropidurus and related genera, to firmly establish the relationships among the providing a new taxonomic scheme. According to species. 38 s. KASAHARA ET AL. Hereditas 125 (I 996)

Until 1978, only three species of the former ing techniques (HOWELL and BLACK 1980). torquatus group were recognized. RODRIGUES Replication R-banding was obtained following (1987), reviewing the forms occurring in regions FPG staining ( DUTRILLAUXand COUTURIER south of the Amazon river, has shown the existence 1981). of eleven species currently known as: T. cocorobensis, T. erythrocephalus, T. etheridgei, T. hispidus, T. hygomi, T. insulanus, T. itambere, T. montanus, T. mucujensis, T. oreadicus, and T. torquatus. Later, Results the 12th species of the group, T.psammonastes, was Conventional staining described (RODRIGUESet al. 1988). T. bogerti from Venezuelan tepuis was not included in the analysis Nine species, T. cocorobensis, T. erythrocephalus, made by RODRTGUES(1987). T. etheridgei, T. hispidus, T. hygomi, T. montanus, The first cytogenetic investigations of lizards T. mucujensis, T. oreadicus, and T. torquatus, pre- identified as Tropidurus torquatus were made on sented a 2n = 36 karyotype, including 12 macro- conventionally stained chromosomes ( GORMANet chromosomes (M) and 24 microchromosomes (m), al. 1967; BECAK et al. 1972). Later, cytogenetic in males and females (Fig. 2a and b). The mecha- analyses of two species assigned to the torquatus nism of sex determination was of the XY:XX type, group, using banding techniques, showed some the Y being a minute microchromosome and the X minor chromosome differences as well as distinct a non-identified microchromosome. mechanisms of sex-determination ( KASAHARAet Two other species, T. itambere and T. psam- al. 1983). Additionally, intraspecific geographical monastes, presented the 2n = 36 (12 M + 24 m) variation was revealed for some species as different karyotype only in females. Males had 2n = 35 populations were karyotyped ( KASAHARAet al. (12 M + 23 m), due to a centric fusion between the 1987a; KASAHARAet al. 1988). These preliminary Y and an autosomal microchromosome (Fig. 2c). results prompted us to characterize the whole The submetacentric chromosome derivative of this torquatus group, for a more accurate comparative rearrangement was not always recognized, but chromosome analysis. the presence of a trivalent in diplotene and metaphase I cells confirmed unequivocally the oc- currence of the X, X,Y:X, X, X,X, mechanism of Material and methods sex determination. The specimens of T. hispidus from 7 localities A sample of 173 specimens representing eleven (Slo Luis, MA, Natal, RN, Jolo Pessoa, PB, species of the torquatus group was obtained from Alagoado, BA, Raso da Catarina, BA, Nova different localities in Brazil (Table 1; Fig. 1). T. Soure, BA, and Santo Amaro das Brotas, SE) insulanus and T. bogerti were not available for exhibited a very prominent secondary constriction cytogenetic analysis because their very restricted at the distal region of the long arm of the distribution did not allow collection of live speci- macrochromosome pair 2, which corresponded to mens. All the specimens were deposited in the a nucleolar organizer region. This cytotype was Museu de Zoologia, Universidade de Slo Paulo referred to as Karyotype A in KASAHARAet al. (MZUSP). Part of the present sample was referred (1987a). The specimens from Santo Inacio, BA, previously in KASAHARAet al. (1983, 1987a, Morro do Chap&, BA, and Mucuj;, BA, pre- 1988), RODRIGUESet al. (1988), YONENAGA-YAS-sented Karyotype B, characterized by a slight sec- SUDA et al. (1988), and PELLEGRINO et al. (1994). ondary constriction in the long arm of pair 2, while Chromosome spreads were obtained from bone those from Jacobina, BA, Grlo Mogol, MG, and marrow, liver, and spleen preparations ( KASA- Diamantina, MG, had Karyotype C, with no sec- HARA et a1 1987a) or from fibroblast cultures ondary constriction in this pair. In both kary- (YONENAGA-YASSUDAet al. 1988). Testes chro- otypes, pair 2 is not the nucleolar organizer. mosome preparations were also made. Some specimens were submitted to in vivo or in vitro NOR staining treatment with 5-bromodeoxyuridine ( BrdU) and 5-fluorodeoxyuridine ( FudR). The species of the torquatus group had, in general, Chromosomes were studied after Giemsa stain- two NORs which were located in the proximal ing, C-banding (SUMNER1972), and NOR stain- centromeric region of an acrocentric pair of mi- Hcrediras 125 (1996) CHROMOSOME BANDINGS IN TROPIDURUS LIZARDS 39

21 I

Fig. 1. Collection localities of Tropidurus torquutus species group in Brazil. 1. SBo Luis (Olho D’Agua), MA; 2. Natal, RN; 3. JoBo Pessoa, PB; 4. Alagoado, BA; 5. Raso da Catarina, BA; 6. Nova Soure, BA; 7. Santo Amaro das Brotas, SE; 8. Arraial do Paulista, BA, Ibiraba, BA; 9. Jacobina (Serra do Ouro), BA; 10. Santo Inacio, BA; 1I. Morro do Chapeu, BA, 12. Mucuje, BA, 13. Guaibim, BA; 14. Pic0 das Almas, BA; Rio das Contas, BA; 15. Reserva Biologica de Aguas Emendadas, N of Brasilia, DF; 16. Buritis, MG; 17. Arinos, MG; 18. Unai, MG; 19. Mocambinho, MG; 20. GrBo Mogol, MG; 21. Pedra Menina, MG; 22. Diamantina, MG; Gouveia, MG; SBo Jo50 da Chapada, MG; 23. Serra do Cip6, MG; 24. Po$os de Caldas, MG; 25. SBo JoBo da Boa Vista, SP; 26. Rio Claro, SP; 27. Piracicaba, SP; 28. Joanopolis, SP; 29. Sorocaba, SP. crochromosomes (Fig. 3a). Nevertheless, some Aguas Emendadas, North of Brasilia, DF, showed variations in the NOR patterns were observed. All the usual single pair of acrocentric microchromo- male specimens of T. itumbere and T. psam- somes bearing NORs, those from Piracicaba, monastes had a single NOR metaphase (Fig. 3b) SP, had two pairs (Fig. 3d and e), and the but its size is twice as large as each of the double sole specimen from Pedra Menina, MG, had. only NORs of the females. T. hispidus with Karyotype one microchromosome with active NOR in all A had active NORs at the distal region of the long metaphases. arm of the macrochromosome pair 2 (Fig. 3~). Intrapopulational variation was observed in T. Interpopulational variability was observed in T. hygomi, with the majority of the specimens show- torquutus. While the specimens from Guaibim, BA, ing two NORs, while the single female of the Buritis, MG, Unai, MG, and Reserva Biologica de sample has only one active NOR per metaphase. 40 s. KASAHARA ET AL. Hereditas I25 (1996)

Table 1. Collection localities, sample size, sex, and C-banding pattern of the specimens of Tropidurus torquarus species group

Species Sample size and sex Localities C-banding pattern Male Female

7.coeorobensis 4 2 Raso da Catarina, BA

7.erythrocephalus 1 1 Santo Inacio, BA II 3 Morro do Chap& BA

7.etheridgei 3 6 Arinos, MG 1 Mocambinho, MG

7.hispidus I SHo Luis (Olho DAgua), MA' 1 Natal, RN 2 JoHo Pessoa, PBb 3 Alagoado, BA 3 Raso da Catarina, BAb 1 I Nova Soure, BA 3 Santo Amaro das Brotas, SEb 3 Santo Inacio, BAb Morro do Chap&, BAb Mucuji, BAb 1 Jacobina (Serra do Ouro), BA' 5 3 GrHo Mogol, MG' 1 Diamantina, MG

T hygomi 4 1 Santo Amaro das Brotas, SE 7'. itambere 2 1 Poqos de Caldas, MG 2 SHo JoHo da Boa Vista, SP 2 Rio Claro, SPa 1 Joanopolis, SP 3 3 Sorocaba, SPa T. monranus 1 1 Pic0 das Almas, BAC I Rio das Contas, BAe 1 Buritis, MG 2 3 GrHo Mogol, MG I Diamantina, MG I 1 Gouveia, MG I 1 SHo JoHo da Chapada, MG 4 2 Sena do Cipb, MG T. mucujensis 4 3 Mucuj;, BA' T. oreadicus 2 I Buritis, MG T. psammonasles 1 Arraial do Paulista, BA I 6 Ibiraba, BA 7'. torqualus 4 2 Guaibim, BA I 2 Reserva Biologics de Aguas Emendadas, N of Brasilia, DF 3 4 Buritis, MG 4 6 Unai, MG 1 Pedra Menina, MG 2 2 Piracicaba, SP Total 99 74

See also: a KASAHARAet al. (1983). KASAHARAet al. (1987a). ' PELLEGRINOet al. (1994)

Fig. 2a-c. Chromosomes after conventional staining. a Karyotype (2n = 36) of Tropidurus oreadicus male with XY sex chromosomes. b Microchromosomes of Tropidurus oreadicus female with XX sex chromosomes. c Karyotype (2n = 35) of Tropidurus itambere male with X, X,Y sex chromosomes. Heredifas I25 (1996) CHROMOSOME BANDINGS rn TROPIDURUS LIZARDS 41 42 s. KASAHARA ET AL Hercdiros 125 (1996)

Fig. 3a-e. Partial metaphases after NOR staining. a NORs of different sizes in a pair of microchromosomes (arrows) of Tropidurus torquaius from Unai, MG. b NOR in one microchromosome (arrow) of Tropidurus itambere male. c NORs in the macrochromosome pair 2 (arrows) of Tropidurus hispidus with Karyotype A. d and e Three and four microchromosomes (arrows) with NORs in Tropidurus torquatus from Piracicaba, SP.

C-banding had C-positive bands, but two or three other microchromosomes in the karyotype occasionally C-banding obtained in all eleven species from the showed faint centromeric C-bands. A variant pat- majority of the collection localities revealed two tern was shown by the specimen of T. torquatus main patterns. One of them (type I) was character- from Pedra Menina, MG, whose metaphases always ized by positive bands in the centromeric region of a variable number of microchromosomes, between exhibited four microchromosomes with remarkable 10 and 14 (Table 1; Fig. 4a and b). In some centromeric C-band (Fig. 4c). populations of T. hispidus, as those from Jolo A C-band polymorphism was found in T. itam- Pessoa, PB, Santo Amaro das Brotas, SE, Santo here. Two specimens, one from Slo Jolo da Boa Inacio, BA, Mucuj2, BA, Grlo Mogol, MG, and Vista, SP, and the other from Joanopolis, SP, had Diamantina, MG, these bands appeared as very one almost completely heterochromatic microchro- conspicuous blocks so that the two classes of mi- mosome in the karyotype (Fig. 4d). One male crochromosomes, with and without C-positive from Po~osde Caldas, MG, was a homozygote bands, were clearly identified (Fig. 4a). The speci- having two of these chromosomes, but the second mens of T. torquatus from Guaibim, BA, had an specimen from Slo Joiio da Boa Vista, SP, and additional C-positive segment at the distal end of two specimens from Sorocaba, SP, did not show the microchromosomes bearing the NORs (Fig. this variant heterochromatic microchromosome at 4b). For this reason, the size of these chromosomes all. was considerably larger than that of the other The macrochromosomes had less definite C- microchromosomes. banding pattern, characterized by slight cen- The alternative C-banding pattern (type 11) was tromeric bands, not always observed in all characterized by a reduced amount of constitutive karyotypes. On the other hand, in T. hispidus and heterochromatin in the microchromosome set sporadically in T. torquatus and T. hygomi, heavy (Table 1; Fig. 4c and d). Only those bearing NORs positive blocks were seen in the centromeric region Heredifns 125 (1996) CHROMOSOME BANDINGS IN TRUPIDURUS LIZARDS 43

Fig. 4a-d. Metaphases with type I (a and b) and type I1 (c and d) C-banding patterns. a Tropidurus hispidus from Jo50 Pessoa, PB. with centromeric C-bands in 13 microchromosomes. Observe C-positive staining bordering the secondary constriction in pair 2. b Tropidurus torquutus from Guaibim, BA, with additional C-positive segment in a pair of microchromosomes (arrows). c Tropidurus rorquatus from Pedra Menina, MG, with centromeric C-bands in four microchromosomes (arrows). d Tropidurus iturnbere from SLo Jo20 da Boa Vista, SP, with centromeric C-bands in one microchromosome (arrow) and one almost completely heterochromatic microchromosome (arrow). of the 5th and, less frequently, of the 2nd pair of exception of T. cucorobensis, allowing the ho- macrochromosomes. The most generalized C- mologous pairing of the macrochromosomes banding pattern in the macrochromosome set of (Fig. 5). In prometaphases some microchromo- the torquutus group was the positive staining of the somes also showed replication bands but their distal region of the long arm of the 2nd pair (Fig. precise identification was not possible due to their 4a and c), although not observed in all C-banded small size. metaphases.

R-banding Discussion Clear replication R-banding patterns were ob- Although all the eleven species of the toryuatus tained in the species of the torquutus group, with group shared the same basic karyotype, a certain 44 S. KASAHARA ET AL. Heredifus 125 (1996)

Fig. 5. R-banding patterns after BrdU incorporation in the macrochromosomes of Tropidurus torquutus male. Observe the pair 2 with R-negative staining in the distal region of the long arms. amount of chromosome variability occurred at in- type I C-banding pattern and a pair of large mi- tra and interspecific levels. crochromosomes, not observed in the karyotypes In the present study, the geographical kary- with the type I1 C-banding pattern of the other five otypic variation in T. hispidus was confirmed populations. According to RODRIGUES( 1987), through the analysis of specimens collected in new who examined characteristics of the external mor- localities, totalling 13 different populations kary- phology of T. torquatus from several localities of otyped up to now. No correlation could be estab- its wide distribution area, there are evidences of lished between each of the three different geographical and ecological differentiation when cytotypes and morphological traits of the animals coastal and inland populations are compared. but, as advanced in KASAHARAet al. (1987a), Considering that Guaibim, BA, is a locality in the some association with habitat is still possible: coast while the other five are far from the coast, while the Karyotype A is characteristic of sandy our data are suggestive that conspicuous kary- habitats on the plains, the Karyotypes B and C otypic differences also distinguish the population are limited to rocky habitats in mountainous ar- of T. torquatus. This fact indicates that cytogenetic eas, near or above 900 meters of altitude. These analyses, if extended to a greater number of popu- data on the geographical distribution of the dis- lations, might elucidate an early hypothesis of tinct cytotypes are consistent with the hypothesis RODRIGUES(1987) that the coastal and inland that more than one species might, in fact, be populations of T. torquatus would be indeed two included under the name hispidus. different species. Tropidurus torquatus also showed geographical The C-banding patterns of T. montanus from differentiation of the karyotypes. The most re- five localities were of type I or of type 11. Although markable discrepant karyotype was that found in more cytogenetic data are necessary to characterize the sample from Guaibim, BA, which exhibited this variability, it is interesting to note that there Hereditas 125 (1996) CHROMOSOME BANDINGS IN TROPIDURUS LIZARDS 45 are evidences that T. montanus shows geographical resolution level. Additionally, there are strong variations in morphological traits. banding homoeologies between the macrochromo- At present, we have no means to infer a phyloge- some sets of the torquatus group and of other netical differentiation pattern of the species of the species of Tropidurus, previously karyotyped by us torquatus group based on the NORs or C-banding, (KASAHARA et al. 1986, 1987b; RODRIGLIESet al. but these data may be useful in further compara- 1989). tive analysis with other tropidurid lizards because The present comparative cytogenetic investiga- our previous studies bf some of the species, as T. tion makes an important contribution to the char- semitaeniatus, T. plica, and T. umbra ( KASAHARA acterization of almost the totality of the species of et al. 1986), T. nanuzae, T. amathites, and T. Tropidurus included in the torquatus group. In spite divaricatus (KASAHARAet al. 1987b), T. strobilurus of consistent karyotypic differences, the cytoge- (RODRIGUESet al. 1989), T. spinulosus, and Ura- netic data are not sufficient for elucidating the noscodon superciliosus (PELLEGRINO et al. 1994), phylogenetic relationships of the species. Much suggested consistent differences. more data covering the geographical ranges of the The most conspicuous interspecific difference species are needed for a correct interpretation of within the torquatus group was related to the the chromosome variations. mechanisms of sex determination which clearly distinguished a major group of species with Acknowledgements. -The authors are very grateful to Dr. Tien XY:XX sex chromosomes and another group, in- Hsi Chu for the fibroblast cultures and to Lucila de L. Segalla cluding T. itambere and T. psammonastes, with Franco, Dayse F. de Oliveira Carneiro, Miriam Romeo, and Cristina M. Barnab&for technical assistance. Gabriel Skuk, Jose XIX2Y:Xl XI X2X2. Considering that in previous Manoel Martins, Alessandra Bizerra, Pedro Luis Bernardo da studies RODRIGUES(1987) could not establish any Rocha. and Alexandre Arahjo helped in the field. evidence of relationship between these two species, This work had financial support from FAPESP and CNPq. their multiple sex chromosomes have more proba- bly evolved independently, from XY:XX mecha- References nism. Although three other species of Tropidurus, T. nanuzae, 7’. amathites, and T. divaricatus, in- BECAK,M. L., BECAK,W. and DENARO,L. 1972. Chromosome cluded in the nanuzae group, also presented the polymorphism, geographical variation and karyotypes in Sauria. - Curyologiu 25: 313-326 XIX,Y:Xl X, X2X, sex chromosomes ( KASAHARA CEI,J. M. 1982. A new species of Tropidurus (Sauria, Iguanidae) et al. 1987b), there are no phylogenetic grounds to from the arid chacoan and western regions of Argentina. ~- Occus. Pup. Mus. Nut. Hist. Uniu. Kansas 97: 1 - 10 interpreting this pattern. Furthermore, the nanuzae DIXON,J. R. and WRIGHT,J. W. 1975. A review of the lizards of group did not share the distribution of constitutive the iguanid genus Tropidurus in Peru. - Nut. Hist. Mus. Los Angeles Contrih. Sci. 271: 1-39 heterochromatin and NOR localization with T. DUTRILI.AUX,B. and COUTURIER,J. 1981. La Pratique de itambere and T. psammonastes, but is is worth to I’Analyse Chromosomique. - Masson, Paris mention that there is an interesting geographic FROST,D. R. 1992. Phylogenetic analysis and taxonomy of the

Tropidurus group lizards (1guania:Tropiduridae). ~ Am. Mus. correlation involving these species: T. psam- Nouit. 3033: 1-68 monastes, T. amathites, and T. divaricatus are re- GORMAN,G. C., ATKINS.L. and HOLZINCER,T. 1967. New stricted to the same sand dune fields of S5o karyotypic data on 15 genera of lizards in the Family Iguanidae, with a discussion of taxonomic and cytological

Francisco River, in the State of Bahia, while T. implications. - Cytogenetics 6 286-299 itambere and T. nanuzae occur along the Espinh- GUDYNAS,E. and SKUK,G. 1983. A new species of the iguanid aGo range. Although the nanuzae group is, lizard genus Tropidurus from temperate South-America (Lacer- tilia:Iguanidae).- C. E. D. Orione Cont. Bid. 10: 1-10 according to FROST(1992), the first out-group HOWELL,W. M. and BLACK,D. A. 1980. Controlled silver-stain- for the species of the torquatus group, the phyloge- ing of nucleolus organizer regions with a protective colloidal developer: a I-step method. - Experientiu 36: 1014-1015 netic relationships in the torquatus group remain KASAHARA,S., YON~NAGA-YASSUDA,Y., SCHINCARIOL, R. A. unsolved. and LABBATE,M. 1983. Chromosome mechanisms of sex It is interesting to remark that almost the whole determination, G- and C-band patterns and nucleolus orga-

nizer regions in Tropidurus toryuutus (Sauria, Iguanidae). ~ of the chromosome variation showed by the species Geneticu 60 151 -156 of the torquatus group involved the microchromo- KASAHARA,S., YON~NAGA-YASSUDA,Y., and RODRIGUES,M. somes. In fact, with few exceptions the macrochro- T. 1986. CaracterizaGio cromosshica das especies Tupinurus sernitueniutus, Plicu plicu e PIicu umbra (Sauria, Iguanidae). - mosomes were highly conserved. This conservatism Cienc. Cult. 38: 944 (Proceedings) was confirmed by comparing their R banding pat- KASAHARA,S., YONENAGA-YASSUDA,Y. and RODRIGUES,M. T. 1987a. Geographical karyotypic variations and chromosome terns, which are very similar, if not identical, in all banding patterns in Tropidurus hispidus (Sauria, Iguanidae) ten species studied of the group, even at the high from Brazil. - Curyologiu 40: 43-57 46 s. KASAHARA ET AL. Hereditas 125 (1996)

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RODRIGUES. M. T. 1986. Um novo Tropidurus com crista dorsal the genus Tropidurus (Sauria, Iguanidae). ~ Cyrugmer. Ce// do Brasil, com comentirios sobre was relacdes. distribuislo e Genet. 48: 68-11