Morphological variation in Gallotia atlantica from the volcanic island of Lanzarote: subspecies designations and recent lava flows

A detailed uni‐ and multivariate analysis of within‐island geographical variation in scalation and body dimensions of the lacertid lizard, Gallotia atlantica, from Lanzarote (and two neighbouring offshore islets) was carried out. Two main morphological groupings were detected: four populations clustered within a putative malpaís group, i.e. from relatively recent volcanic lava fields (seven populations were sampled from these areas), while the other 19 populations (including the three remaining malpaís populations, and those from the offshore islets of Montaña Clara and Alegranza) also clustered together. Thus, while there is a tendency for occupation of malpaís areas to be related to morphological variation, this model does have some inconsistencies. The differentiated malpaís group comprises populations from two geographically isolated areas, one from the central‐western part of the island (Timanfaya) and the other from the north (Malpaís de la Corona). The divergence of these populations is considered to have arisen recently, rather than their being relicts of an ancient, formerly widespread, lineage. The morphological variation partially supports the previous use of two different subspecies to describe the within‐island variation. However, if such a scheme were applied then one of the subspecies would need to encompass populations from the geographically separated southern Timanfaya and Malpaís de la Corona areas, as opposed to just the latter. We reject previous observations that either malpaís individuals in general, or those corresponding to the north‐eastern subspecies, are larger than individuals from other areas. The pattern of morphological variation of G. atlantica within Lanzarote is less pronounced but shows some similarities with patterns of morphological variation in lizards from neighbouring islands. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 85 , 395–406.     

Recent volcanism and mitochondrial DNA structuring in the lizard Gallotia atlantica from the island of Lanzarote

The phylogeography of the lacertid lizard Gallotia atlantica from the small volcanic island of Lanzarote (Canary Islands) was analysed based on 1075 bp of mitochondrial DNA (mtDNA) sequence (partial cytochrome b and ND2) for 157 individuals from 27 sites (including three sites from neighbouring islets). Levels of sequence divergence were generally low, with the most distant haplotypes separated by only 14 mutational steps. MtDNA divergence appears to coincide with formation of the middle Pleistocene lowland that united formerly separate ancient islands to form the current island of Lanzarote, allowing rejection of a two-island model of phylogeographical structure. There was evidence of large-scale population expansion after island unification, consistent with the colonization of new areas. A nested clade phylogeographical analysis (NCPA) revealed significant phylogeographical structuring. Two-step and higher-level clades each had disjunct distributions, being found to the east and west of a common area with a north-south orientation that extends between coasts in the centre-east of the island (El Jable). Other clades were almost entirely restricted to the El Jable region alone. Bayesian Markov chain Monte Carlo analyses were used to separate ongoing gene flow from historical associations. These supported the NCPA by indicating recent (75,000-150,000 years ago) east-west vicariance across the El Jable region. Lava flows covered El Jable and other parts of the central lowland at this time and likely led to population extinctions and temporary dispersal barriers, although present-day evidence suggests some populations would have survived in small refugia. Expansion of the latter appears to explain the presence of a clade located between the eastern and western components of the disjunct clades. Direct relationships between mtDNA lineages and morphology were not found, although one of two morphological forms on the island has a disjunct distribution that is broadly concordant with east-west components of the phylogeographical pattern. This work demonstrates how recent volcanic activity can cause population fragmentation and thus shape genetic diversity on microgeographical scales.     

Within-island geographic variation of the dangerous Taiwanese snake, Trimeresurus stejnegeri, in relation to ecology

Multivariate analysis shows that geographic variation occurs in the scalation and body proportions of the dangerously venomous snake Tritneresurus stejnegeri , within the island of Taiwan. Canonical correlation analysis reveals a high correlation between constellations of morphological characters and ecological factors. Numerical hypothesis testing, using partial Mantel tests, provides evidence of a causal relationship with ecology. Head shape was found to be primarily associated with mean annual temperature and altitude, and head scalation with the annual range of temperature. Body scalation was found to be influenced by altitude, mean annual temperature and mean annual rainfall. Tail length is primarily associated with the range of temperature and rainfall and secondarily with the mean annual temperature and altitude. Geographic proximity was found to be less important in the explanation of the observed geographic pattern than some ecological factors, supporting the hypothesis of an ecogenetic origin of morphological variation.     

Lipophilic compounds in femoral secretions of males and females of the El Hierro giant lizard Gallotia simonyi (Lacertidae)

Many lizards use femoral gland secretions in reliable intraspecific communication. Based on mass spectra, obtained by GC–MS, we found 57 lipophilic compounds in femoral secretions of males and females of El Hierro giant lizards, Gallotia simonyi (fam. Lacertidae). Compounds included steroids (mainly cholesterol) and fatty acids ranging between n-C₁₆ and n-C₂₂ (mainly hexadecanoic and octadecanoic acids), followed by aldehydes, alcohols, ketones, squalene and waxy esters. There were important intersexual differences in the presence, abundance and number of compounds (more numerous in males). Males had higher proportions of the most odoriferous compounds (fatty acids and aldehydes), while females had higher proportions of more stable compounds (steroids, waxy alcohols, waxy esters and terpenoids). This suggests sexual differences in function of femoral secretions. In addition, some compounds could reflect the physiological state, allowing monitoring health of lizards from secretions samples, which is especially important given the critical conservation status of this lizard.     

Distribution of land snails (Mollusca, Gastropoda, Pulmonata) on the island of Gran Canaria (Canary Islands) in relation to protected natural areas

The oceanic island of Gran Canaria, part of the Macaronesian Archipelagos, is highly deteriorated, a situation which has had negative repercussions for endemic species. A recent law, however, established 32 Protected Natural Areas in an attempt to remedy the problem. In the present work we show the distribution of common (non-endemic) land snails on the island. The snails were largely introduced by man over the past five centuries. A comparison between the distribution of the endemic land snails and the protected areas shows the latter to be generally adequate. We also suggest some modifications to the law in two Areas and the addition to the list of two further Areas so as to guarantee protection of several endemic species currently under threat (according to the criteria of the IUCN): 2 critically endangered, Hemicycla saulcyi saulcyi and Napaeus isletae, 2 vulnerable, Theba arinagae and Obelus despreauxii, and 1 lower risk, Hemicycla berkeleyi.     

Fossil lizard from central Europe resolves the origin of large body size and herbivory in giant Canary Island lacertids

The endemic Canary Island lizard clade Gallotia, which includes the largest members of Europe's dominant reptile group, Lacertidae, is one of the classic examples of insular gigantism. For the first time we use fossil data to test the evolutionary reasons for the association between gigantism and herbivory. We describe an almost completely preserved skeleton of J anosikia ulmensis comb. nov. from the early Miocene of Ulm, Germany (MN 2a, ～ 22 Mya). We show that this species and Oligocene Pseudeumeces cadurcensis (Filhol, 1877) are in fact crown lacertids, and the first known pre‐Quaternary record of the total clade of Gallotia. Pseudeumeces confirms the early origin of crown Lacertidae in the Palaeogene of Europe. More importantly, these fossil taxa show that large body size was already achieved on the European mainland by the early Miocene. Furthermore, Pseudeumeces and Janosikia were faunivorous, thus demonstrating that insularity, not large body size, was crucial to the evolution of herbivory in this lineage. Body size change in Gallotia was more complex than previously thought, encompassing size increase [e.g. in the extinct Gallotia goliath (Mertens, 1942)], but more commonly involving miniaturization. The physical environment may play a crucial role in modulating the evolution of body size in this natural laboratory.     

Within-island microgeographic variation in the colour pattern of the skink,Chalcides sexlineatus: Pattern and cause

The colour pattern of the Gran Canarian skink is described with eight independent colour pattern characters. Significant geographic variation occurs in each character. There are generally high levels of congruence between the patterns of geographic variation in each character although some differences exist. In canonical variate analyses, the first canonical variate expresses most of the among-locality variation in colour pattern, indicating a largely unidimensional pattern. Patterns of geographic variation in the colour pattern are portrayed by contouring. This reveals north-east/south-west clines for seven of the individual characters and the generalized pattern (CV1). Four causal hypotheses were erected which predicted four different unidimensional patterns of geographic variation. Mantel tests and partial correlation analyses were used to compare the observed patterns of microgeographic variation with the four hypothesized patterns. This method suggests differential selection occurring between ecotones as the cause of the microgeographic variation. Microgeographic variation in some aspects of the colour pattern can be explained by selection for different anti-predator strategies in the hot, arid southern areas vs the cooler, lusher northern areas.     

Within-island microgeographic variation in body dimensions and scalation of the skink Chalcides sexlineatus, with testing of causal hypotheses

Correlations, univariate analyses, multiple group principal component analyses, canonical analyses, contours and transects arc used to describe microgeographic variation in body dimensions and scalation of the scincid lizard Chalcides sexlineatus within the island of Gran Canaria. Patterns of geographic variation in both character systems show similar north-east/south-west clines, scalation variation is also altitude-related. Canonical variate analyses showed that the generalized patterns of geographic variation in scalation and body dimensions were largely unidimensional. Four hypotheses, each one predicting a unidimensional pattern of microgeographic variation, were tested simultaneously against each described pattern using partial correlation. This method did not reject north/south differences in climate and/or vegetation as a cause of the microevolution in body dimensions. Similarly, altitude and north/south differences in climate/vegetation were not rejected for scalation. Partial correlation rejected the hypothesis that there are two species of Chalcides with different ranges within Gran Canaria. Mantel tests rejected the altitude hypothesis alone as a cause of the generalized geographic variation in body dimensions but did not reject any of the proposed hypotheses as being causal for the generalized geographic variation in scalation. A consideration is made of how habitat variation could cause differential selection and comparisons drawn with a parallel pattern of microgeographic variation found in the colour pattern, scalation and size of a lizard on a neighbouring island.     

Geographic variation in antisnake tactics: the evolution of scent-mediated behavior in a lizard

We used modern comparative methods to examine the evolution of scent-mediated antisnake behavior in the rock-dwelling velvet gecko (Oedura lesueurii). The selective agent is a snake species (broad-headed snake, Hoplocephalius bungaroides) that feeds primarily on velvet geckos by remaining sedentary in rock crevices for days or weeks, waiting to ambush lizards. The past and present distribution of this predator is well documented because of its threatened conservation status. We used this information to sample lizards from three populations distributed with snakes (sympatric) and three populations that appear never to have been distributed with snakes (allopatric) in each of two widespread but geographically distinct genetic groups of velvet gecko (as determined using allozyme electrophoresis). Wild-caught immature geckos from sympatric populations showed higher tongue-flick rates and stronger shifts in locomotion (increased duration of crawling and remaining stationary while pressed against the rock) toward snake-scented rocks than did lizards from allopatric populations. However, predation environment did not significantly affect a lizard's tendency to display other typical antisnake tactics such as tail waving or fleeing. These results were highly repeatable across the two sampled genetic groups of velvet gecko, despite demonstrable genetic divergence between groups. Experiments with hatchling lizards that had no experience with predators indicate that qualitative components of antisnake behaviors are probably inherited. The method of phylogenetically independent contrasts strongly suggests that the presence or absence of snakes has driven the evolution of behavior in velvet geckos. Collectively, these results provide support for an often suggested but speculative expectation that prey can adapt to predation pressure on a local scale.     

Patterns of nocturnal movement of the long-spined sea urchin<Emphasis Type="Italic"> Diadema antillarum</Emphasis> (Philippi) in Gran Canaria (the Canary Islands,central East Atlantic Ocean)

We tagged individuals of the sea urchin Diadema antillarum (Philippi) around the island of Gran Canaria (The Canary Islands) during winter 2001–2002 using a new technique, consisting of the insertion of a hook fastened to a fishing line into the aboral pole (periproctal membrane). This allowed individual identification of tagged sea urchins. The goals were: (1) to quantify nocturnal movements and the homing behaviour of this echinoid on shallow rocky bottoms, and (2) to assess short term spatial and temporal variability of these movements. Tagged sea urchins displayed clear homing behaviour. The mean distance travelled at night was 3.7±1.2 m (range 1.0–5.1 m). Mean speed of nocturnal movement was 33±26 cm h^–1 (range 5–110 cm h^–1). We observed greater movement at midnight than at the beginning and the end of the night.Communicated by H.-D. Franke     

Road edge effect on the abundance of the lizard <Emphasis Type="Italic">Gallotia galloti</Emphasis> (Sauria: Lacertidae) in two Canary Islands forests

Transportation infrastructure is a main cause of environmental change in forest landscapes worldwide. In the Canary Islands, a dense road system fragment the native Canarian pine and laurel forests causing potential changes in population densities of endemic lacertid lizards (genus Gallotia). Our aim was to assess road edge effects on relative abundance patterns of the endemic Gallotia galloti in both forests. We also explored the species–habitat relationships in this road-fragmentation context. We found that lizard relative density in relation to road edges differed between forests. Lizards were more abundant along edges and leeward interior, but virtually absent from the interior of the windward laurel forest. In the pine forest, lizards were present at three distances from edge, with a net decrease in abundance from edge to interior. These patterns may be explained partly by differences in vegetation structure regarding road proximity in each forest that potentially affect the helio- and thigmothermic character of G. galloti, and thus its habitat use. A general suggestion of this study is that road margins create corridors that may be used by native lizards for dispersal through inhospitable forest matrix. The high road density in Tenerife may have negative implications for the conservation of the genetic variability of G. galloti. At the island scale, increased communication between lizard populations through road corridors might increase homogenization of the gene pool. Ecological processes in which this lizard plays important roles may also experience changes along road edges.     

Conservation implications of high genetic variation in two closely related and highly threatened species of Crambe (Brassicaceae) endemic to the island of Gran Canaria: C. tamadabensis and C. pritzelii

We used data from 12 allozyme loci for two endemic Brassicaceae from Gran Canaria (the endangered narrow endemic Crambe tamadabensis and its more widespread congener C. pritzelii) to assess whether their genetic diversity patterns reflect their phylogenetic closeness and contrasting population sizes and distribution areas, and to derive conservation implications. Genetic diversity values are high for both species and slightly higher in C. tamadabensis, despite its narrow distribution in north‐western Gran Canaria. At odds with the generally high interpopulation diversity levels reported in Canarian endemics, values of G_ST in C. tamadabensis and C. pritzelii are rather low (0.067 and 0.126, respectively). We construe that the higher genetic structure detected in C. pritzelii is mainly a result of unbalanced allele frequencies and low population sizes at the edges of its distribution. The overall high allozyme variation detected in C. tamadabensis and C. pritzelii is nevertheless compatible with an incipient but consistent genetic differentiation between the two species, modulated by recurrent bottlenecks caused by grazing and drift. Our data suggest that conservation efforts aimed at maintaining the existing genetic connectivity in each species and ex situ conservation of seeds are the best strategies to conserve their genetic diversity.