Gallotia Simony/ Ma Chado/ and a Comparison with the Extinct Gallotia Simony/ Simony/ from El Hierro (Canary Islands)
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HERPETOLOGICAL JOURNAL, Vol. 8, pp. 85-91 (1998) MORPHOLOGICAL VARIATION IN THE LACERTID GALLOTIA SIMONY/ MA CHADO/ AND A COMPARISON WITH THE EXTINCT GALLOTIA SIMONY/ SIMONY/ FROM EL HIERRO (CANARY ISLANDS) M. A. RODRIGUEZ-DOMINGUEZ1 , C. CASTILL02, J. J. COELL02 AND M . MOLINA-B ORJA2 1 Viceconsejeria de Media Ambiente, Servicio de Planificaci6n de Recursos Natura Les, Centro de Reproducci6n e Jnvestigaci6n de! Lagarto Gigante de El Hierro 38911, Frontera, El Hierro, Canary Islands 2Depto. de Biologia Animal, Facultad de Biologia, Un iversidad de La Laguna, 382 71 Te nerife , Canary Islands Morphological variation was investigated in 56 live adult specimens (3 1 male, 25 female) and ten dead near-hatching embryos of G. simonyi machadoi fromthe "Centro de Reproducci6n e Investigaci6n del lagarto gigante del Hierro" (Frontera, El Hierro). Dimensions, scalation and teeth traits were measured and quantified. Males and females differed significantly (multivariate analysis of variance) in most of these traits, with values for males being greater than those for females. All traits significantly increased with SVL at a greater rate in males than in females, except forbody weight, where no significant difference was found. A qualitative comparison between data from G. s. machadoi and those forten G. s. simonyi showed that formost biometric traits, ranges overlapped forthe two subspecies, but mean and maximum values were higher in G. s. simonyi. Hind limb length increased at a greater rate relative to SVL in G. s. simonyi than in G. s. machadoi. INTRODUCTION per clutch) and the reproductive breeding time occurs between May and June-July (Rodriguez-Dominguez & The lacertids from the Canary Islands are included Molina-Borja, in press). Gallotia 1973) within the endemic genus (Arnold, and Based on immunological and genetic analyses, some are represented by four living species and one or two hypotheses have been suggested to explain the origin that are extinct. Two of the living species are large (up and later diversification of Gallotia forms on different 265 G. stehlini, to mm snout-to-vent length): Schenkel islands of the archipelago (Mayer & Bischoff, 1991; 1901 (Gran Canaria) and G. simonyi Steindachner, Thorpe et al., 1993a,b). Comparing the results of the 1889 G. galloti (El Hierro). The remaining two species: latter papers with those of more recent analyses 1839 G. atlantica 1882 Oudart, and Peters & Doria, are (Gonzalez et al., 1996), different taxonomic relation 145 smaller (up to mm. SVL). ships are shown for Gallotia species. While G. galloti Two extinct species were giant lizards that have is divided into two lineages (G. g. galloti and G. g. been classified on the basis of biometry and teeth caesaris) in the study by Thorpe et al (I993a), the tradi number: G. goliath Mertens 1942 and G. maximaBravo tional subspecies are maintained by Gonzalez et al. 1953. Four extant and extinct subspecies of G. simonyi (I996). Overall, no clear picture yet exists to explain have been described. Now extinct are G. s. simonyi, the present distribution and evolution of the Canarian known only from a small islet near the NW coast of El lizards. Hierro island, and G. s. gomerana, restricted to La The study of endemic Canarian lizards can thus be Gomera island (Hutterer, 1985). A very reduced popu considered as important both from an evolutionary lation of the subspecies G. s. machadoi still persists in point of view and as a means to evaluate diffe rent the NW of El Hierro island (Lopez-Jurado, 1989). A palaeobiogeographic models. With this evolutionary new subspecies of G. simonyi has been recently discov approach in mind, the aims of the present work are: (1) ered in the NW of Tenerife (Hernandez et al., to characterize the morphological variability of living submitted). male and fe male specimens of G. s. machadoi; and (2) The first data on morphological traits of the living G. to compare these parameters with those obtained from s. machadoi were published by Machado (1985) and published and unpublished data of some specimens of Lopez-Jurado ( 1989), but due to danger of extinction of the extinct G. s. simonyi and G. goliath. The present re this population, few specimens were studied and mor sults present the first quantitative data on living phological variation has been reported only in a few specimens of the endangered G. s. machadoi. cases. The general biology, behaviour and ecology of G. s. machadoi is almost unknown. The remaining MATERIAL AND METHODS population lives on a small rocky platform on a very Fifty-six live adult specimens (3 1 males and 25 fe steep inland cliff near Frontera. The species is mostly males) and ten embryos which died close to hatching of herbivorous (Machado, 1985), oviparous ( 4-14 eggs G. s. machadoi from the "Centro de Reproduccion e 86 M. A. RODRIGUEZ-DOMINGUEZ ET AL. Investigaci6n del Iagarto gigante del Hierro" (Frontera) a Males were studied in the present work. Data on some G. s. • 220 simonyi (Bischoff, unpublished) and fossil G. goliath I • (Castillo et al. , 1994) were also compared with those of 200 f - - • -- _o : - ,_- - --- - G. s. machadoi. 1 : - D- ...J _-: Twenty-seven biometric traits were measured in all O r:/l> 180 Females the specimens using the criteria expressed by Perez Mellado & Gosa (1988). The traits were (a) Body 160 ' ' parameters: body weight (BW), snout-to-vent length 140+-�-';'-·��--��-��---.------1 (SVL), head length (HL), head width (HW), head 2 4 6 8 10 12 14 16 18 depth (HD), fore limb length (FLL), hind limb length (HLL), rostral to anterior eye edge (REE), nostril to an terior eye edge (NEE); (b) Scalation traits: masseteric scale diameter (MSD), number of temporal scales (temporalia, T), supraciliary granules (SG), supraciliary scales (supraciliaria, SS), gular scales (gullaria, G), dorsal scales (dorsalia, D), ventral longi tudinal scales (ventralia longitudinalis, VL), ventral transversal scales (ventralia transversalis, VT), femoral pores (FP), fourth finger scales (lamellae, L ); and ( c) dentary traits: number ofpremaxillary teeth (PT), max illary teeth (MAX), dentary teeth (DEN) and pterygoid teeth (PTE). A digital caliper (0.01 mm precision) was used to record the biometric data and body weight was Age (years) obtained by a dynamometer (1 g precision). All indi viduals of nine years old or less were born in captivity FIG. I. Growth curves for male (closed circles) and female in the "Centro de Reproducci6n e Investigaci6n del (open squares) G. s. machadoi. SVL (a) and body weight (b) lagarto gigante del Hierro" and maintained (usually in regressed on age. pairs) in outdoor large terraria. The fiveremaining indi viduals, aged between 12-16 years old (extrapolated presented in Fig. I. The increase in the rate of both from animals of known age and SVL), were captured in traits is greater in the earlier adult stages ( 4-8 years their natural habitat and also maintained in outdoor old), slowing down in the later ones (>8 years old). terraria. The lizards had no predator pressure and were Body traits. Mean values, standard errors, range, fe d ad libitum with plant food obtained fromthe natural sample size and coefficient of variation for different vegetation of the area and with insects and newborn body traits of embryos (E), males (M) and females (F) mice. The diet was supplemented with vitamins and are shown in Table la. Males and females differed sig calcium. nificantly when considering body dimensions (BW, Descriptive statistics were obtained for all traits. SVL, HL, HW, HO, FLL, HLL, REE and MSD) with Measurements were not independent, so male-female SVL as the covariate (MANOVA, F9 45=17.08, . trait comparison was analysed by Multivariate Analy P<0.0001). sis of Variance (MANOVA, with SVL as covariate). Scalation and teeth traits. Considering scale size Least square regression analyses (LSR) were also per and morphology, two zones are distinguished in the formedto explore the relationship between individual temporal region: (1) a central zone around the morphological traits and SVL (as the independent vari masseteric scale showing a great variability in number able) within each sex. For LSR, simple linear and size of scales; and (2) a tympanic region with small regression of log-transformed data was used. Slope size scales and nearly constant scale number. Some fe values for each regression were compared with theo males showed a very reduced scale number in the retical values (3 .0 forbody weight and 1.0 for the other central zone of temporal region, the scale sizes being of traits) by means of a !-test. Analysis of covariance a similar size to the masseteric one. (ANCOVA) was used to compare single body dimen In the embryos, dentition in dentary and maxilar sions among sexes taking into account SVL as the bones was heterodont. The first fiveto six dentary teeth covariate (all data log-transformed). are small and incisor-like. The first two had well-de RESULTS veloped cuspids pointing to the rear of the mouth. Teeth posterior to the sixth are large, wide and tricuspid with MORPHOLOGICAL TRAIT V ARIATI ONS IN G. S. well-developed lateral cuspids. Similar heterodonty is MACHA DO! found in maxillar teeth. Adult dentary and maxillar Age and weight or size relationships. The relation bones also exhibit a similar dentition. Adult pterygoid ships between snout-to-vent length and age, and body teeth are usually situated in one row but were aligned in weight variation and age, of all posthatching lizards are two rows for seven individuals. MORPHOLOGICAL VARIATION IN G. S. MA CHA DO! 87 TABLE I. Mean, standard error, range, sample size and coefficientof variation for: (a) Body Weight (BW), Snout-to-Vent Length (SVL) , Head Length (HL), Head Width (HW), Fore Limb Length (FLL), Hind Limb Length (HLL), Rostral-to-Eye Edge (REE), Nostril-to-Eye Edge (NEE); (b) Masseteric Scale Diameter (MSD), Number of: Temporals (T), Supraciliary Granules (SG), Supraciliary Scales (SS), Gular scales (G), Dorsal Scales (D), Ventral-Longitudinal scales (VL), Ventral-Transversal scales (VT), Femoral pores (F) and fourth finger Lamellae (L) of G.