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IGUANAS: BIOLOGY AND CONSERVATIONAuthor(s): Laurie J. VittSource: Copeia, 2004(4):958-960. 2004.Published By: The American Society of Ichthyologists and HerpetologistsDOI: http://dx.doi.org/10.1643/OT-04-249URL: http://www.bioone.org/doi/full/10.1643/OT-04-249

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q 2004 by the American Society of Ichthyologists and Herpetologists

BOOK REVIEWS

Copeia, 2004(4), pp. 952–960q 2004 by the American Society ofIchthyologists and Herpetologists

LIZARDS: A NATURAL HISTORY OF SOMEUNCOMMON CREATURES—EXTRAORDI-NARY CHAMELEONS, IGUANAS, GECKOS,AND MORE. Text by David Badger. Photo-graphs by John Netherton. 2002. VoyageurPress, Stillwater, Minnesota. ISBN 0-89658-520-4. 160 p., 100 color photos. $29.95 (cloth).—This is a strikingly illustrated coffee table bookintended to whet the interests of general read-ers with vibrant color photographs and interest-ing facts about lizard biology. It follows on theheels of two previous Voyageur Press volumes bythe same writer/photographer team, Frogs(1995) and Snakes (1999). Sadly, photographerJohn Netherton died tragically young beforeLizards appeared in print, and therefore, futureherpetological installments in the series may bein doubt. Netherton’s outstanding photographsremain, however, as a legacy of his obvious tal-ent.

Depending on one’s perspective, this book ei-ther succeeds admirably as a general introduc-tion to lizards and is to be justly lauded, or it isso replete with technical errors and inaccuraciesthat it does even this readership a disservice, forwhich it should be criticized. Frankly, my ownfeelings about the book are ambivalent, and Iwill, therefore, give the case for both points ofview.

As a nontechnical book on lizards, the bookis both easy to look at and easy to read. Neth-erton’s photographs are beautifully composedclose-ups of a variety of taxa that certainly drawa reader in, making a persuasive case that liz-ards are beautiful creatures in a purely aestheticsense. I would fault the photos only in that theyare mostly taken of captive animals and, there-fore, lack the biological context of pictures tak-en in the field. One might compare, for exam-ple, the types of photos that adorn this book tothose in the more technical volume on lizardsby Pianka and Vitt (2003) in which most photoswere taken by various investigators in the field.Nonetheless, Netherton’s images are exquisiteexamples of wildlife photography with animalsin naturalistic, if not natural, environments.They are so crisp one wants to run one’s fingersacross the pebbled skin of their subjects!

The design and layout of the book is pleasing.It is large format (8.5 3 11.5 in.), and many of

the photographs are full page, or nearly so. Itis organized into four chapters, starting with ashort introduction to ‘‘lizards and humans,’’ fol-lowed by the meat of the text on ‘‘physical char-acteristics and behavior’’ and accounts of indi-vidual species under ‘‘families and species,’’concluding with the now obligatory chapter on‘‘lizard conservation.’’ Each section of the sec-ond chapter includes a photograph or two anda one- or two-page text description of ‘‘skin andcoloration,’’ ‘‘smell and taste,’’ ‘‘internal anat-omy,’’ or ‘‘communication,’’ to name just a fewof the topics covered. The third chapter pro-vides individual species accounts of one or twopages each for an eclectic mix of taxa, also ac-companied by color photos. There is no appar-ent rationale for the particular species chosen,nor are species accounts arranged according tophylogenetic or ecological affiliation. Choiceswere probably determined by the availability ofphotographs. The somewhat random assort-ment of species is unlikely to trouble generalreaders and also accounts for the rather awk-ward, run-on subtitle of the book.

Badger’s writing is pleasant and breezy. Thetext is unusually easy to read and is full of in-teresting, catchy information about lizards, em-phasizing at all times their remarkable diversityin nearly all respects. Badger uses quotes fre-quently and effectively, and I especially appre-ciate his generous use of attribution, acknowl-edging by name many of his sources—some-thing unusual and welcome in a general text.This probably reflects Badger’s background as aprofessor of journalism at Middle TennesseeState University.

That the author is a journalist and sciencewriter may be at the heart of the problems withLizards from a technical standpoint. The authoradmits in his introduction that he did not havethe natural affinity for lizards that he had forhis previous subjects of frogs and snakes andhad to be convinced that a book on lizards wasa worthwhile project. Although he obviouslyovercame his ambivalence, the text suffers froman overreliance on secondary and even tertiarysources, perhaps reflecting the author’s lack ofexperience with his subjects. Indeed, many ofthe ‘‘authorities’’ cited or quoted in the text arethemselves science writers rather than scientistsor researchers. The greater the distance be-tween source and statement, the greater thelikelihood that inaccuracies will be introduced,and that is the case here. There is a tendency

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to perpetuate myths that now seem to have tak-en on lives of their own: monitors are primarilycarnivores that can expand their gapes likesnakes; the tongue tips are pressed into theopenings to Jacobson’s organs to deliver chem-icals; lizards ‘‘masticate’’ their food, in some cas-es using ‘‘incisors’’ and ‘‘molars’’; snakes canonly detect vibrations and cannot hear air-borne sound. There are many other inaccura-cies that creep into the text, mostly small, butnonetheless annoying to the professional.There is also a fair amount of redundancy thatcould have been avoided by more careful edit-ing. I will not list the other factual errors I not-ed while reading, but I would be remiss if I didnot point out one: the section entitled ‘‘preda-tion’’ is illustrated by a photograph of a birdeating a skink, as would seem appropriate, butthe text is devoted, not to predation on lizards,but to lizard diet. This might be confusing tothe general reader. In a few other cases there isa disjunction between what is described in thetext (e.g., the tail fin of Hydrosaurus) and theaccompanying photograph (a juvenile Hydrosau-rus lacking a tail fin).

Finally, as a professional I felt that some top-ics were overly simplified. For example, in dis-cussing Anolis dewlaps, expanding throats inChlamydosaurus and Pogona, and the ‘‘wings’’ ofDraco, these structures are described as ‘‘flaps’’or ‘‘folds’’ of skin with no mention of their mus-culo-skeletal underpinnings. Surely even the liz-ard novice would be interested in knowing thatDraco’s wings are supported by ribs and thatdewlaps, frills, and beards are erected by move-ments of bones supporting the tongue andthroat? Along these lines, I was disappointed(and often confused) by the lack of scientificnames everywhere except in the species ac-counts. Curiously, even budding, young herpe-tologists tend to learn these names, just as theylearn the scientific names for dinosaurs. Itwould have been helpful and encouraging tothose who are stimulated to go on in herpetol-ogy, to include the Latin binomials at least onceparenthetically for each species mentioned.

In conclusion, Lizards works well as a brief,readable, and aesthetically striking introductionto a glorious group of animals. I can recom-mend it to anyone looking for a book to give toa young person as a first introduction to lizards,or to amateurs interested in the natural worldfrom the vantage point of an armchair. Thebook is not intended for a professional audi-ence, but even for the more serious amateur, Ithink its technical shortcomings would limit itsutility. I would direct anyone with a more seri-ous interest in lizards to two other recent books

on the topic, both of which I happen to havereviewed recently (Schwenk, 2004a,b): EricPianka and Laurie Vitt’s (2003) scholarly reviewof the taxon, and Wade Sherbrooke’s (2003)small book on horned lizards, which includes anice introduction on lizard biology, generally.Both volumes are also well illustrated with colorphotographs.

LITERATURE CITED

PIANKA, E. R., AND L. J. VITT. 2003. Lizards. Windowsto the evolution of diversity. Univ. of CaliforniaPress, Berkeley.

SHERBROOKE, W. C. 2003. Introduction to horned liz-ards of North America. Univ. of California Press,Berkeley.

SCHWENK, K. 2004a. Leapin’ non-ophidian squamates!Review of: lizards. Windows to the evolution of di-versity by Eric R. Pianka and Laurie J. Vitt. TrendsEcol. Evol. 19:357–358.

———. 2004b. Review of Introduction to horned liz-ards of North America by W. C. Sherbrooke. Copeia2004:190–192

KURT SCHWENK, Department of Ecology and Evolu-tionary Biology, University of Connecticut, Storrs,Connecticut 06269–3043; E-mail: kurt.schwenk@uconn.edu.

GAINING GROUND. THE ORIGIN AND EVO-LUTION OF TETRAPODS. Jennifer A. Clack.2002. Indiana University Press, 601 North Mor-ton Street Bloomington, Indiana, 47404; E-mail:iupress@indiana.edu. ISBN 0-253-34054-3. 369p.$49.95 (hardcover).—The evolution of land-dwelling vertebrates (tetrapods) from aquaticancestors was a gradual, stepwise affair: com-plete limbs with digits evolved in organisms thatretained paddle-like tails with dermal fin raysand fully formed branchial arches. Transitionalsarcopterygians (e.g., osteolepids, Eusthenopter-on) retained a fishlike Gestalt with a full arrayof fins but slightly more ‘‘advanced’’ forms (theup-and-coming crocodile-like tetrapods Elpistos-tege and Panderichthys) had compressed heads,expanded ribs, and no median fins. The ap-pearance of the first bona-fide tetrapods (Acan-thostega, Ichthyostega) was finally manifest by theall-important origin of digits. And this entiremakeover, from ‘‘fish’’ to nascent tetrapod, tookplace underwater. As stated by the author, ‘‘. . .becoming terrestrial was . . . a slow process, . . .[which was] achieved with some difficulty’’ (p.2), and hence Gaining Ground is a shrewd title

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for this outstanding book on the origin and evo-lution of basal tetrapods and their incipient ter-restriality.

The author is a leading authority in thesecomplex and often controversial subjects, buther book is mostly accessible even to nonspe-cialists. However, readers should not be fooledby its easy-reading style—Gaining Ground is asubstantial treatise on the morphology of earlytetrapods, their preservation and paleoenviron-ments, and their anatomical modifications to aprogressively more terrestrial lifestyle. But Gain-ing Ground goes much further, touching uponsuch related topics as the developmental path-ways of digit formation, the role of Hox genesin shaping the fish-tetrapod transition, and evenspeculations as to ‘‘why’’ tetrapods may haveevolved in the first place. The elaborate fabricof paleontology, comparative anatomy, embry-ology, and paleoecology woven in GainingGround is particularly useful for those who teachvertebrate evolution; this book should be kepton the shelf at arm’s reach, alongside Janvier’s(1996) masterwork on the evolution of earlyvertebrates.

Clack’s book begins with a general introduc-tion to geological time, paleontology, taphono-my, and phylogenetic systematics, followed by acomparison (in chapter 2, ‘‘Skulls and skeletonsin transition’’) of the skeleton of an advancedfishlike lobe-fin, the Devonian Eusthenopteron,with that of tetrapods in general. Clack’s pur-pose here is to highlight salient morphologicalaspects of the fish-tetrapod conversion (usuallyassociated with locomotion in water and onland, breathing, feeding, and hearing), settingthe stage for more detailed comparisons andanalysis later in the book. The third chapter isa summary of the groups of basal sarcopterygi-ans (living and fossil) and tetrapodomorphs(stem-group tetrapods), paying special attentionto precisely define what is a tetrapod, and alsoincludes a historical survey of tetrapod phylog-eny. The following two chapters outline the or-igin of early tetrapods in their geological andecological contexts, beginning in the Devonian(Frasnian) with more fragmentary tetrapods(Obruchevichthys, Elginerpeton), then focusing onthe remarkable remains of Acanthostega andIchthyostega from the Famennian of East Green-land, and ending in a summary of Famenniantetrapods known from other localities. Threesubsequent chapters characterize tetrapod ra-diations and their environments during the Car-boniferous, when many tetrapod lineages wereestablished and when, so to speak, the‘‘ground’’ was finally ‘‘gained.’’

In ‘‘From fins to feet: transformation and

transition’’ (chapter 6), Clack further comparesskeletal changes in representative taxa of thefish-tetrapod transformation, from Eusthenopter-on to Panderichthys to the most basal ‘‘complete’’tetrapod Acanthostega. These changes are sum-marized into straightforward ‘‘losses’’ and‘‘gains’’ from one stage to another, a rather sim-plistic but useful portrayal whose phylogeneticunderpinning serves as a framework for under-standing the gradual evolution of terrestriality,the main thread of Gaining Ground. Detailed de-scriptions by the late Erik Jarvik of Eusthenopter-on (e.g., Jarvik, 1980) and Ichthyostega (especiallyhis much-awaited monograph; Jarvik, 1996)have aided greatly in this regard, but the rele-vant taxa have been studied afresh by Clack. Shereinforces that both Acanthostega and Ichthyostegawere highly specialized in their own right andvery distinct from each other, and thus neitherembodied some ‘‘ideal’’ tetrapod Bauplan ( Jar-vik held similar views); the orthogenetic inevi-tability of ‘‘becoming’’ terrestrial is consequent-ly avoided. Modifications in sensory biology (vi-sion, olfaction, lateral line) and in the physiol-ogy of breathing, hearing and reproduction arealso discussed under the fish-tetrapod veil, re-vealing that elucidation of terrestriality requiresthe study of fossil and living organisms in tan-dem (Clack here improves upon the lead of-fered by Schmalhausen, 1968).

One of the most useful features of Clack’sbook is the very clear evaluation of key inno-vations, which are mapped separately onto earlytetrapod phylogenies. Characters highlighted inthis fashion include elements of the dermalskull and braincase, lower jaw, inner ear, verte-brae, ribs, limbs, and pelvic girdle, which areoften discussed in terms of their significance forterrestriality. This structure-by-structure ap-proach is an extremely useful tool for visualizingthe morphological changes that shaped the fish-tetrapod transition, which occurred over a pe-riod of ‘‘only’’ some 25 million years accordingto fossils presently in hand (see also reviews inAhlberg and Milner, 1994; Daeschler and Shu-bin, 1995; Ahlberg and Johanson, 1998).

That the evolution of limbs with digits was de-coupled from an established land-dwelling rolewas known to Alfred Sherwood Romer, who wasmuch impressed by a significant gap in the fossilrecord between the origin and terrestrializationof tetrapods, a gap that until recently spannedsome 30 million years from the Late Devonian(Famennian) to Early Carboniferous (Visean;Romer’s original estimate was much greater atsome 65 million years; Romer, 1958). This Tour-naisian Gap (or ‘‘Romer’s Gap’’; Coates andClack, 1995) has been recently abbreviated by

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the description of Pederpes from Scotland, thefirst Tournaisian tetrapod (Clack, 2002). Themorphological void between fishes and tetra-pods really began to be filled in 1931 with thediscovery of Ichthyostega by G. Save-Soderberghduring the early years of the Danish expeditionsto East Greenland, and it now seems that thevoid is slowly but surely disappearing, both mor-phologically and temporally.

Gaining Ground repeatedly drives home thatnot all early tetrapods were terrestrial; in otherwords, tetrapod characters evolved in succes-sion, which is to be expected as more and morekey ‘‘transitional’’ fossils are found (Darwinwould have been overjoyed!). Much of the fine-tuning that has shed light on this situation iscaused by the discovery of new and better fossilsand the reinterpretation of previously describedmaterial. But it is even more an outcome of theemployment of clear and testable phylogeniesto interpret morphological change, and muchof this effort has Jenny Clack herself at the helm(along with colleagues M. Coates, P. Ahlberg,among others). In summing up Gaining Ground,the lucid words of Colin Patterson, written al-most a quarter of a century ago, appear vision-ary: ‘‘I suggest that we might make progresswith the problem of [the evolution] of tetra-pods by taking that to heart [i.e., focusing oncladograms and not evolutionary scenarios],and going back to the problem that Owen andBischoff argued about—what are the charactersof tetrapods? With a comprehensive answer tothat question, we could make better use of thefossil record, looking in it not for ancestors, butfor the sequence in which those tetrapod char-acters arose’’ (Patterson, 1980:172). Impetus forthis approach grew exponentially after the land-mark study of Rosen et al. (1981), and Clack’sbook, with its manifold cladograms and rationalanalysis of ‘‘intermediate’’ fossils, meets Patter-son’s challenge head-on.

Robert Schelly provided insightful discussion.Financial support from the Fundacao de Am-paro a Pesquisa do Estado de Sao Paulo is grate-fully acknowledged (Fapesp proc. 02/06459–0).

LITERATURE CITED

AHLBERG, P. E., AND Z. JOHANSON. 1998. Osteolepi-forms and the ancestry of tetrapods. Nature 395:792–794.

———, AND A. R. MILNER. 1994. The origin and earlydiversification of tetrapods. Ibid. 368:507–514.

CLACK, J. A. 2002. An early tetrapod from ‘‘Romer’sGap.’’ Nature 418:72–76.

COATES, M. I., AND J. A. CLACK. 1995. Romer’s Gap:tetrapod origins and terrestriality, p. 373–388. In:Studies on early vertebrates (7th International Sym-

posium, 1991, Parc de Miguasha, Quebec), M. Ar-senault, H. Lelievre, and P. Janvier (eds.). Bull.Mus. natl. Hist. nat., Paris, 17 (C) 1–4.

DAESCHLER, E. B., AND N. SHUBIN. 1995. Tetrapod or-igins. Paleobiology 21:404–409.

JANVIER, P. 1996. Early vertebrates. Clarendon Press,Oxford.

JARVIK, E. 1980. Basic structure and evolution of ver-tebrates. Vol. 1. Academic Press, London.

———. 1996. The Devonian tetrapod Ichthyostega. Fos-sils Strata 40:1–206.

PATTERSON, C. 1980. Origin of tetrapods: historical in-troduction to the problem, p. 159–175. In: The ter-restrial environment and the origin of land verte-brates, A. L. Panchen (ed.). Systematics AssociationSpecial Vol. 15. Academic Press, London.

ROMER, A. S. 1958. Tetrapod limbs and early tetrapodlife. Evolution 12:365–369.

ROSEN, D. E., P. L. FOREY, B. G. GARDINER, AND C.PATTERSON. 1981. Lungfishes, tetrapods, paleontol-ogy, and plesiomorphy. Bull. Am. Mus. Nat. Hist.167:159–276.

SCHMALHAUSEN, I. I. 1968. The origin of terrestrialvertebrates. Academic Press, London.

MARCELO R. DE CARVALHO, Departamento de Biol-ogia-FFCLRP, Universidade de Sao Paulo, Av. dosBandeirantes 3900, Ribeirao Preto, Sao Paulo,Brazil, 14040-901; E-mail: mrcarvalho@ffclrp.usp.br.

LIZARDS: WINDOWS TO THE EVOLUTIONOF DIVERSITY. Eric R. Pianka and Laurie J.Vitt. 2003. University of California Press, Berke-ley. ISBN 0-520-23401-4. 346 p. $45.00 (cloth).—This is a beautiful book crammed with a wealthof information, not just about lizards, but aboutecology, anatomy, physiology, biogeography,biomechanics, evolution, and ethnobiology aswell. It really was not written for professionalherpetologists but instead for the serious layperson with an interest in nature. Nevertheless,it is fascinating reading for any herpetologist ornatural historian, no matter how professional.Its unabashed flaunting of the incredible diver-sity of size, morphology, coloration, behavior,and ecology exhibited by squamates is fascinat-ing to the point of incredulousness. All of ushave our own idiosyncratic ways to appreciatenature and none of us knows it all, so everyonewho reads this book will learn a lot. Althoughthe book looks and feels like a coffee-table bookabout lizards, it is much more academic and se-rious than that. If not already there, it will soonbe on the bookshelf of every person who wantsto understand more about these scaly creaturesfor whom the authors have such admiration. Itwas also nice to see a book such as this written

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by a pair of ecologists, and those who betweenthem probably have more first-hand experiencewith the ecology and behavior of lizards in moreparts of the world than any other two herpetol-ogists. As such the book is more alive with thediversity of lizards in their natural habitats andwith their myriad of ways to meet their urgentneeds to warm their bodies, evade predators,compete with others for necessary requisites,find mates, and ultimately pass on their genesto subsequent generations. My, what natural se-lection has laid out across the lizard landscape!

The book is logically organized into threeparts. The first introduces the lay reader to phy-logenetic terminology and the phylogeny ofsquamates. Subsequent chapters draw on the di-versity of lizards to conceptually explore the var-ious ways to skin a cat with respect to thermo-regulation and other physiological processes, lo-comotion, feeding ecology, escaping predators,social behavior, reproductive biology, and com-munity ecology (in a chapter mysteriously enti-tled ‘‘Reflections of the Real World’’). The sec-ond part revisits many of these themes, but inchapters organized systematically by families.The last part ties together some major phylo-genetic trends in one chapter and in the lastchapter relates humans and lizards, how we re-gard them and how we impact them (it is not ahappy chapter, for the most part). Also, there isa useful taxonomic summary of lizard genera, aglossary, list of references, and index.

This work is encyclopedic in scope, althoughin some sections this makes for a bit of tiresomereading (e.g., what exemplar species of eachfamily of lizards eat—and eat and eat—but willbe most useful for those seeking specific infor-mation on diets of particular families). To thisend, the detailed index also will be extremelyuseful. The text is richly referenced, so the read-er knows where to go to find the source of spe-cific information, and professional herpetolo-gists will be comforted to see original authorsrecognized. The authors also are willing toshare some of their personal herpetologicalmusings, notions, and ideas, thoughts that theymight not have published elsewhere. In otherwords, the book is not just an annotated bibli-ography.

Obviously for a book of this nature, the pic-tures are critical for visualizing the diversity ofsquamate evolution. In fact, in a real sense thecolor photographs are the very windows to theevolution of lizard diversity, and they well liveup to the chore. The book’s pictures are excel-lent. I found myself eagerly reaching the end ofevery odd page so I could turn the page andfind another beautiful color picture or two or

three, even feeling disappointment if there werenone. That doesn’t happen often as you gothrough this book; there are many, many pic-tures, representing considerable work in findingand selecting photographs and composingthem into a book. The reader will turn the finalpage and set the book back on the coffee table,but the staggering richness of the photo-graphs—those windows to the evolution of liz-ard diversity—will remain fixed in the mind.Many thanks to all those who contributed pho-tographs for this work; the list in the Acknowl-edgments is long.

The text does an excellent job in document-ing the remarkable diversity of lizard evolutionand what factors constrained or accelerated it.The array of traits that cascade together follow-ing some seemingly simple and rather trivialphylogenetic character change, such as theshape, morphology, and sensory capacity of thetongue, is truly astonishing. Yet for every phy-logenetic generalization that Pianka and Vitt tryto make, they have to subdue it a bit with thetwists, turns, and about-faces that are seen inevery clade and that make for the ecologicalequivalents, character displacements, niche par-titionings, and other machinations of local ad-aptation that speak to the diversity of life in liz-ard form. Just when we get the point of under-standing why we see the broad evolutionary ten-dencies in the lizard tree of life, those broadlimbs are blurred by the countless twigs of in-dividual species doing their best to survive andreproduce and doing so in their own ways intheir own places. Ecologically, we see skinks thatare pygopodids, agamids that are chamaeleon-tids, pygopodids that are snakes, cordylids thatare iguanids, geckos that are skinks, gymnoth-almids that are anguids, skinks that are gerrho-saurids, and lacertids that are teiids, to name afew.

Most who will buy and use this book will likelysearch for specific conceptual or taxonomic top-ics. Those who read it cover to cover will findconsiderable repetition, with even the same an-ecdotes and sentences repeated in different sec-tions of the book. I don’t see a lot wrong withthat, because most readers will use the book asa reference, but for some, the repetition will bedistracting. Others may take issue with the sys-tematic organization of the book, such as thehigher levels of taxonomy adopted by the au-thors, but Pianka and Vitt argue that higher lev-el systematics is a bit arbitrary anyway. Mostwon’t be bothered by whatever system is fol-lowed. Still, I found it odd that Chamaeleo wasused instead of Furcifer for a number of cha-meleon species and Cnemidophorus instead of As-

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pidoscelis for the North American whiptails. Justto be picky for a moment, I was bothered withhow the authors used Neotropics as a habitat orbiome contrasted to deserts, instead of as a bio-geographical realm; the repeated misspelling ofDraco’s wing patagia, even in the Glossary; andthe somewhat careless description of differentniches for a species instead of multiple dimen-sions of the single hypervolume niche per spe-cies (peculiar, coming from Eric Pianka). I wasalso surprised at how few maps were used. Itwould have been useful to have a map of Brazil,for example, with the areas of savanna, cerrado,caatinga, etc., depicted, especially because Vittdraws so heavily on the lizards of Brazil, wherehe has worked extensively. But these are minorcriticisms of this impressive book; the book islucid, lively, personal, scientific, informative, ac-curate, and contagious. Ichthyologists, ornithol-ogists, mammalogists, butchers, bakers, and can-dlestick makers will cease to exist after they haveread this book; they will all convert to be her-petologists.

Pianka and Vitt are not reluctant to sharetheir ideas for future research in lizard diversity.To name a few, they suggest a study of the en-dosymbionts of the herbivorous desert iguanacompared to those of sympatric herbivorousmammals, whose feces they eat in the spring;why phrynosomatine display behavior and mor-phology are not linked phylogenetically; the cal-cium carbonate end caps on eggs of Draco; thesailfin of Hydrosaurus; the knobs of knob-tailedgeckos; or the possible social strategy of a sub-ordinate male anole that incessantly badgers adominant territorial male to flash his dewlap somuch that some nearby predator notices himand eats him, thus leaving the vacant territoryfor the clever subordinate. There are manymore.

Finally, it was amusing to see pop up thefriendly rivalry among herpetologists who studylizards versus those who focus on snakes—thosewho sport ‘‘lizard envy.’’ More than once theauthors point out that snakes are just a distalclade of lizards; thus, all the ‘‘mystery’’ ofsnakes is a subset of lizard diversity, poking a bitof fun at Harry Greene (who wrote the Fore-word for the book) and his equally beautifuland informative book, Snakes: The Evolution ofMystery in Nature (1997), which Pianka and Vitt’sbook resembles in many respects. Even the lastsentence of this book, which appears to havebeen written before the rest of the book, re-minds us to ‘‘. . . bear in mind: snakes are liz-ards too.’’

LITERATURE CITED

GREENE, H. W. 1997. Snakes: the evolution of mysteryin nature. Univ. of California Press, Berkeley.

STANLEY F. FOX, Department of Zoology, OklahomaState University, Stillwater, Oklahoma 74078;E-mail: foxstan@okstate.edu.

ESTUARINE RESEARCH, MONITORING,AND RESOURCE PROTECTION. Michael J.Kennish (ed.). 2004. CRC Press, P.O. Box409267, Atlanta, Georgia. ISBN 0-8493-1960-9.297p. $119.95 (hardcover).—Alteration of theestuarine landscape and deterioration of estua-rine environments is doubtlessly going to con-tinue. To place these anthropogenically derivedlandscape changes into a context from whichwe can learn requires quantitative knowledge ofwhat a pristine watershed of an estuarine envi-ronment should look like. The National Estua-rine Research Reserve System (NERR), whichevolved from the National Estuarine SanctuaryManagement Program of 1972, was developedto address just such issues. There is now a net-work of 25 NERR sites across 21 states andPuerto Rico, covering 15 biogeographic re-gions, and almost 500,000 ha of estuarine, wet-land and upland environments. Selected sitesare typically unique environments that are pris-tine and undisturbed such that they can serveas a reference for comparison to other regionsin terms of abiotic factors and aquatic resourc-es. Their main mission is to collect and providestandardized, quantitative information on na-tional estuarine trends of local and regionalconcern that are used to promote informed re-source management. In addition to this bookproviding a detailed history of the NERR Sys-tem, its mission, goals, and priorities, also pro-vides a valuable watershed management con-cept to those that read it. If done correctly, re-evaluated periodically, and with the use of adap-tive management principles, monitoring can bea powerful tool to provide short- and long-termpatterns in important regional datasets that canbe extrapolated and used to effectively managenational environmental issues. Coupled withthe clear vision on the role of Marine ProtectedAreas and Reserves outlined in Bohnsack(2003), the reader should note that this is prob-ably one of the most important take-home mes-sages of the volume.

The book also provides important data trendsin the form of six case studies: Waquoit Bay

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(MA), Jacques Cousteau (NJ), Delaware (DE),Ashepoo-Combahee-Edisto (ACE) Basin (SC),Weeks Bay (AL), and Tijuana Bay (CA) sites.These case studies represent sites from the Atlan-tic, Gulf, and Pacific coasts of the United States,and each provides an up-to-date understandingof the abiotic and biotic components of the wa-tershed. These will be extremely useful for any-one wishing to work in any of these sites. Datafrom the short- and long-term systemwide mon-itoring program (SWMP) have already been usedto address temporal recovery from storm eventsand assisting with estuarine restoration activities.As the author states in the Preface, the book is‘‘principally designed as a reference volume forestuarine and watershed scientists, resource man-agers, decision makers, and other professionalswho deal with coastal zone issues.’’ There is noquestion that the detailed treatment of the en-vironment of the six case study sites provides awealth of quantitative data.

The only errors noted in the volume are theuse of the old names for members of the Penaei-dae (in the Weeks Bay and ACE Basin NERRchapters; see Perez Farfante and Kensley 1997),and that figure 3.3 has the lower portion missing(Jacques Cousteau NERR chapter). Given theserelatively minor errors, the volume is packed fullof valuable data, is scientifically sound, and I feelthe editor has done a great job meeting the statedgoal of the volume as a reference text for a myriadof professionals. The target audience of this vol-ume is anyone interested in effectively managingcoastal environments with their ‘‘eyes’’ on what apristine estuarine watershed should look like. Ihighly recommend the volume for professionalsand even graduate students who will find the datacurrent and timely.

LITERATURE CITED

BOHNSACK, J. A. 2003. Shifting baselines, marine re-serves, and Leopold’s biotic ethic. Gulf Carib. Res.14:1–8.

PEREZ FARFANTE, I., AND B. KENSLEY. 1997. Penaeoidand sergestoid shrimps and prawns of the world:keys and diagnoses for the families and genera. Me-moires du Museum National d’Histoire Naturelle,Tome 175. Paris.

MARK S. PETERSON, Department of Coastal Sciences,University of Southern Mississippi, 703 East BeachDrive, Ocean Springs, Mississippi 39564; E-mail:mark.peterson@usm.edu.

IGUANAS: BIOLOGY AND CONSERVATION.Allison C. Alberts, Ronald J. Carter, William K.Hayes, and Emılia P. Martins (eds.). 2004. Uni-versity of California Press, Berkeley, California.ISBN: 0-0520-23854-0. 356 p. $65.00 (cloth).—Based on a symposium held in 1997, Iguanas:Biology and Conservation presents summaries andupdates of research conducted on a lizard cladecontaining most of the herbivorous lizard spe-cies on Earth, the Iguanidae. Forty-four speciesin 11 genera are known (several are extinct).The interest these lizards have attracted is im-pressive, considering that extant iguanids rep-resent about 1% of living lizards and 0.6% ofall squamates. Large body size, high densities(at least relictually), and their impressive andeasily studied social behavior contribute to thispopularity. However, the rapid decline of manyspecies has brought them to the forefront ofconservation biology, as reflected by the largeportion of this book dedicated to iguanid con-servation. Like many other large vertebrates,just as we are beginning to learn about thesefascinating beasts, their numbers are diminish-ing to the point of no return. Although onecould easily be misled into thinking that theselizards are doing fine based on the apparenthigh densities of Iguana iguana along rivers inecotourist capitals of the New World or Dipso-saurus dorsalis on the remaining sands of themegadevelopment in the Coachella Valley andother desert regions of the southwest UnitedStates, most species inhabit islands, and mostare considered endangered. Even those on themainland (e.g., Iguana and Ctenosaura) are be-ing eaten into oblivion by humans. The assaultby humans on these animals includes defores-tation of habitat; conversion of habitat to hotels,housing, and tourist attractions; introductionand continued maintenance of predatory catsand dogs; and direct predation by humans forfood.

In his introduction, Gordon Burghardt at-tempts to make the case that we know very littleabout iguanas. His wish list of areas needing em-phasis could be applied to nearly all species onEarth! Considering that two edited researchbooks and hundreds of scientific research arti-cles have been devoted to these lizards and thatthey have received so much attention in thepopular press, more may be known about thebiology of these lizards than nearly all others(with the exception of Anolis). Indeed, one ofthe most thorough lizard studies was conductedon a species of Cyclura (Iverson, 1979). If any-thing, considering the level of funding availablefor many of these endangered lizards, one hasto wonder why more studies like Iverson’s have

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not been conducted. Of course, the more weknow, the better, and considering the imperiledstatus of iguanas, a sense of urgency is warrant-ed.

The book contains three distinct sections: Di-versity; Behavior and Ecology; and Conserva-tion. As it should, the Diversity section first laysout what is known about generic relationships,then species relationships within genera, and,finally, a thorough checklist with synonomies ispresented. This is really useful information!The three chapters that follow use moleculardata to examine evolution within island groups.Two major clades of Cyclura occur in the WestIndies, one of which produced a single species,with the other ultimately diversifying into twolarge clades containing 14 species. The highcorrelation between genetic diversity and islandsize (5 population size) provides insight intoproblems developing realistic management pro-grams designed to maximize genetic diversity.The chapter on sodium and potassium secre-tion seems out of place and outdated. The sum-mary of the distribution of salt glands amonglizards will be useful for physiologists, but a phy-logenetic analysis, identifying origins of differ-ent modes of salt excretion, would have beenmore useful.

The section on Behavior and Ecology is anexceptionally interesting and strong section inthe book. For example, we learn that Cycluracarinata present appeasement displays, derivedfrom signature displays, that reduce aggressionby larger individuals. Among islands, head-bobdisplays vary considerably, partially associatedwith differences among islands in vegetativestructure. Males of Iguana iguana performmuch riskier behaviors than do females, and apossible explanation for this behavior has itsroots in kin selection. An ‘‘altruistic’’ male mayplace himself at risk to reduce the probabilitythat clutch mates (siblings), which often remaintogether, are captured. Although observed inthe lab, this is exciting stuff and should be fol-lowed up in the field. Lekking male marineiguanas (Amblyrhynchus) that are large, havelarge heads, and have fewer parasites achievehigher mating success than those that are small-er, even though many other factors also play im-portant roles in mating success. Female repro-ductive traits provide the impetus for efficientfemale choice, which is facilitated by the lekmating system. In one of two chapters that dealwith ecological determinants of body size, welearn that environmental factors, primarily tem-perature and its effect on warming rate in ma-rine iguanas, directly influence foraging effi-ciency, which in turn influences body size. In

the other chapter (12), two desert iguanids(Dipsosaurus and Sauromalus) respond different-ly to the same environmental gradient in termsof body size patterns. An impressive amount ofdata is brought to bear, showing that diets differamong the species but that ‘‘resource availabil-ity’’ is likely similar. Several hypotheses mightaccount for these differences, but differences ingene flow rates may be most important. Saurom-alus live on isolated islands of rock, whereas Dip-sosaurus are largely distributed continuously inflatland desert habitats. No doubt, as the lasthuman disappears from the planet Earth, achuckwalla will peak out of the crevice that keptit out of the reach of humans and wonder whatall that garbage is!

The final seven chapters center on conserva-tion but only on species of Cyclura. As for mostspecies on Earth, habitat loss is the greatestthreat to these lizards. Quite honestly, it seemsremarkable that any of the Cyclura remain.Translocations, headstarting, repatriation, andhabitat restoration are the key words that de-scribe efforts to save these magnificent animals.Zoos have played a pivotal role in maintainingpopulations of Cyclura but, more important, insupporting research and conservation effortson natural populations. Education and appre-ciation of the value of these lizards as part ofthe natural heritage of their islands have thegreatest potential for allowing these lizards tocoexist with a consumer-driven and increasingpopulation of humans. I was a bit disappointedto not find chapters on conservation of desertiguanas, chuckwallas, green iguanas, or cteno-saurs—must we have to wait until their popula-tions approach the point of no return or domajor programs exist that were not included?

Although a few inconsistencies exist and theintroduction by Burghardt seems to ramble withno real point, editing overall is such that thebook comes across as a well-thought out com-pendium. It is unfortunate that books of thisnature often appear years after the symposiumleading to them. Although authors in some cas-es updated their chapters, the impact of thisbook would have been much greater had it ap-peared while the symposium was still fresh inour minds.

I highly recommend this book to anyone in-terested in evolution, behavior, and conser-vation of reptiles or island species in general.For those of us who have spent most of ourcareers chasing around carnivorous lizards,this book provides many new insights into thefour-legged lepidosaurs that have opted forabundant, low-energy, immobile ‘‘prey’’ rath-er than seeking out often hard to find and

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capture, high-energy (but parasite ridden)prey. It seems to me that now is the time fora follow-up book by a single author, that in-tegrates all that is known about these fasci-nating lizards into a cohesive evolutionary sto-ry pointing to the likely demise of a remark-able evolutionary experiment that is beingdisassembled by the actions of humans.

LITERATURE CITED

IVERSON, J. B. 1979. Behavior and ecology of the RockIguana Cyclura carinata. Bull. Fla. State Mus., Biol.Sci. 24:175–358.

LAURIE J. VITT, Sam Noble Oklahoma Museum of Nat-ural History, 2401 Chautauqua Avenue, Norman,Oklahoma 73072–7029; E-mail: vitt@ou.edu.