Preliminary Investigations on the Reproduction of the Frillneck Lizard Chlamydosaurus Kingii in the Northern Territory

Preliminary investigations on the reproduction of the Frillneck Lizard Chlamydosaurus kingii in the Northern Territory

Gavin S. Bedford, Keith A. Christian and Anthony D. Griffiths School of Biological Sciences, Northern Territory University, P.O. Box 40146, Casuarina, Northern Territory 0811

ABSTRACT The oviposition of nine female frillneck lizards Chlamydosaurus kingii was induced in the laboratory, and the mean clutch size of 15.1 (range 12-23) was larger than previously reported for this species. There was significant variation in egg mass among clutches. Three females were relocated 3 months later using radiotelemetry and two were gravid for a second time in the same breeding

season. The soil temperatures adjacent to a natural nest ranged between 28.8 and 33.7°C with a mean Downloaded from http://meridian.allenpress.com/rrimo/book/chapter-pdf/2644360/rzsnsw_1993_019.pdf by guest on 01 October 2021 of 29.5°C in the mornings (between 0900 and 1000 h) and 31.9°C in the afternoons (between 1600 and 1700 h). The incubation period of the natural nest was 69 days compared to laboratory incubation periods of 73-80 days at 30°C and 54 days at 33°C.

INTRODUCTION dry season (32.8°C) (Chrisdan and Bedford, The frillneck lizard Chlamydosaurus kingii is submitted). one of the most conspicuous reptiles of the Shine and Lambeck (1989) found that breed- wet-dry tropical woodlands of northern Aust- ing activities of frillneck lizards begin during ralia. These lizards are among the largest the "build-up" to the wet season (October- agamids in Australia. They are sexually dimor- November), and an examination of museum phic (Shine 1990) with adult males reaching a specimens revealed that reproduction snout vent length (SVL) of 290 mm, with a extended into the wet season. They found mass up to at least 870 g. Females are much clutch sizes ranging from 4—13 (mean = 8.88; smaller, reaching a SVL of 235 mm and a mass n = 24), however the sample included animals of 400 g. A study of the ecology of the frillneck from both the Northern Territory and lizards (Shine and Lambeck 1989) revealed Queensland museums. There are differences that they are primarily arboreal spending more in the populadons of frillneck lizards between than 90 per cent of the time in trees. these two geographic regions, with those in the Terrestrial locomotion of this species is mainly Northern Territory being significantly larger bipedal (Shine and Lambeck 1989), although than those in Queensland. bipedal locomotion may be less common in juveniles which, when run on a treadmill, used The aim of this report is to provide quadrupedal locomotion almost exclusively addidonal reproductive data from frillneck lizards from the Northern Territory with (Christian and Bedford, unpublished data). respect to the timing of reproduction and Despite living in a relatively stable thermal clutch size. In a summary of the reproduction environment, frillneck lizards in the tropical of Australian reptiles, Heatwole and Taylor regions of the Northern Territory display (1987) noted that it is not known whether any markedly seasonal differences in behaviour. tropical Australian lizard species produce They are inconspicuous in the dry season multiple clutches per year. Here we report the (May-August) when they spend much of the results of preliminary investigations designed dme high in trees, and they move from tree to to determine whether the frillneck lizards in tree less frequently than in the wet season the "Top End" of the Northern Territory pro- (December to April) (Chrisdan and Bedford, duce more than one clutch per year. submitted). This change in behaviour is reflected in seasonal differences in field METHODS metabolic rates as determined by isotopic Gravid C. kingii were caught by hand in the techniques: dry season metabolic rates are wet seasons (December-March) of 1991—92 reduced to 23 per cent of wet season levels and 1992-93 in the Gunn Point area, 60 km (Christian and Green 1994). There are also east of Darwin, Northern Territory. These seasonal differences in thermoregulatory animals were temporarily housed in outdoor behaviour with the lizards selecdng signifi- pens (3 m long, 1.8 m wide and 2 m high) at the cantly higher mean midday body temperatures Northern Territory University, Darwin. In an in the wet season (36.6°C) than during the attempt to induce oviposition, animals which

Herpetology in Australia 127 were suspected of having fully formed eggs were injected with either 0.5 ml Oxytocin (Heriot Agvet Pty Ltd) or 0.5 ml Syntocin (Ilium, Troy Laboratories Pty Ltd). The injected animals were then placed in warm (approximately 30°C) 80 L plastic containers with a substrate of 50 mm damp Vermiculite. On three occasions oviposition occurred within a few minutes of injecting the hormone, but in the other cases oviposition did not occur immediately and the animals were reinjected several days later. Once oviposition occurred the eggs were immediately placed in a small plastic container with 200 g of Vermiculite and 190 ml of water Fig. I. Frillneck Lizard Chlamydosaurus kingii with a radio- to give a water potential of -300 kPa using the transmitter attached at the base of the tail. Downloaded from http://meridian.allenpress.com/rrimo/book/chapter-pdf/2644360/rzsnsw_1993_019.pdf by guest on 01 October 2021 methods of Tracy ei al. (1978). The containers were covered with perforated "Cling Wrap" RESULTS thin plastic sheeting. The containers were then placed in an incubator (Clayson) at 30°C in In March 1993 all three telemetered female 1991-92 and at 33°C in 1992-93. frillneck lizards were taken back to the laboratory. In order to determine whether more than One was not gravid, and had declined sub- one clutch of eggs can be produced during a stantially in body condition. Its transmitter was breeding season, three lizards which oviposited removed and the animal was released in the laboratory during November 1992 were immediately. The other two had gained mass fitted with location transmitters (Biotelemetry, compared to their November post-oviposition Adelaide, SA) and released at the point masses, but not enough to bring them up to of capture within one week of oviposition. their initial pre-oviposition masses. In March Transmitters weighed about 15 g (from 5 to 8 the mass of frillneck lizard number 255 was % of the non gravid animals' mass ) and were 231 g and that of number 241 was 296 g com- glued to the base of the tail using 5 minute pared to 243 and 305 g respectively before they epoxy resin (Araldite, Selleys Chemical). laid their first clutches in November (see Table Adhesive bandage was then wrapped around 1 for masses at other times). They were the tail and transmitter to hold the transmitter injected with hormone as described above, but in place. The transmitter was fitted to the tail without any result. After two weeks in captivity approximately 50 mm posterior of the vent in the animals were X-rayed then released at the an attempt to prevent it from impeding copu- point of capture. The X-rays revealed approxi- lation. However, other agamids apparently do mately 8 eggs in the larger of the two (number not need to mate a second time in a year for a 241) and approximately six eggs in number second clutch after a first fertile clutch has 255. been laid {Ctenophorusfionni, G. Johnston, pers. Table I shows the female sizes and clutch comm.; Physignathus lesueurii and Amphibolurus sizes for all the clutches obtained in the labora- diemensis, P. Harlow, pers. comm.; Pogona vitticeps, Chris Banks, pers. comm.). After tory. The clutch size for these 9 clutches being released at their site of capture the tele- ranged from 12 to 23 with a mean of 15.1 (sd metered lizards were recaptured every two = 3.5). The mass of the eggs at the time of weeks and palpated for eggs. laying ranged from 2.41 to 4.53 g. A one-way ANOVA was performed on the egg masses Staff at the Territory Wildlife Park (at Berry from different clutches, and there was a signifi- Springs, 65 km south of Darwin) observed a cant difference among them (Fg gs = 168.5, P female frillneck lizard digging a natural nest < 0.0001). A comparison of the means with on the Park grounds on November 17, 1992. Fisher's PLSD (P = 0.05) revealed that the We buried a thermocouple temperature probe mean egg mass of each of the six clutches was (type K) adjacent to the eggs which the female significandy different from that of the other buried at a depth of 20 cm. Park staff checked clutches reflecting little variation in egg mass the nest daily and read the nest temperature at within a clutch, but considerable variation irregular intervals, but usually near 0900 h among clutches. The clutch with the largest and 1600 h, using a Fluke thermocouple eggs had a mean mass of 4.4 g per egg, and thermometer in conjunction with the buried the clutch with the smallest eggs had a mean probe. mass of 2.7 g per egg. There was no significant

128 Herpetology in Australia Table 1. Data from female Chlamydosaurus kingii and their clutches. As described in the text, three females were recaptured using radiotelemetry in order to determine whether they produced a second clutch. The size of the second clutch for those two females known to have produced two clutches is given in parentheses. Non-gravid masses were measured after oviposition.

Female mass (g) Mean egg Lizard Number SVL (mm) (non-gravid) Mass(g)(sd) Clutch size Second Clutch?

FN60 225 308 4.4 (0.09) 16 FN79 202 237 2.9 (0.12) 12 FN89 220 254 3.9(0.33) 12 FN90 189 170 3.6(0.19) 12 FN74 232 294 3.4(0.13) 23 FN 84 220 272 2.7(0.20) 17 FN243 185 186 15 No FN255 203 188 13 Yes (6) FN241 217 254 16 Yes (8) Downloaded from http://meridian.allenpress.com/rrimo/book/chapter-pdf/2644360/rzsnsw_1993_019.pdf by guest on 01 October 2021 correlation between mean egg mass within a never observed aggregations of adults. Hatch- clutch with any of the following parameters: lings of the iguanid lizards Iguana iguMna female SVL (P = 0.72), non-gravid female (Burghardt et al. 1977, Greene et at. 1978) and mass {P = 0.61), clutch mass {P = 0.65), or Conolophus pallidus (Christian and Tracy 1982) clutch size (P = 0.91). Thus, there are no have been reported to remain close together obvious factors related to female size or repro- for several days after leaving the nest. Greene ductive effort that account for the large varia- et al. (1978) suggest that the aggregations of tion in egg mass. hatchling iguanas may serve a function related to defense against predation. The incubation period in the laboratory at 30°C was 73-80 days, and at 33°C it was 54 days. The incubation period in the natural nest DISCUSSION was 69 days. Sixteen of the hatchlings Clutches were laid in the laboratory from incubated in the laboratory were measured at late November to mid March. We observed the time of hatching, and the mean SVL was gravid females as late as April in the field. 48 mm (range: 42—51 mm), and the mean mass Although the sample is small, the two teleme- was 3.6 g (range: 2.4-4.6 g). tered females that were gravid for a second The natural nest was located on an area of time in one season indicate that multiple flat, coarse-grained sandy soil. The site had clutches are possible for this species. A frillneck some sparse grass and leaf litter, but there was lizard originally from the Northern Territory no vegetation directly over the nest and it but held in captivity at the Melbourne Zoo has received direct sun most of the day. A stringy- also laid two clutches per season during two bark gum Eucalyptus tetrodonta was 1.5 m from years (Christ Banks, pers. comm.). A sample of the nest, and there were several sparse shrubs 14 species of agamids indicates that all but one (Calytrix) also 1.5 m from the nest. The other have more than one clutch per year (Dunham characteristics of the natural nest are listed in et al. 1988). In some lizard species the number Table 2. After the eggs from the natural nest of clutches varies from year to year, presum- hatched there were 10 empty eggshells in the ably in response to factors such as rainfall and nest and eight hatchlings sitting in Calytrix food availability (Ballinger 1977, 1983; Turner shrubs 1.5 m from the nest site. On six other 1977; DeMarco 1989). Whether two clutches occasions we have observed groups of hatch- per year is the norm for frillneck lizards is not lings perched in shrubs or small trees. We have known, but given what is known from other

Table 2. Data collected from a natural nest of the Frillneck Lizard Chlamydosaurus kingii. Soil temperatures were measured adjacent to the eggs at the same depth, and they were collected across the entire incubation period.

Depth of nest 10-15 cm Date eggs laid 17-11-92 Date eggs hatched 24-01-93 Clutch size 10 Maximum soil temperature 33.7°C at 1715 h on 13-12-92 Minimum soil temperature 28.8°C at 0910 h on 13-12-92 Mean soil temperature at 0900-1000 h 29.5°C(n= 10,sd= 1.2) Mean soil temperature at 1600-1700 h 31.9°C(n= 13,sd= 1.3)

Herpetology in Australia 129 species it is poissible that the number of ACKNOWLEDGEMENTS clutches per year depends on the availability of We thank Brian Cande and the other staff resources. at the Territory Wildlife Park for locating the natural nest and collecting temperature data. The clutch sizes of the two telemetered We thank Mike Thompson for advice on eggs females dropped from 13 and 16 for the first and Roger Haensel for the X-rays. This work clutch to six and eight respectively for the was done under permit from the Conservation second clutch. A number of species of lizards Commission of the Northern Territory and the are known to produce second clutches of equal Northern Territory University Animal size to the first (Vitt 1977), but in other species Experimentation Ethics Committee. The Aust- late season clutches are smaller than first ralian Research Committee and the NTU have clutches (Nussbaum 1981, and references provided financial support for studies of therein). In those species in which clutch size frillneck lizards. decrease, however, the decrease is much smaller than the 50 per cent decrease shown REFERENCES

by the two frillneck lizards (Jackson and Downloaded from http://meridian.allenpress.com/rrimo/book/chapter-pdf/2644360/rzsnsw_1993_019.pdf by guest on 01 October 2021 Telford 1974; Nussbaum 1981). BALLINGER, R. E., 1977. Reproductive strategies: food availability as a source of proximal variation in a lizard. 58: 628-35. The clutch sizes we observed were markedly Ecology larger than those reported by Shine and BALLINGER, R. E., 1983. Life history variations. Pp. 241-260 in Lizard Ecology: Studies of a Model Organism ed by R. B. Lambeck (1989) from a sample of 24 lizards Huey, E. R. Pianka, and T. W. Schoener. Harvard from the Northern Territory and Queensland. University Press: Cambridge, Massachusetts.

Although the inclusion of animals from BURGHARDT, G. M., GREENE, H. W. AND RAND, A. S., 1977. Queensland could possibly account for a lower Social behavior in hatchling green iguanas: Life at a mean clutch size (assuming the smaller reptile rookery. Science 195: 689-91. Queensland frilled lizards produce smaller CHRISTIAN, K. A. AND TRACY, C. R., 1982. Reproductive clutches), but that would not account for the behavior of the Galapagos land iguana Conolophtts pallidus on Isla Santa Fe, Galapagos. Pp 366-79 in fact that the maximum clutch size of 13 Iguanas of the World; Behavior, Ecology and Conservation reported by Shine and Lambeck (1989) is ed by G. M. Burghardt and A. S. Rand. Noyes smaller than the mean clutch size we observed. Publishers: Park Ridge, New Jersey. It seems unlikely that a sample of lizards from CHRISTIAN, K. A. AND BEDFORD, G. S. Seasonal changes in museums would be biased toward animals con- thermoregulation by the Frillneck Lizard, taining their second clutch of the season. As Chlamydosaurus kingii, in Tropical Australia, (submitted). stated above, there can be large variations CHRISTIAN, K. A. AND GREEN, B., 1994. Seasonal energetics among years with respect to lizard fecundity, and water turnover of the frillneck lizard, and it is possible that the two years of this study Chlamydosaurus kingii, in the wet-dry tropics of Australia. Herpetologica (in press). were unusually productive compared to the years represented by the sample of gravid CHURCHILL, S. K., 1991. The ecology of the orange horse- shoe bat, Rhinonycteris aurantius. MSc Thesis, University females in museum collections. of Queensland. Many, if not the vast majority of lizards fall DEMARCO, V. G., 1989. Annual variation in the seasonal shift in egg size and clutch size in Sceloporus woodi. into two groups with respect to their repro- Oecologia 80: 525-32. ductive strategies: (1) early maturing, multiple- DUNHAM, A. E., MILES, D. M. AND REZNICK, D. N., 1988. Life brooded, and (2) late-maturing, single-brooded history patterns in squamate reptiles. Pp. 441-522 in (Tinkle et al. 1970). Lizards that reproduce at Biology of the Reptilia ed by C. Gans and R. B. Huey. an age greater than one year are termed "late- Alan R. Liss: New York. maturing" (Tinkle et al. 1970). Frillneck lizards GREENE, H. W., BURGHARDT, G. M., DUGAN, B. A. AND RAND, reach a SVL of 80-120 mm at the end of their A. S., 1978. Predation and the defensive behavior of first year (unpublished data), but females in the green iguanas (Reptilia, Lacertilia, Iguanidae). J. Her- Northern Territory are not sexually mature petol. 12: 169-76. until they reach a SVL of 175 mm (Shine and HEATWOLE, H. F. AND TAYLOR, J., 1987. Ecology of Reptiles. Lambeck 1989). Thus, frillneck lizards appear Surrey Beatty & Sons, Sydney. to be late-maturing but possibly multiple- JACKSON, J. F. AND TELFORD, S. R. Jr., 1974. Reproductive brooded. This is also the case for water dragons ecology of the Florida scrub lizard, Sceloporus woodi. 1974: 689-94. (P. Harlow, pers. comm.) and bearded dragons Copeia (Badham 1971, as cited in Heatwole and NUSSBAUM, R. A., 1981. Seasonal shifts in clutch size and Taylor 1987). It appears that these large Aust- egg size in the side-blotched lizard, Uta stansburiana Baird and Girard. Oecologia 49: 8-13. ralian agamids have a reproductive strategy that differs from the lizards considered in the SHINE, R., 1990. Function and evolution of the frill of the frill neck lizard, Chlamydosaurus kingii (Sauria: early review by Tinkle et al. (1970). Agamidae). /. Linn. Soc. 40: 11-20. 130 Herpetology in Australia SHINE, R. AND LAMBECK, R., 1989. Ecology of frillneck TURNER, F. B., 1977. The dynamics of populations of lizards, Chlamydosaurus kingii (Agamidae), in tropical squamates, crocodilians and Rhynchocephalians. Australia. Anst. Wildl. Res. 16: 491-500. Pp. 157-264 in Biology of the Reptilia ed by C. Gans and D. W. Tinkle. Academic Press: New York. TINKLE, D. W., WILBUR, H. M. AND TILLEY, S. G., 1970. Evolutionary strategies in lizard reproduction. Evolution ViTT, L. J., 1977. Observations on clutch and egg size and 24: 55-74. evidence for multiple clutches in some lizards of south- western United States. Herpetologica 33: 333-38. TRACY, C. R., PACKARD, G. C. AND PACKARD, M. J., 1978. Water relations of chelonian eggs. Physiol Zool. 51: 378-87. Downloaded from http://meridian.allenpress.com/rrimo/book/chapter-pdf/2644360/rzsnsw_1993_019.pdf by guest on 01 October 2021

BOOK REVIEWS

Skinks of the Northern Territory The skinks of the Northern Territory must qualify for P. Homer this description par excellence, and I freely admit to abject failure in many of my own attempts to identify these Northern Territory Museum of Arts and Sciences scaly little Territorians over the years. Unfortunately, Handbook Series Number 2: 1-174 pages, 1992 reliable identification is the first step upon which any Recommended retail price 119.95 plus postage other scientific endeavour rests: there is litde point in ISBN 0 7245 2681 1, ISSN 1034-4403 carrying out detailed behavioural, ecological or physiological work if you do not even know what species The Australian continent has a remarkably distinctive you are using, or (even worse) whether you are fauna, and this statement is just as true of its reptiles as inadvertently "lumping" two or more separate species for the better-known mammals and birds. Indeed, one under the same name. I fell into this trap myself once in of the most extraordinary examples of co-existence of the Territory with goannas, and am not alone in my closely-related species anywhere on the face of our planet embarrassment. The history of Australian herpetology occurs among the spinifex grasslands of arid Australia. contains many sorry tales of detailed studies later invali- The animals in question are scincid lizards, and especially dated by the discovery that the original work had failed the genus Ctenotus. Although these elegant speedsters are to distinguish species correctly. rarely seen by casual tourists, detailed studies have shown that even quite small areas may contain several Ctenotus This book is a superb effort. Perhaps it is only those species. Even to an expert, these little lizards are difficult of us who have struggled in vain to identify nondescript to tell part, and new species are still being described at a tropical skinks who will fully appreciate the quality of the rapid rate. Although the phenomenon of high species- scholarship in this work, or recognize the years of richness has attracted wide international scientific atten- original research necessary for a compilation such as this. tion to these cryptic little skinks, we are still a long way But even someone with only a passing interest in reptiles, from understanding why Australian deserts support so who wants to find out the identity of a particular animal many species of lizards. The 1992 Whitley Award Best encountered in the Territory, should be able to do so Field Guide will help considerably to resolve questions with this beautifuly produced little book. Paul Horner like this one, because at a single stroke it makes this very has spent almost twenty years in the Northern Territory, confusing group of animals much more accessible for and has seen the level of understanding of the reptile study. fauna advance enormously over that period. Indeed, he has been one of the most active participants in that Any field naturalist knows that a good field guide is advance, and has now synthesized the available indispensable, especially when one is dealing with a information (much of it from his own observations) to diverse and complex group where the characters produce a book that could not even have been imagined distinguishing different species are often very subtle. twenty years ago. Herpatology in Australia 131 The general organization of the book is straight- The Biology and Evolution of Australian Lizards forward, and it is clearly written with the aim of utility Allen E. Greer in the field. The opening chapters are brief and thank- Surrey Beatty & Sons Pty Ltd, Sydney fully, we are spared the kind of rambling discourse on 264 pages, Recommended retail price $60 biology of the organisms that occupies the first part of many field guides. The author has wisely avoided this temptation. The relevant protective legislation is briefly Australian snakes and lizards have become increasingly reviewed, the characters explained, and then the book popular as the topic for general publications. Invariably, dives straight into a key to the genera of Northern these books or magazines focus on the photogenecity of Territory skinks. The key is designed for someone who Australian reptiles. Large format, full colour, glossy folios have been the preferred mode to present Australian is prepared to take on the full scientific nomenclature reptiles to a buying public. While this form of publication but this is inevitable given the nature of the traits that raises people's appreciation of these creatures they are distinguish taxa in this confusing group. Any character- rarely backed up by accurate or up-to-date biological infor- istics that may be problematic are illustrated, and illus- mation. trated well. This is an enormous boon to the non-expert reader. Overall, keying out a lizard in your hand still will Unfortunately, the convention has been to dish up the not be EASY, but anyone with perseverance and same hackneyed collection of folk lore, supposedly to give common sense should make it through. I managed to the publication some biological credibility. Rarely is there key out a Ctenotus in a couple of minutes, and that is a any attempt to verify the information that is portrayed. feat that I have rarely achieved in my professional life. The reason for this has been that often the authors of Downloaded from http://meridian.allenpress.com/rrimo/book/chapter-pdf/2644360/rzsnsw_1993_019.pdf by guest on 01 October 2021 these articles have little expertise or experience with the The heart of the book is the specific account of the animals that they write about. Myths and misinformation species. For each taxon, the author gives a detailed continued to be perpetuated in a time-honoured cycle. description and definition, a distribution map (with specific points, not shading!), and some brief notes on 1990 has seen a severing of the cycle. Allen Greer's "The habitat preferences (often very useful in identifying these Biology and Evolution of Australian Lizards" ignores the animals) and on behaviour, ecology, reproduction, myths and concentrates on verifiable biological data. It is abundance and the like. Paul's firsthand experience with a fantastic compilation of information, gleaned from these animals is very obvious here, and much of the various sources including the normally unheralded band information has not been published previously. Colour of amateur herpetologists who are bound to relish the con- photographs of each species are provided, and the tents of this book. Greer has attempted to synthesize date general standard of the photographs is excellent. In from widely different scientific planes and to gel this into many cases a lizard can be identified simply from the a realistic presentation about the biology of Australian photograph. The photographs are combined in double- lizards.. Gaps in the data are pointed out. page spreads, facilitating comparisons of similar species. "The Biology and Evolution of Australian Lizards" is Finally, the book finishes with a glossary, bibliography, not going to be everyone's "cup of tea". Not too many index, and appendices on geographic distributions. people, even those with a zoological bent, would lie awake Attention to detail is obvious throughout the volume, at night thumbing through tables detailing lizard snout- and I could not find any significant mistakes at all. One vent lengths or listing the number of pre-sacral vertebrae interesting sideline, that show the quality of the book for an impressive number of species. The book is heavily relative to some others, comes from the fact that two of factual but it is surprisingly readable. Amateur and pro- the photographs used in this book {Ctenotus amhemensis fessional herpetologists will devour the contents with glee. and Egernia greeri) appeared in print simultaneously in Greer's penchant for phylogenetics underlies his treat- another book, the Ehmann (1992) reptile volume of the ment of each major reptile group. In the same chapters prestigious Australian Museum series "Encyclopedia of are parcels of basic but important morphological details Australian Animals". Unfortunately, both photographs that give this publication the appearance of a textbook. To are misidentified in Ehmann's book, as C. tanamiensis and some extent that is what this book is. The author is clearly E. richardi. Paul Horner's book not only has the correct trying to set a standard for subsequent herpetological liter- identifications, but provides locality data for the speci- ature. Being true to his science, Allen Greer has liberally mens in the photographs. We have seen an enormous laced the text with extensive reference citations. The explosion of field guides on Australian animals over the bibliography alone is informative and current. It is a pity last few years, but few of them can stand comparison that a number of small editing and printing errors exist in with this volume. It deals with an enormous group of the text, but these have been detailed in the author's errata. species that are very difficult to distinguish from each other, are poorly-known ecologically, and that occur over Some hackles will be raised through Greer's recognition a huge geographic area. It is a landmark effort, and ably and use of certain sensitive generic names. I do not see this sets the stage for more detailed research on these as a drawback, but rather as a practical attempt to deal with magnificent litde animals. a messy saga in herpetological systematics. Allen Greer has not been shy about recognizing the source of his data (or Dr Rick Shine his systematic tags). This,speaks heavily of his determina- Department of Zoology tion to be as objective as possible throughout the book. University of Sydney In summary, Allen Greer has done Australian (Reprinted from Aust. Zool. Vol. 28(1-4) herpetology an enormous service. The book is certainly an accurate documentation of current knowledge in this area. It is bound to be a significant reference now and many years in the future. The contribution that his book makes to Australian zoology was recently recognized by the awarding of the 1990 Whitley Medal to Allen Greer and Surrey Beatty & Sons. No Australian herpetologist can afford to be without a copy.

Arthur White (Reprinted from Avst. Zool. Vol. 26(3-4) 132 Herpetology in Australia