Insular shifts and trade-offs in life-history traits in pond frogs in the Zhoushan Archipelago, China

Island and mainland populations of animal species often differ strikingly in life-history traits such as clutch size, egg size, total reproductive effort and body size. However, despite widespread recognition of insular shifts in these life-history traits in birds, mammals and reptiles, there have been no reports of such life-history shifts in amphibians. Furthermore, most studies have focused on one specific life-history trait without explicit consideration of coordinated evolution among these intimately linked life-history traits, and thus the relationships among these traits are poorly studied. Here we provide the first evidence of insular shifts and trade-offs in a coordinated suite of life-history traits for an amphibian species, the pond frog Rana nigromaculata . Life-history data were collected from eight islands in the Zhoushan Archipelago and neighboring mainland China. We found consistent, significant shifts in all life-history traits between mainland and island populations. Island populations had smaller clutch sizes, larger egg sizes, larger female body size and invested less in total reproductive effort than mainland populations. Significant negative relationships were found between egg size and clutch size and between egg size and total reproductive effort among frog populations after controlling for the effects of body size. Therefore, decreased reproductive effort and clutch size, larger egg size and body size in pond frogs on islands were selected through trade-offs as an overall life-history strategy. Our findings contribute to the formation of a broad, repeatable ecological generality for insular shifts in life-history traits across a range of terrestrial vertebrate taxa.     

A model for the species–area–habitat relationship

Aim To propose a model (the choros model) for species diversity, which embodies number of species, area and habitat diversity and mathematically unifies area per se and habitat hypotheses. Location Species richness patterns from a broad scale of insular biotas, both from island and mainland ecosystems are analysed. Methods Twenty‐two different data sets from seventeen studies were examined in this work. The r² values and the Akaike's Information Criterion (AIC) were used in order to compare the quality of fit of the choros model with the Arrhenius species–area model. The classic method of log‐log transformation was applied. Results In twenty of the twenty‐two cases studied, the proposed model gave a better fit than the classic species–area model. The values of z parameter derived from choros model are generally lower than those derived from the classic species–area equation. Main conclusions The choros model can express the effects of area and habitat diversity on species richness, unifying area per se and the habitat hypothesis, which as many authors have noticed are not mutually exclusive but mutually supplementary. The use of habitat diversity depends on the specific determination of the ‘habitat’ term, which has to be defined based on the natural history of the taxon studied. Although the values of the z parameter are reduced, they maintain their biological significance as described by many authors in the last decades. The proposed model can also be considered as a stepping‐stone in our understanding of the small island effect.     

From sampling stations to archipelagos: investigating aspects of the assemblage of insular biota

Aim To investigate the formation of nestedness and species co‐occurrence patterns at the local (sampling station), the intermediate (island group), and the archipelago scale. Location The study used data on the distribution of terrestrial isopods on 20 islands of the central Aegean (Greece). These islands are assigned to two distinct subgroups (Kyklades and Eastern islands). Methods The Nestedness Temperature Calculator was used to obtain nestedness values and maximally nested matrices, the EcoSim7 software and a modified version of Sanderson (2000 ) method were used for the analysis of species co‐occurrences. Idiosyncratic temperatures of species and the order of species placement in the maximally nested matrices were used for further comparisons among spatial scales. The relationships of nestedness values with beta‐diversity, habitat diversity and a number of ecological factors recorded for each sampling station were also investigated. Results Significant nestedness was found at all spatial scales. Levels of nestedness were not related to beta‐diversity or habitat diversity. Nestedness values were similar among spatial scales, but they were affected by matrix size. The species that contributed most to the nested patterns within single islands were not the same as those that produce nestedness at the archipelago scale. There was significant variation in the frequency of species occurrence among islands and among spatial scales. There was no direct effect of ecological factors on the shaping of patterns of nestedness within individual islands, but habitat heterogeneity was crucial for the existence of such patterns. Positive associations among species prevailed at all scales when species per station were considered, while negative associations prevailed in the species per island matrices. All associations resulted from the habitat structure of sampling stations and from particularities of geographical distributions. Conclusions There was no clear‐cut distinction between nestedness patterns among spatial scales, even though different species, and partially different factors, contributed to the formation of these patterns in each case. There was a core of species that contributed to the formation of nested patterns at all spatial scales, while the patterns of species associations suggested that biotic interactions are not an important causal factor. The results of this study suggest that locally rare species cannot be widespread at a higher spatial scale, while locally common species can have a restricted distribution.     

Correlates of extinction proneness in tropical angiosperms

Rapid losses and degradation of natural habitats in the tropics are driving catastrophic declines and extinctions of native biotas, including angiosperms. Determining the ecological and life-history correlates of extinction proneness in tropical plant species may help reveal the mechanisms underlying their responses to habitat disturbance, and assist in the pre-emptive identification of species at risk from extinction. We determined the predictors of extinction proneness in 1884 locally extinct ( n  = 454) and extant ( n  = 1430) terrestrial angiosperms (belonging to 43 orders, 133 families, and 689 genera) in the tropical island nation of Singapore (699.4 km²), which has lost 99.6% of its primary lowland evergreen rainforest since 1819. A wide variety of traits such as geographical distribution, pollination system, sexual system, habit, habitat, height, fruit/seed dispersal mechanism, and capacity for vegetative re-sprouting were used in the analysis. Despite controlling for phylogeny (as approximated by family level classification), we found that only a small percentage of the variation in the extinction probability could be explained by these factors. Epiphytic, monoecious, and hermaphroditic species and those restricted to inland forests have higher probabilities of extinction. Species dependent on mammal pollinators also probably have higher extinction probabilities. More comparative studies that use species traits to identify extinction-prone plant species are needed to guide the enormous, but essential task of identifying species most in need of conservation action.     

Allozyme diversity and history of distribution expansion in the maritime perennial plant Hedyotis strigulosa (Rubiaceae), distributed over the wide latitudes in the Japanese Archipelago

Allozyme diversity was examined in 30 populations of the maritime perennial plant Hedyotis strigulosa var. parviflora , which is distributed from subtropical islands to the central mainland of Japan. Genetic diversity within populations tended to be larger in southern island populations than in northern mainland populations. In the southern part of the distribution, the population size is generally large and populations are distributed more continuously than in the northern area, resulting in the larger effective size of southern populations as a whole. These factors play a major role in maintaining greater genetic diversity in the southern populations. By contrast, genetic diversity in the northern populations is very low, probably resulting from bottlenecks of population establishments during recolonization from refugial area to the northern areas. Geographically close populations were located near each other in the multidimensional scaling and the phenogram based on genetic distances, suggesting that gene flow among remote populations is rather limited. The pattern of genetic diversity in H. strigulosa var. parviflora is likely caused by the distribution expansion of the species; in the last glacier era, the species was restricted to the southern area; its advance to the northern area is relatively recent. Another variety endemic only to the Daito Islands, H. strigulos a var. luxurians , has lower genetic diversity than H. strigulosa var. parviflora and has genetically diverged from H. strigulosa var. parviflor a.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 679–688.     

How Predator Food Preference can Change the Destiny of Native Prey in Predator–Prey Systems

The aim of this work is to develop and analyse a mathematical model for a predator-2 preys system arising in insular environments. We are interested in the evolution of a native prey population without behavioural traits to cope with predation or competition, after the introduction of alien species. Here, we consider a long living bird population with low fertility rate. We point out the effects of the preference of the predator for either juvenile or adult stages. In addition, we study the impact of alien prey introduction in such a model. We use a reaction-diffusion system with a singular logistic right hand side. The aim of this work is to bring interesting dynamics to the fore. As a first example, oscillatory behaviour takes place in the model without alien preys and when predators have an average preference coefficient. Introduction of alien preys can lead to species extinction.     

Evolution of distribution and habitat patterns in endemic millipedes of the genus Dolichoiulus (Diplopoda: Julidae) on the Canary Islands, with notes on distribution patterns of other Ganarian species swarms

Forty-six species of the genus Dolichoiulus , all endemic, occur on the Canary Islands The highest number of species occur on the largest, highest island (Tenerife); fewest occur on Lanzarote, Fuerteventura (low, xeric), El Hierro and La Palma (small, remote). Most of the Dolichoiulus species live on one island only, as in other endemic Canarian species swarms. The scarcity of pluri-insular Dolichoiulus species, in connection with information on phylogeny, suggests that speciation has mainly taken place within individual islands. Distribution patterns are partly governed by habitat differences between species, but vicariance patterns between species living in the same kind of habitat are evident on La Gomera and Tenerife. Dolichoiulus species occur in all kinds of natural habitats. Laurisilva and cave species are generally paler than other species. In the laurisilva of eastern Tenerife, microhabitat differentiation between species is pronounced. In some, but far from all, cases, species coexisting in the same microhabitat are of different sizes. The ancestral colonizing species of Dolichoiulus is/are hypothesized to have been small and to have lived in coastal habitats. Colonization of higher altitudes was usually accompanied by an increase in body size. Invasion of the laurisilva was usually accompanied by a habitat shift from the ground layer to logs.     

Adaptive variation in head size in Vipera berus L. populations

To prove that predators are morphologically adapted to the size of their prey one has to demonstrate that the morphological variation in the trophic apparatus is related to the prey size distribution and that the variation in the trait has some effect on individual fitness. I have studied geographic variation in relative head length (RHL) of adders, Vipera berus , on the Swedish mainland and on groups of islands in the Baltic Sea, and the relationship between RHL and physical condition, a character related to fitness. I also examined the relationship between RHL and sex and colour morph. Relative head length of adders was smallest on the mainland and increased on the islands with increasing body size of the main prey, Microtus agrestis , suggesting stabilizing selection for head size within each population. There was no difference in RHL between sexes or colour morphs. However, physical condition was positively correlated with RHL, indicating directional selection for larger heads. The observed pattern is interpreted as an evolutionary response to the geographic variation in body size of the main prey species and the smaller number of alternative prey species available on islands.