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Populations of the same species occupying different microhabitats can either exhibit generalized traits across them or display intraspecific variability, adapting to each microhabitat in order to maximize performance. Intraspecific variability contributes to the generation of diversity, following selection and adaptation, and understanding such variability is important for comprehending how individuals choose their microhabitats. Compared with interspecific variability, however, intraspecific variability in functional morphology and its relationship with microhabitat preference and use have been relatively little studied. Here we examined whether populations of the gecko Mediodactylus kotschyi that differ in the substrates they occupy display habitat-specific behaviors and differing morphologies associated with functional adaptation to their microhabitats. We collected 207 geckos from under or on rocks or on trees from seven populations in Greece. On large islands individuals occupy both substrates; whereas small islets are devoid of trees and the geckos are restricted to rocks, while on the mainland they are only found on trees. We determined gecko substrate preferences in the laboratory, together with their clinging abilities to the different substrates. We measured their limbs, digits, and claws and assessed how these measurements relate to clinging ability. Geckos from all populations preferred the tree made available to them, but this preference was not statistically significant. Geckos from both large and small islands clung better to the tree than to the rock in the laboratory, while those from the mainland clung similarly to both substrates. Geckos collected from trees had longer manual digits and hind limbs. Geckos collected from large and small islands had taller (longer on the dorso-ventral axis; henceforth deeper ) claws. Longer digits and deeper but shorter claws were associated with a better ability to cling to rocks. Our findings suggest that while M. kotschyi is potentially preferentially arboreal, due to the great variation and plasticity it possesses, it can successfully also exploit the habitats available on the smallest, treeless islets in the Aegean Sea. Our study suggests that the dichotomous use of generalist versus specialist in describing species habitat use is oversimplified, and we suggest the use of a generalist specialist gradient instead. © 2021 Oxford University Press. All rights reserved.

Aim: Isolation is a key factor in island biology. It is usually defined as the distance to the geographically nearest mainland, but many other definitions exist. We explored how testing different isolation indices affects the inference of impacts of isolation on faunal characteristics. We focused on land bridge islands and compared the relationships of many spatial and temporal (i.e., through time) isolation indices with community-, population- and individual-level characteristics (species richness, population density and body size, respectively). Location: Aegean Sea islands, Greece. Time period: Current. Taxon: Many animal taxa. Methods: We estimated 21 isolation indices for 205 islands and recorded species richness data for 15 taxa (invertebrates and vertebrates). We obtained body size data for seven lizard species and population density data for three. We explored how well indices predict each characteristic, in each taxon, by conducting a series of ordinary least squares regressions (controlling for island area when needed) and a meta-analysis. Results: Isolation was significantly (and negatively) associated with species richness in 10 of 15 taxa. It was significantly (and positively) associated with body size in only one of seven species and was not associated with population density. The effect of isolation on species richness was much weaker than that of island area, regardless of the index tested. Spatial indices generally out-performed temporal indices, and indices directly related to the mainland out-performed those related mainly to neighbouring islands. No index was universally superior to others, including the distance to the geographically nearest mainland. Main conclusions: The choice of index can alter our perception of the impacts of isolation on biological patterns. The nearly automatic, ubiquitous use of distance to the geographically nearest mainland misrepresents the complexity of the effects of isolation. We recommend the simultaneous testing of several indices that represent different aspects of isolation, in order to produce more constructive and thorough investigations and avoid imprecise inference. © 2019 The Authors. Global Ecology and Biogeography published by John Wiley & Sons Ltd

Insular animals are thought to be under weak predation pressure and increased intraspecific competition compared with those on the mainland. Thus, insular populations are predicted to evolve 'slow' life histories characterized by fewer and smaller clutches of larger eggs, a pattern called the 'island syndrome'. To test this pattern, we collected data on egg volume, clutch size and laying frequency of 31 Aegean Island populations of the closely related geckos of the Mediodactylus kotschyi species complex. We tested how predation pressure, resource abundance, island area and isolation influenced reproductive traits. Isolation and predation were the main drivers of variation in life-history traits. Higher predator richness seemed to promote faster life histories, perhaps owing to predation on adults, whereas the presence of boas promoted slower life histories, perhaps owing to release from predation by rats on the eggs of geckos. Insular geckos followed only some of the predictions of the 'island syndrome'. Predation pressure seemed to be more complex than expected and drove life histories of species in two opposing directions. Our results highlight the importance of considering the identity of specific predators in ecological studies. © 2020 The Linnean Society of London, Biological Journal of the Linnean Society.

Aim: Animal body sizes are often remarkably variable across islands, but despite much research we still have a poor understanding of both the patterns and the drivers of body size evolution. Theory predicts that interspecific competition and predation pressures are relaxed on small, remote islands, and that these conditions promote body size evolution. We studied body size variation across multiple insular populations of 16 reptile species co-occurring in the same archipelago and tested which island characteristics primarily drive body size evolution, the nature of the common patterns, and whether co-occurring species respond in a similar manner to insular conditions. Location: Aegean Sea islands. Time period: 1984–2016. Major taxa studied: Reptiles. Methods: We combined fieldwork, museum measurements and a comprehensive literature survey to collect data on nearly 10,000 individuals, representing eight lizard and eight snake species across 273 islands. We also quantified a large array of predictors to assess directly the effects of island area, isolation (both spatial and temporal), predation and interspecific competition on body size evolution. We used linear models and meta-analyses to determine which predictors are informative for all reptiles, for lizards and snakes separately, and for each species. Results: Body size varies with different predictors across the species we studied, and patterns differ within families and between lizards and snakes. Each predictor influenced body size in at least one species, but no general trend was recovered. As a group, lizards are hardly affected by any of the predictors we tested, whereas snake size generally increases with area and with competitor and predator richness, and decreases with isolation. Main conclusions: No factor emerges as a predominant driver of Aegean reptile sizes. This contradicts theories of general body size evolutionary trajectories on islands. We conclude that overarching generalizations oversimplify patterns and processes of reptile body size evolution on islands. Instead, species’ autecology and island particularities interact to drive the course of size evolution. © 2018 John Wiley & Sons Ltd

Tail autotomy is mainly considered an antipredator mechanism. Theory suggests that predation pressure relaxes on islands, subsequently reducing autotomy rates. Intraspecific aggression, which may also cause tail loss, probably intensifies on islands due to the higher abundance. We studied whether tail autotomy is mostly affected by predation pressure or by intraspecific competition. We further studied whether predator abundance or predator richness is more important in this context. To test our predictions, we examined multiple populations of two gecko species: Kotschy's gecko (Mediodactylus kotschyi; mainland and 41 islands) and the Mediterranean house gecko (Hemidactylus turcicus; mainland and 17 islands), and estimated their abundance together with five indices of predation. In both species, autotomy rates are higher on islands and decline with most predation indices, in contrast with common wisdom, and increase with gecko abundance. In M. kotschyi, tail-loss rates are higher on predator and viper-free islands, but increase with viper abundance. We suggest that autotomy is not simply, or maybe even mainly, an antipredatory mechanism. Rather, such defence mechanisms are a response to complex direct and indirect biotic interactions and perhaps, in the case of tail autotomy in insular populations, chiefly to intraspecific aggression. © 2016 The Authors. Journal of Animal Ecology © 2016 British Ecological Society

We recently studied whether, on islands, predation or intraspecific aggression is the main driver of tail-loss, a common defense mechanism among lizards. We concluded the latter was the stronger driver (Itescu et al. 2017). Werner (2017) suggested that we failed to falsify an alternative hypothesis. He claims that on low-predation islands lizards live longer. Thus while tail loss is caused by predators, it accumulates over longer periods, resulting in overall higher tail-loss rates in populations experiencing weak predation. Here we test this hypothesis and three other arguments he presented, and fail to support them. We therefore adhere to our original conclusion that intraspecific aggression is the main driver of lizard tail loss on islands. © 2017 by Koninklijke Brill NV, Leiden, The Netherlands.

The gecko Mediodactylus kotschyi is considered rare in mainland Greece, yet it is very abundant on the Aegean islands. It has been thought to be saxicolous throughout much of its range. In a recent survey on the Peloponnese peninsula, however, we encountered it mainly on trees, and with higher frequency than previously reported. We combined our observations of localities in which we detected this gecko, and places where we failed to detect it, with data about its occurrence from the literature and museum collections. We posited two hypotheses as possible causes for the apparent relative scarcity of M. kotschyi in the Peloponnese: that it is associated with low precipitation and that it has an aversion to limestone rock. We predicted that M. kotschyi would be more likely to be found in arid places and where limestone is not the dominant type of rock, since it has been reported that this substrate is less suitable for this species. Moreover, we predicted that geckos occurring in limestone regions would be found on trees rather than under rocks. Geckos were indeed found mainly in the more arid parts of the Peloponnese, but not exclusively so. We found no evidence of limestone avoidance. We suggest that, because M. kotschyi is better known as being mostly saxicolous over most of its range, and exclusively so on the Greek islands, in the Peloponnese the search for this species has been restricted to a single habitat type, i.e., under rocks and not on trees. It may thus inhabit more localities in the Peloponnese and be more abundant there than has previously been thought. © Firenze University Press.

The island syndrome describes the evolution of slow life history traits in insular environments. Animals are thought to evolve smaller clutches of larger offspring on islands in response to release from predation pressure and interspecific competition, and the resulting increases in population density and intraspecific competition. These forces become more pronounced with diminishing island size, and life histories are thus expected to become slowest on small, isolated islands. We measured clutch sizes in 12 insular populations of Mediodactylus kotschyi, a small gecko from the Cyclades Archipelago, a set of land-bridge islands in the Aegean Sea (Greece). We analyse variation in clutch size in relation to island area, island age, maternal body size, the presence of putative competitors and nesting seabirds (which increase resource abundance in the form of marine subsidies), and richness of predators. Clutch size of M. kotschyi decreases with increasing island area, in departure from classic island syndrome predictions, suggesting the evolution of faster life histories on smaller islands. There are no relationships between clutch size and island age, maternal size, the presence of competitors or predator richness. Instead, larger clutches on small islands could simply reflect the beneficial effect of marine subsidies derived from resident seabird colonies. Indeed, populations of M. kotschyi on islands with nesting seabirds have clutch sizes 30.9 % larger (1.82 vs. 1.39 eggs) than populations on islands without nesting seabirds. Thus, our data suggest that bottom-up effects of marine subsidies may supersede the expression of a simple island syndrome in the Aegean M. kotschyi. © 2015, Springer Science+Business Media New York.

Sexual size dimorphism (SSD) can allow males and females of the same species to specialize on different sized food items and therefore minimize intraspecific competition. Interspecific competition, however, is thought to limit sexual dimorphism, as larger competitors in the community will prevent the larger sex from evolving larger size, and smaller species may prevent the smaller sex from becoming even smaller. We tested this prediction using data on the sexual size dimorphism of lizards, and mammalian carnivores, on islands world-wide. Because insular communities are depauperate, and guilds are species-poor, it is often assumed that enhanced sexual size dimorphism is common on islands. The intensity of interspecific competition, hindering enhanced dimorphism, is thought to increase with competitor richness. We tested whether intraspecific sexual size dimorphism of mammalian carnivores and lizards decreases with increasing island species richness. We further computed the average sexual dimorphism of species on islands and tested whether species-rich islands are inhabited by relatively monomorphic species. Within families and guilds across carnivores and lizards, and with both intraspecific and interspecific approaches, we consistently failed to find support for the notion that species-poor islands harbour more sexually dimorphic individuals or species. We conclude that either interspecific competition does not affect the sexual size dimorphism of insular lizards and carnivores (i.e. character displacement and species sorting are rare in these taxa), or that the number of species in an assemblage or guild is a poor proxy for the intensity of interspecific competition in insular assemblages. © 2014 British Ecological Society.

Fil: Raia, Pasquale. Università degli Studi di Napoli Federico II; Italia Fil: Passaro, Federico. Università degli Studi di Napoli Federico II; Italia Fil: Carotenuto, Francesco. Università degli Studi di Napoli Federico II; Italia Fil: Maiorino, Leonardo. Università degli Studi Roma Tre; Italia Fil: Piras, Paolo. Università degli Studi Roma Tre; Italia Fil: Teresi, Luciano. Università degli Studi Roma Tre; Italia Fil: Meiri, Shai. Tel Aviv University; Israel Fil: Itescu, Yuval. Tel Aviv University; Israel Fil: Novosolov, Maria. Tel Aviv University; Israel Fil: Baiano, Mattia A. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Fil: Baiano, Mattia A. Instituto Catalán de Paleontología Miquel Crusafont; España Fil: Baiano, Mattia A. Universidad Nacional de Río Negro; Argentina Fil: Martínez, Ricard. Universitat Autònoma de Barcelona; España Fil: Fortelius, Mikael. University Of Helsinski; Finlandia Luxuriant, bushy antlers, bizarre crests, and huge, twisting horns and tusks are conventionally understood as products of sexual selection. This view stems from both direct observation and from the empirical finding that the size of these structures grows faster than body size (i.e., ornament size shows positive allometry).We contend that the familiar evolutionary increase in the complexity of ornaments over time in many animal clades is decoupled from ornament size evolution. Increased body size comes with extended growth. Since growth scales to the quarter power of body size, we predicted that ornament complexity should scale according to the quarter power law as well, irrespective of the role of sexual selection in the evolution and function of the ornament. To test this hypothesis, we selected three clades (ammonites, deer, and ceratopsian dinosaurs) whose species bore ornaments that differ in terms of the importance of sexual selection to their evolution.We found that the exponent of the regression of ornament complexity to body size is the same for the three groups and is statistically indistinguishable from0.25.We suggest that the evolution of ornament complexity is a by-product of Cope’s rule. We argue that although sexual selection may control size in most ornaments, it does not influence their shape.