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Differences were related to their contrasting courtship behaviours: horizontal head movements and severe biting in T. marginata, vertical head bobs and carapace butting in T. graeca, and mounting and tail thrusting in T. hermanni. There was no difference in the frequency of observations of courtship or fighting among the three species, but courtship was about 10 times more common than combat in males. All species showed greatest courtship activity in autumn; copulation was rarely observed in T. hermanni (only 0.36\% of courting males) and not seen in the other species in the field. Observations made throughout the activity season indicated that feeding was equally common in males and females in all three species. Differences in shape were more likely to be the result of sexual selection than of natural selection for fecundity. Detailed predictions are made for sexual dimorphism of other characters in these species.}, keywords = {Courtship, Reptilia, Sexual dimorphism, sexual selection, Shape, Testudo}, author = {Willemsen, R.E. and Hailey, A.} } @article {14179, title = {A latitudinal cline of dark plastral pigmentation in the tortoise Testudo hermanni in Greece}, journal = {Herpetological JournalHerpetological Journal}, volume = {9}, number = {3}, year = {1999}, note = {Cited By (since 1996): 7Export Date: 6 September 2006Source: ScopusReferences: Ballasina, D., Report on the repatriation of tortoises to Greece by the Carapax centre (Italy) (1992) Bull. Brit. Herpetol. Soc., 40, pp. 2-4; Bartholomew, G.A., Physiological control of body temperature (1982) Biology of the Reptilia, 12, pp. 167-211. , Gans, C. and Pough, F. H. (Eds). 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The most likely explanation is selection for thermoregulation, with decreased dark pigmentation in the north reducing heat loss to the substrate by infra red radiation during activity. This hypothesis was supported by data on body (T(b)) and substrate (T(s)) temperatures in populations from northern, central and southern Greece. T(b) was generally above T(s), showing that heat would generally be lost rather than gained through the plastron, and the mean difference T(b)-T(s) was greatest in the north: +6.6 {\textdegree}C, compared to +2.4 {\textdegree}C in the south. Mean T(b) was lowest in the south (26.9{\textdegree}C, compared to 29.3{\textdegree}C in the north) and the slope of T(b) on T(s) was about 1 (compared to 0.5 in the north). Thermoregulation in southern Greece is similar to that of tropical tortoises, with avoidance of overheating being the major problem, rather than elevation of T(b) for activity.}, keywords = {Cline, Pigmentation, Reptilia, Testudo, thermoregulation, Tortoise}, author = {Willemsen, R.E. and Hailey, A.} } @article {14180, title = {Variation of adult body size of the tortoise Testudo hermanni in Greece: Proximate and ultimate causes}, journal = {Journal of ZoologyJournal of Zoology}, volume = {248}, number = {3}, year = {1999}, note = {Cited By (since 1996): 34Export Date: 6 September 2006Source: ScopusReferences: Allee, W.C., Park, O., Emerson, A.E., Park, T., Schmidt, K.P., (1949) Principles of Animal Ecology, , Philadelphia: W. B. Saunders; Archer, W.H., The mountain tortoise (1948) Afr. Wildl., 2 (2), pp. 75-78; Archer, W.H., The mountain tortoise (1948) Afr. 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The age at maturity ({\'a}) increased with body size in the 17 populations, while the Bertalanffy growth constant (k) decreased with body size. The quantities {\'a}M and M/k (where M is the instantaneous mortality rate) were invariant with body size, suggesting that differences between populations were adaptive rather than the result of short-term disturbance. Body size was greater in cooler areas, and increased with both latitude and altitude. This pattern is opposite to that found in most ectotherms (the reverse Bergmann{\textquoteright}s rule), and to that which occurs between tortoise species. Several hypotheses about the possible ultimate causes of variation of body size were rejected, including adaptation to long-term habitat disturbance (land use), character displacement, social factors, energetics, thermoregulation, r-K selection, the length of the season available for incubation, or differences in juvenile mortality. The most likely ultimate cause of size variation between sites is differences in adult mortality, the correlation with environmental temperature being through the frequency of fires.}, keywords = {Body size, Geographic variation, growth, Life history invariants, Reptilia, temperature, Testudo hermanni}, author = {Willemsen, R.E. and Hailey, A.} } @article {14181, title = {Variation in adult survival rate of the tortoise Testudo hermanni in Greece: Implications for evolution of body size}, journal = {Journal of ZoologyJournal of Zoology}, volume = {255}, number = {1}, year = {2001}, note = {Cited By (since 1996): 8Export Date: 6 September 2006Source: ScopusReferences: Adolph, S.C., Porter, W.P., Temperature, activity, and lizard life histories (1993) Am. Nat., 142, pp. 273-295; Avery, R.A., Thermoregulation, metabolism and social behaviour in Lacertidae (1976) Morphology and biology of reptiles, pp. 245-259. , Bellairs, A. d{\textquoteright}A. \& Cox, C. B. (Eds). 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The average survival rate from the two methods was significantly positively related to mean adult body size (r = 0.815) and negatively related to effective temperature (r = -0.880). A life table model was used to investigate the consequences of adult survival rate for optimum age at maturity and mean adult size of females. The basis of the model was the trade-off between increased reproduction from large adult size, and decreased survival to maturity from delayed age at maturity. The model showed that the observed variation in adult survival rates was more than sufficient to account for the variation of adult size among populations. The model thus predicted an even greater range of mean adult size than that observed. A negative correlation between growth rate and adult survival rate or age at maturity, as observed, was needed to fit the size data. The model supports the hypothesis that variation in adult size of T. hermanni in Greece is adaptive (rather than a non-adaptive environmental or phylogenetic effect) and due to variation in adult survival rates (rather than other factors such as juvenile survival).}, keywords = {Body size, Growth rate, life history, Life table, Reptilia, Survival rate, Testudo hermanni}, author = {Willemsen, R.E. and Hailey, A.} } @article {14183, title = {Body mass condition in greek tortoises: Regional and interspecific variation}, journal = {Herpetological JournalHerpetological Journal}, volume = {12}, number = {3}, year = {2002}, note = {Cited By (since 1996): 3Export Date: 6 September 2006Source: ScopusReferences: Avery, H.A., Vitt, L.J., How to get blood from a turtle (1984) Copeia, 1984, pp. 209-210; Blaxter, K., (1989) Energy Metabolism in Animals and Man, , Cambridge: Cambridge University Press; Blood, D.C., Henderson, J.A., (1968) Veterinary Medicine, 3rd Edition, , London: Baillie?re, Tindall \& Cassell; Bonnet, B., Influence of nutritional conditions on the organic composition of blood and urine in the juvenile sea turtle Chelonia mydas L. 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Mass-length relationships differed significantly between the sexes (females being heavier) and among the species (T. marginata being least heavy). Mass-length relationships for each species and sex were used to calculate the condition index (CI) log (M/M?), where M is observed mass and M? is mass predicted from length, which is equal to residuals from the regression of log M on log length. It was possible to use the empirical mass-length relationships from one population of T. hermanni to calculate CI in other populations of substantially different adult size. The seasonal pattern of the CI varied with latitude, with a sharper and later peak further north, and habitat, declining more in summer at a xeric coastal site. The seasonal patterns of CI in T. graeca and T. marginata were similar, with sharper and later peaks compared to T. hermanni. These seasonal patterns of CI were related to differences in activity and food availability among species and sites. The variability of the CI was similar in all three species, with most values between -0.1 and +0.1; seasonal variation was of relatively low amplitude, with a range of about 0.05 between the highest and lowest monthly means.}, keywords = {Condition index, Reptilia, season, Testudo graeca, Testudo hermanni, Testudo marginata}, author = {Willemsen, R.E. and Hailey, A.} } @article {6163, title = {Population density and adult sex ratio of the tortoise Testudo hermanni in Greece: Evidence for intrinsic population regulation}, journal = {Journal of ZoologyJournal of Zoology}, volume = {251}, number = {3}, year = {2000}, note = {Cited By (since 1996): 12Export Date: 6 September 2006Source: ScopusReferences: Andrewartha, H.G., Birch, L.C., (1954) The Distribution and Abundance of Animals, , Chicago: University of Chicago Press; Arnqvist, G., Multiple mating in a water strider: Mutual benefits or intersexual conflict (1989) Anim. 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(Lond.), 248, pp. 379-396; Williams, G.C., (1966) Adaptation and Natural Selection, , Princeton, NJ: Princeton University Press; Wright, J., Steer, E., Hailey, A., Habitat separation in tortoises and the consequences for activity and thermoregulation (1988) Can. J. Zool., 66, pp. 1537-1544; Wynne-Edwards, V.C., (1962) Animal Dispersion in Relation to Social Behaviour, , Edinburgh: Oliver \& Boyd; Wynne-Edwards, V.C., (1986) Evolution Through Group Selection, , Oxford: Blackwell Scientific Publications.}, month = {2000///}, pages = {325-338}, abstract = {The idea that the size of animal populations may be regulated by factors intrinsic to them, such as behaviour, has a long history in ecology. Although this idea is now rejected as a general mechanism, it may apply in some species where females are damaged during courtship attempts, such as the Mediterranean tortoise Testudo hermanni. The sex ratios (adult males: adult females) of most dense Greek populations were more extreme (over 1.5:1) than could be accounted for by the earlier age at maturity of males, and some ranged to over 6:1. The sex ratio was correlated with the population density of adult males and the frequency of courtship attempts on females, and negatively correlated with the longevity of females. A high population density of males limits the density of adult females. This intrinsic regulation of population size is a consequence, rather than a goal, of a particular type of courtship behaviour (persistent thrusts by the male using a tail with a sharp terminal spur) in T. hermanni. Existing population densities at many Greek sites are probably unnaturally high, owing to the reduction of natural predators.}, keywords = {Courtship behaviour, density, KAPE, Population regulation, Reptilia, Sex ratio, Testudo hermanni, Tortoise}, author = {Hailey, A. and Willemsen, R.E.} } @article {5933, title = {Spatial heterogeneity in the effects of climate change on the population dynamics of a Mediterranean tortoise}, journal = {Global Change BiologyGlobal Change BiologyGlobal Change Biology}, volume = {17}, number = {10}, year = {2011}, note = {ISI Document Delivery No.: 816ASTimes Cited: 0Cited Reference Count: 86}, month = {Oct}, pages = {3075-3088}, type = {Article}, abstract = {Climatic shifts may increase the extinction risk of populations, especially when they are already suffering from other anthropogenic impacts. Our ability to predict the consequences of climate change on endangered species is limited by our scarce knowledge of the effects of climate variability on the population dynamics of most organisms and by the uncertainty of climate projections, which depend strongly on the region of the earth being considered. In this study, we analysed a long-term monitoring programme (1988-2009) of Hermann{\textquoteright}s tortoise (Testudo hermanni) aimed at evaluating the consequences of the drastic changes in temperature and precipitation patterns predicted for the Mediterranean region on the demography of a long-lived species with low dispersal capability and already suffering a large number of threats. Capture-recapture modelling of a population in the Ebro Delta (NE Spain) allowed us to assess the effect of climate variability on the survival of tortoises. Winter rainfall was found to be the major driver of juvenile and immature survival, whereas that of adults remained high and constant across the study. Furthermore, local climate series obtained ad hoc from regional climate simulations, for this and 10 additional Mediterranean locations where tortoises occurred, provided us with reliable future climate forecasts, which were used to simulate the fate of these populations under three precipitation scenarios (mean, wet and dry) using stochastic population modelling. We show that a shift to a more arid climate would have negative consequences for population persistence, enhancing juvenile mortality and increasing quasiextinction risk because of a decrease in recruitment. These processes varied depending on the population and the climate scenario we considered, but our results suggest that unless other human-induced causes of mortality are suppressed (e. g. poaching, fire, habitat fragmentation), climate variability will increase extinction risk within most of the species{\textquoteright} current range.}, keywords = {ADULT SURVIVAL, animals, climate projections, CONSERVATION STRATEGIES, demography, DESERT TORTOISES, extinction risk, GOPHERUS-AGASSIZII, Greece, impacts, northern, population matrices, precipitation, reptiles, survival, Testudo hermanni, TESTUDO-HERMANNI, TURTLE POPULATION}, isbn = {1354-1013}, author = {Fernandez-Chacon, A. and Bertolero, A. and Amengual, A. and Tavecchia, G. and Homar, V. and Oro, D.} } @conference {2014, title = {Comparison of Reptile Communities in Three Types of Thermophilous Mediterranean Forest in Southern Greece}, booktitle = {10th International Congress on the Zoogeography and Ecology of Greece and Adjacent Regions}, year = {2006}, month = {26-30 June}, publisher = {Hellenic Zoological Society, Hellenic Ornithological Society, Hellenic Ministry of Rural Development, Section of Animal Biology (Dept. Of Biology, Univ. of Patras)}, organization = {Hellenic Zoological Society, Hellenic Ornithological Society, Hellenic Ministry of Rural Development, Section of Animal Biology (Dept. Of Biology, Univ. of Patras)}, address = {Patras}, keywords = {communities, forests, reptiles, thermophilous, δάση, ερπετά, κοινότητες}, author = {Ioannidis, Y and Cheiras, G and Kardakari, N} }