Nestedness of desert bat assemblages: species composition patterns in insular and terrestrial landscapes

Nested patterns of community composition exist when species at depauperate sites are subsets of those occurring at sites with more species. Nested subset analysis provides a framework for analyzing species occurrences to determine non-random patterns in community composition and potentially identify mechanisms that may shape faunal assemblages. We examined nested subset structure of desert bat assemblages on 20 islands in the southern Gulf of California and at 27 sites along the Baja California peninsula coast, the presumable source pool for the insular faunas. Nested structure was analyzed using a conservative null model that accounts for expected variation in species richness and species incidence across sites (fixed row and column totals). Associations of nestedness and island traits, such as size and isolation, as well as species traits related to mobility, were assessed to determine the potential role of differential extinction and immigration abilities as mechanisms of nestedness. Bat faunas were significantly nested in both the insular and terrestrial landscape and island size was significantly correlated with nested structure, such that species on smaller islands tended to be subsets of species on larger islands, suggesting that differential extinction vulnerabilities may be important in shaping insular bat faunas. The role of species mobility and immigration abilities is less clearly associated with nestedness in this system. Nestedness in the terrestrial landscape is likely due to stochastic processes related to random placement of individuals and this may also influence nested patterns on islands, but additional data on abundances will be necessary to distinguish among these potential mechanisms.     

Of mice and mammoths: evaluations of causal explanations for body size evolution in insular mammals

Aim We investigated the hypothesis that the insular body size of mammals results from selective forces whose influence varies with characteristics of the focal islands and the focal species, and with interactions among species (ecological displacement and release). Location Islands world‐wide. Methods We assembled data on the geographic characteristics (area, isolation, maximum elevation, latitude) and climate (annual averages and seasonality of temperature and precipitation) of islands, and on the ecological and morphological characteristics of focal species (number of mammalian competitors and predators, diet, body size of mainland reference populations) that were most relevant to our hypothesis (385 insular populations from 98 species of extant, non‐volant mammals across 248 islands). We used regression tree analyses to examine the hypothesized contextual importance of these factors in explaining variation in the insular body size of mammals. Results The results of regression tree analyses were consistent with predictions based on hypotheses of ecological release (more pronounced changes in body size on islands lacking mammalian competitors or predators), immigrant selection (more pronounced gigantism in small species inhabiting more isolated islands), thermoregulation and endurance during periods of climatic or environmental stress (more pronounced gigantism of small mammals on islands of higher latitudes or on those with colder and more seasonal climates), and resource subsidies (larger body size for mammals that utilize aquatic prey). The results, however, were not consistent with a prediction based on resource limitation and island area; that is, the insular body size of large mammals was not positively correlated with island area. Main conclusions These results support the hypothesis that the body size evolution of insular mammals is influenced by a combination of selective forces whose relative importance and nature of influence are contextual. While there may exist a theoretical optimal body size for mammals in general, the optimum for a particular insular population varies in a predictable manner with characteristics of the islands and the species, and with interactions among species. This study did, however, produce some unanticipated results that merit further study – patterns associated with Bergmann’s rule are amplified on islands, and the body size of small mammals appears to peak at intermediate and not maximum values of latitude and island isolation.     

The tempo and mode of evolution: body sizes of island mammals

The tempo and mode of body size evolution on islands are believed to be well known. It is thought that body size evolves relatively quickly on islands toward the mammalian modal value, thus generating extreme cases of size evolution and the island rule. Here, we tested both theories in a phylogenetically explicit context, by using two different species-level mammalian phylogenetic hypotheses limited to sister clades dichotomizing into an exclusively insular and an exclusively mainland daughter nodes. Taken as a whole, mammals were found to show a largely punctuational mode of size evolution. We found that, accounting for this, and regardless of the phylogeny used, size evolution on islands is no faster than on the continents. We compared different selection regimes using a set of Ornstein-Uhlenbeck models to examine the effects of insularity of the mode of evolution. The models strongly supported clade-specific selection regimes. Under this regime, however, an evolutionary model allowing insular species to evolve differently from their mainland relatives performs worse than a model that ignores insularity as a factor. Thus, insular taxa do not experience statistically different selection from their mainland relatives.     

Small mammals on Massachusetts islands: the use of probability functions in clarifying biogeographic relationships

Summary Thirty-three insular small mammal communities along the coast of Massachusetts (USA) were surveyed to investigate the biogeographic relationships of the insular communities and to examine the distribution patterns of individual species. Nine species of terrestrial small mammals were observed in the total insular fauna, whereas thirteen occurred on the mainland. The species-area relation yielded a z value of 0.06, which is the lowest value yet reported for insular mammal communities.Multiple logistic regression was used to calculate probability functions for each species in order to identify variables potentially important in determining a species' occurrence on islands and to estimate probabilities of occurrence on islands. Statistically significant and ecologically interpretable functions were obtained for all but one species. Occurrence on islands was positively related to increasing island size in four species and to decreasing island isolation in four species. The extremely low z value, negative correlations of species number with isolation variables, and the inclusion of an isolation variable in the logistic functions of four species indicated that immigration was an important determinant of small mammal occurrence on these islands. There was a positive relationship between population density and number of islands occupied.Logistic regression has several advantages over linear discriminant function analysis, and we suggest that it may be useful in other ecological studies and in the preservation of endangered species.     

Size evolution in island lizards

Aim  The island rule, small animal gigantism and large animal dwarfism on islands, is a topic of much recent debate. While size evolution of insular lizards has been widely studied, whether or not they follow the island rule has never been investigated. I examined whether lizards show patterns consistent with the island rule.
Location  Islands worldwide.
Methods  I used literature data on the sizes of island–mainland population pairs in 59 species of lizards, spanning the entire size range of the group, and tested whether small insular lizards are larger than their mainland conspecifics and large insular lizards are smaller. I examined the influence of island area, island isolation, and dietary preferences on lizard size evolution.
Results  Using mean snout–vent length as an index of body size, I found that small lizards on islands become smaller than their mainland conspecifics, while large ones become larger still, opposite to predictions of the island rule. This was especially strong in carnivorous lizards; omnivorous and herbivorous species showed a pattern consistent with the island rule but this result was not statistically significant. No trends consistent with the island rule were found when maximum snout–vent length was used. Island area had, at best, a weak effect on body size. Using maximum snout–vent length as an index of body size resulted in most lizard populations appearing to be dwarfed on islands, but no such pattern was revealed when mean snout–vent length was used as a size index.
Main conclusions  I suggest that lizard body size is mostly influenced by resource availability, with large size allowing some lizard populations to exploit resources that are unavailable on the mainland. Lizards do not follow the island rule. Maximum snout–vent length may be biased by sampling effort, which should be taken into account when one uses this size index.     

Body size evolution in insular vertebrates: generality of the island rule

Aim My goals here are to (1) assess the generality of the island rule – the graded trend from gigantism in small species to dwarfism in larger species – for mammals and other terrestrial vertebrates on islands and island‐like ecosystems; (2) explore some related patterns of body size variation in insular vertebrates, in particular variation in body size as a function of island area and isolation; (3) offer causal explanations for these patterns; and (4) identify promising areas for future studies on body size evolution in insular vertebrates. Location Oceanic and near‐shore archipelagos, and island‐like ecosystems world‐wide. Methods Body size measurements of insular vertebrates (non‐volant mammals, bats, birds, snakes and turtles) were obtained from the literature, and then regression analyses were conducted to test whether body size of insular populations varies as a function of body size of the species on the mainland (the island rule) and with characteristics of the islands (i.e. island isolation and area). Results The island rule appears to be a general phenomenon both with mammalian orders (and to some degree within families and particular subfamilies) as well as across the species groups studied, including non‐volant mammals, bats, passerine birds, snakes and turtles. In addition, body size of numerous species in these classes of vertebrates varies significantly with island isolation and island area. Main conclusions The patterns observed here – the island rule and the tendency for body size among populations of particular species to vary with characteristics of the islands – are actually distinct and scale‐dependent phenomena. Patterns within archipelagos reflect the influence of island isolation and area on selective pressures (immigration filters, resource limitation, and intra‐ and interspecific interactions) within particular species. These patterns contribute to variation about the general trend referred to as the island rule, not the signal for that more general, large‐scale pattern. The island rule itself is an emergent pattern resulting from a combination of selective forces whose importance and influence on insular populations vary in a predictable manner along a gradient from relatively small to large species. As a result, body size of insular species tends to converge on a size that is optimal, or fundamental, for a particular bau plan and ecological strategy.     

Risk-taking behavior of bull-headed shrikes that recently colonized islands

Individuals which have invaded urbanized environments are reported to engage in riskier behaviors, possibly influenced by the scarcity of predators in urbanized areas. Here, we studied the risk-taking behavior of birds which had invaded a new natural environment, rather than an artificial urban environment, using recently established populations of the bull-headed shrike Lanius bucephalus, which naturally colonized three subtropical islands in Japan. We compared flight initiation distance (FID), the distance at which an individual approached by a human initiates flight, between the islands and the temperate mainland. FID was longer for the insular shrikes compared with the mainland shrikes after controlling for other factors, indicating that the individuals which had invaded a new natural environment had a lower propensity for risk-taking. A possible explanation for these results is that low risk-taking behavior might be adaptive on the islands due to predation by the black rat Rattus rattus, an unfamiliar predator not found in shrike habitats on the temperate mainland. Further studies are needed to examine the nest predation rate, predator species, and nest site selection of these insular shrike populations.     

Does relaxed predation drive phenotypic divergence among insular populations?

The evolution of striking phenotypes on islands is a well‐known phenomenon, and there has been a long‐standing debate on the patterns of body size evolution on islands. The ecological causes driving divergence in insular populations are, however, poorly understood. Reduced predator fauna is expected to lower escape propensity, increase body size and relax selection for crypsis in small‐bodied, insular prey species. Here, we investigated whether escape behaviour, body size and dorsal coloration have diverged as predicted under predation release in spatially replicated islet and mainland populations of the lizard species Podarcis gaigeae. We show that islet lizards escape approaching observers at shorter distances and are larger than mainland lizards. Additionally, we found evidence for larger between‐population variation in body size among the islet populations than mainland populations. Moreover, islet populations are significantly more divergent in dorsal coloration and match their respective habitats poorer than mainland lizards. These results strongly suggest that predation release on islets has driven population divergence in phenotypic and behavioural traits and that selective release has affected both trait means and variances. Relaxed predation pressure is therefore likely to be one of the major ecological factors driving body size divergence on these islands.     

A unified model of island biogeography sheds light on the zone of radiation

Islands acquire species through immigration and speciation. Models of island biogeography should capture both processes; however quantitative island biogeography theory has either neglected speciation or treated it unrealistically. We introduce a model where the dominance of immigration on small and near islands gives way to an increasing role for speciation as island area and isolation increase. We examine the contribution of immigration and speciation to the avifauna of 35 archipelagoes and find, consistent with our model, that the zone of radiation comprises two regions: endemic species diverged from mainland sister-species at intermediate isolation and from insular sister-species at higher levels of isolation. Our model also predicts species-area curves in accord with existing research and makes new predictions about species ages and abundances. We argue that a paucity of data and theory on species abundances on isolated islands highlights the need for island biogeography to be reconnected with mainstream ecology.     

Ecomorphological pattern in Bermuda birds: The influence of competition and implications for nature preserves

Summary Previous ecomorphological studies of introduced land birds on oceanic islands have revealed several patterns consistent with the hypothesis that interspecific competition influenced the assembly process of that community. We extend one of these analyses to Bermuda, which differs from the previous islands in its size, latitude and isolation from the mainland. We tested for the presence of morphological overdispersion in surviving introduced passeriforms. Despite a possible predominance of random events in the assembly process due to Bermuda's small size and the presence of migrant species, the surviving passeriforms were morphologically overdispersed. The presence of this pattern supports the hypothesis that the assembly of the Bermuda passeriform community has been influenced by interspecific competition. This marks the third distinct community of island land birds in which this competitive pattern has been identified. This is particularly interesting because certain attributes of Bermuda and its avifauna make it more equivalent to communities in mainland refuges than avian assemblages on islands studied previously. This suggests that interspecific competition may have important effects on the structure of refuge communities.     

Sexual size dimorphism in island plants: the niche variation hypothesis and insular size changes

The niche variation hypothesis predicts insular populations exhibit increased sexual size dimorphism (SSD), to minimize intraspecific competition. Although many animal taxa conform to this prediction, insular patterns of SSD have yet to be investigated in plants. Here, we tested for differences in SSD of dioecious plants that colonised four island groups (Kermadec, Three Kings, Chatham and Auckland Islands) from New Zealand. Using herbarium collections, we quantified leaf and stem sizes of 263 individuals from 28 dioecious taxa. We developed a novel analytical technique to explore changes in the direction of SSD on islands. Lastly, we tested for evolutionary size changes of male and female plants on islands. The degree of SSD did not vary predictably between insular and mainland taxa, contrary to predictions of the niche variation hypothesis. Furthermore, the direction of SSD was not predictable on islands, while it was consistently female biased on the mainland. Our results suggest that selection favours increased size of both sexes on islands and that SSD is unpredictable for insular plants.     

Evolutionary size changes in plants of the south‐west Pacific

Aim To investigate evolutionary changes in the size of leaves, stems and seeds of plants inhabiting isolated islands surrounding New Zealand. Location Antipodes, Auckland, Campbell, Chatham, Kermadec, Three Kings and Poor Knights Islands. Methods First, we compared the size of leaves and stems produced by 14 pairs of plant taxa between offshore islands and the New Zealand mainland, which were grown in a common garden to control for environmental effects. Similar comparisons of seed sizes were made between eight additional pairs of taxa. Second, we used herbarium specimens from 13 species pairs to investigate scaling relationships between leaf and stem sizes in an attempt to pinpoint which trait might be under selection. Third, we used herbarium specimens from 20 species to test whether changes in leaf size vary among islands located at different latitudes. Lastly, we compiled published records of plant heights to test whether insular species in the genus Hebe differed in size from their respective subgenera on the mainland. Results Although some evidence of dwarfism was observed, most insular taxa were larger than their mainland relatives. Leaf sizes scaled positively with stem diameters, with island taxa consistently producing larger leaves for any given stem size than mainland species. Leaf sizes also increased similarly among islands located at different latitudes. Size changes in insular Hebe species were unrelated to the average size of the respective subgenera on the mainland. Main conclusions Consistent evidence of gigantism was observed, suggesting that plants do not obey the island rule. Because our analyses were restricted to woody plants, results are also inconsistent with the ‘weeds‐to‐trees’ hypothesis. Disproportionate increases in leaf size relative to other plant traits suggest that selection may favour the evolution of larger leaves on islands, perhaps due to release from predation or increased intra‐specific competition.     

Are island and mainland biotas different? Richness and level of generalism in parasitoids of a microlepidopteran in Macaronesia

Island communities are exposed to several evolutionary and ecological processes that lead to changes in their diversity and structure compared to mainland biotas. These phenomena have been observed for various taxa but not for parasitoids, a key group in terms of community diversity and functioning. Here we use the parasitoid communities associated with the moth Acroclita subsequana (Lepidoptera: Tortricidae) in the Macaronesian region, to test whether species richness differs between islands and mainland, and whether island parasitoid faunas are biased towards generalist species. Host larvae were collected on several islands and adjacent mainland, carefully searched for ectoparasitoid larvae and dissected to recover any endoparasitoids. Parasitoids were classified as idiobionts, which usually have a wide host range (i.e. generalists), or koinobionts that are considered specialists. Mainland species richness was lower than expected by chance, with most of the species being koinobionts. On the other hand, island communities showed a greater proportion of idiobiont species. Overall parasitism rates were similar between islands and mainland, but islands had higher rates of parasitism by idiobionts than expected by chance, and mainland areas showed the highest koinobiont parasitism rates. These results suggest that island parasitoid communities are dominated by generalists, in comparison to mainland communities. Several hypotheses may explain this pattern: (1) generalist parasitoids might have better dispersal abilities; (2) they may be less constrained by ‘sequential dependencies’; and (3) island parasitoids probably have fewer competitors and/or predators, thus favouring the establishment of generalists. New studies including multiple hosts, other habitats, and/or more islands are necessary to identify which of these processes shape island parasitoid communities.     

Evidence of connectivity between continental and differentiated insular populations in a highly mobile species

Aim Genetically differentiated insular populations are candidates for independent units for conservation. However, occasional immigration to reduced island populations may occur and potentially have important consequences in their future viability and evolutionary potential. In this study, we investigate the conservation implications of population structure and connectivity of insular and continental populations of a migratory raptor as determined using genetic tools and satellite tracking. Location Western European populations in the Iberian Peninsula and two insular populations in the Mediterranean Sea (Balearic Islands) and Atlantic Ocean (Canary Islands). Methods We genotyped 22 microsatellite loci in 96 Egyptian vultures (Neophron percnopterus) from the Iberian Peninsula, 36 from Menorca (Balearic archipelago) and 242 (85% of the current population) from Fuerteventura (Canary Islands). We analysed genetic variation to estimate structure, gene flow, genetic diversity, effective size and recent demographic history of the populations. Additionally, 19 vultures were marked with satellite transmitters to track their migration routes. Results Insular populations were genetically differentiated from those of the mainland. We detected immigration in the insular populations and within the continental counterpart. We found similar levels of genetic variability between the continent and the islands, and a bottleneck analysis indicated recent sharp population declines in both archipelagos but not on the continent. Main conclusions Our study provides evidence that, in spite of significant differentiation, insular populations of highly mobile species may remain connected with the mainland. Conservation programmes should take into account population connectivity and integrate differentiated units of management within complex units of conservation that can best maintain processes and potential for evolutionary change.     

Terrestrial bird community patterns on the coralline islands of the Dahlak Archipelago, Red Sea, Eritrea

Aim This study aims to explain the patterns of species richness and nestedness of a terrestrial bird community in a poorly studied region. Location Twenty‐six islands in the Dahlak Archipelago, Southern Red Sea, Eritrea. Methods The islands and five mainland areas were censused in summer 1999 and winter 2001. To study the importance of island size, isolation from the mainland and inter‐island distance, I used constrained null models for the nestedness temperature calculator and a cluster analysis. Results Species richness depended on island area and isolation from the mainland. Nestedness was detected, even when passive sampling was accounted for. The nested rank of islands was correlated with area and species richness, but not with isolation. Idiosyncrasies appeared among species‐poor and species‐rich islands, and among common and rare species. Cluster analysis showed differences among species‐rich islands, close similarity among species‐poor and idiosyncratic islands, and that the compositional similarity among islands decreased with increasing inter‐island distance. Thus, faunas of species‐poor, smaller islands were more likely to be subsets of faunas of species‐rich, larger islands if the distance between the islands was short. Main conclusions Species richness and nestedness were related to island area, and nestedness also to inter‐island distances but not to isolation from the mainland. Thus, nestedness and species richness are not affected in the same way by area and distance. Moreover, idiosyncrasies may have been the outcome of species distributions among islands being influenced also by non‐nested distributions of habitats, inter–specific interactions, and differences in species distributions across the mainland. Idiosyncrasies in nested patterns may be as important as the nested pattern itself for conservation – and conservation strategies based on nestedness and strong area effects (e.g. protection of only larger islands) may fail to preserve idiosyncratic species/habitats.     

A comparison of relict versus equilibrium models for insular mammals of the Gulf of Maine

For the mammalian faunas of 24 landbridge islands in the Gulf of Maine (0.003–279 km² in size), area accounts for 86% of variance in species richness. The slope, z , of the species-area curve is 0.247. For the seven largest islands (>10km²), the non-equilibrium hypothesis of relaxation following saturation in the post-Pleistocene is suggested by (1) elevated slope of the species-area curve (0.353), (2) correlation of species richness with island age ( r =- 0.81) and water depth to mainland ( r = -0.70), (3) highly non-random nested subsets of species ranked by island area, and (4) discontinuity with the extremely depauperate faunas of oceanic islands of the eastern North Atlantic. The alternative hypothesis of a dynamic equilibrium determined by recurrent immigration and extinction is supported by (1) documented turnover in 16 species, (2) correlation of species-area residuals with distance ( r = - 0.90), (3) distribution dependent upon vagility with reduction or absence of hibernators and other poor dispersers, (4) low levels of endemism, and (5) congruence of community structure with that of mainland fauna for both trophic level and body size.
I conclude that while some insular populations may be relictual, the faunal composition of most of these islands is dependent on recurrent colonization, much of which takes place over ice bridges. However, true equilibrium is perturbed by climatic shifts, range expansions, and human disturbance.     

Mixed patterns of morphological adaptation to insularity in an aerial displaying bird,the Common Snipe Gallinago gallinago

On islands, colonizing birds may evolve behavioural and morphological adaptations to the new environment, often resulting in changes in body size and reduction or even total loss of flight. These island populations have therefore been used to test hypotheses related to adaptations for flight. However, in certain species in which flight is used not only in foraging and migration but also in mating displays, disentangling the effects of natural and social selection is difficult. Thus, sedentary populations of species that perform aerial displays (such as the Common Snipe Gallinago gallinago that breed in the Azores archipelago) may offer an opportunity to separate the effects of natural and social selection on morphology. If insular Common Snipe respond to the characteristic ecological context of oceanic islands, we expect them to differ from migratory conspecifics in body size and by having relatively smaller and more rounded wings. On the other hand, if social selection exerts a more powerful force over the morphology of this species, we expect that sedentary and migratory birds will not differ in flight‐related characters. We tested these hypotheses by comparing morphological characters measured on live Common Snipe captured in the Azores during the breeding season with those measured on migratory specimens hunted during autumn/winter in mainland Portugal. Sedentary Azorean birds were smaller and had relatively shorter tails but did not show the tendency for insular birds to possess more rounded wings as described in other taxa, including in the Azores. Bergman's rule might explain the difference in body size and shorter tails may be responsible for behavioural differences between populations. The lack of difference in wing shape might be explained by the need of the Common Snipe to perform aerial displays during courtship, suggesting an effect of social selection on the migratory strategy of this species.     

Patterns of Genetic and Reproductive Traits Differentiation in Mainland vs. Corsican Populations of Bumblebees

Populations on islands often exhibit lower levels of genetic variation and ecomorphological divergence compared to their mainland relatives. While phenotypic differentiation in characters, such as size or shape among insular organisms, has been well studied, insular differentiation in quantitative reproductive traits involved in chemical communication has received very little attention to date. Here, we investigated the impact of insularity on two syntopic bumblebee species pairs: one including species that are phylogenetically related (Bombus terrestris and B. lucorum), and the other including species that interact ecologically (B. terrestris and its specific nest inquiline B. vestalis). For each bumblebee species, we characterized the patterns of variation and differentiation of insular (Corsican) vs. mainland (European) populations (i) with four genes (nuclear and mitochondrial, 3781 bp) and (ii) in the chemical composition of male marking secretions (MMS), a key trait for mate attraction in bumblebees, by gas chromatography-mass spectrometry (GC-MS). Our results provide evidence for genetic differentiation in Corsican bumblebees and show that, contrary to theoretical expectations, island populations of bumblebees exhibit levels of genetic variation similar to the mainland populations. Likewise, our comparative chemical analyses of MMS indicate that Corsican populations of bumblebees are significantly differentiated from the mainland yet they hold comparative levels of within-population MMS variability compared to the mainland. Therefore, insularity has led Corsican populations to diverge both genetically and chemically from their mainland relatives, presumably through genetic drift, but without a decrease of genetic diversity in island populations. We hypothesize that MMS divergence in Corsican bumblebees was driven by a persistent lack of gene flow with mainland populations and reinforced by the preference of Corsican females for sympatric (Corsican) MMS. The impoverished Corsican bumblebee fauna has not led to relaxation of stabilizing selection on MMS but to consistent differentiation chemical reproductive traits on the island.     

If hippopotamuses cannot swim,how did they colonize islands?

Owing to their aquatic lifestyle, hippopotamuses are normally believed to have reached islands by swimming. Yet, some studies suggest they cannot swim due to their relatively high density. If so, this raises the question of how hippopotamuses would have reached some islands. Their immigration into the British Isles, Sicily, Malta, Zanzibar and Mafia can be accounted for, because these islands sit on continental shelves and were often linked to the mainland during the Pleistocene glacio‐eustatic sea‐level falls. In contrast, their occurrence in Crete, Cyprus and Madagascar would be more difficult to explain. Available geological evidence does not seem to rule out that the latter islands might have been connected with the nearest mainland areas in very recent times. This study intends to consider possibilities about how hippopotamuses reached islands and to show that more effective collaboration is required among specialists involved with the study of insular evolution, colonization and speciation.