Ecology of lacertid lizards in a desert area of eastern Morocco

Some basic aspects of the ecology of five lacertid lizards inhabiting the desert area of Tafilalet, in south-eastern Morocco, are considered: Mesalina guttulata , occupying almost exclusively the hammada habitat; Mesalina olivieri, Acanthodactylus boskianus and Acanthodactylus scutellatus found on sand and bare-ground reg ; and Acanthodactylus longipes , inhabiting the sand-dune habitat, the erg.
Acanthodactylus scutellatus, A. boskianus, A. longipes and M. olivieri had a bimodal diel activity pattern; in M. guttulata daily activity was centred on the morning hours. All species exhibit fairly precise thermoregulation with a strong relation between body and substratum temperature, particularly for species occupying softer substrata. Mesalina guttulata had a diverse diet during both seasons observed, while in the remaining species termites and ants formed the bulk of the food.
The foraging behaviour, diet and ecological characteristics of Lacertidae from the Tafilalet, especially the genus Acanthodactylus , seem to be optimal for the challenging environmental conditions of this desert area. The implications of the foraging strategy of Tafilalet's Acanthodactylus for other species of the genus are discussed.     

Survey of the reptilian fauna of the Kingdom of Saudi Arabia. V. The lizard fauna of Turaif region

Turaif area located in the Northern border region of Saudi Arabia is one of the most important regions of the Kingdom. This work was proposed to throw light on the diversity of lizard fauna investigated through the collection and subsequent identification of specimens from different localities of Turaif region of Kingdom of Saudi Arabia. Sixteen species of lizards belonging to 5 families (Agamidae, Gekkonidae, Lacertidae, Scincidae and Varanidae) were recorded. Lacertidae was the most common family. Three species of lizards namely Acanthodactylus orientalis, Acanthodactylus scutellatus and Acanthodactylus grandis were reported for the first time in the Turaif region of Saudi Arabia. The geographical distribution of the collected species within this province was mapped.     

A multivariate analysis of the fringe-toed lizards of the Acanthodactylus scutellatus group (Squamata: Lacertidae): systematic and biogeographical implications

The taxonomy of the fringe-toed lizards of the Acanthodactylus scutellatus group has long been unstable and no consensus exists on the systematic status of its various forms. A multivariate analysis of morphological characters, performed on over 1000 specimens from most of the African range of this group, allowed us to clarify the specific allocation of most of the Saharan populations included in this species group. Based on comparisons of morphology between allopatric and sympatric populations of this complex, we propose the recognition of six biological species. Our results confirm the specific status of Acanthodactylus aureus , A. dumerili , A. scutellatus, A. longipes and the recently described A. taghitensis . In addition, we re-validate A. senegalensis (occurring from Mauritania and Mali south to Senegal), which has been treated as a synonym of A. dumerili by previous authors. Acanthodactylus longipes is reported for the first time from coastal Mauritania, and A. taghitensis (previously known only from a very small region in Algeria) is reported from continental Mauritania. The systematic section of this paper includes a full list of examined material, diagnosis and known distribution of each species, in addition to some information on geographical variation and ecology. A key for specific identification is provided as an appendix.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 137 , 117−155.     

Studies of Acanthodactyla in Tunisia. II. Biometrical relations

The four species of the g. Acanthodactylus Wieg., 1834 occuring in Tunisia are quite individualized by their biometrical relations which, moreover, allow to bring out their relationships. The size of the upper quarter in our sampling oppose two groups of two species each one. The distribution of the ratio between the length of the body and that of the tail place apart A. boskianus from the three other species in a more marked way for adults of both sexes than for juveniles.     

Influence of a broad temperature range on the oxygen consumption rates of three desert lizard species

Oxygen consumption rates of three desert lizard species: (Acanthodactylus boskianus, Lacertidae; Scincus mitranus, Scincidae; Diplometopon zarudnyi, Amphisbaenidae) were determined in relation to ambient temperatures ranging from 10 to 35 degrees C using a double-chamber, volumetric closed system. The interspecific differences in the oxygen consumption rates were found to have a close relationship between the characteristics of the M-T curves, the thermoregulatory behaviour and the ecology of the respective species. The M-T curves of the studied lizards were specifically modified and adjusted to their climatic region.     

Systematics and phylogeography of Acanthodactylus schreiberi and its relationships with Acanthodactylus boskianus (Reptilia: Squamata: Lacertidae)

Acanthodactylus is a widespread lacertid genus occurring from the Iberian Peninsula and western North Africa to western India including the Middle East, Cyprus, and the Arabian Peninsula. The genus is in dire need of a taxonomic revision, and the phylogenetic relationships amongst and within its species remain unclear. In particular, the taxonomy and relationship of the allopatric, narrow‐ranged Acanthodactylus schreiberi and its close relative, the widespread Acanthodactylus boskianus asper, are poorly understood. We estimated the phylogenetic and phylogeographical structure of A. schreiberi across its distribution range, and evaluated its relationships to A. b. asper, using mitochondrial and nuclear data. The phylogenetic results indicate that both species are paraphyletic, with A. schreiberi nested within A. b. asper, and the subspecies A. schreiberi syriacus nested within a distinct lineage of A. b. asper. We suggest that the group is in need of a taxonomic revision because the identified lineages and genetic diversity are incongruent with the currently recognized taxonomy. We tentatively conclude that A. schreiberi is restricted to Cyprus and Turkey, reduced to a single form, and that the populations in Lebanon and Israel belong to A. b. asper. © 2014 The Linnean Society of London     

Skeletochronological determination of age and demographic analysis of a population of Acanthodactylus pardalis (Lischenstein, 1823) of the Kerkennah islands (Tunisia)

The skeletochronological analysis of 34 adult lizards from population of Acanthodactylus pardalis of Kerkennah islands in Tunisia enabled to: determine the individual age of the animals; delineate four various cohorts making up the studied population; evaluate around 4 years life span in nature of this species; point out that the renewal rate accounts for 41% of its population. Moreover the method of "maximums successifs" is applied on sizeable sample in order to shore up the obtained results.     

Spatial and reversal learning in congeneric lizards with different foraging strategies

Environmental demands that require intensive search for mates, food and nest sites are correlated with efficient spatial memory in many mammalian and avian species. This convergence of evidence has led to the view that spatial memory, and the neurological structures associated with it, have been selected in niches that require memory for the location of goal objects. Whether such evolutionary demands are also correlated with nonspatial abilities that require flexible use of associations similar to those required for spatial memory has not been well studied. In addition, correlations between niche types and the use of spatial or nonspatial memory have not been investigated in nonmammalian, nonavian taxa. In this study, we investigated the relationship between foraging strategies and performance on two tasks, one spatial and the other nonspatial, in congeneric lizard species: Acanthodactylus boskianus, an active forager that collects clumped sedentary prey, Acanthodactylus scutellatus, a sit-and-wait predator that collects distributed mobile prey. The two species did not differ in their performance of a spatial memory task, but A. boskianus, the active forager, performed better on the reversal of a visual discrimination, a nonspatial task. These findings question the generality of the spatial adaptation model for vertebrates. We present the pliancy hypothesis, which we developed to account for these results. Copyright 1999 The Association for the Study of Animal Behaviour.     

Mise en évidence de la présence d'acide ribonucléique dans le noyau des spermatides âgées d'Acanthodactylus erythrurus lineomaculatus Dum. & Bibr. (Lacertidae)

Résumé Nous avons abservé au moyen de méthodes cytochimiques, l'élimination progressive de l'ARN nucléaire dans les spermatides âgées d'un Lézard. Il ressort de cette étude que l'ARN lié à la chromatine s'observe pendant toute la spermiogenèse et n'est éliminé qu'au terme ultime de cette spermiogenèse c'est à dire au moment de la transformation de la spermatide en spermatozoïde.Ces résultats diffèrent de ceux obtenus chez un Insecte où l'ARN nucléaire disparaît aux tous premiers stades de la spermiogenèse, en même temps que les nucléoles.

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Demonstration of nuclear RNA in the mature spermatids of Acanthodactylus erythrurus lineomaculatus Dum. & Bibr. (Lacertidae)

Summary Cytochemical methods have shown a progressive elimination of nuclear RNA from mature spermatids in the lizard Acanthodactylus erythrurus. The RNA bound to chromatin has been observed throughout spermiogenesis and is eliminated only at the final stage, i. e., the stage of transformation from the spermatid to the spermatozoon.These results differ from those obtained in insects where the nuclear as well as the nucleolar RNA disappears during the early stages of spermiogenesis.

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Zusammenfassung Mit cytochemischen Untersuchungsmethoden haben wir bei der Eidechse Acanthodactylus erythrurus festgestellt, daß die Kern-RNS aus den älteren Spermatiden eliminiert wird. So wird die chromatingebundene-RNS, die während der gesamten Spermiogenese beobachtet werden kann, erst im letzten Stadium der Spermiogenese ausgestoßen, das heißt im Stadium der Umwandlung der Spermatiden in Spermien.Diese Ergebnisse stehen nicht im Einklang mit denen, die bei Insekten gewonnen wurden. Bei den letzteren verschwindet die RNS des Zellkerns und der Nukleolen in den Anfangsstadien der Spermiogenese.

     

Age related differences in lipophilic compounds found in femoral gland secretions of male spiny-footed lizards, Acanthodactylus erythrurus

Although chemoreception plays an important role in social organization of many lizards, only a few studies have examined the chemicals found in secretions used for intraspecific communication. We report the composition of the secretion of the femoral glands of males of the spiny-footed lizard (Acanthodactylus erythrurus). On the basis of mass spectra, obtained by GC/MS, we identified 45 lipophilic compounds, including several alcohols ranging from 10 to 29 carbon atoms (mainly hexacosanol and tetracosanol), steroids (mainly cholesterol and dehydrocholesterol), n-C9 to n-C20 carboxylic acids, esters of carboxylic acids, and minor components such as lactones, ketones, squalene and a-tocopherol. Some of these compounds are reported for the first time in lizards. Adult and subadult males differed in the composition of secretions, with C9 to C15 carboxylic acids being more abundant in younger than in older lizards, whereas C16 to C20 carboxylic acids were more abundant in older lizards. Also, older lizards had significant lower proportions of cholesterol and campesterol but higher proportions of dehydrocholesterol.     

Change your diet or die: predator-induced shifts in insectivorous lizard feeding ecology

Animal feeding ecology and diet are influenced by the fear of predation. While the mechanistic bases for such changes are well understood, technical difficulties often prevent testing how these mechanisms interact to affect a mesopredator’s diet in natural environments. Here, we compared the insectivorous lizard Acanthodactylus beershebensis’ feeding ecology and diet between high- and low-risk environments, using focal observations, intensive trapping effort and fecal pellet analysis. To create spatial variation in predation risk, we planted “artificial trees” in a scrubland habitat that lacks natural perches, allowing avian predators to hunt for lizards in patches that were previously unavailable to them. Lizards in elevated-risk environments became less mobile but did not change their microhabitat use or temporal activity. These lizards changed their diet, consuming smaller prey and less plant material. We suggest that diet shifts were mainly because lizards from risky environments consumed prey items that required shorter handling time.     

New compounds, sexual differences, and age-related variations in the femoral gland secretions of the lacertid lizard Acanthodactylus boskianus

Integumental gland secretions in lizards have been postulated to play a role as semiochemicals, but few studies have analysed the chemical nature of the gland secretions used in communication. We analysed the femoral gland secretions of Acanthodactylus boskianus using GC–MS, compared secretions of both sexes and different ages of males. For the first time in reptiles monoglycerides of fatty acids and glycerol monoethers of long chain alcohols were identified. In addition, alcohols, steroids, carboxylic acids, alkanes, amides, aldehydes, carboxylic acid esters, and squalene occurred. Sexual differences and age correlation in the amount of all major groups of compounds occurred, as such these results strengthen the theory that these secretions are used as semiochemicals. This work lays the foundations to test in future the role of chemical cues in mate choice and dominance hierarchies in lizards and to test the activity of compounds in behavioural assays to eventually identify the pheromones.     

Density-dependent aposematism in the desert locust

The ecological processes underlying locust swarm formation are poorly understood. Locust species exhibit phenotypic plasticity in numerous morphological, physiological and behavioural traits as their population density increases. These density-dependent changes are commonly assumed to be adaptations for migration under heterogeneous environmental conditions. Here we demonstrate that density-dependent nymphal colour change in the desert locust Schistocerca gregaria (Orthoptera: Acrididae) results in warning coloration (aposematism) when the population density increases and locusts consume native, toxic host plants. Fringe-toed lizards (Acanthodactylus dumerili (Lacertidae)) developed aversions to high-density-reared (gregarious-phase) locusts fed Hyoscyamus muticus (Solanaceae). Lizards associated both olfactory and visual cues with locust unpalatability, but only gregarious-phase coloration was an effective visual warning signal. The lizards did not associate low rearing density coloration (solitarious phase) with locust toxicity. Predator learning of density-dependent warning coloration results in a marked decrease in predation on locusts and may directly contribute to outbreaks of this notorious pest.     

Bird predation alters infestation of desert lizards by parasitic mites

Theory predicts that predators can reduce parasite abundance on prey by reducing prey density and through disproportionate predation on heavily infested individuals. We experimentally tested this prediction by examining the effects of bird predation on parasitic mite infestation of the prey lizard Acanthodactylus beershebensis. We manipulated predation by adding perches to arid scrubland, allowing avian predators to hunt for lizards in a habitat the birds would not normally use. Host density influenced parasite abundance in hatchlings, but not in older aged individuals and parasite abundance did not affect lizard host survival. Contrary to expectation mite abundance on adult lizards increased under low predation intensities. We explain these results by suggesting a novel hypothesis based on the assumption that the two components of predation, i.e. actual removal of prey and risk, exert contradictory effects on macroparasite abundance.     

Effect of testosterone on T cell-mediated immunity in two species of Mediterranean lacertid lizards

One of the primary assumptions of the immunocompetence handicap hypothesis is that testosterone has an immunosuppressive effect, but conflicting results have been reported in a variety of bird species concerning the effect of testosterone on the humoral and the T cell-mediated components of the immune system. The T cell-mediated component of the immune system is particularly important during the breeding season, because the likelihood of injury during sexual competition is high and T cell-mediated immunity is essential for healing wounds and resisting infection. In this study we examined the effect of experimentally increased levels of testosterone during breeding season on T cell-mediated immunity in male lizards of two Mediterranean lacertid species, Psammodromus algirus and Acanthodactylus erythrurus. The hormonal treatment significantly increased testosterone of the experimental individuals. T cell-mediated responses to phytohemagglutinin stimulation were significantly suppressed in testosterone-treated males of both species. Furthermore, there was a significant negative relationship between individual variability in T cell-mediated responsiveness and plasma testosterone concentration. The present study is the first to demonstrate testosterone-induced suppression of T cell-mediated immunity in lizards.     

Blue tail and striped body: why do lizards change their infant costume when growing up?

Ontogenetic changes in color and pattern that are not directlyrelated to reproduction are very common yet remain a poorlyunderstood phenomenon. One example is conspicuous colors inthe tails of fish, amphibians, and reptiles that fade out laterin life. We suggest a novel hypothesis: conspicuous tail colorsthat appear only in juveniles compensate for an increased activitylevel, deflecting imminent attacks to the tail. We observedblue-tailed, newly hatched lizards (Acanthodactylus beershebensis)in the field and compared 5 behavioral parameters with thoseof older individuals that had already lost their neonate coloration.In addition, we explored whether tail displays, often assumedto direct a predator's attention to the tail, disappear withthe color change. Striped blue-tailed hatchlings foraged moreactively than 3-week-old juveniles, spent a longer time in openmicrohabitats, and performed deflective tail displays. In comparison,2 other lacertids that do not undergo ontogenetic change didnot switch to safer foraging when growing up. The results suggestthat activity alteration may be a major factor affecting theontogenetic color and pattern change. Active lizards that foragein open habitats increase their probability of attack by ambushpredators. Conspicuous colors and deflection displays may shiftattacks to the expendable tail, increasing the prey's overallprobability of surviving attacks. The persistence of both stripedbody pattern and blue tail fits the active foraging period ofneonates and hence may be appropriate for other species thatdisplay a conspicuous tail accompanied by a striped pattern.     

Avoiding injury and surviving injury: two coexisting evolutionary strategies in lizards

Congenital morphological differences between injured and intact individuals in a population may reflect adaptations to avoid injury, to survive injury, or both. We explore the possible occurrence of such adaptations, analysing the relationship between tail state (original, O; regenerated, R) and morphology (20 scale characters) in 23 taxa of Lacertidae. In some taxa, such as Acanthodactylus opheodurus , morphologies of O and R lizards differed significantly. In these, usually O individuals were morphologically typical (modal), while R individuals were extreme; in others, the contrary occurred. Indices of pattern similarity detected fewer differences between O and R lizards than those based on absolute values. We developed unbiased estimates of classification rates of discriminant analysis. The order of inclusion of characters in the discriminant functions of males and females of the same species differed more than the order for the same sex in other species. Some morphological differences between O and R lizards seem adaptive or linked by pleiotropy to adaptive traits. Congenital morphological differences between O and R lizards are probably more frequent than detected because we show that age and geographical heterogeneity of our samples are confounding factors to O–R differences. R-morphologies might reflect microevolution in favour of paedotypic forms, possibly possessing greater regenerating capacities than relatively more peratypic forms.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78 , 307–324.     

Influence of heat transmission mode on heating rates and on the selection of patches for heating in a mediterranean lizard

Heliothermy (heat gain by radiation) has been given a prominent role in basking lizards. However, thigmothermy (heat gain by conduction) could be relevant for heating in small lizards. To ascertain the importance of the different heat transmission modes to the thermoregulatory processes, we conducted an experimental study where we analyzed the role of heat transmission modes on heating rates and on the selection of sites for heating in the Mediterranean lizard Acanthodactylus erythrurus (Lacertidae). The study was conducted under laboratory conditions, where two situations of different operative temperatures (38 degrees and 50 degrees C) were simulated in a terrarium. In a first experiment, individuals were allowed to heat up during 2 min at both temperatures and under both heat transmission modes. In a second experiment, individuals were allowed to select between patches differing in the main transmission mode, at both temperatures, to heat up. Experiences were conducted with live, nontethered lizards with a starting body temperature of 27 degrees C. Temperature had a significant effect on the heating rate, with heat gain per unit of time being faster at the higher operative temperature (50 degrees C). The effect of the mode of heat transmission on the heating rate was also significant: at 50 degrees C, heating rate was greater when the main heat transmission mode was conduction from the substrate (thigmothermy) than when heating was mainly due to heat gain by radiation (heliothermy); at 38 degrees C, heating rates did not significantly differ between transmission modes. At 38 degrees C, selection of the site for heating was not significantly different from that expected by chance. However, at 50 degrees C, the heating site offering the slowest heating rate (heliothermic patch) was selected. These results show that heating rates vary not only with environmental temperature but also with different predominant heat transmission modes. Lizards are able to identify and exploit this heterogeneity, selecting the source of heat gain (radiation) that minimizes the risk of overheating when temperature is high.     

Microstructure of scales in selected lizard species

In the present study, it was hypothesized that micromorphology of the surface of many lizard scales appears to mimic the topography of the habitat in which they live. Many authors have suggested that the microstructure of the superficial surface of scales have undergone important adaptations and have functional value in lizards. In this study, we investigated the variation and adaptation of the micromorphology and microstructure of the superficial surface of the dorsal and ventral scales from the mid-body region of Stellagama stellio (Agamidae), Stenodactylus petrii (Gekkonidae), Acanthodactylus boskianus (Lacertidae), Eumeces schneideri (Scincidae), Trachylepis quinquetaeniata (Scincidae), Scincus scincus (Scincidae), Varanus griseus (Varanidae), Chameleo chameleon (Chamaeleonidae). Skin specimens were prepared and analyzed using scanning electron microscopy. The dorsal and ventral scale surfaces had microstructure in the studied species and they exhibited unique patterns that somewhat resembled the topography of the microhabitats in which they lived. Similarity was detected in the three most related species, those having a common family, Scincidae. Ecomorphological relationships were detected between the dorsal and ventral scale microstructures and microhabitats. We conclude that environmental factors have observable influences on the microstructure of lizard scales.     

Ontogenetically stable dimorphism in a lacertid lizard (Acanthodactylus boskianus) with tests of methodology and comments on life-history

Recent arguments in the literature prompted us to compare methods for assessing sexual dimorphism in body proportions of lacertid lizards, using Acanthodactylus boskianus . Although expressing body-part measurements as proportional to head length was the most effective method, we recommend using trunk length for the baseline as a general method for lizards. We also argue that, when aiming to assess sexual dimorphism in body proportions of lizards, if the context is ecological, all available adults should be included. However, for morphology and taxonomy, small sub-samples of the largest individuals that maximally express their genetic morphological potential should be used. In A. boskianus , the sexual dimorphism of mensural characters in adults was typical: males were larger, with relatively larger head and appendages. However, the ontogeny of this dimorphism was unusual in that the differences existed already in youth and thereafter persisted isometrically. The sexual dimorphism of meristic characters was male-biased in numbers of femoral pores and of caudal vertebrae, and female-biased in numbers of ventral plates along the trunk and of precaudal vertebrae. Size dimorphism may conceivably play a role in sex recognition because two potential visual cues (i.e. size dimorphism and dichromatism) appear to complement each other.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society 2009, 97 , 275–288.