Varanoid Lizards of the World, edited by Eric R. Pianka and lications focused on the husbandry of these forms. This new book, Dennis King with Ruth Allen King. 2004. Indiana Univ. Press. edited by Pianka, King, and King, which purports to be a “com- prehensive account of virtually everything important that is known about monitor lizards and their allies,” is divided into three major sections. The first is focused largely on phylogeny and biogeogra- phy, the second and longest is committed to species by species descriptions, and the third discusses the evolution of body size and its importance to reproduction and also examines captive care. In their introduction Eric Pianka and Dennis King briefly ex- plain that their goal was to compile a single volume reference work about monitor lizards and their allies that touches not only on the systematics of individual species, but also examines their collec- tive biogeography, paleontology, and other important features of their biology. A significant discussion of the paleontology of varanoids by Australian paleoherpetologist Ralph Molnar follows the introductory chapter and occupies 58 pages of this 602-page book. It is the most detailed and up-to-date paleontological re- view of the group and it also contains a very useful annotated inventory of terrestrial fossil varanoids, updating that of Estes (1983). There are occasional confusing or erroneous statements in Molnar’s review. For example, he notes that the oldest “necrosaur”

traces back to the Albian of Utah (Early Cretaceous, about 112-99 million years mya). This can only be the fossil originally described by Cifelli and Nydam in 1995. However, this fossil was later shown Herpetological Review, 2005, 36(3), 345Ð347. by Nydam (2000) to be that of Primaderma, which is a © 2005 by Society for the Study of Amphibians and Reptiles monstersaurian not a “necrosaur.” Molnar at first refers to Varanoid Lizards of the World, edited by Eric R. Pianka and Primaderma as the oldest land-dwelling platynotan. Later when Dennis King with Ruth Allen King. 2004. Indiana University Press, he discusses monstersaurs in detail, Molnar claims that members 601 North Morton Street, Bloomington, Indiana 47404-3797, USA of this taxon date back to at least the Late Cretaceous, and that the ([email protected]). 602 pp. Hardcover. US $89.95. ISBN 0- oldest forms were found in Asia. However, a mere paragraph af- 253-34366-6. terward, Molnar states: “Primaderma nessovi is the oldest known monstersaur…its remains were found in the [Early Cretaceous] MICHAEL J. BALSAI Cedar Mountain Formation of Central Utah” (page 28). Unfortu- Department of Biology, University of Pennsylvania Philadelphia, Pennsylvania 19104-6018, USA nate oversights such as this should not have escaped the editors. e-mail: [email protected] The first section of the book concludes with two relatively brief chapters. One, by Pianka and Molnar, is concerned with the bio- Monitor lizards and their close geography and phylogeny of varanoids generally, although its fo- relatives have always been a cus is on the genus Varanus. The final chapter in the first section, source of interest and fascination contributed by Jennings and Pianka, summarizes hypotheses about among herpetologists and laymen the tempo and timing for the radiation of Australian monitor spe- alike. Despite this, until the 1980s cies. Their analysis suggests that Australian goannas have under- general works on varanids were gone repeated and episodic speciation, that the increasing aridity largely limited to those by of Australia during the later Tertiary apparently helped drive this Mertens (1942a; 1942b; 1942c) diversification and radiation, and that the Australian varanid ra- for the living species and diations appear to parallel those of pygopods. Fejérváry (1918; 1935) for the In Part II, the individual species accounts are broken up into extinct taxa. Recently, interest in four sub-groupings: African varanid species, Asian varanid spe- these lizards has undergone a re- cies, Australian varanid species, and other varanoids. Each sub- naissance with the publication of section contains its own bibliography, and virtually all species important volumes devoted to in- descriptions contain all or most of the following subdivisions: dividual taxa, for example: nomenclature, geographic distribution, fossil record, diagnostic Auffenberg (1981; 1988: 1994), characteristics, description, size, habitat and natural history, re- Lenz (1995), and Murphy et al. (2002), as well as general works production, movement, population genetics, physiology, fat bod- and symposia, such as: Böhme and Horn (1991), Bennett (1998), ies, testicular cycles, and parasites. Range maps varying in their Horn and Böhme (1999), and Green and King (1999), and books level of detail are also usually included. Some species acounts are on varanid paleontology (Molnar 2004). This brief listing does supplemented by tables summarizing morphometric, physiologi- not even begin to account for the enormous wealth of current pub- cal, or breeding data, or graphs, such as that showing the relation-

Herpetological Review 36(3), 2005 345 ship between body mass and snout-vent length in Varanus gouldii. leads the chapter, is the only fossil taxon treated in its own indi- Nearly all species accounts provide a black and white photo above vidual segment (about two pages) despite the fact it was discussed the text for each description, along with duplicate, but separately earlier in Molnar’s chapter. Not even the gigantic Australian organized color print, showing a typical representative, usually an varanid, Varanus priscus [Megalania prisca] warranted its own adult. These photographs vary in quality, size, and detail. chapter, although this latter species went extinct much more re- The African varanid section had a wide range of authors and, as cently in the Pleistocene. All subspecies for Heloderma are dis- with all the other sections, the amount of detail varied from spe- cussed within the chapters about the two species and their range cies to species. All known African species (Varanus albigularis, maps also include subspecies information (both chapters are by V. exanthematicus, V. griseus, V. niloticus, V. ornatus) and the rela- Heloderma expert, Daniel Beck). Eric Pianka’s short chapter on tively recently described Middle Eastern V. yemenensis (Böhme Lanthanotus is notable for demonstrating how little is known about et al. 1987, 1989 [note: these two citations are incorrectly listed in this important taxon, particularly concerning its diet and times of the book’s bibliography and are corrected here]) are discussed. activity, as well as reproductive, thermoregulatory, and foraging The account by Michael Stanner describing the desert monitor, behaviors. Varanus griseus, consumes the greatest percentage of this section, The book concludes with a two-chapter section. The first, by about 45%, but fails to resolve whether or not this species is actu- Pianka, draws an important relationship between body size and ally venomous as some earlier reports claimed (Ballard and Anto- reproductive tactics, and basically argues that body size influences nio 2001; Sopiev et al. 1987). reproduction more strongly than phylogeny, particularly: egg mass, The Asian monitor section describes 23 species, with that for clutch size, clutch mass, neonate snout-vent length, and neonate Varanus salvator split into two separate segments: one devoted to body mass. Pianka also notes that clutch sizes for larger species the nominate form and the other to the various subspecies. It is are normally smaller than those produced by the smaller species heavily dominated by German authors and, therefore, many of the and that maternal snout-vent length is a more potent influence on species descriptions include diagnostic characters containing ref- clutch size within a given species than between any two species. erences to hemipeneal (and sometimes hemiclitoral) characters that The final chapter, by Hans-Georg Horn, examines the captive care are derived from Böhme (1988). Unfortunately, no general dis- of monitors with an eye toward biological, technical, and legal cussion of the specialized terms regarding these important genital difficulties, and he discusses many relevant issues including: dif- characters is provided in this volume and the book’s glossary also ficulties providing the proper diets, distinguishing males from fe- lacks any definitions to assist unfamiliar readers. Considering that males, the problems involved with distinguishing between two these and many other authors place great significance on these species showing very similar appearance, problems involving the morphological characters to differentiate many varanid species, proper lighting and construction of enclosures, and the effects of this deficiency must be corrected in any future editions to ensure various laws, including CITES legislation, on the captive conser- comprehensiveness. vation of monitors. In the Asian monitor section several individual species mem- This book will likely be an important single-volume source about bers of the so-called “Varanus indicus group” are described sepa- varanoid lizards for some time to come. Despite some mostly mi- rately from the nominate taxon and similar partitioning was also nor editorial oversights, including those mentioned above, some made for members of the “V. prasinus group” (which in this vol- of which may have resulted from the sudden and unfortunate death ume includes: V. kordensis, and V. macraei; but not V. beccari, and of Dennis King during the production of this volume, I believe V. bogerti). None of the Asian monitor chapters are as detailed as this book to be a welcome and important addition to the library of that for V. griseus, however, the two segments concerned with V. any herpetologist. Its importance lies not only in what informa- salvator and its subspecies, when combined, come close. The oth- tion it manages to include within its covers, but also in its indica- ers, all succinct reviews, average about four or five pages each. tion of the significant work still remaining for the more complete The section on Australian monitors has the greatest number of understanding and conservation of these important reptiles. authors and, not surprisingly, the majority are Australians (the one dealing with V. keithhornei was even contributed by “Croc Hunter” LITERATURE CITED Steve Irwin). One significant difference between this section and those for the African and Asian monitors is that each Australian AUFFENBERG, W. 1981. The Behavioral Ecology of the Komodo Monitor. species contains a list of specimens in the major Australian mu- University of Florida Press, Gainesville. 406 pp. ––––––. 1988. Gray’s Monitor Lizard. University of Florida Press, seum collections. Many of these specimen lists appear quite ex- Gainesville. 419 pp. haustive and all are potentially very useful to any varanid researcher ÐÐÐÐÐÐ. 1994. The Bengal Monitor. University of Florida Press, doing work on these species. Unlike for the previous two sec- Gainesville. 560 pp. tions, I could not detect any missing or incorrectly cited refer- BALLARD, V., and F. B. ANTONIO. 2001. Varanus griseus (desert monitor): ences, but similar to these earlier sections, the amount of detail toxicity. Herpetol. Rev. 32:261. provided for any given Australian species varies. Generally, these BENNETT, D. 1998. Monitor Lizards: Natural History, Biology and Hus- descriptions were all concise and averaged somewhere between bandry. Edition Chimaira, Frankfurt am Main. 352 pp. six and ten pages. BÖHME, W. 1988. Zur Genitalmorphologie der Sauria: Funktionelle und The “Other Varanoids” section contains accounts for stammesgeschichteliche Aspekte. Bonn. Zool. Monogr. 27:1Ð176. ÐÐÐÐÐÐ, J. P. FRITZ, AND F. SCHÜTTE. 1987. Neuentdeckung einer Gro§echse Lanthanotus, both living species of Heloderma, and the Late Cre- (Sauria: Varanus) aus der Arabischen Republik Jemen. Herpetofauna. taceous monstersaur, Estesia mongoliensis. Estesia, which did not 46:13Ð20. merit a color photo, although a black and white print of the skull

346 Herpetological Review 36(3), 2005 ÐÐÐÐÐÐ, and H.-G. HORN (eds.). 1991. Advances in Monitor Research, reptiles and amphibians were bit players in these studies. A few Mertensiella 2:1Ð266. species were notable exceptions, of course (e.g., coral snakes, egg- ÐÐÐÐÐÐ, U. JOGER, AND B. SCHÄTTI. 1989. A new monitor lizard (Reptilia: eating snakes; reviewed in Pough 1994); but these involved one Varanidae) from Yemen, with notes on ecology, phylogeny and zooge- herp mimicking another. Cases of a herp mimicking an inverte- ography. Fauna of Saudi Arabia 10:433Ð448. brate are rare (Autumn and Han 1989; Gans 1987; Parker and CIFELLI, R. L., AND R. L. NYDAM. 1995. Primitive helodermatid-like platynotan from the early Cretaceous of Utah. Herpetologica. 51:286Ð Pianka 1974; Vitt 1992) and often anecdotal. 291. In 1977 we proposed that a ground-dwelling lizard in the ESTES, R. 1983. Sauria terrestria, Amphisbaenia. Gustav Fischer Verlag, Kalahari Desert mimicked a beetle (Huey and Pianka 1977). The Stuttgart, Germany. 249 pp. possibility that a lizard could mimic a beetle must have seemed FEJÉRVÁRY, G. J. 1918. Contributions to a monograph on fossil Varanidae ludicrous to many, and no doubt some of our colleagues won- and on Megalanidae. Ann. Mus. Natl. Hung. 16:341Ð467. dered whether we’d spent too much time out in the Kalahari sun. ÐÐÐÐÐÐ. 1935. Further contributions to a monograph of the Megalanidae Our evidence was circumstantial; but we were convinced that this and fossil VaranidaeÐwith notes on recent varanians. Ann. Hist.-Nat. was mimicry, and that it involved both color and locomotor be- Mus. Natl. Hung. 29:1Ð130. havior. GREEN, B., AND D. KING. 1999. Monitors: The Biology of Varanid Liz- ards, 2nd Edition. Krieger Publishing Co., Malabar, Florida. 116 pp. The model is the juvenile lacertid lizard, Heliobolus lugubris, HORN, H.-G., AND W. BÖHME (eds.). 1999. Advances in Monitor Research and the mimic is a carabid beetle, Anthia spp. Adult H. lugubris II. Mertensiella 11:1Ð 366. are sand colored and reasonably cryptic, but juveniles are decid- LENZ, S. 1995. Zur Biologie und Ökologie des Nilwarans, Varanus niloticus edly conspicuous against the red Kalahari sand. Juveniles have (Linnaeus 1766) in Gambia, Westafrika. Mertensiella 5:1Ð256. jet-black bodies, with broken white stripes—only their tails are MERTENS, R. 1942a. Die Familie der Warane (Varanidae) Erster Teil: sand colored (Fig. 1). No other lacertid in the Kalahari changes Allgemeines. Abh. Senckenberg. Naturforsch. Ges. 462:1Ð116. color so dramatically during ontogeny. Nor does any other lacertid ÐÐÐÐÐÐ. 1942b. Die Familie der Warane (Varanidae) Zweiter Teil: Der of which we are aware. Schädel. Abh. Senckenberg. Naturforsch. Ges. 465:117Ð234. Both juvenile and adult H. lugubris are wide foragers (Huey ÐÐÐÐÐÐ. 1942c. Die Familie der Warane (Varanidae) Dritter Teil: Taxonomie. Abh. Senckenberg. Naturforsch. Ges. 466:235Ð391. and Pianka 1981), but they differ strikingly in the way they move. MOLNAR, R. E. 2004. Dragons in the Dust: The Paleobiology of the Giant Adults move like normal lacertids (with lateral undulations), but Monitor Lizard Megalania. Indiana University Press, Bloomington, juveniles often move stiff legged, with their backs arched and their Indiana. 210 pp. tails pressed to the ground (Fig. 1). When the juveniles ‘metamor- MURPHY, J. B., C. CIOFI, C. DE LA PANOUSE, AND T. WALSH. 2002. Komodo phose’ to the adult coloration, they switch from arch-walking to a Dragons: Biology and Conservation. Smithsonian Institution Press, normal walking style. Juvenile H. lugubris are the only lizards Washington, D.C. 267 pp. known to use arch-walking. NYDAM, R. L. 2000. A new taxon of helodermatid-like lizard from the We didn’t do a formal phylogenetic analysis, but we realized Albian-Cenomanian of Utah. J.Vert. Paleontol. 20:285Ð294. that two unique features of the juveniles (coloration, arch walk- SOPIEV, O., V. M. MAKEYEV, S. V. KRUDRJSTEV, AND A. N. MAKAROV. 1987. A case of intoxification from the bite of Varanus griseus [in Russian]. ing) must be evolutionarily derived and thus called for explana- Izv. Akad. Nauk Turmen. SSR, Ser. Biol, Nauk. 87:78. tion. When we looked around the Kalahari, we soon noticed cara- bid beetles (Anthia). These beetles are black-and-white, a classic aposematic pattern, and often abundant. Locals refer to them as “oogpisters” (which translates euphemistically as “eye squirter”), Herpetological Review, 2005, 36(3), 347Ð348. © 2005 by Society for the Study of Amphibians and Reptiles because these beetles squirt from their abdomen a noxious mix- ture of formic acid, tiglic acid, and other compounds (Scott et al. Die Mimikry zwischen Eidechsen und Laufkafern (Mimicry 1975). As far as we were aware, no vertebrate predator ate these Between Lizards and Ground Beetles), by Almuth D. Schmidt. noxious beetles. 2004. Edition Chimaira, Frankfurt am Main, Germany To us the observed patterns strongly suggested that the beetles (www.chimaira.de). 374 pp. Hardcover. ¤ 58.00 (approximately were noxious models and that juvenile H. lugubris (approximately US $75.00). ISBN 3-930612-69-0. the same size as the beetles) were Batesian mimics, involving both RAYMOND B. HUEY color and movement. We noted that juveniles of this species had a Department of Biology, Box 351800, University of Washington Seattle, Washington 98195-1800, USA e-mail: [email protected]

and ERIC R. PIANKA Integrative Biology C0930, The University of Texas Austin, Texas 78181-0253, USA e-mail: [email protected]

For more than a century, warning coloration and mimicry have fascinated evolutionary biologists and natural historians. In fact, these phenomena helped convince late 19th Century biologists of the power of natural selection. FIG. 1. An arch-walking juvenile Heliobolus lugubris (photograph cour- During the 20th Century, studies of mimicry were common; but tesy of A. Schmidt).

Herpetological Review 36(3), 2005 347