Society for the Study of Amphibians and Reptiles Crocodilurus amazonicus Spix, 1825: The Valid Name for Crocodilurus lacertinus Auctorum (nec Daudin, 1802) (Squamata: Teiidae) Author(s): Jean-Christophe de Massary and Marinus S. Hoogmoed Source: Journal of Herpetology, Vol. 35, No. 2 (Jun., 2001), pp. 353-357 Published by: Society for the Study of Amphibians and Reptiles Stable URL: http://www.jstor.org/stable/1566133 Accessed: 10/06/2009 18:35 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=ssar. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit organization founded in 1995 to build trusted digital archives for scholarship. We work with the scholarly community to preserve their work and the materials they rely upon, and to build a common research platform that promotes the discovery and use of these resources. For more information about JSTOR, please contact support@jstor.org. Society for the Study of Amphibians and Reptiles is collaborating with JSTOR to digitize, preserve and extend access to Journal of Herpetology. http://www.jstor.org SHORTER COMMUNICATIONS of predatory fish on amphibian species richness and distribution. Biol. Conserv. 79:123-131. HEYER,W. R., R. W. MCDIARMID, AND D. L. WEIGMANN. 1975. 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D. 1988. Allotopic distribution of two salamanders: effects of fish predation and competitive interactions. Copeia 1988:290-298. SMITH, G. R., J.E. RETTIG,G. G. MITTELBACH,J.L. VALIULIS,ANDS. R. SCHAACK.1999. The effects of fish on assemblages of amphibians in ponds: a field experiment. Freshwater Biol. 41:829-837. WARBURTON, K. 1990. The use of local landmarks by foraging goldfish. Anim. Behav. 40:500-505. WAKE,D. B. 1991. Declining amphibian populations. Science 253:860. WARD, D., AND 0. J. SEXTON. 1981. Anti-predator role of salamander egg membranes. Copeia 1981:724726. ZARET, T. M. 1980. Predation and freshwater communities. Yale Univ. Press, New Haven, CT. Accepted: 24 May 2000. Vol.35, No. 2, pp. 353-357,2001 Journalof Herpetology, Copyright2001 Societyfor the Study of Amphibiansand Reptiles Crocodilurus amazonicus Spix, 1825: The Valid Name for Crocodilurus lacertinus Auctorum (nec Daudin, 1802) (Squamata: Teiidae) JEAN-CHRISTOPHEDE MASSARY1AND MARINUS S. HOOGMOED2 1Museumnational d'Histoire naturelle,Laboratoirede Zoologie (Reptilesand Amphibiens),25 rue Cuvier 75005 Paris, France;E-mail: massary@mnhn.fr 2NationaalNatuurhistorisch Museum (formerlyRijksmuseum van NatuurlijkeHistorie), Departmentof Vertebrates (Reptilesand Amphibians),Postbus 9517 2300 RA Leiden, The Netherlands;E-mail: hoogmoed@nnm.nl Neusticurus bicarinatus (Linnaeus, 1758; Gymnophthalmidae) and Crocodiluruslacertinusauctorum (nec Daudin, 1802; Teiidae) are two sympatric lizard species in eastern Amazonia. Both have semiaquatic habits and live near creeks or damp areas. Moreover, even though C. lacertinus is distinctly larger than N. bicarinatus, they look somewhat similar because they both have a "crocodile-like" tail with a pair of dorsal crests formed by continuous rows of keeled scales. Daudin's (1802) diagnosis of C. lacertinus clearly reveals that he redescribed N. bicarinatusand certainly not C. lacertinus sensu auctorum. In this paper, we provide detailed information supporting this interpretation, explain the origin of this confusion, and the evolution in the use of those two names. The abbreviations used in this paper are MNHN for the Museum national d'Histoire naturelle, Paris, RMNH for the Nationaal Natuurhistorisch Museum, Leiden, UUZM for the Uppsala University, and ZSMH for Zoologische Staatssammlung, Munchen. 354 SHORTER COMMUNICATIONS Linnaeus (1758:201) described Lacertabicarinata,a lizard species now known as N. bicarinatus. Subsequent authors such as Daubenton (1782), La Cepede (1788), Bonnaterre (1789), and Gmelin (1789) always cited this name referring to Linnaeus'sL. bicarinata;the vernacular names "le sillonne" (sometimes spelled "sillone") or "le lezard sillonne" were often used. In addition, their descriptions showed that they all had the same concept of L. bicarinataas Linnaeus. In a systematic revision of the genus Neusticurus,Uzzell (1966: 281) examined the type specimen of L. bicarinata Linnaeus (UUZM 70). He also studied the specimen on which Dumeril and Bibron (1839) based their description of N. bicarinatus(MNHN 4181) and concluded: "The holotype of Lacertabicarinataand the specimen on which Dumeril and Bibron (1839) based the genus Neusticurus (MNHN 4181, probably from French Guiana) are conspecific. The description and figure given by Dumeril and Bibron are excellent. Later workers have applied the name bicarinatacorrectly." We have not personally examined Linnaeus's type specimen. Nevertheless, both of us examined MNHN 4181 so that we have no doubt that our own concept of L. bicarinataagrees with that of Linnaeus (1758). Daudin (1802:85) described a lizard species he named Tupinambislacertinus;he provided both a short Latin diagnosis and a detailed French description; he also used the vernacular name "le Lezardet" or "le tupinambis lezardet" to designate this species. Moreover, Daudin (1802:87) added "Je regarde comme synonyme du tupinambis lezardet, le Lacerta bicarinatade Linnaeus, et le sillonne des naturalistes francais" and thus indicated he was just redescribing and renaming a previously known and officially named species. Now the question is whether Daudin was right to consider his T lacertinus as a synonym of L. bicarinata. In a footnote on page 85, Daudin (1802) gave the following Latin diagnosis of his T. lacertinus: "Tupinambis lacertinus: caudd longiore, compressa bicarinata usque ad basin, squamis dorsalibusquibusdamcarinatis; seriebus longitudinalibusocto scutellorum in abdomine" (long tail, compressed with a continuous double ridge till pelvis, some dorsal scales keeled, abdomen with eight longitudinal rows). This short diagnosis should theoretically be sufficient to preclude confusion between N. bicarinatus(Linnaeus, 1758), which usually has eight longitudinal rows (Hoogmoed, 1973), and Crocodilurus,which has at least 19 (Avila-Pires, 1995); and at least sufficient to know that T lacertinuscannot be identical to Crocodilurus.However, Daudin's description is slightly ambiguous. For instance, concerning the tail, he wrote "compressabicarinatausque ad basin":in this description, Daudin did not mention the fact that the two rows of keeled scales occurring on the tail, do continue on the back. On the contrary, it could be concluded that they would stop at the pelvis, as indeed is the case in Crocodilurus.It is an unfortunately worded detail in the description and in fact, when describing the tail, he apparently just wanted to point out there were two rows of continuous keeled scales along the length of the tail, forming a double ridge. This point of view could be contested, if it were not for the French description which is more detailed, and unambiguously confirms our interpretation. Daudin wrote: "On voit de tres petites ecailles, semblables a celles des vrais lezards, sur le milieu du dos et des flancs; et entre le dos et les flancs, il y a des rangees transversales d'ecailles de moyenne grandeur et un peu carenees. Les deux rangees du milieu forment chacune une carene un peu plus marquee, legerement crenelee en petites dents de scie, et prolongee sur toute la queue". In this phrase, Daudin mentioned some transverse (sic, instead of longitudinal) rows of enlarged, slightly keeled scales between the back and the flanks, and emphasized that two rows of slightly keeled scales in a median position extended along the length of the tail. This character does not occur in Crocodilurus but exactly describes the situation in N. bicarinatus.In addition, some lines further down, Daudin repeated in French that his T lacertinushad eight longitudinal rows of ventral scales; and added that there were 24 pores under each thigh (= 48 in total), which clearly corresponds to a male N. bicarinatus, whereas Crocodilurushas only six to 12 pores in total (Avila-Pires, 1995). Daudin also mentioned small, slightly keeled scales on both sides of the occipital area. This character only applies to N. bicarinatus.Finally, Daudin described the coloration of his species as follows: "Le tupinambis lezardet paroit d'un gris roussatre clair, assez semblable au nankin." The nankin coloration is a sort of reddish-brown to yellow tan. This coloration fits certain specimens of N. bicarinatus in preservative well but disagrees with the dark brown of preserved Crocodilurus.Moreover, Daudin indicated that the ventral coloration was whitish. This condition often occurs in N. bicarinatus,whereas the ventral surface in Crocodilurusis always cream, with irregular, well marked grey and black spots. Most of the scalation and coloration characters given by Daudin do not occur in Crocodilurus.However, all the characters he gave are present in N. bicarinatus.The source of the confusion between Crocodilurusand L. bicarinata can now be considered. Cuvier (1816) seems to be the first to fall in error. He gave (p. 26) a short but erroneous French diagnosis of Daudin's Lezardet: "depourvu d'ecailles larges et carenees sur le dos." Cuvier clearly indicated that the species he was dealing with had no enlarged keeled scales on the back unlike Daudin's specimen. Spix (1825:19) defined a new lizard genus, Crocodilurus;in the same paper, he described two species which he placed in this new genus: C. amazonicus(p. 19 and pl. XXI) and C. ocellatus(p. 20 and pl. XXII, fig. 1), which is just a young C. amazonicus,as Boie (1826:119) suggested. Hoogmoed and Gruber (1983) reviewed the type specimens of the Leiden and Munich Museums. They indicated that Spix's specimen of C. amazonicus still exists (ZSMH 638/0) as do those of C. ocellatus (RMNH 3394 and ZSMH 639/0). Cuvier (1829) erroneously considered C. amazonicusthe same species as Daudin's lezardet and L. bicarinataLinnaeus, 1758. In addition, there is proof confirming Cuvier's wrong interpretation since we found in the Laboratoire d'Anatomie compar6e (MNHN) a skull prepared by Cuvier with a hand written label "le lezardet," which, however, is a skull of Crocodilurus,and not of Daudin's lezardet (= N. bicarinatus).Wagler (1830:153) placed Daudin's lezardet in the genus Crocodilurus,but he no longer considered L. bicarinataa synonym of T. lacertinus and placed it in his new genus Thorictis. One year later, Gray (1831:29), without commenting on Wagler's point of view (it seems he did not yet know SHORTER COMMUNICATIONS Wagler's book), considered L. bicarinata,T lacertinus, and Crocodilurus amazonicus as identical. Dum6ril & Bibron (1839) considered T lacertinus and L. bicarinata two different species. They placed Daudin's Lezardet in the genus Crocodilurusand defined the genus Neusticurus, taking as type species Linnaeus' L. bicarinata (based on MNHN 4181); in addition, they gave a very accurate description that showed they had exactly the same concept of L. bicarinata as Linnaeus. From that moment, C. lacertinus and N. bicarinatusappeared side by side for the first time in the literature. Today, those two names are still commonly used to designate two different species despite the fact that L. bicarinataand T. lacertinus originally designated the same taxon (see Appendix). In short, first Daudin consciously described the same species as Linnaeus's L. bicarinata.Thus T lacertinus Daudin, 1802 has simply to be considered a nomen novum for L. bicarinata Linnaeus, 1758. Third, MNHN 8272, cannot be the type-specimen for T lacertinus as Brygoo (1989:21-22) stated because it is a true Crocodilurus.In addition, because the description of T lacertinus, was actually a redescription of L. bicarinataLinnaeus, 1758 by Daudin (1802) who expressly stated this, the type of L. bicarinata(UUZM 70) automatically also become the type for T. lacertinus Daudin, 1802 according to the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature 1999: Arts. 67.8, 72.7). Nevertheless, we tried to identify the specimen on which Daudin based his description of T lacertinus. In 1839, when describing N. bicarinatus, Dum6ril and Bibron indicated that they had two specimens: the existing MNHN 4181 and an old nearly entirely discolored specimen, probably that of Daudin, which now has to be considered as lost. Finally, C. amazonicusSpix, 1825 is the oldest available name in the literature to designate the taxon previously known as C. lacertinussensu auctorum; it appears to be the correct name to apply to this taxon. We are conscious that the use of C. amazonicus instead of the well-known C. lacertinus is disruptive of stability. Under the plenary powers, the International Commission of Zoological Nomenclature has the authority to set aside the type status of the existing type specimen of a validly described taxon to maintain current and long established usage of a name. In some cases, the instability generated by changing a commonly employed name may warrant such action; however, in the case at hand, we propose that the first available name, viz. C. amazonicus Spix, 1825, be applied to the taxon formerly known as C. lacertinus (Daudin, 1802). Acknowledgments.-The authors are very grateful to all colleagues who reviewed previous drafts of this paper: T. C. S. Avila-Pires, P. David, A. 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In Gray, 1831. A synopsis of the species of the class reptilia [reprint edition], pp. i-xx. Herprint international, Bredell, South Africa. UZZELLJR., T. M. 1966. Teiid lizards of the Genus Neusticurus (Reptilia, Sauria). Bull. Am. Mus. Nat. Hist. 132:277-327. VANZOLINI, P. E. 1972. Miscellaneous notes on the ecology of Brazilian lizards (Sauria). Pap. Avul. Zool. 26:83-115. . 1981. Introduction. The scientific and political contexts of the Bavarian expedition to Brazil. In J. B. von Spix and J. G. Wagler, Herptology of Brazil, pp. ix-xxix. Society for the Study of Amphibians and Reptiles, Athens, OH. VANZOLINI, P. E., AND J. VALENCIA. 1965. The genus Dracaena,with a brief consideration of macroteiid relationships (Sauria, Teiidae). Arq. Zool. 13:7-35. WAGLER, J. [G.] 1830. Naturliches System der Amphibien, mit vorangehender Classification der Saugethiere und Vogel. Ein Beitrag zur vergleichenden Zoologie. J. G. Cotta'schen Buchhandlung, Miinchen, Germany. WIEGMANN, A. F A. 1834. Herpetologia Mexicana, seu decriptio amphibiorum Novae Hispaniae, quae itineribus comitis de Sack, Ferdinandi Deppe et Chr. Guil. Schiede in Museum Zoologicum Berolinense pervenerunt. Pars prima, Saurorum species amplectens. Adiecto systematis saurorum prodro- SHORTER COMMUNICATIONS mo, additisque multis in hunc Amphibiorum ordinem observationibus. Sumptibus C. G. Luderitz, Berolini. Accepted: 20 August 2000. APPENDIX List of synonyms for both Neusticurus bicarinatus and Crocodilurusamazonicus. It includes the Latin binomen and references mentioned in the text plus all references we found where the Latin binomen Crocodilurus lacertinus was used. It is not exhaustive for N. bicarinatus. Neusticurus bicarinatus(Linnaeus, 1758) Lacerta bicarinata Linnaeus, 1758:201 (holotype UUZM 70; Type locality "Indiis" in error); Gmelin, 1789: 1060. LacertaBicarinata;Daubenton, 1782;680, 706; La Cepede, 1788: Synopsis methodica QuadrupedumOviparum, 266; Bonnaterre, 1789:39. Tupinambislacertinus Daudin, 1802:85. Thorictis bicarinatus;Wagler, 1830:153. Teius A.[da] Bicarinata;Gray, 1831:29. Neusticurus bicarinatus;Dumeril and Bibron, 1839:64; Boulenger, 1885:381; Goeldi, 1902:537-548; Cunha, 1961:118; Uzzell, 1966:281; J. A. Peters and DonosoBarros, 1970:206; Hoogmoed, 1973:28, 330; Hoogmoed and Lescure, 1975:160; Hoogmoed, 1979:278; Gasc, 1990:54; Avila-Pires, 1995:421; Jorge da Silva and Sites, 1995:900; Ulber, 1996:iv; Gorzula and Senaris, 1999:157. Neusticurus rudis; Gasc, 1990:55 (part.). Crocodilurusamazonicus Spix, 1825 "Le lezardet" (non Daudin, 1802); Cuvier, 1816:26, 1824:263, 1829:27. Crocodilurus amazonicus Spix, 1825:19, pl. XXI (holotype ZSMH 638/0; type-locality: Sao Paulo de Olivenqas, Rio Solimoes, Brazil); Boie, 1826:119; Fitzinger, 1827:746; Wiegmann, 1834:8; Gray, 1838:278. CrocodilurusocellatusSpix, 1825:20, pl. XXII, fig. 1 (syntypes RMNH 3394 and ZSMH 639/0; type-locality: Sao Paulo de Olivenqas, Rio Solimoes, Brazil). Crocodilurus lacertinus (non Daudin, 1802); Wagler, 1830:153; Peters, 1877:411, 414; Guichenot, 1855:29; Boulenger, 1885:380; Goeldi, 1902:546; Burt and Burt, 1931:326, 1933:60; Hellmich, 1960:81; Cunha, 1961:116; Vanzolini and Valencia, 1965:20; DonosoBarros, 1968:118; Rand and Humphrey, 1968:3; J. A. Peters and Donoso-Barros, 1970:102; Vanzolini, 1972: 105; Hoogmoed, 1973:28; Hoogmoed and Lescure, 1975:157; Muller, 1976:540; Hoogmoed, 1979:278; Vanzolini, 1981:XXI; Hoogmoed and Gruber, 1983: 392; Lescure, 1977:55; 1986:114; Ayala, 1986:571; Gasc, 1990:54, 75; Avila-Pires, 1995:535; Jorge da Silva and Sites, 1995:900; Ulber, 1996:iv; Gorzula and Senaris, 1999:150. "Le Crocodilure lezardet" (non Daudin, 1802); Cocteau, 1835:586. "C.[rocodilure] des Amazones"; Cocteau, 1835:586. Crocodilurus Lacertinus (non Daudin, 1802); Dumeril and Bibron, 1839:46; Gray, 1845:25. Crocodilurus lacertina (non Daudin, 1802); Crump, 1971:20. Crocodilus lacertinus (non Daudin, 1802); Jegu and Keith, 1998:33. 357 Vol.35, No. 2, pp. 357-360,2001 Journalof Herpetology, Copyright2001Societyfor the Study of Amphibiansand Reptiles Barycholos savagei: A Junior Synonym of Paludicola ternetzi, with Notes on Development ULISSES CARAMASCHI1AND JOSE P. POMBAL JR.2 Departamento de Vertebrados, Museu Nacional/UFRJ, Quinta da Boa Vista, 20940-040 Rio de Janeiro,RJ,Brasil IE-mail:ulisses@acd.ufrj.br;2E-mail:pombal@acd.ufrj.br Alipio de Miranda-Ribeiro (1874-1939) was one of the foremost Brazilian naturalists of his era and published extensively on all vertebrate groups. Regarding anurans, several of his papers appeared in a Brazilian farm journal, 0 Campo,which had a very limited circulation. Although most of these papers were reprinted in 1955 (Arq. Mus. Nac. Rio de Janeiro, 42), many published species names have been overlooked, misinterpreted, or not associated with any biological entity, as seen below. Paludicolaternetzi was described by Miranda-Ribeiro (1937) based on a female specimen at the Museu Nacional, Rio de Janeiro (MNRJ 488; Fig. 1), collected at "Vao do Maranhao," State of Goias, Brazil, on March 1923, by Carlos Ternetz. The relatively detailed description, among other characters, refers to an sternal apparatus with double xiphisternum, each half ending in a curved, cartilaginous plate. Bokermann (1966) transferred the species to the genus Physalaemuswithout comment and probably only because the genus Paludicolawas synonymized with Physalaemusby Parker (1927). The combination Physalaemus ternetzi was used again by Lynch (1970, 1971), who associated the species with the Physalaemus cuvieri group. The genus Barycholoswas proposed by Heyer (1969) to accommodate one species, B. pulcher (Boulenger, 1898), known from the Pacific lowlands of Ecuador (Frost, 1985). This genus was considered most closely related to Eleutherodactylusbut characterized by the absence of a groove on the outer circumference of the toe discs (present in Eleutherodactylus)and by having a posteriorly bifid mesosternum with a calcified style, each half bearing a cartilaginous xiphisternum (the mesosternum is a cartilaginous plate in Eleutherodactylus). A second species in the genus, Barycholossavagei, was described by Lynch (1980) based on specimens housed at the Mus6um national d'Histoire naturelle, Paris (MNHNP) and at the National Museum of Natural History, Smithsonian Institution, Washington, DC (USNM). The specimens were collected between the Rio Tapirape and Conceicao (MNHNP 1946-328, holotype; USNM 130184, paratype), and at Bananal and Rio Vermelho (MNHNP 1946-327 plus 327 a-k, USNM 130182-130183, paratypes), all localities in the State of Goias, Brazil. Barycholossavagei and B. pulcher were diagnosed from Adenomera,the fuscus group of Leptodactylus,Lithodytes,and Vanzoliniusby having calcified, stylelike sterna which bifurcate posteriorly and support separate xiphisternal elements. Barycholossavagei differs from B. pulcherby having a tarsal fold instead of a tarsal tubercle, a sharp canthus rostralis