BOOK REVIEW

Caribbean Journal of Science, Vol. 36, No. 1-2, 171-173, 2000 Copyright 2000 College of Arts and Sciences University of Puerto Rico, Mayaguez

The Iguanid Lizards of Cuba. L. Rodrguez Schettino (ed.). 1999. University Press of Florida, Gainesville, Florida, USA. 488 pp., 58 color plates, 21 line drawings, 66 distribution maps. ISBN 0-8130-1647-9. $85.00 (cloth). This most impressive reference catches the eye immediatelybut the dust jacket, profusely decorated with brightly colored illustrations of Cuban lizards, is only the beginning. Inside, the reader discovers that Lourdes Rodrguez Schettino has assembled not only the expected chapters on systematics, morphology, ecology, behavior, and biogeography, but also a great deal of detailed information on parasitology and genetics, which are topics rarely addressed in faunal guides. In her introduction (Chapter 1), Rodrguez Schettino addresses the importance of iguanian lizards in Antillean faunas and then provides a brief history of iguanian taxonomy. Coverage of the latter topic will attract some criticism because she and the other contributors to this volume chose to retain the family Iguanidae sensu lato instead of the seven families proposed by Frost and Etheridge (1989). Given the current state of the dispute, and some remaining unresolved relationships, that decision is conservative and easily justified. Some readers, however, might quibble with the reasons cited. The first cited objection to the recognition of seven iguanian families is that of Lazell (1992), but more relevant are the objections by Macey et al. (1997), who demonstrated the monophyly of iguanians and recommended reversion to the traditional system. I cannot criticize the choice made by Rodrguez Schettino and her colleagues, although I wish that she had used iguanian rather than iguanid in the title of this volume. Either is correct but the former is less confrontational toward those who hold different opinions and note that taxa above that of the family should also be monophyletic. In contrast to the decision to lump the iguanian families, Rodrguez Schettino retained the generic dis tinction of Chamaeleolis and Anolis. Justification for this choice is a desire for more evidence to support the synonymy of these genera, despite the impressive data set presented by Hass et al. (1993). More recently, Jackman et al. (1999) and Poe (1999) strongly supported the inclusion of Chamaeleolis within Anolis. Since the latter contributions were not available when this book was written, the decision to retain Chamaeleolis can be defended as the conservative alternative in a situation that had not been resolved conclusively. Still, the failure to address at this time the use of the anoline genera proposed by Guyer and Savage (1986) leaves the reader wondering if the preferences of Rodrguez Schettino and her colleagues are arbitrary and reflect solely a reluctance to give up a familiar taxonomy. After quick discussions of the distributions of iguanian lizards and descriptions of Cuban members of 171

the family Iguanidae sensu lato, Rodrguez Schettino addresses in a section on classification some of the issues raised above (including the anoline genera proposed by Guyer and Savage, 1986). Although it was not clear why taxonomy and classification are treated separately, or why some of the choices were made, by the time this section ended with a list of Cuban iguanians, I had accepted the taxonomic treatment chosen and was ready to move on. The introduction continues with a brief survey of the relevant fossil record and a very nice but exceedingly short history of studies on Cuban lizards. Inexplicably included with the latter is a commentary on legends, beliefs, and traditions associated with Cuban lizards. This discussion certainly deserved its own section. The last topic in this chapter deals with conservation and concludes with a listing of species presumed extinct, critically endangered, endangered, vulnerable, or at lower risk. Although these categories are adequately defined and the lack of accurate data in many cases is noted, I would have liked to see more information on this subject. Notable by their absence are discussions of the roles in conservation of various Cuban agencies and institutions, current efforts to conserve these lizards (and other wildlife), and projections of what should be done and probably will be done. Morphology is the subject of the second chapter, also by Rodrguez Schettino. Here she relates external features to function and uses excellent line drawings by Lazaro Gonzalez Pino to illustrate relevant fea tures. Treatment of internal anatomy is rather superficial. Sections introducing morphological variation and ecomorphology are excellent. Nevertheless, I longed for more detail on these topics, particularly the latter, which is so inextricably associated with West Indian anoles. Chapter 2 concludes with a section on identification and dichotomous keys to genera and species of Cuban iguanians. I had only a few Cuban specimens with which to test the keys, but my efforts to identify them and the subjects of photographs I had taken during a visit to the U.S. Naval Base at Guantanamo were re warded. A few couplets are confusing, but a quick review of the section on external morphology or the individual species accounts addresses the problems. I would have liked to see some drawings with the keys or at least a closer placement of the illustrations and the keys. Chapter 3, on ecology and behavior, was coauthored by Rodrguez Schettino, Mercedes Martnez Reyes, and Luis V. Moreno Garca. It has to be one of the most informative chapters in the book. The inclusion of many references to work on Cuban species is of universal utility and the many citations speak to the diligence of the Cuban herpetological community. Regretfully, they also serve as reminders of the difficulty of acquiring many of these very relevant papers outside of Cuba. Many of the cited references undoubtedly will become more available solely as a consequence of the publication of this volume. The first section of this chapter is devoted to spatial distribution, which includes tables associating species

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BOOK REVIEW excluding the glossary, literature cited, and index). An introduction provides criteria for descriptive terms used in the accounts and maps of Cuba show provincial borders and many cities. These are essential for visualizing species distributions but are often and inexplicably omitted from works of this type. Accounts for genera include a synonymy (with types indicated), content (number of species, total and in Cuba), and a brief description. The account for Leiocephalus is accompanied by a line drawing illustrating line patterns and the account for Cyclura includes a lateral view indicating important features of scalation. Those for Chamaeleolis and Anolis do not include line drawings, although general features are illustrated in the earlier chapter on morphology. Species accounts are arranged by group and by date of description within groups. This complicates substantially any effort to find a given account without resorting to the index and I would have preferred a more utilitarian alphabetical arrangement. Each account provides vernacular names in Spanish and English, a synonymy (with types indicated), a notation of geographic range, a specific list of localities from which specimens are known, content (currently recognized subspecies in polytypic forms), a description, a list of bibliographic sources organized by topic, notes on morphological variation (sometimes divided into geographic, sexual, and age variation), and a summary of natural history. The quantity of information varies widely depending on how much is known about each species. As a test, I examined the literature about Anolis porcatus, a species for which I have made a concerted effort to accumulate references. The list of citations is amazingly complete, although the description of a subspecies was omitted (Perez-Beato, 1997). Range maps accompany each account and most species are illustrated in plates. Plates are of varying sizes and often correspond to the actual size of the species illustrated. Most plates figure adult males (50 species), but females are figured for 4 species, as are a few juveniles (3 species). Paintings by Lazaro Gonzalez Pino are used instead of pho tographs. Some are striking. I was particularly taken by the adult males of Cyclura nubila nubila (plate 7) and Anolis rubribarbus (plate 55), both of which also appear on the dust jacket. Paintings can be of great utility, often illustrating diagnostic characters more effectively than photographs (e.g., Conant and Collins, 1998), but they can be mere portraits if care is not taken to illustrate important features. Many of the plates included in this volume fall into the latter category. This would not be a problem if line drawings, inserts, or additional paintings were included. Unless one is attempting to identify an adult male, with dewlap extended (in the case of anoles), the plates will be of marginal help. Complicating the problem of illustrating Cuban species is the fact that many are polytypic and considerable variation exists. This is more than adequately addressed in the text but it further reduces the utility of the plates beyond their considerable decorative value. An adequate glossary, a bibliography of more than 350 sources, and an exceedingly useful index complete the volume. The value of the bibliography cannot be overestimated because the inclusion of many Cuban

with vegetation types (which are mapped in color as Plate 58), substrates (largely ecomorphological associations), and altitude. The discussion of ecomorphs provides a somewhat different perspective (Cuban based?), but it does not represent the most widely accepted views and the most recent work on the subject (e.g., Losos et al., 1998). I am not aware of anyone else still using indices of locomotion. Equally startling is the lack of any discussion on the use of principal components analysis of morphological attributes to define ecomorphs. Subsequent sections address climatic relationships, activity, trophic relationships, reproduction, and community structure. Although I wanted more, the authors are to be applauded for including so much information in so few pages. Genetics is the subject of Chapter 4, written by Georgina Espinosa Lopez and Ada R. Chamizo Lara. A brief survey of relevant genetic studies is followed by more detailed discussions relating genetic variability to aspects of natural history and examining the utility of extant data in systematic studies. These discussions are organized by groups, first anoles and then the genus Leiocephalus. A short concluding section on karyology includes some references to Cyclura. Inclusion of this topic in a faunal guide is laudable, if only because the subject will be increasingly important as more information and new applications come to light. Chapter 5, on parasites, by Alberto Coy Otero, addresses another topic rarely included in faunal guides. However, it is a topic about which more is known on Cuban species than on any other West Indian fauna. The abundance of information is largely attributable to the author, who with various collaborators has made the study of reptilian parasites a major focus of his research career. Most of the information is presented in a list of parasites of Cuban iguanians and a table of parasites, their geographic distribution, and their Cuban hosts. The next chapter, by the editor, deals with historical and ecological biogeography of the Caribbean region. Rodrguez Schettino nicely summarizes the abundant literature on historical biogeography and also deals nicely with the various interpretations for existing faunas. She concludes that definitive evidence supporting land bridges (e.g., MacPhee and Iturralde-Vinent, 1994), vicariance (e.g., Crother and Guyer, 1996), or dispersal (e.g., Hedges, 1996) is lacking. However, she does not evade the question, noting rightly that a combination of factors may be responsible, and explaining why in most instances the dispersal hypothesis is widely accepted as the most plausible explanation. A section addressing the zoogeography of Cuban iguanians, which includes a table plotting their distribution by biogeographic region, precedes the final section on ecological biogeography. This treatment is too brief, but nevertheless categorizes lizard assemblages by habitat. She concludes that climatic factors and vegetative structure are the most important variables. In contrast, trophic requirements appear inconsequential, as no evidence links any Cuban species with particular prey. The concluding chapter, also by Rodrguez Schet tino, provides systematic accounts of species, and constitutes the bulk of the book (277 of 380 pages of text,

BOOK REVIEW publications not in general circulation is a tremendous resource. Like any other text dealing with organisms being actively studied, a time lag between completion and publication can quickly render at least taxonomic considerations obsolete. For example, Powell and Hendersons (1999) addenda to the checklist of West Indian amphibians and reptiles includes two Cuban iguanians that do not appear in this book, Anolis (=Chamaeleolis) agueroi (Daz et al., 1998) and Anolis incredulus (Garrido and Moreno, 1998). Additional new taxa and taxonomic changes are in press. The book has the high quality I have come to expect from the University Press of Florida, which also published the comprehensive volume on West Indian amphibians and reptiles by Schwartz and Henderson (1991). The price is also reasonable for a volume such as this. Typographical errors are almost non-existent (I found two) and, although the translation by Janet Perodn Hernandez is on rare occasions a bit stilted, the text is clear and readable. The plates are nicely executed and the line drawings are clear and detailed. I was impressed by the fact that maps are of an appropriate size, although locality dots are too tiny. In summary, I really like this book and wish that others like it were available for the remaining West Indian islands. My criticisms are minor and mostly deal with problems that could have been avoided only by delaying publication. These problems are more than offset by the evident strengths of a volume that brings together so much information. Certainly, herpetologists working in the region must have a copy on their shelves, but other biologists, hobbyists, and many non-specialists with an interest in natural history will want to own this book as well. Acknowledgment.Richard E. Glor made several exceedingly helpful comments on an earlier draft of this review.

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LITERATURE CITED
Conant, R., and J. T. Collins. 1998. A Field Guide to Reptiles & Amphibians, Eastern and Central North America. 3rd ed., expanded. Houghton Mifflin Co., Boston and New York. Crother, B. I., and C. Guyer. 1996. Caribbean historical biogeography: was the dispersal-vicariance debate eliminated by an extraterrestrial bolide? Herpetologica 52:440465. Daz, L. M., N. Navarro, and O. H. Garrido. 1998. Nueva especie de Chamaeleolis (Sauria: Iguanidae) de la Meseta de Cabo Cruz, Granma, Cuba. A new species of Chamaeleolis (Sauria: Iguanidae) from Meseta de Cabo Cruz, Granma, Cuba. Avicennia 8/9:2734.

Frost, D. R., and R. Etheridge. 1989. Phylogenetic analysis and taxonomy of iguanian lizards (Reptilia: Squamata). Univ. Kansas Mus. Nat. Hist. Misc. Publ. (81):165. Garrido, O. H. and L. V. Moreno. 1998. Nueva especie de Anolis (Lacertilia: Iguanidae) del Pico Turquino, Sierra Maestra, Cuba. A new species of Anolis (Lacertilia: Iguanidae) from Pico Turquino, Sierra Maestra, Cuba. Avicennia 8/9:3540. Guyer, C., and J. M. Savage. 1986. Cladistic relationships among anoles (Sauria: Iguanidae). Syst. Zool. 35:509531. Hass, C. A., S. B. Hedges, and L. R. Maxson. 1993. Molecular insights into the relationships and biogeography of West Indian anoline lizards. Biochem. Syst. Ecol. 21:97114. Hedges, S. B. 1996. The origin of West Indian amphibians and reptiles. In R. Powell and R. W. Henderson (eds.), Contributions to West Indian Herpetology: A Tribute to Albert Schwartz, pp. 95128. Soc. Study Amphib. Rept. Contrib. Herpetol., vol. 12, Ithaca, New York. Jackman, T. R., A. Larson, K. de Queiroz, and J. B. Losos. 1999. Phylogenetic relationships and tempo of early diversification in Anolis lizards. Syst. Biol. 48:254285. Lazell, J. D., Jr. 1992. The family Iguanidae: disagreement with Frost and Etheridge (1989). Herpetol. Rev. 23:109112. Losos, J. B., T. R. Jackman, A. Larson, K. de Queiroz, and L. Rodrguez Schettino. 1998. Contingency and determinism in replicated adaptive radiations of island lizards. Science 279:21152118. MacPhee, R. D. E., and M. A. Iturralde-Vinent. 1994. First Tertiary land mammal from Greater Antilles: an Early Miocene sloth (Xenarthra, Megalonychidae) from Cuba. Amer. Mus. Novit. (3094):113. Perez-Beato, O. 1997 (dated 19961997). A new sub species of Anolis porcatus (Sauria: Polychrotidae) from western Cuba. Rev. Biol. Trop. 44(3)/45(1): 295299. Poe, S. 1999. In J. B. Losos and M. Leal (eds.), Anolis Newsletter V. Washington Univ., St. Louis, Missouri. Powell, R., and R. W. Henderson. 1999. Addenda to the checklist of West Indian amphibians and reptiles. Herpetol. Rev. 30:137139. Schwartz, A., and R. W. Henderson. 1991. Amphibians and Reptiles of the West Indies: Descriptions, Distributions, and Natural History. University of Florida Press, Gainesville. ROBERT POWELL, Department of Biology, Avila College, Kansas City, Missouri 64145-1698, USA