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Studies on Lizards and Tuataras in Zoos and Aquariums. Part I—Introduction, History, Families Iguanidae, Agamidae, Chamaeleonidae, and Infraorder Gekkota

Studies on Lizards and Tuataras in Zoos and Aquariums. Part I—Introduction, History, Families Iguanidae, Agamidae, Chamaeleonidae, and Infraorder Gekkota

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Herpetological Review, 2015, 46(3), 464–482. © 2015 by Society for the Study of and Studies on and in and . Part I—Introduction, History, Families , , Chamaeleonidae, and Infraorder

chapters written by experts in herpetology (Hutchins et al. 2003). Some families have been only superficially investigated in People often ask, “What good are lizards?” to which we re- zoos: , Agamidae, , , Biped- spond with “What good are people?” Such anthropocentrism is idae, , , , Gymnophthalmi- abhorrent. Lizards have as much of a place on the planet as any dae, , , and Xantusiidae. living creature, including humans. Indeed, they have successfully inhabited for much longer than humans have – lizards will History undoubtedly persist long after humans and most other have gone extinct. Lizards are spectacular products of natural se- In May 1849, the London , founded in 1828, opened the lection and have diversified to fill amazing variety of ecologi- first zoo building in the world, a modified carnivorous cal niches. They are extremely good “model” organisms for study, facility which used the dens as enclosures for lizards and understanding their ecology and diversity can be exceedingly and other reptiles (Keeling 1992). The second zoo reptile build- informative. What we have learned about lizards is applicable to ing at London, designed and constructed specifically for reptiles, nearly every conceptual area in modern biology; indeed, in many was opened in August 1883 (Guillery 1993) and is currently a cases development of entire fields of biology had their origins in the study of lizards. Because many lizards are quite beautiful, they are very popular as pets among herpetoculturists around the world, and some people make their living by breeding many of lizards in captivity for resale. Eric R. Pianka & Laurie J. Vitt, Lizards: Windows to the of Diversity, 2003

Curator of herpetology at the New York Zoological Park, Ray- mond L. Ditmars in his book, Reptiles of the World (1933), mar- veled at the diversity of lizards, “Looking down upon the vast Or- der now engaging the student’s attention, even the most passive

of observers cannot refrain from expressing amazement at the DC WASHINGTON LIBRARIES, INSTITUTION SMITHSONIAN array of varied forms. In a subdivision, the Sauria, we shall con- sider creatures twelve feet long, with claws as long as those of a leopard— strong and active enough to leap at the throat of a young gazelle, tear, dismember and devour the prey; and passing such we stop to realize that tiny, limbness and worm- like, slow-moving things, burrowing their life away deep in the ground where they need no eyes—in fact, have none—are also true lizards.” (Figs. 1, 2) In this two-part paper, I follow the organizational framework of McDiarmid et al. (2012) as a taxonomic review of extant lizard groups. Part II will appear in the next issue. Original standard and scientific names are used. To update these names, use The (Uetz and Hošek 2013) and Standard English and Scientific Names: Checklist of the Standard English Common Names of Amphibians & Reptiles (Crother 2012) if occurring in North America. A complete reference called Grzimek’s Life Encyclopedia was published on all reptiles in 2003, with

JAMES B. MURPHY Division Amphibians & Reptiles, National Museum of , 10th and Ave NW, Washington, DC 20013-7012, USA Fig. 1. Chromolithograph from Kunstformen der Natur [Art Forms in e-mail: [email protected] Nature], by (1904).

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Fig. 2. Illustration in Gotthilf Heinrich von Schubert’s Naturgeschichte des Thierreichs. . ., published in 1869.

tropical building today. The newest London Reptile House London Zoo curator E. G. Boulenger published an excellent was designed by curator Joan Beauchamp Procter in 1926–1927 book with many photographs called Reptiles and Batrachians and is still in use. The building included a center island to house (1914). He included many observations on lizards living in the venomous and other potentially dangerous reptiles and a collection. “Reptiliary” housing . New zoo design features were in- Jon Coote (2001) speculated that many reptiles, especially corporated: differential heating, “ principle lighting” lizards, arrived at the London Zoo in good condition in the nine- (illuminating enclosures from above), elaborate naturalistic dis- teenth century because those responsible for transporting them plays, and better visitor traffic. were given specific instructions for their care: “Correspondents Philip Lutley Sclater, Secretary for the Society (1859‒1902), should engage some individual of the ship’s company to take compiled nine editions of an inventory (“List of the vertebrat- charge of the animals on board and guarantee him a handsome ed animals now or lately living in the gardens of the Zoologi- recompense on bringing them safely to their destination...” Food cal Society of London”) beginning in 1862. In 1872, his revised was also important for reptiles on the long sea journeys and list included tuataras and many lizard taxa such as the Moloch, correspondents were advised: “ , which are abundant Bearded Dragon, mastigure (), Shingleback in tropical climates, may be preserved in a jar, well tied down and other , several , as well as lacertids and and with the addition of the Blattae or cock-roaches so gener- iguanids to round out the collection. Books on the husbandry of ally obtainable on board in all their stages of growth, and of meal lizards began to appear during the Victorian era in greater num- worms, which are equally abundant in the bread room …” bers (Murphy and McCloud 2010). For instance, A. D. Bartlett During that time, there was closer cooperation between zoo (1899) had written Wild Animals in Captivity Being an Account workers and private herpetoculturists than there is now. Until of the Habits, Food, Management and Treatment of the Beasts 1903, keepers at the London Zoo were able to trade or sell sur- and at the ‘Zoo,’ which treated the habits of lizards such as plus animals to the general public and they supplied herps to the Stump-tailed Skink, , , and . amateur British herpetoculturists. Reverend Gregory Bateman These books had a practical application in that private fanciers (1897) purchased a pair of from the Zoo for £2.00 ($3.00) used the zoo protocols, particularly those from the London Zoo, and other species were offered for sale in London. Prices were though it must be stressed that these Victorian fanciers, mostly reasonable: small monitor lizards—5 shillings (38 cents US), women, went on to develop many innovative techniques. Later, Common —about £2.00 ($3.00), Red Tegus—£3.00 to £8.00

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Stanley Smyth published monographs and papers on and reptile longevities. In his 1937 publication, he mentioned that varanids and large iguanids are often kept in captivity but it is surprising that only two lizard taxa had reached twenty years in captivity—Slow-Worm ( fragilis) for 32 years and Cunningham’s Skink (Egernia cunninghami) for 20 years. In North American collections, Snider and Bowler (1992) listed one anguid, one cordylid, three gekkonids, four iguanids, three skincids, one varanid, and a number of helodermatids passing twenty years in captivity. Carey (1973) provided some notable longevity records for captive iguanas. SMITHSONIAN INSTITUTION LIBRARIES, WASHINGTON DC WASHINGTON LIBRARIES, INSTITUTION SMITHSONIAN In the early twentieth century, papers on lizard husbandry and captive management began to regularly appear, written by London Zoo curators such as E. G. Boulenger (1913, 1920) and Joan Proctor (1928) and many European and herpe- tologists (see Card and Murphy 2000, Murphy 2007, 2009‒2010 for descriptions). In 1955, an extraordinary four-volume set by Wilhelm Klingelhöffer named Terrarienkunde appeared and was THE husbandry reference book for decades in . Toward the end of the century, zoo herpetologists became better at rec- ognizing signs of reproductive and other behaviors and recording these observations in scientific journals. Many books focused on captive lizards: Lizards in the Terrarium (Jes 1987), a two-volume work called Lizards (Rogner 1997), Encyclopedia of Terrarium (Bru-

JESSIE COHEN, NATIONAL ZOO NATIONAL COHEN, JESSIE ins 1999), Biology, Husbandry, and Medicine of the (Jacobson 2003), Life (Leutscher 1952), Firefly Encyclo-

PHOTO BY BY PHOTO pedia of the Vivarium (Alderton 2007), Reptiles, Amphibians, and Invertebrates (Bartlett et al. 2001), Handbuch der Terrarienkunder (Stettler 1978), Das Terrarium (Kahl et al. 1980), two-volume Die Terrarientiere (Nietzke 1969, 1972), Guide to Lizards, All about Liz- ards, and Giant Lizards (Sprackland 1977, 1992, 2010), Breeding Terrarium Animals (Zimmermann 1983), The Completely Illus- trated Atlas of Reptiles and Amphibians for the Terrarium (Obst et al. 1988), Reptiles and Amphibians: Their Care and Behavior (Vogel 1964), and The Lives of Captive Reptiles (Petzold 2008). Ex- Fig. 3. Two of the rarest lizards in the world are shown here. Top: The Jamaican Land Iguana ( collei) was described by John Edward citing new possibilities appeared in the literature such as inves- Gray as Colley’s Cyclura in 1845. Illustration from Catalogue of the tigating the evolution of social cooperation in lizards (Dickinson specimens of lizards in the collection of the . This spe- and Koenig 2003; Sinervo and Clobert 2003) and manipulating cies was thought to be extinct until rediscovered in 1970. It now faces clutch and offspring size (Sinervo 1994). Ethologists and other re- another threat due to plans to alter the . searchers from the academic community began working jointly Bottom: This male Island Iguana ( with zoo colleagues on projects in zoos and aquariums, resulting lewisi) allows itself to be groomed with a toothbrush by standing im- in improved studies (see Sajdak 1983; Chiszar et al. 1993; Chiszar mobile as its entire body is brushed. and Smith 2005; Garrett 2005 for reviews). Reviews focusing on the nutrition of carnivorous and herbivorous lizards were pub- ($4.50 to $12.00) depending upon size, Bearded Dragons—10 lished (Allen and Oftedal 1994; Baer 1994). Crews and Les- shillings to £1.00 (75 cents to $1.50), Green Anoles—as low as lie Garrick (1980) provided many examples for inducing lizards half-a-crown (16 cents), European Chameleons—3 shillings and to reproduce in captivity. In the United States, researchers such 6 pence to 7 shillings and 6 pence (30 to 65 cents), British Com- as Gordon Burghardt, Charles Carpenter, David Chiszar, William mon Lizards—4 to 6 pence each (6 to 9 cents). Cooper Jr., Gary Ferguson, William Gehrmann, James Gillingham, Beginning in 1882, Johann von Fischer from Vienna began Harry Greene, Thomas Jenssen, Hobart Smith, and Paul Weldon, to publish a large number of papers on captive lizards: Ringed were pioneers who used zoo and aquarium collections to broad- ( torquatus), European Chameleon ( en their studies. William Cooper Jr. has used zoo collections as a vulgaris), Cape (Uromastix capensis auct.), Stum- resource for his studies on lizard feeding behavior for many years. melschwanz Lizard (Trachydosaurus asper), iguana (Iguana tu- Examples include varanoid lizards ( suspectum and berculata), Cylindrical Lizard (Gongylus ocellatus), Pygmy Lat- Varanus exanthematicus), (), and skinks eral Fold Lizard (Ablepharus pannonicus), Schleuderschwanz (Tiliqua scincoides and Trachydosaurus rugosus). Susan Barnard (Uromastix acanthinurus), Common Skink (Scincus officinalis), from Zoo Atlanta and her associates have published a number of Spotted Lizard, pardalis), Worm Lizard (Trogonophis papers on parasitism and husbandry in zoo lizards (see Murphy wiegmanni), and various lacertids. His seminal book on captive 2007). Backues and Ramsey (1994) performed an ovariectomy for management was called Das Terrarium, seine Bepflanzung und treatment of follicular stasis in lizards. Two important overviews Bevölkerung (1884) and includes recommendation for many by Stamps (1997) and et al. (2003) examined social behavior mixed lizard combinations. and spacing patterns in lizards.

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The late Thomas Huff from Reptile Breeding Foundation in

Canada identified a condition that he called “captive stagnancy.” (1924) He believed that changing environmental factors was key to im- proving reproductive behavior and his solution was to change exhibit furniture, potential breeders, temperature, and so on to induce courtship and copulation (see Huff 1980). Warwick et al.

(1995) addressed the issue of stress. Methods for incuba- END WORLD’S GALAPAGOS. tion and raising neonates became more refined (see Packard and Phillips 1994). Earlier, reptile eggs were discarded in many instances because it was believed that hatching them was im- possible. The importance and provision of light for reptiles held in indoor enclosures were highlighted in key papers WILLIAM FROM BEEBE’S by Gary Ferguson and associates (1996; Jones et al. 1996), Wil- liam Gehrmann (1971, 1987, 1994), Carol Townsend (1979) and Townsend and Charles Cole (1985), and Jozsef Laszlo (Murphy ILLUSTRATION 2007). Philip Regal (1980) examined temperature and light re- quirements. From Toledo Zoo, R. Andrew Odum (1984) stressed the importance of water quality. Lawson et al. (2008) stressed the importance of science in zoos and aquariums. Dave Chiszar, Hobart Smith, and I argued that assessing the competence of captive-produced herps is as important as breeding the animals (Chiszar et al. 1993). We were surprised to discover that captive and wild Komodo Dragons have significantly different body temperatures (Wikramanay- ake et al. 1999; Walsh et al. 1999). If released into the wild, would Fig. 4. Curator Perkins from moved 60 Galápagos Land captives with lower temperatures adjust thermoregulatory pat- Iguanas ( subcristatus) from , (known also upward? If deficits brought on by captivity are identified, as South Seymour Island) to , which contained can these be remediated? How can this be done? What necessary no iguanas in the 1940s. Had this not been done, this species would studies will be in place to assess survivability in the wild? These now be extinct, thanks to human activities. See text. challenging issues have only been minimally addressed but there are some impressive exceptions: West Indian iguanas (Cyclura) remained in 2000. Two years later, the population crashed pre- serve as examples of successful field studies and reintroductions. cipitously and only 15‒25 lizards were left. In 2003, five iguanas Family Iguanidae.—West Indian Land Iguanas (Cyclura) face were hatched at the and two more lizards two a perilous future. Jeffrey Lemm and Allison Alberts from San Di- years later. Described in 1940, the lizard was widely distributed ego Zoo Global in have worked for many years with in dry over most of the island but is now restricted to a these iguanid lizards. Their recent comprehensive book (Lemm few remnant populations, due to human development, ani- and Alberts 2012) covers evolution and , species mals, and . Be sure to read Burton’s Little Blue accounts, natural history, husbandry, nutrition, health and Book (2010) about the challenges of conserving this species. He medical management, and conservation. The San Diego Zoo has highlights the contribution by retired National Zoo curator Dale been an integral player in iguana research and conservation over Marcellini in developing the conservation strategy. Marcellini many years through publication of books and papers by Alberts suggested to on many occasions that fieldwork should always and colleagues. be planned in a place where one could sit on the veranda of a The Jamaican Land Iguana (Cyclura collei) was described by four-star hotel with a refreshing drink in hand and watch lizards as Colley’s Cyclura in 1845 (Fig. 3). Four years with binoculars—Dale called this plan “Resort Research”! An- later, one was donated to London Zoo and lived in the Zoo for 3 other amazing field account was written by Quentin Bloxam, who years, 4 months, and 29 days (Coote 2001). Concern about the nearly died while working on conserving this on Grand iguana’s future accelerated following the introduction of the In- Cayman Island (see Murphy 2007:63). Sarah Kuppert (2013) de- dian on in 1872 and things were critical by scribed interactive male and female head bob displays at Smith- 1916 when this iguana was only found on a few small islets off the sonian National Zoological Park (SNZP). This adult male allowed coast of Jamaica where the predaceous mongoose had not been itself to be groomed with a toothbrush by standing immobile to introduced. In 1940, a total of 22 specimens were brought into allow its entire body to be brushed. The keeper pulled off dried captivity to try to save the species from but none ever skin by spraying these areas beforehand with water (Fig. 3). reproduced. The lizard was later thought to be extinct. A hunt- Noegel (1989) reproduced five species of West Indian rock er’s dog discovered a carcass in 1970 and a later survey located iguanas (Cyclura) at the Life Fellowship Bird Sanctuary, Seffner, about 100 lizards left in the wild in the Hellshire Hills. There is Florida. Duval and Christie (1990) outlined care for the Cuban now a headstarting program at the Hope Zoo in Kingston, which Ground Iguana (Cyclura n. nubila), based on their experiences involves other zoos as well (Gibson 1993; Vogel et al. 1996). How- at the Indianapolis Zoo. The reproductive biology of Cuban Rock ever, today there exists a sizeable threat to irrevocably develop Iguanas (C. nubila) has been covered in detail at San Diego Zoo the Hellshire Hills, the only place where the lizard is found. (Shaw 1954; Alberts 1995, 2002; Alberts et al. 1997) and Prague The Grand Cayman (Cyclura nubila lewisi) is one Zoo (Rehák and Král 1993; Rehák 1994, 1995 [1997]; Rehák and of the most rock iguanas and Fred Burton Velenský 1997, 2001). Haast (1969) hatched Rhinoceros - (2000) mentioned that perhaps only a few hundred individuals nas (C. cornuta) at the Miami Serpentarium and Rehák (1996)

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documented the first captive breeding of the his observations made during hatching and raising of a Green in the Czech Republic. In two papers, Boylan (1984, 1985) de- Iguana in the Berlin Tierpark (Fig. 5.). Mendelssohn (1980) pro- scribed captive management of a population at the Taronga Zoo, vided recommendations on care, based on reproduction in a Sydney, and Petzold (1967) at Tierpark Berlin. Otten- colony at Tel Aviv University in Israel. Howard (1980) bred them walder (1980) mentioned that Zoodom released rhino iguanas at the Twycross Zoo and Banks (1984) at Melbourne Zoo. Wallach into the wild and described reproduction in captivity (Sanlley (1966) described hypervitaminosis D in Green Iguanas. Bosch Castro and Duval 1979). Charles Knapp from the Department and Werning (1996) discussed the captive management of Green of Conservation at the John G. Shedd Aquarium in Chicago has Iguanas and other iguanids. Wallach and Hoessle (1968) docu- published field studies on several West Indian iguanas (Knapp mented fibrous osteodystrophy in Green Iguanas. Zwart and Van et al. 1999; Knapp 2000, 2001; Knapp and Malone 2003;). Duval de Watering (1969) described pathology and etiology of abnor- (1982) offered suggestions for the captive management of West mal bone formation in this species. The reproductive biology of Indian rock iguanas. Bosch (1991) discussed ground iguanas (Cy- Green (Iguana) and (Dipsosaurus) iguanas has received clura cychlura figginsi) in the Löbbecke Museum (Dusseldorf). attention (Judd et al. 1976). Day et al. (2000) provided a species See also Bosch and Boscheinen (2001). Murphy (1969) described account of the (Iguana delicatissima). conspecific grooming behavior in this species. Bosch and Werning (1996) wrote a manual on iguana care. C. B. “Si” Perkins, Curator of Reptiles at San Diego Zoo, and Arnett (1979) reproduced the Banded Iguana (Brachylo- G. Allan Hancock mounted trips in 1932 and 1933 to the Galápa- phus fasciatus) at Knoxville Zoo and over 100 young have been gos Islands to study and collect animals for the Zoo (see Banning produced at San Diego Zoo (Fig. 6.). In 2008, a new species was 1933 for additional information). Hancock’s ship was the state- described (B. bulabula) and a breeding program has been - of-the-art Valero III, nearly 200 feet in length. He was a bit of a tiated by San Diego Zoo. Boylan (1989) bred the Fijian Crested prig, insisting that his fellow passengers—a group of scientists Iguana (B. vitiensis) at Taronga Zoo in Sydney, Australia. Carpen- including Perkins—dress for dinner and moreover, cease smok- ter and Murphy (1978) noticed aggressive behavior and color ing and drinking. For those of us who are not focused on the lat- change between males in the Fiji Island Iguana at Dallas Zoo. est in apparel and enjoy a bit of whisky and strong cigarettes in Marcellini and Jenssen (1991) documented avoidance learn- the evening, this trip would have been unendurable. The nota- ing by the Curly-tailed Lizard (Leiocephalus schreibersi). Petzold tion “No cocktails tonight” appeared daily in the Perkins diaries (1962) successfully bred the Northern Curly-tailed Lizard (Leio- (Campbell 1978). cephalus carinatus) at Berlin Tierpark. Gibson and Buley (1996) During the first trip, Hancock and Perkins decided to move 40 described captive management and breeding of Galápagos Land Iguanas (Conolophus subcristatus; Fig. 4.) from Spiny Iguanas (Oplurus c. cuvieri) at Durrell Wildlife Conserva- Baltra Island (known also as South Seymour Island) to North Sey- tion Trust. Vogel (1992) outlined care and breeding of the Ser- mour Island which contained no iguanas. As Perkins wrote in his rated Casque-headed Iguana ( serratus) over several diaries, their reason: “. . . in a few years come down and see if generations in the Exotarium of Frankfurt Zoo. At Dallas Zoo, we anything has happened. A good idea, I believe.” (Campbell 1978). received a group of lizards confiscated by U.S. & Wildlife Twenty more iguanas were translocated on the second trip. Service, which included Casque-headed Iguanas (Laemanctus Baltra was an American airbase during World War II in the longipes), Smooth-helmeted Iguanas (Corytophanes cristatus), 1940s and several thousand military and support personnel were Hernandez Smooth-helmeted Iguanas (Corytophanes hernande- stationed there, in part to guard the Canal. Several fac- sii), Common Monkey Lizards ( marmoratus) (Fig. 7), tors contributed to the disappearance of these lizards—habitat and Tree Runners (). All but the last taxon was kept at destruction, introduction of feral animals, and using the lizards ca. 72°F (22°C) ambient temperature with a basking spot. A col- for target practice. league told me that he found a Smooth-helmeted Iguana resting There appeared to be virtually no successful reproduction on a tree trunk in the field and did not move for over a week until or recruitment on North Seymour for the subsequent 47 years a large passed by. At Dallas Zoo, a group of Baja California so a pair of adults was brought to the Research Rock Lizards (Petrosaurus thalassinus) and Banded Rock Lizards Center on Santa Cruz Island to begin a captive colony; addition- (P. mearnsi) were displayed in a large enclosure (Fig. 8). These al iguanas were included in this potential breeding group later lizards were hardy and constantly active. Bayard Brattstrom vis- (Cayot et al. 1994). These iguanas reproduced and survived, par- ited and suggested that we arrange two hotspots above two plat- ticularly because feral and dogs had been reduced on Baltra. forms where these lizards would learn to shuttle back and forth In June 1991, 35 five-year old iguanas were repatriated to Baltra between them. A timer would be used for tripping a relay switch and 24 were released the next year. This head-starting program to turn one or the other off; both would not be on simultane- was truly an accomplishment deserving praise, thanks to Perkins ously. An intriguing display showing shuttling thermoregulation and Hancock. (see Regal 1980:83 for another view) but we never got around to Ray Pawley from Chicago Zoological Park (Brookfield Zoo) doing it. maintained a colony of Marine Iguanas (Amblyrhynchos crista- Reproduction and behavior of the Green Crested Basi- tus) in a large exhibit in the reptile building (1965, 1966, 1969, lisk ( plumifrons) were described by Pawley (1972) 1971). I visited the Zoo on a number of occasions and was im- at Brookfield Zoo, Blake and Stewart (1980) at Edinburgh Zoo, pressed by the overall health and activity of these lizards. This Olexa (1976) at Prague Zoo, and Banks (1983) at Melbourne Zoo. species is challenging to maintain successfully in captivity but Kuppert (2013) recorded head bob displays at National Zoo. one of his saurians lived for over six years. Stunkard and Gandal (1966) identified a digenetic trematode, Dennis Desmond told me that his adult female Green Iguana Parahaplometroides basiliscae from the mouth of a basilisk at (Iguana iguana) would sleep with a furry toy virtually every night New York Zoological Park. At Dallas Zoo, a mixed exhibit of Com- and the lizard often wrapped its forelimbs around this unusual mon Basilisks (Basiliscus basiliscus), Western Basilisks (B. galeri- object (see photo in Murphy 2007). Dedekind (1977) recorded tus), and Brown Basilisks (B. vittatus) were visible to the visitor

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Fig. 5. Illustration of Common or Green Iguana (Iguana iguana) in Baron ’s Le règne animal…, published in 1836‒9. An adult female slept with a furry toy virtually every night and the lizard often wrapped its forelimbs around this unusual object.

Fig. 7. Illustration of Common Monkey Lizard (top right) () by H. R. Schinz (1833), Naturgeschichte und Abbildun- gen der Reptilien. I saw one in the wild in and it was virtually invisible in a bush. It remained immobile when I reached for it and only moved when I grasped it.

Fig. 8. Illustration of Baja California Rock Lizard (Petrosaurus thalas- Fig. 6. The Fiji Island Iguana ( fasciatus) here pictured in “Proceedings of Scientific Meetings of the London Zoological So- sinus) and Banded Rock Lizard (P. mearnsi) [above] from Nouvelles ciety” is certainly impressive but it is at risk due to human factors. archives du Muséum d’ de Paris, 1865‒1908. An The San Diego Zoo is playing a significant role in developing captive interesting exhibit using these lizards could be developed to show assurance colonies. shuttling thermoregulation. See text.

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Fig. 11. Illustration of Egyptian Spiny-tailed Lizard (Uromastyx ae- gyptius) in Baron Georges Cuvier’s Le règne animal…, published in 1836‒9. The Zoo was successful at keeping members of this in large enclosures with a sandy substrate and high temperatures.

Fig. 9. There are beautiful images in “Proceedings of Scientific Meet- ings of the London Zoological Society” between 1861‒1929. An adult male Frilled Lizard ( kingi) from Australia flared its frill with open mouth, stood bipedally, made a series of grunts and rushed toward the keeper at Dallas Zoo. New employees found this display to be unsettling when witnessed the first time and, of course, were never forewarned by their perverse co-workers.

Fig. 12. A small group of Common Flying Lizards ( volans) was kept at Dallas Zoo where males regularly displayed to each other by extending dewlaps and partially extending wings, and bobbing the body. This illustration represents a different species (D. lineatus) but is included to show the head and body of this unique lizard. Publica- tions of London Zoological Society between 1860–1921 had beautiful

SMITHSONIAN INSTITUTION LIBRARIES, WASHINGTON DC WASHINGTON LIBRARIES, INSTITUTION SMITHSONIAN lizard illustrations such as Flying Lizards (Draco). To see a complete list of all amphibians and reptiles, consult Transactions of the Zoo- logical Society of London: an index to the artists, 1835–1936, compiled by Nina J. Root and Bryan R. Johnson, and Proceedings of the Zoologi- cal Society of London: an index to the artists, 1848–1900, compiled by Nina J. Root and Bryan R. Johnson (1986).

Fig. 10. Illustration of Bearded Dragon ( barbata) in Baron Georges Cuvier’s Le règne animal…, published in 1836‒1839. This liz- ard is hardy in captivity and is highly variable in color.

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when entering the reptile building. The enclosure was large and Gamble et al. (1996) investigated plasma Itraconazole phar- high; the lizards were active, climbing branches and the rock fa- macokinetics in spiny lizards (Sceloporus spp.) at Dallas Zoo. A çade. number of spiny lizards, , and other southwestern At Dallas Zoo, a number of beautiful Sceloporus were kept U.S. desert lizards were kept in a large outdoor exhibit with a over the years: Adler’s Spiny Lizard (S. adleri), Yarrow’s Spiny sizeable hibernaculum and rocky crevices. The herbivorous Lizard (S. jarrovii), Mexican Emerald Spiny Lizard (S. formosus), chuckwallas were fed fruits and vegetables in large metal trays, Blue Spiny Lizard (S. serrifer cyanogenys), Crevice Swift (S. tor- attracting a number of hymenopterans. Granite Spiny Lizards (S. quatus), Crevice Spiny Lizard (S. poinsettii), and Eastern Fence orcutti) would congregate around the trays, carefully snatch the Lizard (S. undulatus). Brasfield et al. (2008) examined reproduc- at mid-body and immediately and violently bang them tive and thyroid hormone profiles after a period of brumation against the rocks. I watched hundreds of feeding episodes and and Rich and Talent (2008) recorded the effects of prey species as far as I could , no lizard was ever stung. Keeper Mark Doles on food conversion efficiency and growth in the Western Fence studied maternal care in this colony (Doles 1998). He asked two Lizard (S. occidentalis). The Coachella Valley Fringe-toed Lizard questions: how long would a female continue to rebury her nest (Uma inornata) and Colorado Desert Fringe-toed Lizard (U. no- if excavated and could she discriminate between her nest and tata) were maintained in a large outdoor exhibit with an exten- another dummy hole nearby? Read his paper to answer both sive sandy area. The lizards were removed to an indoor holding questions. A hint—females excavated multiple exploratory holes area in the winter. Foster et al. (2015) maintained and reproduced but abandoned all but one for oviposition. the Yuman Fringe-toed Lizard (U. rufopunctatus) at Phoenix Knight Anoles ( equestris) at Dallas Zoo displayed to Zoo. In central Texas, semi-arid places with limestone ledges and smaller anoles (A. sagrei, A. carolinensis) that were used as food. slabs called Texas holey rock (popular in cichlid aquaria) have Instead of trying to flee from the larger lizards, the food anoles the most beautiful male Eastern Collared Lizards (Crotaphytus sometimes displayed head bobs, lateral compression, and dew- collaris) which were kept on display at Dallas Zoo. The Reticu- lap extension. On rare occasions, they continued the display lated Collared Lizard (C. reticulatus) from the Rio Grande Valley even while being swallowed. Rosenberg (1941) covered behavior in south Texas was also exhibited. Both species fed well on small and husbandry in A. equestris and A. ricordii. One of the most lizards (Uta, Urosaurus, Sceloporus) and insects. I was surprised interesting anole exhibits showing their lifestyle was at Lincoln when a cage mate (Sceloporus serrifer cyanogenys) larger than Park Zoo in Chicago during the 1950s where a large planted ex- the Eastern Collared Lizard had been killed and swallowed, later hibit housed Brown and Green Anoles from Florida. This stately to be regurgitated after being partially digested. This predaceous enclosure was remindful of a Victorian terrarium or Wardian saurian reminded me of a miniature Tyrannosaurus rex. case, viewable from all sides, with intricate and ornate ironwork In 1890 and 1897, three specimens of Ditmars’ Horned Liz- and was the first display visible as one entered the reptile house. ard (Phrynosoma ditmarsi) were collected somewhere near the These lizards were very active: fighting, courting, ovipositing, United States–Mexican border but the site was not recorded. hatching, and feeding. One day at Dallas Zoo, I felt a tremendous The first lizard was deposited in the collection of the American pain on my left earlobe as I was speaking on the telephone. One Museum of Natural History in New York. The second and third of my beloved employees had attached a large male A. carolinen- specimens were donated to Raymond L. Ditmars who was able sis to the lobe and it insisted on hanging on for some minutes. My to observe one for about a year until it died (Ditmars 1951). Both gentle nature toward my co-workers was seriously challenged. were preserved and sent to Leonhard Stejneger at United States In my view, the most productive zoo scientist studying igua- National Museum in Washington, DC. Stejneger described the nid lizards is Allison Alberts, Chief Conservation and Research lizard in honor of Ditmars (Stejneger 1906). No additional speci- Officer for San Diego Zoo Global. She is the co-founder and past mens were found alive until 1970. In a brilliant plan to discover co-chair of IUCN Iguana Specialist Group and president of Inter- the lizards’ location, two of the preserved lizards’ stomachs were national Iguana Foundation. She has studied endangered igua- examined, and remnants, fragments, and small peb- nas in , , , Fiji, and also bles were saved and sent to specialists for identification and a Komodo Dragons. To prove my point, consult the bibliography at possible locale was pinpointed, based on known distributions the end of this paper. of these animal, vegetable, and mineral clues. With this addi- Family Agamidae.—Murphy et al. (1978) observed defensive tional information, Ditmars’ Horned Lizard was rediscovered. To behavior in the Angle-headed Dragon (Goniocephalus dilophus, pursue this amazing story, consult Ditmars (1951), Roth (1971, now Hysilurus dilophus) at Dallas Zoo. This spectacular adult liz- 1997), Lowe et al. (1971), Montanucci (1989), and Sherbrooke et ard emitted a series of grunts and hissing sounds as it leaped bi- al. (1998). Montanucci (1984, 1989) outlined breeding, captive pedally toward an intruder. Specialized motor patterns included care, and longevity of the Short-horned Lizard (P. douglassi) and lateral presentation with open mouth, tongue rolled forward, gu- Ditmars’ Horned Lizard. In 2005, the Texas Horned Lizard (P. cor- lar sac expanded, sidehopping, tail lashing, and color change. An nutum) reproduced at Fort Worth Zoo. adult male Frilled Lizard (Chlamydosaurus kingi) from Australia Lawler et al. (1994) outlined the ecological and nutritional flared its frill, stood bipedally, made a series of grunts and rushed management and Densmore et al. (1994) described a molecular toward the keeper at this Zoo (Fig. 9). In 2004, nine Javan Hump- approach for determining genetic variation in captive and natu- headed Lizards (= Chameleon Forest Dragon, ral populations of the Piebald (Sauromalus varius) chamaeleontinus) hatched at Denver Zoo. at -Sonora Desert Museum. Lawler and Jarchow (1986) Almandarz (1969) placed Bearded Dragon ( presented a captive management plan, At Metro Toronto Zoo, barbatus) eggs in a portable baby incubator and seven hatched Barker and Cranfield (1988/1989) identified Schellackia (Lainso- at Lincoln Park Zoo (Fig. 10). Van Aperen (1969) bred them at nia) sp. in chuckwallas. Aucone et al. (2003) compared two artifi- the Melbourne Zoo. Sherriff (1989) outlined the maintenance cial ultraviolet light sources used for Chuckwalla (S. obesus, now and breeding of the Inland Bearded Dragon (Pogona vitticeps) S. ater) husbandry. at Edinburgh Zoo. Morley (1992) bred the Nobbi (Amphibolurus

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Dallas Zoo in the 1970s, two Sri Lankan lizards were displayed: Mountain Horned Iguana ( stoddartii) and Sri Lanka Horned Agama (C. aspera). At the same time at Fort Worth Zoo, a spectacular adult pair of -headed Lizards ( scu- tatus) was exhibited in a large planted enclosure. In 2004, 20 Mali Spiny-tailed Lizards (Uromastyx maliensis) hatched at Detroit Zoological Institute. Watson (1969) outlined the care of Mastigure Lizards (Uromastyx) at Jersey Wildlife Pres- ervation Trust. Wheeler (1990, 1991) described husbandry of Mastigures at Oklahoma City Zoo. Morales and Dunker (2001) found tuberculosis, Mycobacterium marinum, in a group of Egyptian Mastigures (Uromastyx aegyptius) at Steinhart Aquar- ium (Fig. 11). Raphael et al. (1999) evaluated vitamin D concen- trations in Uromastyx spp. with and without radiographic evi- dence of dystrophic mineralization at New York Zoological Park. MAYR LIBRARY, MUSEUM OF COMPARATIVE , HARVARD UNIVERSITY HARVARD ZOOLOGY, COMPARATIVE OF MUSEUM LIBRARY, MAYR Coburn (1975) described post-mortem removal and artificial

ERNST ERNST hatching of Rainbow Lizard eggs () at the Cots- wold Wildlife Park, Oxfordshire UK. A small group of Common Flying Lizards (Draco volans) was kept in a large planted exhibit at Dallas Zoo where they lived for several years (Fig. 12). Males regularly displayed to each other by extending dewlaps and par- tially extending wings, and bobbing the body but I never saw them lock jaws or bite each other. Some colonies lived in the enclosure and were intermittently picked off by the lizards. Family Chamaeleonidae.—Most chameleons have been dif- ficult to keep for extended periods in captivity (Fig. 13). In 1818, the Menagerie of the Imperial Cabinet (Vienna) had chameleons from North and eggs were laid in 1845 but did not hatch. The menagerie received many chameleons as gifts from various sources between 1818‒1851. Duméril (1854–1855) found that captive chameleons fared poorly between 1851 and 1855 at the Menagerie Jardin des Plantes, Paris for although more than 160 were kept, none lived more than 13 months. Females often died from reproductive complications—no eggs were ever hatched. See Andreone et al. (2005), Andrews (2005, 2008), Andrews and Fig. 13. Illustration of Common Chameleon (Chamaeleo vulgaris, Donoghue (2004), Andrews et al. (2004), Andrews et al. (2008), now C. chamaeleon) in Albert E. Brehm’s Illustrirtes Thierleben für Ferguson et al. (2003, 2007), Karsten et al. (2008, 2009) for recent Volk und Schule. Brehm’s Illustrirtes Thierleben für Volk und Schule; success stories. bearb. von Friedrich Schödler. Mit, Abbildungen nach der Natur, aus- Johann von Fischer was the first to document copulatory geführt unter leitung von R. Kretschmer. Neue Stereotypausg. Bibliog- behavior and reproduction in Common Chameleons (Chamae- raphisches institut, Leipzig, 1875–1883 (Illustrated Animal Life for the People and the Schools; edited by Friedrich Schödler. With Illustra- leo vulgaris, now C. chamaeleon) in 1882 and 1884 (see Murphy tions from Nature, Made under the Direction of R. Kretschmer. New 2005 for English translation; Fig. 14). He attempted to incubate Stereotypical Edition, Bibliographic Institute, Leipzig.). nearly 800 eggs but most never hatched. The first Veiled Chame- leon (Chamaeleo calyptratus) obtained by the London Zoo was nobbi) at Adelaide Zoo. Montali et al. (1975) diagnosed derma- caught in Aden, South Yemen, on 15 March 1885 and donated on tophilosis in Australian bearded lizards at National Zoo. Weigel 3 June 1885 by Lt.-Col. J. W. Yerbury (Coote 2001). Sarah Kuppert (1989) maintained and bred the Superb Dragon ( su- (2013) described color changes during courtship in this species perba) at Australian Reptile Park, Gosford. He has also published at Smithsonian National Zoological Park (SNZP) (Fig. 15). Gary a helpful book on the care of Australian lizards (1988). Ferguson and colleagues published a book on care of Parson’s, Demeter (1981) reported on husbandry and breeding of the Panther, Veiled, and Jackson’s Chameleons in 2007. (Physignathus cocincinus) at National At the now-defunct Belle Vue Zoo in England (see Bennett Zoo, Dedekind and Petzold (1982) bred the species in the Ber- and Barnaby 1989 for this sad story), average longevity for 24 lin Tierpark, and Frolov and Sanoyan (1986) in Moscow Zoo. The chameleons between the years 1898–1901 was slightly over three National Aquarium in Baltimore has a large exhibit devoted to months and none survived for one year (Flower 1925). Raymond Lesueur’s Water Dragon (Physignathus lesueurii). Both Hydro- L. Ditmars had problems with chameleons which rarely survived saurus and Physignathus should be kept in large, semi-aquatic longer than five or six months (Ditmars 1933). Flower (1925) exhibits or the lizards will likely abrade their rostrals by banging found that chameleons rarely lived one year in captivity. An ex- into the exhibit glass. Lederer (1931) published an early paper on ception was the South African Dwarf Chameleon (Bradypodion ethology of the Sailfin Lizard ( amboinensis). Viss- pumilum) which have live young. In 55 years, only 15 chame- er (1984) described husbandry and reproduction of the Sailfin leons of many species out of nearly 770 survived over one year, Lizard at Rotterdam Zoo. Mitchell (1986) reproduced the Philip- a rather depressing situation. By 1928, sixteen different species pine Water Dragon (Hydrosaurus pustulatus) at Dallas Zoo. At of chameleons had been exhibited alive in London but mortality

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Fig. 14. Illustration of Common Chameleon (Chamaeleo chamae- Fig. 15. Illustration of Veiled Chameleon (Chamaeleo calyptratus) leon) from John Anderson’s Zoology of Egypt Volume First. Reptilia from John Anderson’s Zoology of Egypt Volume First. Reptilia and - and Batrachia (1898). Early records show that this species was very trachia (1898). This species is very hardy in captivity. difficult to keep in captivity. rates were high. At the Giza Zoological Gardens in Egypt where On the floor of this exhibit, a number of large Ficus trees were Flower was director, 200 chameleons were maintained at the zoo placed—each with a single chameleon—and encircled with a between 1898‒1924 but none lived for as long as four years. sheet metal barrier at the base to keep the lizards separated as See Ferguson et al. (2004) for color variation, natural history, these highly territorial lizards were incompatible. Daily feeding conservation, and captive management of the Malagasy Panther demonstrations were held and were exceedingly popular with Chameleon (Furcifer pardalis). This work included reproductive visitors. Occasionally, sarcophagid larvae would be missed by and nutritional data on five captive generations at Texas Chris- the chameleons and would escape into the flower pots where- tian University; an overall history of chameleons in captivity was upon these insects hatched in the exhibit and were later plucked included. Gary Ferguson has worked in a zoo setting for many off by the lizards. One day, a fly landed on the inside of the glass years to investigate the captive biology of chameleons (Ferguson at an adult human’s eye level and a visitor witnessed the chame- 1994; Ferguson et al. 1993, 1996, 2004). Early on when he brought leon tongue up front and personal. She emitted a scream rivaling his herpetology class to Dallas Zoo for the annual visit I asked the decibel level of a jet plane and fell backwards into the crowd, him if he had ever seen a living chameleon. He replied no, which taking a few small children to the floor with her. he said was saddening for an avid lizard ecologist. Luckily, we Gary Ferguson, Rick Hudson from Fort Worth Zoo, and I trav- had a female Jackson’s Chameleon ( jacksonii) that had eled to to collect specimens for Texas zoos and preserve a recently given birth and the neonates were being maintained series of Jackson’s Chameleons for a description of what became outside in a large screened cage. When he viewed these tiny liz- a new taxon, now Triceros jacksonii xantholopus Eason, Fergu- ards—going into I would characterize as a catatonic state, staring son, Hebrard 1988 (see Ferguson et al. 1991). We were hosted by motionless on his haunches with no perceptible movement—I Nairobi , which includes a park. knew he was doomed. I offered some to keep in his lab in Fort Chameleons were regularly collected as food for mambas and Worth and he jumped at the opportunity—the rest is history! boomslangs living in a large pit. Jim Hebrard, then in the zoology These young Jackson’s Chameleons were inaccurate at first department at University of Nairobi, kindly allowed us to stay at as they tried to catch live crickets, often missing wide right or his apartment in the city but the space was so packed with other left. Occasionally, the tongue tip stuck firmly to the cage so the herpetologists—Butch Brodie, Ron Nussbaum, Tom Madsen, Ken chameleon had to walk to the tip to reel in the tongue. The adults Dodd, and a couple of other persons—that the living quarters were kept in a large planted enclosure that covered the entire were mindful of a homeless shelter. Gary, Rick, and I spent much lobby of the reptile building at Dallas Zoo. Some of the trees time collecting on Mt. Kenya. Gary brought along some tree- were over 20 feet (6 m) high. When new males were first put into climbing equipment to scale trees but the first trial was less than the enclosure, they spotted each other immediately and quickly impressive—he ascended about six feet, lost his balance, and fell moved toward each other to engage in male-male combat (Kom- to the ground. The equipment remained unused for the rest of mentkampf). After the struggle, the dominant chased the sub- the trip. We hired local villagers to help collect for us but virtually dominant around the entire enclosure and eventually drove it all were terrified of chameleons, believing that the lizard’s sharp to the very top of the highest tree. As the dominant lizard came claws were deadly poisonous. They were clearly conflicted, a few closer, the loser released its grip, fell to the substrate, and quickly Kenyan shillings per lizard versus instantaneous death. Only one dashed away. We tested this multiple times with a series of males person actually held a chameleon—all of the others carried these and the outcome was always the same; no loser suffered inju- saurians on the end of very long branches, both holding on for ries. Females did not flee but rejected male advances by opening dear life. The villagers were stunned when we allowed these crea- mouths, lateral body compression, color change, and rocking tures to crawl on us—women screaming and children running back-and-forth. away at full speed. One surprising and unexpected fact was the

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- sity of when sleeping they become much lighter in color and stand out chame- leons against the vegetation. As we walked on the trail, we could hear in the popu- large mammals crashing through the bush. A Tree Hyrax sudden- lous city of ly emitted a call that was blood-curdling and nearly caused car- Nai- robi; diac arrest for all of us. At that moment, we realized that this was al- though our first field experience where we potentially could be killed terri- torial, and eaten by animals and so we decided to retire from collect- ing at night in dangerous settings. When we reached the lodge for a few drinks, elephants, cape buffalos, rhinos, wart hogs, and other mega-mammals were gathered around a pool illuminated at night for the tourists, and a leopard ran in front of our car as we left. Returning to our small hut for another well-needed drink on the patio after a starry night of chameleon stalking, a group of hyenas encircled us and they were much too close; we decided to retire earlier than planned. What was interesting was these predators were absolutely silent in their approach and we only knew of their presence when we turned on a flashlight and spot- ted eye-shine. Brodie and Nussbaum were studying (Afrocaecilia = taitana) in the Taita Hills near Nairobi and sug- gested that we enlist the villagers to help with collecting because they were not afraid of chameleons but rather viewed these liz- ards as a source of income. When we arrived at the village, adults and children rushed the car and gave us a verbal price list using scientific names, reproductive condition, health, and sizes; for example, a gravid female Böhme’s Two-horned Chameleon (Kin- yago boehmei) was a steal at two Kenyan shillings. Soon, we were inundated with hundreds of herps, made our choices and pur- chased them, and were frantic to leave since the locals insisted that we buy all of the remaining ones. As we left the village, peo- ple ran after the car and threw more animals through the open windows on us, free of charge! Once we were out of sight, these animals were released. The introduced Hawaiian stock of Jackson’s Chameleon was definitely Chamaeleo j. xantholophus, the taxon described by Ferguson and colleagues. At Dallas Zoo, chameleons from this place seemed to adapt more easily to captivity when compared to the Kenyan lizards. Perhaps one reason is the daily tempera- ture fluctuation is much less dramatic in Hawaii. It was so cold at night on Mt. Kenya, we slept in our jackets inside of heavy sleep- ing bags. At San Diego Zoo, Charles Shaw recorded his observations on

Fig. 16. Illustration of Flying (Ptychozoon kuhli) from A con- the reproductive biology of Chamaeleo basiliscus (now C. africa- tribution to the zoögeography of the East Indian islands, by Thomas nus) (Shaw 1959). Castle (1990, 1991) described husbandry and Barbour (1912). At Dallas Zoo, we tested the gliding behavior but we breeding of several chameleons at the Oklahoma City Zoo. Bar- were rather unimpressed with the outcome. The lizards did not crash rie et al. (1993) documented diseases of chameleons at this Zoo. to the ground when released from the top of a high ladder but the Scott Stahl (1996) covered medical management. trajectory was not dramatic. Why do some chameleons do poorly in captivity? Here is a hint from a section in one of my earlier papers (Murphy 2009): High-casqued Chameleons (Trioceros hoehnelii) were spaced “A new challenge is the recent study by Karsten and associates about 3 m (10 ft) apart within juniper bushes and hedge-rows at which explains why some imported animals could probably the vet station a few miles out of town and the bongo acclimation never have been successfully kept in captivity for extended pe- center at the base of Mt. Kenya. Jackson’s Chameleons were usu- riods. They found that Labord’s Chameleon (Furcifer labordi), ally found higher in trees, especially at night, and several males from arid southwestern Madagascar, is an “annual” lizard living might be found in the same tree. An exciting followup to our re- mostly as an egg for about nine months and then only four or search in the 1980s was recently published by Jan Stipala. He has five months outside the egg. These lizards reach sexual maturity completed a really interesting book called Mountain Dragons: In in two months. This may mean that the brief life span of chame- Search of Chameleons in the Highlands of Kenya (2014). He has leons in captivity might, for some species, actually represent the found two additional on the west side of Mount Kenya natural adult life span. So questions are asked and suggestions and at higher elevations than where we looked. given by the authors in their abstract: ‘Consequently, a new ap- Studying chameleons in Africa is unique. One night we praisal may be warranted concerning the viability of chameleon were using flashlights and headlamps to spot chameleons since breeding programs, which could have special significance for

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species of conservation concern. Additionally, because F. labor- di is closely related to other perennial species, this chameleon group may prove also to be especially well suited for compara- tive studies that focus on life history evolution and the ecologi- cal, genetic, and/or hormonal determinants of aging, longevity, and senescence.’ Flower’s statement about chameleons being short-lived reptiles written over 80 years ago was prophetic.” Final thought: When Gary Ferguson and I traveled to Mada- gascar to do research for our books (2004, 2007), we spent one evening looking at a list of all chameleons in the country in order to predict the likely future for all of them. This was not a detailed scientific study but rather based on our observations in the field, discussions with colleagues, and literature. We concluded, much to our horror, that probably two-thirds were at serious risk of ex- tinction and these were mostly taxa that lived in primary forest or fragile areas modified by humans. Weedy species such as the Panther Chameleons were common even in places with high hu- man densities so they may well provide a starting point for a new chameleon radiation. Infraorder Gekkota.—Dale Marcellini from National Zoo published several papers on acoustics, vocal and visual displays of geckos (1977, 1978). Demeter and Marcellini (1981) recorded courtship and aggressive behavior of the Streak Lizard (Go- natodes vittatus) at National Zoo. Heinrich Dathe (1942) record- ed the birth of a Panther or (Gecko gecko) at Leipzig Zoo, Brodský (1969) at Prague Zoo, and Petzold (1963) bred this species at Berlin Tierpark. I witnessed a Tokay Gecko’s bit- ing power in person as I watched the San Antonio Zoo’s curator pick one up whereupon it turned and bit him on the finger. The pressure was so great, the lizard’s eyes sank into their orbits. It held on for one hour and nothing was successful at dislodging it, including holding it under water, until it released my colleague, probably from boredom. Frolow (1981, 1987) outlined reproduc- tion of the Skink Gecko ( scincus) in Moscow Zoo. As part of initial keeper training at Dallas Zoo, new employees were told to clean this gecko’s enclosure and the impressive de- fensive display was startling to all—we were accused of being sa- Fig. 17. Illustration of Common Flat-tailed Gecko (Uroplatus dists. Benefield et al. (1981) propagated Western Banded Geckos fimbriatus) [above] and Mountain Horned Lizard (Phyrnosoma (Coleonyx variegatus) at Tulsa Zoo. From the Houston Zoo, Neit- orbiculare) from Histoire naturelle de Lacépède. Nouvelle Édition par man (1983) described captive husbandry of the tuberculate Cuvier in 1841. Many of the gekkonid species in Madagascar are at geckos of the genus Coleonyx. Risley (1989) observed breeding serious risk of extinction due to human causes. of the Gecko (Chondrodactylus angulifer) at London Zoo. Astreiko and Popovskaya (1999) from Tula Exotarium in Russia (1980) bred the Flat-tailed Day Gecko (P. laticauda) at Twycross investigated the potential of breeding and raising the African Zoo. Rundquist (1980) reproduced Koch’s Day Gecko (P. mada- Fat-tailed Gecko (Hemitheconyx caudicinctus) under labora- gascariensis kochi) at Oklahoma City Zoo. Malagasy Flat-tailed tory conditions. At Dallas Zoo, we kept the Namib Sand Gecko Geckos (P. serraticauda) reproduced at Central Park Zoo in New ( rangei) and a group of Flying Geckos (Ptychozoon York in 2003. In 2005, the endangered Yellow-throated Day Gecko kuhli; Fig. 16). Both taxa did well in captivity. We also maintained (P. flavigularis) hatched at Society. Linnetz several Cayman Island geckos with the strange standard name of et al. (1996) found gout in Giant Day Geckos. The late Sean McKe- Woodslave ( praesignis) in the family Sphaerodactyli- own from Fresno Zoo published many studies on Malagasy and dae. We displayed the lovely Marbled Bow-fingered Gecko (Cyr- Indian Ocean geckos (1982, 1984, 1985, 1989, 1992, 1994; McKe- todactylus marmoratus) from Peinsular , Marbled Velvet own and Miller 1984). Tytle (1994) presented an overview of the Gecko (Oedura marmorata) from Australia, and Walberg’s Gecko genus Phelsuma. Scott Pfaff and Sean Foley have developed an (Homopholis walbergii) from Africa. impressive program at Riverbanks Zoo, Columbia, South Caroli- Several papers on the long-term Madagascar Giant Day na, breeding many species of the endangered group of flat-tailed Gecko (Phelsuma madagascariensis) colony at National Zoo have geckos (Uroplatus) from Madagascar (Fig. 17). One realized goal been published by Béla Demeter and colleagues (1976, 1994). of this initiative is the distribution of many lizards to various zoos. Bloxam and Townson (1980), Bloxam and Vokins (1978), Lange- Leopard Geckos ( macularius) are so prolific baek (1979), Wheler and Fa (1995) and Cooper et al. (1998) de- in captivity, one could consider them a laboratory animal (see scribed breeding, research, and medical management at Jersey Thorogood and Whimster 1979; de Vosjoli et al. 2005). Author Wildlife Preservation Trust with the endangered Round Island Philippe de Vosjoli considers this species to be the first domes- Gecko (P. guentheri), the largest species in the genus. Howard ticated lizard, like goldfish and parakeets. An array of color and

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cage furniture such as vertical tree limbs for a large display. Soon the geckos began to lay eggs, which we incubated at ca. 80°F (26°C). At least 60 hatched but all were males, suggesting Tem- perature Dependent Sex Determination. The adults vocalized when threatened, a deep guttural croaking noise. I long to see a living example of the world’s largest gecko, named Kawekaweau ( delcourti) of Māaori legend, the presumably ex- tinct giant from New Zealand (Bauer and Russell 1986). The specimen was found in the basement of the Marseille Museum of Natural History, unlabeled and likely there for over a century. What was intriguing was this lizard was 54% larger than R. lea- chianus—about two feet long and thick as a man’s wrist. What an awesome display that creature would make! I tried to convince my bosses to fund a collecting trip to try and find this gecko but I was summarily dismissed as a lunatic, who had taken complete leave of his senses. At Dallas Zoo, we kept two pygopodids: Burton’s Legless Liz-

Fig. 18. Illustration of Burton’s ( burtonis) in The ard (Lialis burtonis) and Common Scaly-foot (Pygopus lepidopo- Lizards of Australia and New Zealand by John Edward Gray in 1867. dus) (Fig. 18). We fed anoles, geckos, and skinks to the Lialis and This lizard suffocates lizard prey by encircling the thoracic region I was constantly amazed by the that allowed the with its flexible jaws. It is an extremely variable species. jaws to wrap around the prey in the thoracic region to prevent breathing—this flexibility was very efficient in causing death. pattern morphs are now available on dealers’ pricelists—a pair Banks et al. (1999) managed and bred the Striped Legless Lizard of amelanotic lizards was being offered for US $1000 each. Ernie (Delma impar) at Melbourne Zoo. Wagner from Seattle Zoo wrote an interesting and important pa- per in 1980 showing skewed sex ratios based on egg incubation temperatures. See also his other papers (1974, 1979). Literature Cited Kaverkin et al. (1994) described husbandry and reproduc- tion of a eublepharid gecko, Goniurosaurus kuroiwae splendens, Alberts, A. C. 1989. Ultraviolet visual sensitivity in desert iguanas: at Moscow Zoo. We kept the African Fat-tailed Gecko in the late implications for pheromone detection. Anim. Behav. 38:129‒137. 1960s at Dallas Zoo, which has now become a staple with herpe- ———. 1990. Chemical properties of femoral gland secretions in the toculturists. One of the most spectacular eublepharids at Dallas , Dipsosaurus dorsalis. J. Chem. Ecol. 16:13‒25. Zoo was the Gecko (Aeluroscalabotes felinus). A small group ———. 1991. Phylogenetic and adaptive variation in lizard femoral was kept in a cool room ca. 72°F (22°C) ambient temperature gland secretions. Copeia 1991:69‒79. with a damp paper towel substrate. These lizards were very se- ———. 1992. Constraints on the design of chemical communication cretive and could not be placed on exhibit. As they grew, they systems in terrestrial vertebrates. Amer. Nat. 139S:62‒89. ———. 1992. Density dependent scent gland activity in desert igua- all regularly pulled off skin with their mouths and swallowed it. nas. Anim. Behav. 44:774‒776. Some of the rarest geckos in zoo collections are any of the ———. 1993. Chemical and behavioral studies of femoral gland se- eight species of the genus from New Zealand. There cretions in iguanid lizards. Brain Behav. Evol. 41:255‒260. was a colony of eleven male and ten female Green Tree Geckos ———. 1993. Relationship of space use to population density in an (N. elegans) at the National Zoo maintained by Béla Demeter and herbivorous lizard. Herpetologica 49:469‒479. colleagues. These geckos were wild-caught and sent by Auckland ———. 1994. Off to see the lizard: Lessons from the wild. Vivarium Zoo in 1985. All of these smallish lizards in the genus are spectac- 5(5):26‒28. ularly colored and relatively easy to maintain in captivity, much ———. 1995. Use of statistical models based on radiographic mea- like day geckos. All of the Naultinus survived over a minimum of surements to predict oviposition date and clutch size in rock igua- several years. The National Zoo specimens were kept as groups nas (Cyclura nubila). Zoo Biol. 14:543‒553. ———. (ed.). 2000. West Indian Iguanas: Status Survey and Conser- of four in cages measuring ca. 3 ft 3 2 h (1 m 1 m 0.6 m) × × × × vation Action Plan. IUCN-The World Conservation Union, Gland, with an ambient temperature of 72°F (22°C) and a small lamp Switzerland. provided as a thermal gradient. No courtship was observed so ———. 2002. Ten years of conservation research on Cuban rock igua- the lizards were watched throughout the night for two consecu- nas. Herpetol. Rev. 33:119‒120. tive days to see if things improved—they did not and no repro- ———. 2003. Conserving the remarkable reptiles of Guantanamo duction occurred. Many examples are pictured in Sharell (1966). Bay. In R. W. Henderson and R. 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