Variation in body temperatures of the Common Chameleon Chamaeleo chamaeleon (Linnaeus, 1758) and the African Chameleon Chamaeleo africanus

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Variation in body temperatures of the Common Chameleon Chamaeleo chamaeleon (Linnaeus, 1758) and the African Chameleon Chamaeleo africanus Laurenti, 1768 MARIA DIMAKI', EFSTRATIOS D. VALAKOS² & ANASTASIOS LEGAKIS ³ 1. The Goulandris Natural History Museum, 13, Levidou St., 145 62 Kifissia, Greece. 2. Section of Animal & Human Physiology, Dept. of Biology, Univs. of Athens, 157 84 Athens, Greece. 3. Zoological Museum, Dept. of Biology, Univ. of Athens, 157 84 Athens,Greece. 8th International Congress on the Zoogeography and Ecology of Greece and Adjacent Regions, Kavala, Greece, 17-21/5/1999

Introduction Data on the thermal ecology of the African Chameleon Chamaeleo africanus Laurenti, 1768 and the Common Chameleon Chamaeleo chamaeleon (Linnaeus, 1758) are reported from Greece. The African Chameleon is a new species for Greece (Böhme et al., 1998). There are few field records of Chameleon body temperatures. Stebbins (1961) gives body temperatures records of captive Chamaeleo dilepis and C. namaquensis. Burrage (1973) has done most of the work on both field and laboratory thermoregulation of C. pumilus and C. namaquensis. The preferred body temperatures in a laboratory temperature gradient of the African Chameleon and the Common Chameleon were measured and compared with field body temperatures. A comparison was made between spring and summer and between the two sexes.

Distribution The distribution of the African Chameleon ranges from the Red Sea to western Mali (Central Africa) (Böhme, 1985). To the north it has reached Egypt (Joger, 1981). The presence of the species at Ramleh, close to Alexandria in Egypt, was first recorded by Anderson (1898).

Methods A total of 78 body temperatures were taken in the field. Body temperatures (Tb) were obtained using Weber quick-reading cloacal thermometer. Air temperature (Ta) 1-1.5m above the ground and substrate temperature (Ts) were measured at the site of capture. Tb was measured within 10 seconds of capture. Fieldwork at Pylos was carried out in August 1997 and April, May, June and August 1998. At Samos island, fieldwork was done in May 1997, October 1997 and June 1998. Most of the fieldwork was undertaken between 18:30-3:00 while few data were collected during the daytime. When it is dark it is more effective to find the animals using flashlights (Cuadrado, 1997). For analysis of the data, the year was divided into two seasons: spring (April- May-June) and summer (August). Seven individuals of the African Chameleons (2 males and 5 females) and two of the Common Chameleon (males) were housed in temperature gradient for studying the preferred body temperatures. Four hundred and sixteen cloacal temperatures were recorded in individuals from laboratory.

Seasonal variation in body (Tb), substrate (Ts) and air temperatures (Ta) for Chamaeleo africanus spring summer 36 36 32 32 28 28 24 24 16 16 12 Min-Max 12 Min-Max 8 Tb Ts Ta 25%-75% Medianvalue 8 Tb Ts Ta 25%-75% Medianvalue Cloacal temperatures differed significantly between spring and summer and so did substrate temperatures and air temperatures. As the months became hotter the animals reached higher temperatures (t-test, p<0.05).

Body temperatures of Chamaeleo chamaeleon and Chamaeleo africanus 36 32 28 24 The Tb of the chameleons at the field ranged from 10.4C to 31.6C while associated substrate temperatures varied from 10.2C to 33C and air temperatures varied from 11.6C to 28.1C. The minimum temperature we measured was 10,4C in C. africanus in April at 24:40, and the maximum was 31.6C in C. africanus in June. The mean Tb -at the field- of C. africanus was 23.4C while for C. chamaeleon it was 24.2C. 16 12 8 TbC.africanus TbC. chamaeleon Min-Max 25%-75% Medianvalue SUMMER Valid N Mean Minimum Maximum Std.Dev. TB 35 26,23429 23,80000 31,00000 1,376044 TS 35 27,62286 21,40000 33,00000 2,903151 TA 35 25,53714 21,40000 28,10000 1,392646 SPRING TB 43 21,11163 10,40000 31,60000 3,368779 TS 43 19,78837 10,000 32,60000 3,717283 TA 43 19,49535 11,60000 25,00000 2,958760

Field temperatures of Chamaeleo africanus 36 spring TB= 4,0178+,8571* TS Correlation: r =,9621 32 There was a significant correlation between Tb and Ts only in the spring. The slope of the equation was 0,85. There was a significant correlation between Tb and Ta in spring and summer. The slope of the equation was 0,83 in the summer and 1 in the spring. Tb 28 24 16 12 8 8 12 16 24 28 32 36 Regresion 95%confid. spring Ts 36 TB=,2430+ 1,0690* TA Correlation: r =,9327 summer TB= 5,0370+,8306* TA Correlation: r =,8407 32 32 31 28 30 24 29 28 Tb Tb 27 16 26 12 8 10 12 14 16 18 2 24 26 Regresion 95%confid. 25 24 23 21 2 23 24 25 26 27 28 29 Regresion 95%confid. Ta Ta

Noof obs Laboratory temperatures of C. africanus & C. chamaeleon C. africanus 180 31,0 160 140 1 10 Noof obs 80 60 40 0 30 10 15 25 30 35 40 Uper Boundaries(x<= boundary) C. chamaeleon 32,7 Expected Normal The preferred body temperature at the laboratory gradient ranged from 26.0C to 36.0C for C. chamaeleon and from 25.0C to 35.0C for C. africanus. The median Tb for C. africanus was 31C while for C. chamaeleon was 32.7C. 25 The mean Tb for C. africanus was 31C while for C. chamaeleon was 31.6C. 15 10 5 0 2 24 26 28 30 32 34 36 38 40 Expected Normal Uper Boundaries(x<= boundary)

Discussion There was no difference between the two sexes neither between the two species, both in field and laboratory. The Tb at the field ranged from 10.4C to 31.6C for C. africanus and from 23.5 C to 25.0 C for C. chamaeleon. Stebbins (1961) gives 21.0-36.5C for C. dilepis from South West Africa. For C. pumilus the annual range was 3.5-37.0C and for C. namaquensis was 14.0-39.7C (Burrage, 1973). The mean Tb -at the field- of C. africanus was 23.4C while for C. chamaeleon it was 24.2C, similar to other chameleon species. For C. pumilus it was 22.4C and for C. namaquensis 28.7C (Burrage, 1973). For C. dilepis from South West Africa the mean Tb was 31.2C (Stebbins, 1961). The mean Tb -at the lab- for C. africanus was 31C while for C. chamaeleon was 31.6C, which is similar to other chameleon species. For C. pumilus body temperature ranged from 7.0-30.0C (mean 25.0C), while for C. namaquensis was 18.5-36.2C (mean 29.3C) (Burrage, 1973). Stebbins (1961) gives a preferred body temperature range of 28.5-36.5C (mean 33.5C) for C. namaquensis. According to Huey and Slatkin (1976) both chameleon species are thermoconformers because the slope of the equation Tb vs Ta is near to 1.

References Anderson, J. 1898. Zoology of Egypt. I. Reptilia and Batrachia. London. Quaritch, 371 pp. Böhme, W. 1985. Zoogeographical patterns of the lizard fauna of the African Subsaharan Savanna belt, with preliminary description of a new chameleon. Proc. Intern. Symp. African Vertebr., Bonn. pp. 471-478. Böhme, W., Bonetti, A. & Chiras, G. 1998. The chameleons of the Greek mainland: taxonomic allocation and conservation needs of a second European species (Squamata: Sauria: Chamaeleonidae). Herpetozoa 11 (1/2): 87-91. Burrage, B.R. 1973. Comparative ecology and behavior of Chamaeleo pumilus pumilus (Gmelin) and C.namaquensis A.Smith (Sauria: Chamaeleonidae). Ann.S.Afr. Mus. 61, 1973: 1-158. Cuadrado, M. 1997. Efectividad de los censos nocturnos de camaleon comun. Bol. Asoc. Herpetol. Esp. No 8: 27-28. Huey, R.B. & Slatkin, M. 1976. Costs and benefits of thermal thermoregulation. Quarterly Review of Biology 51: 363-384. Ioannides, Y., Dimaki, M. & Dimitropoulos, A. 1994. The herpetofauna of Samos (Eastern Aegean, Greece). Ann. Musei Goulandris 9:445-456. Joger, U. 1981. Zur Herpetofaunistik Westafrikas. Bonn. zool. Beitr. 32 (3-4): 297-350. Ondrias, J.C. 1968. Liste des amphibiens et des reptiles de la Gréce. Biologia Gallo-Hellenica Vol. I (2): 111-135. Stebbins, R.C. 1961. Body temperature studies in South African lizards. Koedoe 4: 54-67. Acknowledgments We wish to thank the Hellenic Ornithological Society for their hospitality during some of the trips. Also many thanks to I. Schogolev, Y. Ioannidis, G. Chiras and A. Bonetti for assistance in the field work.

The African Chameleon Chamaeleo africanus from Pylos (photography: M. Dimaki).

The Common Chameleon Chamaeleo chamaeleon from Samos (photography: M. Dimaki).