Metapopulations of moths on islands: a test of two contrasting models

1. We describe a generalized mainland-island metapopulation model which includes migration among the island populations. We test model predictions with quantitative data on more than 200 species of moths in two contrasting networks of small islands. The data include a direct measure of migration rate, based on trapping of moths on rocky skerries with no local populations of the vast majority of species.
2. We predicted that moths which are strong fliers but uncommon on the islands have a higher incidence on scattered islands than on islands in a group, because the latter 'compete' for immigrants from the mainland. In contrast, we predicted that weakly flying species with potentially large local populations on the islands occur more frequently on islands in a group due to enhanced colonization rate.
3. Both predicted patterns were observed. Island occupancy increased significantly with the number of individuals caught on the rocky skerries, which is our measure of migration rate from the mainland, supporting the basic assumption that the species occur on the islands in a balance between colonizations and extinctions.
4. These results demonstrate that the moth metapopulations on islands represent a mixture of Levins's and mainland-island metapopulations, and that the mixture is different for different species in the same landscape.     

Community-wide character displacement in New Zealand skinks

Aim  To test for community-wide character displacement in New Zealand skinks.
Location  Four small islands in the New Zealand archipelago.
Methods  (1) We conducted a field experiment on a single island to evaluate whether prey size selection is correlated with lizard body size. We pitfall trapped 69 skinks from three species, measured several aspects of their morphology and presented each animal with a variety of different-sized prey in a food choice experiment. (2) We tested whether the morphological characteristics associated with prey size selection were evenly partitioned in four island skink communities using null models.
Results  Prey size selection was associated with skink morphology; larger skinks consumed larger prey. Null model analyses showed support for evenly displaced body sizes on one island, weak support on one island and no support on two islands.
Main conclusions  Results showed mixed support for community-wide character displacement in New Zealand skinks. Differences in body sizes appear to reflect the use of different-sized prey. Even differentiation in body sizes on one island suggests that species coexistence is facilitated by interspecific differences in prey size selection. However, little support was found on other islands, suggesting that other factors, such as interspecific differences in habitat selection and/or diurnal activity patterns, may interact with differences in prey size selection to promote coexistence among New Zealand skinks.     

Anagenetic evolution in island plants

Aim  Plants in islands have often evolved through adaptive radiation, providing the classical model of evolution of closely related species each with strikingly different morphological and ecological features and with low levels of genetic divergence. We emphasize the importance of an alternative (anagenetic) model of evolution, whereby a single island endemic evolves from a progenitor and slowly builds up genetic variation through time.
Location  Continental and oceanic islands.
Methods  We surveyed 2640 endemic angiosperm species in 13 island systems of the world, both oceanic and continental, for anagenetic and cladogenetic patterns of speciation. Genetic data were evaluated from a progenitor and derivative species pair in Ullung Island, Korea, and Japan.
Results  We show that the anagenetic model of evolution is much more important in oceanic islands than previously believed, accounting for levels of endemic specific diversity from 7% in the Hawaiian Islands to 88% in Ullung Island, Korea, with a mean for all islands of 25%. Examination of an anagenetically derived endemic species in Ullung Island reveals genetic (amplified fragment length polymorphism) variation equal or nearly equal to that of its continental progenitor.
Main conclusions  We hypothesize that, during anagenetic speciation, initial founder populations proliferate, and then accumulate genetic variation slowly through time by mutation and recombination in a relatively uniform environment, with drift and/or selection yielding genetic and morphological divergence sufficient for the recognition of new species. Low-elevation islands with low habitat heterogeneity are highly correlated with high levels of anagenetic evolution, allowing prediction of levels of the two models of evolution from these data alone. Both anagenetic and adaptive radiation models of speciation are needed to explain the observed levels of specific and genetic diversity in oceanic islands.     

Patterns in the assembly of an island plant community

Aim  To test for patterns in the assembly of an island plant community.
Location  Islands off the west coast of Vancouver Island, British Columbia, Canada.
Methods  Twenty-seven islands were visited by boat, and the abundance of six woody angiosperm species was quantified. Null models were then used to test whether: (1) some species co-occur less than expected by chance (i.e. co-occurrence assembly rule), (2) the incidence and abundance of some species are inversely related to the abundance of other species (i.e. incidence assembly rule), and (3) support for assembly rules precludes evidence for nestedness, which refers to a pattern in species composition in which the species present on depauperate islands form regular subsets of those occurring on progressively more diverse islands.
Results  Most species co-occurred with other species at frequencies expected by chance. However, one species ( Sambucus racemosa ) co-occurred with other species less frequently than randomized expectations. The observed incidence and abundance patterns of most species were also consistent with randomized patterns. However, the incidence and abundance of S. racemosa declined with the abundance of other plant species. Weak, variable support was found for nestedness of the total plant community. However, stronger, consistent support was found after removing S. racemosa from the matrix prior to analyses.
Main conclusions  Most species were assembled on islands in a manner consistent with randomized expectations. However, non-random distributional patterns were observed in one species whose distribution was consistent with the hypothesis that competition limits the assembly of island communities.     

Transit towards floristic homogenization on oceanic islands in the south-eastern Pacific: comparing pre-European and current floras

Aim  To quantify the occurrence of processes of homogenization or differentiation in the vascular flora of six oceanic islands.
Location  Six islands in the south-eastern Pacific drawn from the Desventuradas Archipelago, Easter Island and the Juan Fernández Archipelago.
Methods  Using published floristic studies, we determined the floristic composition of each island at two different stages: (1) pre-European colonization and (2) current flora. We compared changes in the number of shared plants and the floristic similarity among islands for each stage.
Results  The number of plant species doubled from 263 in pre-European flora to 531 species currently. Only three native species became extinct, four natives were translocated among the islands and 271 plant species were introduced from outside. The frequency of plant species shared by two or more islands is higher in the post-European floras than prior to European contact, and the level of floristic similarity between islands increased slightly.
Main conclusions  Considering the low naturalization rate of alien plants, the small number of extinctions and the meagre increase in floristic similarity, these islands are undergoing a slow process of floristic homogenization.     

Ecological traits influence the current distribution and range of an island endemic bird

Aim  The aim of this paper is to investigate the causes of the current restricted distribution of a narrow-range endemic bird species, the Canary Islands stonechat, Saxicola dacotiae .
Location  Eastern islands of the Canary Islands archipelago.
Methods  We compared climatic patterns (temperature and rainfall), habitat and microhabitat structure, food availability during a full annual cycle, and the abundance of native avian competitors and predators inside and outside the species' range. Three study areas, located in similar habitats on nearby islands, were studied: northern Fuerteventura, close to the northern border of the species' range; southern Lanzarote, 22 km from the nearest site occupied by stonechats; and the Lobos islet, 10 km from the nearest occupied site and 2 km from the coast of Fuerteventura.
Results  The cover of suitable habitats (slopes with high cover of large shrubs, stony fields and ravines) and microhabitats (shrubs and boulders) and the abundance of arthropods during the breeding period of Canary Islands stonechats were lower outside than inside the species' range. Temperature, rainfall and the abundance of competitors and predators inside and outside the species' range did not differ significantly.
Main conclusions  Ecological requirements explaining the distribution of the Canary Islands stonechat within its range seem to be the main factor hindering its settlement on nearby islands. Geological and palaeoclimatic processes, as well as past and current human impact, could also have constrained the distribution of this narrow-range endemic bird species.     

Habitat diversity, ecological requirements of species and the Small Island Effect

Aim  To explore the causal factors leading to a significant Small Island Effect (SIE), that is, the absence of the commonly found species–area relationships below an island size, on the terrestrial isopod communities from a large number of islands.
Location  Ninety islands of the Aegean Sea (Greece).
Methods  The detection of a significant SIE is assessed through the application of all three methods available in the literature. Species are divided into generalists and specialists. We tested if the minimum area and the area range of each species' occurrences differ between generalists and specialists. Next, we searched for differences in the ratios of specialists to generalists above and below the SIE threshold, and tested their cumulative ratios when islands are arranged according to increasing area, altitude or habitat diversity in order to identify the threshold where they become statistically indistinguishable from the ratio of the total set of islands.
Results  Our results indicate a strong effect of habitat availability on the SIE. Communities of islands within the SIE range, host a higher percentage of generalists. An analysis of the specific habitat requirements shows that, for isopods, the crucial factor is the lack of habitats related to inland waters from small islands.
Main conclusions  The distribution of habitats on islands of different size is of major importance for the occurrence of a SIE. The relative representation of specialist and generalist species on islands of different size plays an important role in shaping SIE-related patterns. Conservation efforts should pay special attention on freshwater habitats, especially on small Aegean islands. Identifying the causal factors of SIE, combined with a thorough knowledge of the ecological requirements of species can offer insights into identifying habitat types and groups of species that are more vulnerable to alterations of the environment.     

Enhanced niche opportunities: can they explain the success of New Zealand's introduced bird species?

Aim  The niche hypothesis could explain why some species introduced to new locations reach higher densities than in their native range: it posits that the new environment provides more abundant or higher quality resources or habitat, a more suitable physical environment or both. We investigate whether 11 bird species occur at higher densities in their introduced range than in their native range and whether the differences can be explained by the availability of preferred habitat or the suitability of climatic conditions in their introduced range relative to their native range.
Location  South Island, New Zealand (the introduced range); UK (the native range).
Methods  We first develop a series of models that accurately predict the density of 11 bird species at 54 UK farmland sites, which are closely matched to our New Zealand sites, from habitat and climatic variables. We then use these models to predict the density of the 11 species at 54 New Zealand farmland sites and compare the predicted and observed values.
Results  Actual densities at New Zealand sites were on average (median) 22 times (range: 1–6361) higher than predicted from the UK models and similarly higher than actually observed at comparable UK sites. Habitat and climatic variables can accurately predict bird densities in the UK but grossly underestimate densities for all species except Turdus merula in New Zealand.
Main conclusions  These findings indicate that factors other than the measured habitat and climatic variables must differ between the two regions and explain the much higher densities of New Zealand birds. We suggest that introduced birds, other than T. merula , in New Zealand may still experience enhanced niche opportunities due to greater availability of higher quality resources within habitats, release from natural enemy regulation, less exposure to extreme weather events, particularly during winter, or some combination of these processes.     

High local species richness of parasitic wasps (Hymenoptera: Ichneumonidae; Pimplinae and Rhyssinae) from the lowland rainforests of Peruvian Amazonia

Abstract.  1. The parasitic wasp family Ichneumonidae (Hymenoptera) is of great interest because it has been claimed that its species richness does not increase with decreasing latitude.
2. No extensive studies of the family have been conducted in South American localities.
3. Arthropods were sampled using 27 Malaise traps in the Allpahuayo–Mishana National Reserve (56 000 ha) in the north-eastern Peruvian Amazonian lowland rainforest. The total duration of the sampling programme was 185 Malaise trap months.
4. Altogether, 88 species were collected. This is one of the highest local pimpline and rhyssine species numbers ever recorded. A comparison with results from Mesoamerica revealed that at equal numbers of individuals sampled, the number of Pimplinae and Rhyssinae species in Peruvian Amazonia is at least twofold compared with lowland locations in Mesoamerica and somewhat higher than in the most species-rich Costa Rican higher altitude localities.
5. Non-parametric methods of estimating species richness were applied. These suggest that additional sampling would yield a considerable number of new Pimplinae and/or Rhyssinae species.     

Echoes of the whispering land: interacting roles of vicariance and selection in shaping the evolutionary divergence of two <Emphasis Type="Italic">Calceolaria</Emphasis> (Calceolariaceae) species from Patagonia and Malvinas/Falkland Islands

A key to understanding the origin and identity of young species lays on the knowledge of the Quaternary climatic oscillations’ effect on gene flow and vicariance. Even though the effect of climatic fluctuations is relatively well understood for southern hemisphere plant species, little is known about their effect on the evolutionary histories of species from mainland and islands. Thus, we investigated whether Quaternary climate-driven fluctuations translated into lineage divergence and speciation, followed or not by climatic niche differentiation, in two allopatric plant species, Calceolaria uniflora and C. fothergillii from Patagonia and Malvinas/Falkland islands, respectively. We sampled the range of both species, and sequenced two chloroplastic (cpDNA; trnS–trnG and trnH–psbA), and one single copy “anonymous” non-coding nuclear region (nDNA). We performed phylogeographic and dating analyses, and adjusted spatio-temporal diffusion models. We complemented molecular evidence with climatic niche differentiation analyses and species paleo-distribution projections. A species coalescent reconstruction based on multi-locus data retrieved both species as monophyletic. Estimates from cpDNA indicated the species diverged during the Great Patagonian Glaciation. Chloroplast and nuclear DNA showed east–west distribution of the main genetic groups but with contrasting spatial genetic diversity. The spatio-temporal diffusion analyses showed that between 1–0.8 Mya and 570 Kya the lineage leading to C. fothergillii diverged from C. uniflora and arrived to the islands. Climatic niche projections hindcasted range expansions during glaciations, and contractions during the interglacial periods. Comparisons of climatic niches between the two study species indicated that temperature variables show evidence of niche conservatism while precipitation regimes supported niche divergence, even when considering the background environmental divergence. Our study indicates that glacial fluctuations affected the mainland/islands connections favouring speciation mediated not only by isolation, but also by climatic niche differentiation.     

Distributional patterning of terrestrial herpetofauna on the Wessel and English Company Island groups,northeastern Arnhem Land,Northern Territory,Australia

Forty-four species of terrestrial reptiles and eight species of frogs were recorded from 60 continental islands of the Wessel and English Company groups off northeastern Arnhem Land, Northern Territory. Two gecko species, Oedura rhombifer and Heteronotia binoei, were present on the most islands (34 and 31, respectively), and occurred on islands < 5 ha. In contrast, agamids, pygopodids and varanids were absent from islands < 18 ha, and snakes and frogs were not reported from islands < 240 ha. Island size explained 82% of the variation in species richness for terrestrial reptiles, and 84% of that for lizards. The relationship was less good for (i) groups with generally uncommon species (notably snakes), for which sampling effort explained more variation, and (ii) groups with species which had relatively specific habitat requirements (notably frogs), for which island size and isolation factors were not especially relevant. For most taxonomic groups considered, isolation factors added little to the relationship between species richness and island size. Across all reptiles, larger species were found on fewer islands, and had larger island size thresholds. This relationship broke down with analysis restricted to the single most species-rich family, Scincidae. Only 6 of the 20 most frequently recorded species showed significant variation in abundance among 8 vegetation types sampled by 226 quadrats across 40 islands. The number of species (alpha-diversity) and total abundance of herpetofauna within quadrats was generally unrelated to island size; however, (with analysis restricted to islands on which they occurred) six individual species were significantly more abundant on smaller islands than on larger islands, with no species showing the opposite pattern. The islands’ herpetofauna is largely a relatively depauperate subset of that of the far more complex sandstone massif and escarpment of western Arnhem Land, especially missing species associated with rugged sandstone gorges, riparian areas, open forests, swamps and clay soils. Patterns in species richness and composition are explained by greater range of environments on larger islands allowing better retention of species since isolation and/or richer tallies at the time of isolation. The evidence suggests that there has been relatively little colonization, although at least two gecko species and one varanid may have moved reasonably frequently.     

The island biogeography of exotic bird species

Aim   A recent upsurge of interest in the island biogeography of exotic species has followed from the argument that they may provide valuable information on the natural processes structuring island biotas. Here, we use data on the occurrence of exotic bird species across oceanic islands worldwide to demonstrate an alternative and previously untested hypothesis that these distributional patterns are a simple consequence of where humans have released such species, and hence of the number of species released.
Location   Islands around the world.
Methods   Statistical analysis of published information on the numbers of exotic bird species introduced to, and established on, islands around the world.
Results   Established exotic birds showed very similar species–area relationships to native species, but different species–isolation relationships. However, in both cases the relationship for established exotics simply mimicked that for the number of exotic bird species introduced. Exotic bird introductions scaled positively with human population size and island isolation, and islands that had seen more native species extinctions had had more exotic species released.
Main conclusion   The island biogeography of exotic birds is primarily a consequence of human, rather than natural, processes.     

The effect of island area on body size in a primate species from the Sunda Shelf Islands

Aim  We examine the effect of island area on body dimensions in a single species of primate endemic to Southeast Asia, the long-tailed macaque ( Macaca fascicularis ). In addition, we test Allen's rule and a within-species or intraspecific equivalent of Bergmann's rule (i.e. Rensch's rule) to evaluate body size and shape evolution in this sample of insular macaques.
Location  The Sunda Shelf islands of Southeast Asia.
Methods  Body size measurements of insular macaques gathered from the literature were analysed relative to island area, latitude, maximum altitude, isolation from the mainland and other islands, and various climatic variables using linear regression.
Results  We found no statistically significant relationship between island area and body length or head length in our sample of insular long-tailed macaques. Tail length correlated negatively with island area. Head length and body length exhibited increases corresponding to increasing latitude, a finding seemingly consistent with the expression of Bergmann's rule within a single species. These variables, however, were not correlated with temperature, indicating that Bergmann's rule is not in effect. Tail length was not correlated with either temperature or increasing latitude, contrary to that predicted by Allen's rule.
Main conclusions  The island rule dictating that body size will covary with island area does not apply to this particular species of primate. Our study is consistent with results presented in the literature by demonstrating that skull and body length in insular long-tailed macaques do not, strictly speaking, conform to Rensch's rule. Unlike previous studies, however, our findings suggest that tail-length variation in insular macaques does not support Allen's rule.     

Associational resistance mediated by natural enemies

Abstract.  1. Associational resistance theory suggests that the association of herbivore-susceptible plant species with herbivore-resistant plant species can reduce herbivore density on the susceptible plant species. Several casual mechanisms are possible but none has so far invoked natural enemies. Associational resistance mediated by natural enemies was tested for by examining densities of a gall fly, Asphondylia borrichiae (Diptera: Cecidomyiidae), and levels of parasitism on two closely related seaside plants, Borrichia frutescens and Iva frutescens , when alone and when co-occurring.
2. Both Borrichia and Iva grow alone or together on small offshore islands in Florida. Each host plant species has its own associated race of fly, but both races of fly are attacked by the same four species of parasitoids. Borrichia normally has a higher density of galls than Iva , and galls are larger on Borrichia than on Iva .
3. Gall size, gall abundance, parasitism levels, and parasitoid community composition were quantified on both Borrichia and Iva on islands where each species grew alone or together. Some islands were then manipulated by adding Borrichia to islands supporting only Iva , and by adding Iva to islands supporting only Borrichia . Subsequent gall densities and gall parasitism levels on the original native species were then examined.
4. On both natural and experimentally manipulated islands, gall densities on Iva were significantly lowered by the presence of Borrichia . This is because bigger parasitoid species that were common on Borrichia galls, which are bigger, spilled over and attacked the smaller Iva galls. Thus, parasitism rates on Iva were higher on islands where Borrichia co-occurred than on islands where Borrichia were absent. Most parasitoids from Iva were too small to successfully attack the large Borrichia galls and so gall density on Borrichia was unaffected by the presence of Iva .     

Soil nutritional factors improve models of plant species distribution: an illustration with Acer campestre (L.) in France

Aim  To estimate the relative importance of climate and soil nutritional variables for predicting the distribution of Acer campestre (L.) in French forests.
Location  France.
Methods  We used presence/absence information for A. campestre in 3286 forest plots scattered all over France, coupled with climatic and edaphic data. More than 150 climatic variables (temperature, precipitation, solar radiation, evapotranspiration, water balance) were obtained using a digital elevation model (DEM) and a geographical information system (GIS). Six direct soil variables (pH, C/N ratio, base saturation rate, concentrations of calcium, magnesium and potassium) were available from EcoPlant, a phytoecological data base for French forests. Using a forward stepwise logistic regression technique, we derived two distinct predictive models for A. campestre ; the first with climatic variables alone and the second with both climatic and edaphic variables.
Results  The distribution of A. campestre was poorly modelled when including only climatic variables. The inclusion of edaphic variables significantly improved the quality of predictions for this species, allowing prediction of patches of presence/absence within the study region.
Main conclusion  Soil nutritional variables may improve the performance of fine-scale (grain) plant species distribution models.     

Sequential colonization and diversification of Galapágos endemic land snail genus Bulimulus (Gastropoda, Stylommatophora)

Species richness on island or islandlike systems is a function of colonization, within-island speciation, and extinction. Here we evaluate the relative importance of the first two of these processes as a function of the biogeographical and ecological attributes of islands using the Galápagos endemic land snails of the genus Bulimulus, the most species-rich radiation of these islands. Species in this clade have colonized almost all major islands and are found in five of the six described vegetation zones. We use molecular phylogenetics (based on COI and ITS 1 sequence data) to infer the diversification patterns of extant species of Bulimulus, and multiple regression to investigate the causes of variation among islands in species richness. Maximum-likelihood, Bayesian, and maximum-parsimony analyses yield well-resolved trees with similar topologies. The phylogeny obtained supports the progression rule hypothesis, with species found on older emerged islands connecting at deeper nodes. For all but two island species assemblages we find support for only one or two colonization events, indicating that within-island speciation has an important role in the formation of species on these islands. Even though speciation through colonization is not common, island insularity (distance to nearest major island) is a significant predictor of species richness resulting from interisland colonization alone. However, island insularity has no effect on the overall bulimulid species richness per island. Habitat diversity (measured as plant species diversity), island elevation, and island area, all of which are indirect measures of niche space, are strong predictors of overall bulimulid land snail species richness. Island age is also an important independent predictor of overall species richness, with older islands harboring more species than younger islands. Taken together, our results demonstrate that the diversification of Galápagos bulimulid land snails has been driven by a combination of geographic factors (island age, size, and location), which affect colonization patterns, and ecological factors, such as plant species diversity, that foster within-island speciation.     

Freshwater paths across the ocean: molecular phylogeny of the frog Ptychadena newtoni gives insights into amphibian colonization of oceanic islands

Aim Amphibians are a model group for studies of the biogeographical origins of salt‐intolerant taxa on oceanic islands. We used the Gulf of Guinea islands to explore the biogeographical origins of island endemism of one species of frog, and used this to gain insights into potential colonization mechanisms. Location São Tomé and Príncipe, two of the four major islands in the Gulf of Guinea, West Africa, are truly oceanic and have an exceptionally high biodiversity. Methods Mitochondrial DNA is used to test the endemic status of a frog from São Tomé and compare it with congeneric taxa from tropical Africa. Existing data on surface currents, surface salinity, atmospheric circulation and bird migration in the Gulf of Guinea are summarized to address hypotheses concerning colonization mechanisms. Results The endemic status of Ptychadena newtoni (Bocage) is supported here by mitochondrial DNA sequences, and analysis of this and other molecular data indicates that an East African species close to Ptychadena mascareniensis (Duméril and Bibron) is its nearest relative. We refute the possibility that this population was anthropogenically introduced, in favour of a natural dispersal mechanism. Main conclusions With six endemic frogs and one caecilian, the Gulf of Guinea islands harbour a diverse amphibian fauna. Five of these species appear to have their closest relatives in East Africa. Insufficient evidence exists for transportation by storms, birds or rafts alone. However, we propose a synergy of rafting, favourable surface currents and a reduction in salinity of surface waters. Catastrophic events, or wet periods in climatic history, could allow freshwater paths to open far enough to enable continental flora and fauna to reach these and other isolated oceanic islands.     

Why Do Cryptic Species Tend Not to Co-Occur? A Case Study on Two Cryptic Pairs of Butterflies

As cryptic diversity is being discovered, mostly thanks to advances in molecular techniques, it is becoming evident that many of these taxa display parapatric distributions in mainland and that they rarely coexist on islands. Genetic landscapes, haplotype networks and ecological niche modeling analyses were performed for two pairs of non-sister cryptic butterfly species, Aricia agestis-A. cramera and Polyommatus icarus—P. celina (Lycaenidae), to specifically assess non-coexistence on western Mediterranean islands, and to test potential causes producing such chequered distribution patterns. We show that the morphologically and ecologically equivalent pairs of species do not coexist on any of the studied islands, although nearly all islands are colonized by one of them. According to our models, the cryptic pairs displayed marked climatic preferences and ‘precipitation during the driest quarter’ was recovered as the most important climatic determinant. However, neither dispersal capacity, nor climatic or ecological factors fully explain the observed distributions across particular sea straits, and the existence of species interactions resulting in mutual exclusion is suggested as a necessary hypothesis. Given that the studied species are habitat generalists, feeding on virtually unlimited resources, we propose that reproductive interference, together with climatic preferences, sustain density-dependent mechanisms like “founder takes all” and impede coexistence on islands. Chequered distributions among cryptic taxa, both sister and non-sister, are common in butterflies, suggesting that the phenomenon revealed here could be important in determining biodiversity patterns.     

Attributes of the seed bank after a century of primary succession on islands in Lake Hjälmaren, Sweden

1 A large number of islands was created when the water table of Lake Hjälmaren, south central Sweden, was lowered between 1882 and 1886. We have complete lists of vascular plant species for 40 of these islands from 1886, 1892, 1903–04, 1927–28 and 1984–85.
2 We have investigated the seed bank on nine of these islands and compared species composition at different soil depths with the species lists from the islands in 1886–1985, and with the present vegetation in the area of seed bank sampling. We have also investigated the distribution in the soil of seeds from species with different ecological attributes, including seed longevity, successional status, seed weight, seed form and species longevity.
3 Seeds in soil samples were allowed to germinate over the course of two summers with an intermediate cold storage. We found 1944 seeds representing 65 taxa. The mean seed density was 84 seeds dm ^–2 .
4 The similarity between the surface soil (0–3 cm) seed bank and the vegetation at the different vegetation analyses increased from 1886 to 1993. The similarity between the present vegetation and the seed bank decreased with increasing soil depth, and the soil at 12–15 cm had no species in common with the present vegetation. Several species now absent from the vegetation were found in the seed bank.
5 Deeply buried seeds came from early successional, annual species with long-term persistent and low-weight seeds, as expected from seed bank theories, but were slightly elongated, which was in contrast to theories. Spherical seeds were associated with the surface soil, as were short-lived and high-weight seeds from late successional, perennial species.     

Effects of site and species characteristics on nested patterns of species composition in sedge meadows

Biotas of both geographical islands and habitat islands are often nested subsets of the biotas of successively more species-rich islands within the same system. The life history characteristics of a species may determine how that species contributes to the general pattern of species nestedness. Here, I investigate the floras of 56 sedge meadow wetlands in northern Illinois (USA) in order to characterize the degree of nestedness in these communities, determine which individual plant species contribute to the nested pattern, and investigate species characteristics that might be related to nonrandom patterns of distribution in individual plant species. The entire assemblage of species at all sedge meadows was significantly nested. Species richness and area were significantly correlated, and the nested pattern was closely related to site area, suggesting that species drop out of the assemblage in a predictable order as site area decreases. Some individual species exhibited nonrandom distributions across the sites, occurring more often in large, species-rich sites. Large sites were more likely than smaller sites to contain conservative species, i.e., those typical of pristine natural habitat, whereas nonconservative species were distributed more randomly among sites. Nested patterns of distribution of conservative species with respect to site area may result from their high probability of extinction on small sites or from a tendency for required habitats to co-occur on the same large sites. This revised version was published online in June 2006 with corrections to the Cover Date.