Johannes Foufopoulos

Reptile Extinctions on Land‐Bridge Islands: Life‐History Attributes and Vulnerability to Extinction

One of the central questions of conservation biology is what life‐history traits render a species prone to extinction. We addressed this problem by calculating extinction rates for 35 species of turtles and squamates (lizards and snakes) occurring on 87 land‐bridge islands in the Mediterranean Sea. We calculated extinction rates in two ways: first, by incorporating the known sequence of historical island separations and second by ignoring history and assuming that the islands became isolated simultaneously. The second procedure is simpler and more frequently used in the literature and produces estimates of extinction rates that are similar to the first, more complex procedure. We then determined the relationship between extinction rates (calculated using both methods) and body mass, longevity, habitat specialization, and population abundance using two methods: first, by accounting for the phylogenetic relationships among species and, second, by ignoring them. Only population abundance and habitat specialization explained a significant amount of the observed variation in species extinction rates. Body mass itself did not explain variation in extinction rates, although it was strongly correlated with abundance. These conclusions were obtained using both procedures for calculating extinction rates and both procedures for correlating extinction rates with life‐history traits.

Galápagos Birds and Diseases: Invasive Pathogens as Threats for Island Species

Exotic diseases and parasites have caused extinctions on islands and continents, particularly when they spread through assemblages of immunologically naive species. Hawaii has lost a substantial part of its endemic bird fauna since the introduction of avian malaria at the beginning of the 20th century. In contrast, the Galapagos archipelago still possesses its entire endemic avifauna. Several of these Galapagos bird populations are in decline, however, and wildlife managers seek guidance to counteract a potential man-made ecological disaster. We recommend that endemic birds be tested for susceptibility to disease outside the Galapagos so that protection efforts can be better designed to deal with actual threats. At present, the best and perhaps only management option is to protect the isolation of these island communities because treating or vaccinating wild bird populations against diseases is almost impossible. If the isolation of the Galapagos Islands is successful, we will preserve the complete avifauna of an archipelago for the first time in the history of human colonization in the Pacific eco- region.

Immunological investments reflect parasite abundance in island populations of Darwin's finches

The evolution of parasite resistance can be influenced by the abundance of parasites in the environment. However, it is yet unresolved whether vertebrates change their investment in immune function in response to variation in parasite abundance. Here, we compare parasite abundance in four populations of small ground finches (Geospiza fuliginosa) in the Galapagos archipelago. We predicted that populations exposed to high parasite loads should invest more in immune defence, or alternatively use a different immunological defence strategy. We found that parasite prevalence and/or infection intensity increased with island size. As predicted, birds on large islands had increased concentrations of natural antibodies and mounted a strong specific antibody response faster than birds on smaller islands. By contrast, the magnitude of cell–mediated immune responses decreased with increasing parasite pressure, i.e. on larger islands. The data support the hypothesis that investments into the immune defence are influenced by parasite–mediated selection. Our results are consistent with the hypothesis that different immunological defence strategies are optimal in parasite–rich and parasite–poor environments.

Intraspecific competition and high food availability are associated with insular gigantism in a lizard

Resource availability, competition, and predation commonly drive body size evolution. We assess the impact of high food availability and the consequent increased intraspecific competition, as expressed by tail injuries and cannibalism, on body size in Skyros wall lizards (Podarcis gaigeae). Lizard populations on islets surrounding Skyros (Aegean Sea) all have fewer predators and competitors than on Skyros but differ in the numbers of nesting seabirds. We predicted the following: (1) the presence of breeding seabirds (providing nutrients) will increase lizard population densities; (2) dense lizard populations will experience stronger intraspecific competition; and (3) such aggression, will be associated with larger average body size. We found a positive correlation between seabird and lizard densities. Cannibalism and tail injuries were considerably higher in dense populations. Increases in cannibalism and tail loss were associated with large body sizes. Adult cannibalism on juveniles may select for rapid growth, fuelled by high food abundance, setting thus the stage for the evolution of gigantism.

Tail Shedding in Island Lizards [Lacertidae, Reptilia]: Decline of Antipredator Defenses in Relaxed Predation Environments

The ability of an animal to shed its tail is a widespread antipredator strategy among lizards. The degree of expression of this defense is expected to be shaped by prevailing environmental conditions including local predation pressure. We test these hypotheses by comparing several aspects of caudal autotomy in 15 Mediterranean lizard taxa existing across a swath of mainland and island localities that differ in the number and identity of predator species present. Autotomic ease varied substantially among the study populations, in a pattern that is best explained by the presence of vipers. Neither insularity nor the presence of other types of predators explain the observed autotomy rates. Final concentration of accumulated tail muscle lactate and duration of movement of a shed tail, two traits that were previously thought to relate to predation pressure, are in general not shaped by either predator diversity or insularity. Under conditions of relaxed predation selection, an uncoupling of different aspects of caudal autotomy exists, with some elements (ease of autotomy) declining faster than others (duration of movement, lactate concentration). We compared rates of shed tails in the field against rates of laboratory autotomies conducted under standardized conditions and found very high correlation values (r > 0.96). This suggests that field autotomy rates, rather than being a metric of predatory attacks, merely reflect the innate predisposition of a taxon to shed its tail.

MITES ASSOCIATED WITH THE SMALL GROUND FINCH, GEOSPIZA FULIGINOSA (PASSERIFORMES: EMBERIZIDAE), FROM THE GALÁPAGOS ISLANDS

In collections of ectoparasites from 368 small ground finches, Geospiza fuliginosa, in populations from the islands of Isabela, Santa Cruz, San Cristóbal, and Santa Fé, in the Galápagos Archipelago, Ecuador, we found 8 species of mites. Four mite species were common on all islands sampled, i.e., Mesalgoides geospizae Mironov and Pérez (Psoroptoididae), Xolalges palmai Mironov and Pérez (Xolalgidae), and 2 new species, Trouessartia geospiza n. sp. (Trouessartiidae) and Proctophyllodes darwini n. sp. (Proctophyllodidae). Four other species were represented by single collections from G. fuliginosa, i.e., Pterodectes atyeoi n. sp. (Proctophyllodidae), Strelkoviacarus sp. (Analgidae), Dermoglyphus sp. (Dermoglyphidae), and Dermanyssus sp. (Dermanyssidae). Authorship of new species names is attributed to the 3 authors who prepared the descriptions (B.M.O.C., J.F., D.L.). Trouessartia geospiza and P. atyeoi were also found on previously collected specimens of other Geospiza species in museum collections. For the 4 common species, we found no differences in prevalence among the 4 island populations, but infection prevalence differed among the 4 species. The mean infection prevalence was high for T. geospizae (89%), moderate for M. geospizae (58%) and X. palmai (44%), and low for P. darwini (26%) in all populations. The feather mite fauna of G. fuliginosa was similar to that of other Geospiza species, and generally related to communities found on other emberizid finches.

Effects of fragmentation on genetic diversity in island populations of the Aegean wall lizard Podarcis erhardii (Lacertidae, Reptilia).

Landbridge islands offer unique opportunities for understanding the effects of fragmentation history on genetic variation in island taxa. The formation of islands by rising sea levels can be likened to a population bottleneck whose magnitude and duration is determined by island area and time since isolation, respectively. The Holocene landbridge islands of the Aegean Sea (Greece) were formed since the last glacial maximum and constitute an ideal system for disentangling the effects of island area, age and geographic isolation on genetic variability. Of the many reptile species inhabiting this island system, the Aegean wall lizard Podarcis erhardii is an excellent indicator of fragmentation history due to its widespread distribution and poor over-water dispersal abilities. In this study, we utilize a detailed record of Holocene fragmentation to investigate the effects of island history on wall lizard mitochondrial and nuclear microsatellite diversity. Findings show that the spatial distribution of mitochondrial haplotypes reflects historical patterns of fragmentation rather than geographic proximity per se. In keeping with neutral bottleneck theory, larger and younger islands retain more nuclear genetic variation than smaller, older islands. Conversely, there is no evidence of an effect of isolation by distance or effect of distance to the nearest larger landmass on genetic variability, indicating little gene flow between islands. Lastly, population-specific measures of genetic differentiation are inversely correlated with island area, suggesting that smaller islands exhibit greater divergence due to their greater susceptibility to drift. Taken together, these results suggest that both island area and time since isolation are important predictors of genetic variation and that these patterns likely arose through the progressive fragmentation of ancestral diversity and the ensuing cumulative effects of drift.

Reproductive Biology of Insular Reptiles: Marine Subsidies Modulate Expression of the “Island Syndrome”

Abstract Differences in ecological conditions can result in the evolution of dramatic inter-population shifts in whole suites of traits. We studied variation in reproductive output in three lizard populations of the Skyros Wall Lizard (Podarcis gaigeae, Lacertidae) endemic to the Skyros Archipelago (Greece), which live under similar climatic conditions but differ in predation pressure and food availability. Based on the “island syndrome” hypothesis, we predicted that females from island populations would produce larger, but fewer offspring. The study populations differ conspicuously in average body size, with males from the satellite Lakonissi and Diavates islets being respectively 20% and 39% larger than males from the main Skyros Island. Lizards from these predator-free islets produced eggs of larger size than the main Skyros population; however, they also produced significantly larger clutches than the Skyros population (2.31±0.83 and 2.73±1.0 vs. 1.97±0.58 eggs). All inter-population differences in clutch size, clutch volume, and egg size were explained by corresponding differences in average body size of the dams, revealing that across all populations, reproductive effort scaled similarly with maternal body size. There was no evidence of trade-offs between egg size and clutch size as generally encountered in many reptile taxa. The occurrence of this unusual pattern of reproductive investment among islet populations of giants is probably best explained by the occurrence of two underlying drivers: first, the substantial marine subsidies by resident seabird colonies and second, the existence of intense cannibalistic behaviors in the form of attacks to the tail and severe intraspecific predation on juveniles. This suggests that subsidies-driven gigantism in island endemics may free species from such trade-offs and allow a population to maximize reproductive output in multiple, normally conflicting dimensions.

Diversity, ecological structure and conservation of herpetofauna in a Mediterranean area (Dadia National Park, Greece)

The current study attempts to explore the diversity patterns and the ecological structure of herpetofauna in order to provide a guideline for its conservation management, taking as case study the Dadia national park, in Greece. We surveyed 36 sites by conducting time constraint visits and random transects to sample semi-aquatic (amphibians and freshwater terrapins) and terrestrial (lizards and terrestrial tortoises) herpetofauna respectively. We recorded 20 herpetofauna taxa, including five protected species. The park authorities should maintain brooks as a high priority habitat for semi-aquatic herpetofauna, periodically flooded land and lowland streams as important habitats, and to a lesser degree anthropogenic wet habitats. Semi-aquatic species have narrow habitat requirements related to substrate type and humidity. Terrestrial herpetofauna species are influenced by the degree of shade and the type of substrate; they favour a diversity of semi-open habitats (open oakwoods, heaths), located in the buffer zone rather than in the core of the park that contains densely forested habitats. Management focusing on raptors, the initial conservation priority of the national park, is compatible with the conservation of the terrestrial herpetofauna, as the proposed maintenance of an open forest structure is beneficial for terrestrial herpetofauna as well. Our research indicates that future management programs will have to address the needs of herpetofauna species independently and that they should also be integrated in the monitoring programme of the park, with emphasis on the conservation status and trend of the five protected species.

Digestive performance in five Mediterranean lizard species: effects of temperature and insularity.

Temperature sensitivity of digestive processes has important ramifications for digestive performance in ectothermic vertebrates. We conducted a comparative analysis of temperature effects on digestive processes [gut passage times (GPTs) and apparent digestive efficiencies (ADEs)] in five lacertid lizards occurring in insular (Podarcis erhardii, P. gaigeae), and mainland (P. muralis, P. peloponnesiaca, Lacerta graeca) Mediterranean environments. GPTs were negatively correlated to temperature with mainland taxa having 10–20% longer GPTs than island taxa. In contrast to previous studies that estimate ADEs using bomb calorimetry, we compare ADEs by analyzing discrete efficiencies for lipids, sugars and proteins at three temperature regimes (20, 25, and 30°C); each of these categories produces different results. ADEs for lipids and sugars showed a monotonic increase with temperature whereas ADEs for proteins decreased with temperature. Island taxa had consistently higher ADEs than their mainland counterparts for lipids and for proteins but not for sugars. They are characterized by superior energy acquisition abilities despite significantly shorter GPTs. Their increased digestive performance relative to the mainland species appears to allow them to maximize energy acquisition in unproductive island environments where food availability is spatially and seasonally clustered.