Testing for MacArthur-Wilson Equilibrium with the Arthropods of the Miniature Spartina Archipelago at Oyster Bay, Florida

SYNOPSIS. The unique features of the MacArthur-Wilson modelof equilibrium biogeography are the immigration and extinctioncurves. In the model, the immigration rate for each island isnegatively correlated with species number, and island extinctionrate is positively correlated with species number. Direct tests of the sign and statistical significance of immigrationand extinction curves were done, with data on arthropods oftiny Spartina alterniflora islets, in north Florida. Rey (1981)defaunated six islets in the spring, then censused the recolonizationweekly, for one year. Two control islets and a mainland plotwere also censused. Distinct tendencies in favor of MacArthur-Wilsoncorrelations were present for both immigration and extinction.But sufficient extraneous variation occurs in immigration andextinction rates to make correspondence to MacArthur-Wilsoncurves statistically non-significant, unless probabilities arecombined among islands. Thus, mathematical deductions from theMacArthur-Wilson model, which treat the curves as deterministiclines with no variance, are less appropriate than approachesthat incorporate stochastic variation. The study of community patterns, by models such as the MacArthur-Wilsonmodel, are not a substitute for careful autecological studies.Mechanisms of colonization and co-existence are only dimly hintedat by gross community patterns.     

Effects of isolation,logging and dispersal on woody-species richness of islands

Isolation effects on species richness of woody plants were investigated in a system of islands that were created by the filling of the Clarks Hill Reservoir, Georgia. This reservoir was built between 1946–1954. Some islands were logged and cleared of woody plants prior to the filling of the reservoir and others were not logged. The presence of logged versus unlogged islands in the same system allowed us to test whether and how geographical isolation interacts with island history and species-specific dispersal properties in determining patterns of among-island variation in species number. Thirty-six years after the islands were created, logged islands had significantly fewer species of woody plants than unlogged ones. On logged islands, total number of woody species was negatively correlated with distance to the closest mainland (r=–0.95). On unlogged islands, variation in species number was very low (CV=4.9%) and was not correlated with distance to the mainland. These results indicate that the studied system as a whole has not yet reached equilibrium. However, the mean number of species on unlogged islands was very close to the intercept of the regression obtained for logged islands, suggesting that islands close to the mainland have already reached their equilibrium species richness. This conclusion is consistent with predictions of island biogeography theory. When species representing different dispersal properties were analyzed separately, statistically significant distance effects were obtained for bird-dispersed species (r=0.88) and for species with no adaptations to bird or wind dispersal (r=0.81). Wind-dispersed species did not show a decrease in species number with increasing isolation, but their relative frequency was positively and significantly correlated with distance to the mainland (r=0.94). Historical factors, as well as differences among species in dispersal properties, are important in explaining patterns of among-island variation in species number.     

The form of the curves: a direct evaluation of MacArthur & Wilson's classic theory

1. We calculate the yearly numbers of bird species immigrating to – and becoming extinct on – 13 small islands of the British Isles, using a long and relatively complete data record.
2. We estimate the size of the colonist pool for each island using four methods.
3. We assume that immigrations and extinctions are distributed binomially, and use a maximum likelihood method to fit concave immigration and extinction functions to the data, utilizing all four species pool estimates.
4. Extinction rates increase significantly and consistently with increasing numbers of breeding species on each island. For nine of the 13 islands the extinction functions are significantly concave.
5. Immigration rates decrease consistently with increasing numbers of breeding species on each island. Seven islands have significantly concave immigration functions.
6. Immigration rates and extinction rates decline consistently, but not significantly, with island distance and island size, respectively. The number of breeding species does not always reflect the number of species likely to have reached an island. Moreover, some species may choose not to breed when their chance of extinction is high. These factors, plus the modest range of island areas and distances in our database, reduce our chances of finding the theoretically predicted effects of area and distance on extinction and immigration rates.     

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.     

Nested distribution of amphibians in the Zhoushan archipelago, China: can selective extinction cause nested subsets of species?

Because of their poor dispersal ability, amphibians are well suited for testing the selective extinction theory on islands. Amphibian fauna in the Zhoushan archipelago, China, exhibit a high level of nestedness (C = 0.893), and the species number is lower on islands than on similar sized areas on the mainland. No correlation was found between island-specific species richness and the nearest distance from a larger island, distance from the mainland or density of human population. These results suggest that no amphibian colonisation has occurred in the archipelago since island isolation 7000–9000 years ago. Furthermore, the results imply that selective extinction contributes to the nestedness of amphibians in the Zhoushan archipelago. The incidence of a species on the islands is significantly correlated with log area of the smallest island occupied by the species and the number of provinces on the Chinese mainland in which the species occur. However, there is no correlation with average body length of adults and island occurrence. It is concluded that (1) the area of the smallest island occupied by a species is a good estimate of the minimum area for a viable population of the species and a good predictor of species incidence on islands, (2) species with a restricted distribution range are more vulnerable to extinction from islands than those with a wide distribution range and (3) the effect of body size on occurrence on the islands is uncertain, and may be specific to the archipelago and taxa studied. The observed nestedness of amphibian assemblages has two implications for conservation: (1) not only can all the species found in several small reserves be found on a large reserve of the same total size, but additional species can be found on the single large reserve; (2) for a reserve to maintain viable populations of all species in a region it should be at least as large as the smallest island occupied by the most vulnerable species. Received: 16 December 1996 / Accepted: 22 September 1997     

Genetic and phylogenetic consequences of island biogeography

Abstract.— Island biogeography theory predicts that the number of species on an island should increase with island size and decrease with island distance to the mainland. These predictions are generally well supported in comparative and experimental studies. These ecological, equilibrium predictions arise as a result of colonization and extinction processes. Because colonization and extinction are also important processes in evolution, we develop methods to test evolutionary predictions of island biogeography. We derive a population genetic model of island biogeography that incorporates island colonization, migration of individuals from the mainland, and extinction of island populations. The model provides a means of estimating the rates of migration and extinction from population genetic data. This model predicts that within an island population the distribution of genetic divergences with respect to the mainland source population should be bimodal, with much of the divergence dating to the colonization event. Across islands, this model predicts that populations on large islands should be on average more genetically divergent from mainland source populations than those on small islands. Likewise, populations on distant islands should be more divergent than those on close islands. Published observations of a larger proportion of endemic species on large and distant islands support these predictions.     

Colonization of Sphagnum on land uplift islands in the Baltic Sea: time, area, distance and life history

Aim To test whether species richness of Sphagnum mosses on islands in a land uplift archipelago is related to island age, area or connectivity, and whether the frequency of different species can be predicted by their life history and autecology. Location The northern Stockholm archipelago in the Baltic Sea, east‐central Sweden, with a current land uplift rate of 4.4 mm year^?1. Methods We sampled 17 islands differing in area (0.55–55 ha), height (3.6–18 m, representing c. 800–4000 years of age) and distance from mainland (1.6–41 km). For each Sphagnum patch we measured area, height above sea level, horizontal distance from the shore and shading from vascular plants. Factors affecting island species richness, species frequency and habitats on the islands were tested by stepwise regressions. Species frequency was tested on nine life history and autecological variables, including estimated abundance and spore output on the mainland, habitat preference and distribution. Results We recorded 500 patches of 19 Sphagnum species, distributed in 83 rock pools on 14 islands. Island species richness correlated positively with island area and with degree of shelter by surrounding islands, while distance from the mainland, connectivity, height or age did not add to the model. Species frequency (number of colonized islands and rock pools) was mainly predicted by spore output on the mainland and by habitat preference (swamp forest species were more frequent than others), while spore size, for example, did not add to the model. Species differed in mean height above and horizontal distance from the shore, area of occupied rock pools and in the degree of shading of patches. The mean horizontal distance from the shore and the area of occupied rock pools correlated positively with the normal growth position above the water table among species. Spore capsules were found in only 2% of patches, mostly in the bisexual Sphagnum fimbriatum. Main conclusions The presence of Sphagnum in the Stockholm archipelago seems to be governed by regional spore production and habitat demands. Sphagnum does not appear to be dispersal limited at distances up to 40 km and time spans of centuries. Species with a high regional spore output have had a higher colonization rate, which, together with the rarity of spore capsules on the islands, indicate the mainland as a source for colonization rather than dispersal among islands. Swamp forest species seem more tolerant to the island conditions (summer droughts and some salt spray) than open mire species. The different distances from the sea occupied by the species indicate a slow, continuous succession and species replacement towards the island interior as islands are being uplifted and thus expand in area. This partly explains why larger islands harbour more species. Our results thus support some of the island biogeographical theories related to the species–area relationship.     

Patterns of endemic extinctions among island bird species

The relationship between island biogeography and the vulnerability of island biota to extinction as a result of human activities was examined. In particular, this study analyzed whether island area, maximum elevation of an island, isolation from the nearest continental landmass, or date of human colonization had statistically significant relationships with the proportion of endemic island bird species that have become endangered or extinct. The study examined islands or island groups with endemic bird species, and which have never been connected to a continental landmass. Both modern and fossil bird species were incorporated into the analysis. Islands that were colonized by humans earliest had the lowest proportion of modern species alone, and modern and fossil species combined, that have gone extinct. However, date of human arrival was not correlated with the proportion of modern species that are endangered. Maximum elevation of an island was negatively correlated with the proportion of modern species that are extinct, and was positively correlated with the proportion of modern species that are endangered. Area was negatively correlated with the proportion of modern species that are endangered. Isolation of islands was not significantly correlated with the proportion of modern species extinct or endangered, but was positively correlated with the proportion of modern and fossil species combined that have gone extinct. These results indicate that the initial spasm of island bird extinctions due to human contact may have, in part, passed. They also indicate that bird species on islands colonized earliest by humans may have had more time to adapt to the presence of man and his commensal species, resulting in reduced extinction rates.     

Species immigration, extinction and turnover of vascular plants in boreal lakes

Dictated by limited resource availability for land acquisition, a central question in conservation biology is the ability of areas of different size to maintain species diversity. The selected reserves should not only be species rich at the moment, but should also maintain species diversity in the long run. We used two sets of data on vascular plant species in boreal lakes collected in 1933/34 and 1996 to test the relationships between lake area and the extinction, immigration and turnover rates of the species. Moreover, we investigated, whether the number of species in 1933/34 or water connection between lakes was related to extinction, immigration and turnover rates of species. We found that lake area or shoreline length was not correlated with immigration or turnover rate. But extinction rate was slightly negatively correlated with shoreline length. The original number of species was positively related to the number of species extinctions and to the absolute turnover rate in the lakes, which indicates that species richness does not create stability in these communities. Species number was not correlated with immigration rate. Upstream water connections in the lakes did not affect immigration, extinction or turnover rates. We conclude that length of the shoreline is a better measure of suitable area for water plants than the lake area, and that because the correlation between shoreline length and extinction rate was slight, also small lakes can be valuable for conservation.     

Factors explaining increased body size in common shrews (Sorex araneus) on Scottish islands

Aim  Island populations of small mammals are often characterized by a larger body size compared with neighbouring mainland or continental populations of the same species. A number of reasons have been put forward to explain this phenomenon. The aim of this study was to test which of these hypotheses can best explain the increase of body size in common shrews ( Sorex araneus ) on islands.
Location  The fieldwork for this study was carried out on the islands of the Inner Hebrides, Clyde Islands and the west coast of Scotland.
Methods  This study compared body sizes of common shrews from mainland and island sites on the west coast of Scotland, based on measurements of hind foot lengths. On 10 of the 13 islands sampled, common shrews were significantly larger than on the mainland. Body size did not vary significantly among mainland populations. We used the directional contrasts method to test the relative contributions of possible factors explaining the large body size observed in the island populations.
Results  We found that body size of common shrews on islands was positively related to distance from mainland, negatively related to average annual temperature, negatively related to island size, and may also be influenced by the presence or absence of pygmy shrews ( Sorex minutus ) on the island.
Main conclusions  Our results suggest a role for founder events, Bergmann's rule and K -selection in determining body size of common shrews on islands.     

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.     

Landscape structure and land use history influence changes in island plant composition after 100 years

Aim We investigated how current and historical land use and landscape structure affect species richness and the processes of extinction, immigration and species turnover. Location The northern part of the Stockholm archipelago, Baltic Sea, Sweden. We resurveyed 27 islands ranging from 0.3 to 33 ha in area. Methods We compared current plant survey data, cadastral maps and aerial photographs with records obtained from a survey in 1908, using databases and a digital elevation model to examine changes in plant community dynamics in space and time. We examined the effects of local and landscape structure and land use changes on plant species dynamics by using stepwise regression in relation to eight local and three landscape variables. The eight local variables were area, relative age, shape, soil heterogeneity, bedrock ratio, number of houses, forest cover change, and grazing 100 years ago. The three landscape variables were distance to mainland, distance to closest island with a farm 100 years ago, and structural connectivity. Hanski’s connectivity measure was modified to incorporate both connectivity and fragmentation. Results The investigated islands have undergone drastic changes, with increasing forest cover, habitation, and abandonment of grassland management. Although the total species richness increased by 31% and mean island area by 23%, we found no significant increase in species richness per unit area. Local variables explain past species richness (100 years ago), whereas both local and landscape variables explain current species richness, extinctions, immigrations and species turnover. Grazing that occurred 100 years ago still influences species richness, even though grazing management was abandoned several decades ago. The evidence clearly shows an increase in nitrophilous plant species, particularly among immigrant species. Main conclusions This study highlights the importance of including land use history when interpreting current patterns of species richness. Furthermore, local environment and landscape patterns affect important ecological processes such as immigration, extinction and species turnover, and hence should be included when assessing the impact of habitat fragmentation and land use change. We suggest that our modified structural connectivity measure can be applied to other types of landscapes to investigate the effects of fragmentation and habitat loss.     

Turnover of passerine birds on islands in the Aegean Sea (Greece)

Aim We wish to determine the effect of migratory status on turnover rates in island birds. Because turnover is influenced by factors other than migratory status, we also considered the influence of body size and physical characteristics of the islands inhabited on the probabilities of extinction and immigration. Location The Mediterranean islands of Delos, Astypalea, Paros, Naxos and Lesvos in the Aegean Sea, Greece. Methods The passerine birds of these islands were surveyed between 1954 and 1961 by G.E. Watson, and were resurveyed between 1988 and 1992. The effects of migratory status and body size on the probabilities of extinction and immigration were examined by G‐tests of linear trend in proportion, and analysis of variance, respectively. A combined analysis of migratory status, body size and physical characteristics of the islands was carried out using logistic regressions of the probabilities of extinction and immigration on these factors. Results Species number on each island changed little between surveys, with no island's species number changing by more than one species. Twelve population extinctions and 11 immigrations were recorded. The smallest island, Delos (6 km²), had the highest annualized relative turnover rate (1.08), while the four larger islands (96–1614 km²) had lower and mutually similar rates (0.21–0.27). Populations on higher elevation islands were less likely to go extinct. There is no evidence for an effect of body size on the probabilities of extinction or immigration. Migratory status affected extinction and immigration probabilities differently: migratory species were more likely to immigrate, but less likely to go extinct. Main conclusions The position of the Aegean islands along a major north–south flyway may account for the observed effects of migratory status. The annual passage of large numbers of migrants may, via the rescue effect, decrease the chances of extinction, while at the same time increasing the chances of colonization of unoccupied islands. The likelihood of both extinction and immigration involves a complex interaction between life‐history traits and island characteristics. The effects of migratory status will depend not only on consideration of vagility, vulnerability and stochasticity identified by previous authors, but also upon the location of the islands in relationship to migratory pathways.     

Island incidence and mainland population density: mammals from Mediterranean islands

Abstract. The reasons why some species are resistant to extinction or are better invaders of islands than others remain unexplained. In this study, we test the hypothesis that mammals living on the mainland at higher density than predicted by the density/body mass relationship have a much greater chance to colonize a small island successfully, and/or that they are less likely to become extinct when living on small islands. For this, we used data compiled on mammals from a number of Mediterranean islands. We show a nested pattern for mammals on western Mediterranean islands, which suggests that the distribution of mammals on these islands is not the result of a random process. Using two comparative methods, we show that mammal density on the mainland, corrected for body mass, is negatively correlated with island size. Mammals with a high density compared to the density/mass relationship are the best invaders and/or probably have less chance of going extinct on small islands when population size is small.     

Lost and found: Short-term dynamics of the flora on 100 small islands in the White Sea

The aim was to uncover factors that influence short-term (decade) flora dynamics and species richness of northern marine islets characterized by poor flora and weak anthropogenic pressure. The study used presence–absence data of vascular plant species on 100 small uprising islets of the Kandalaksha Gulf of White Sea (Northern Karelia, Russia). We investigated the influence of islands' attributes on species richness and rates of flora dynamics. Two island types were analyzed separately: younger, stone-like and older, islet-like (which generally are larger and have higher diversity of habitats). Sampled islands were studied via classical biogeographical per island approach and metapopulation per species approach. Stone-like islands had noticeably poorer flora with higher rates of immigration and extinction when compared to those of islet-like islands. The species number for islet-like islands correlated positively with number of habitats, abundance of different habitat types and island area. Species richness of stone-like islands correlated positively only with number of habitat types. Plant species associated with birds, crowberry thickets and coastal rocks were the most stable, and the species of disturbed habitats were significantly less stable. Floristic changes that have occurred have been caused by the massive establishment of new species rather than the extinction of pre-existing taxa. Thus, most of these islands are still in the colonization (assortative) stage. While we found no relationship between island area and species number for stone-like islands, this relationship was seen on islet-like islands.     

Factors influencing vascular plant diversity on 22 islands off the coast of eastern North America

Aim The influence of physiographic and historical factors on species richness of native and non‐native vascular plants on 22 coastal islands was examined. Location Islands off the coast of north‐eastern USA and south‐eastern Canada between 41° and 45° N latitude were studied. Island size ranges from 3 to 26,668 ha. All islands were deglaciated between 15,000 and 11,000 yr bp ; all but the four New Brunswick islands were attached to the mainland until rising sea level isolated them between 14,000 and 3800 yr bp . Methods Island species richness was determined from floras compiled or revised since 1969. Simple and multiple regression and rank correlation analysis were employed to assess the relative influence of independent variables on species richness. Potential predictors included island area, latitude, elevation, distance from the mainland, distance from the nearest larger island, number of soil types, years since isolation, years since deglaciation, and human population density. Results Native vascular plant species richness for the 22 islands in this study is influenced most strongly by island area, latitude, and distance from the nearest larger island; richness increases with island area, but decreases with latitude and distance from the nearest larger island as hypothesized. That a similar model employing distance from the mainland does not meet the critical value of P confirms the importance of the stepping‐stone effect. Habitat diversity as measured by number of soil types is also an important predictor of native plant species richness, but at least half of its influence can be attributed to island area, with which it is correlated. Two historical factors, years since deglaciation and years since isolation, also appear to be highly correlated with native species richness, but their influence cannot be separated from that of latitude for the present sample size. Non‐native vascular plant species richness is influenced primarily by island area and present‐day human population density, although human population density may be a surrogate for the cumulative effect of several centuries of anthropogenic impacts related to agriculture, hunting, fishing, whaling, tourism, and residential development. Very high densities of ground‐nesting pelagic birds may account for the high percentage of non‐native species on several small northern islands. Main conclusions Many of the principles of island biogeography that have been applied to oceanic islands apply equally to the 22 islands in this study. Native vascular plant species richness for these islands is strongly influenced by physiographic factors. Influence of two historical factors, years since deglaciation and years since isolation, cannot be assessed with the present sample size. Non‐native vascular plant species richness is influenced by island area as well as by human population density; human population density may be a surrogate for other anthropogenic impacts.     

Mammals of Australian islands: factors influencing species richness

Distribution patterns of indigenous non-volant terrestrial mammals on 257 Australian islands were examined in relation to environmental parameters and the effects of human-induced disturbance during prehistoric and historic times on island species numbers. Species occurrence for individual species, for taxonomic and trophic groups, and for all species together was related to environmental parameters using regression analysis and the extreme-value function model. Patterns of occurrence were examined separately within three major biogeographic regions derived by pattern analysis. The number of species known to have occurred on these islands during historic times was adequately predicted from area alone. No statistically significant improvement in predicted species number was gained by including island elevation, mean annual rainfall, isolation from the mainland or the number of potentially competing species present on the island. Similarly, no single factor other than area was found to influence consistently the presence of individual species. We conclude that the occurrence of indigenous non-volant terrestrial mammal species on these islands indicates a relictual rather than equilibrial fauna. Visitation by Aboriginal people during prehistoric times did not significantly increase mammal extinctions on islands. Examination of patterns of species richness for a given area on a regional basis showed that islands in and around Bass Strait and Tasmania (Bass Region) were the most species-rich, islands off the northern coasts were slightly less rich, and islands off the south western coasts had fewest species. This is in contrast to the usual latitudinal gradient in species richness patterns. However, islands off the northern and eastern coasts had an overall greater number of different species. When considered in relation to the number of different species of mammals occurring within each region, islands of a given size in Bass Region typically bore a higher proportion of this species pool than other regions. The Bass Region was found to be particularly rich in macropoid herbivores and dasyurid carnivores and insectivores. Analyses indicated that there is a very strong relationship between the presence of exotics as a whole and the local extinction of native mammals. Many mammal species formerly widespread on the Australian mainland are now restricted totally to islands (nine species) or are threatened with extinction on the mainland and have island populations of conservation significance (ten species). In all, thirty-five islands protect eighteen taxa of Australian threatened mammals. The land-use and management of these islands is of considerable importance to nature conservation. The introduction of exotic mammals to these islands should be prevented; any introductions that occur should be eradicated immediately.     

Determinants of floristic diversity and vegetation composition on the islands of Lake Burollos,Egypt

Abstract. A floristic and environmental survey was carried out on 22 uninhabited islands (0.1‐8.4 ha) in Lake Burollos, Egypt. A total of 58 vascular plant species was recorded. The number of habitats on each island was counted. There was a positive correlation between island area and number of habitat types. Island area was significantly positively correlated with various measures of floristic diversity, including the total number of species present, and the numbers of annual, herbaceous, and shrublet species. Perennial and shrub species numbers did not differ significantly with island area. In addition to island area, elevation and soil salinity, as well as distance to the Mediterranean Sea, all contributed significantly to variation in species composition in the terrestrial habitats. Water salinity and transparency accounted for 69% of the variation in aquatic species numbers. There was a weak effect of isolation on similarity of species composition on islands. Eight vegetation types, represented by 13 indicator species identified after TWINSPAN analysis, were distinguished by soil characteristics. Species richness was inversely correlated with clay, organic carbon and total nitrogen in the soil, but positively correlated with calcium carbonate content. From a management perspective, long‐term monitoring of threatened habitats in the lake is urgently required as a starting point to preserve biodiversity. Finally, we conclude that the present study supports the hypothesis indicating that larger areas feature higher species richness due to increased numbers of habitats.     

Diversity–invasibility relationships across multiple scales in disturbed forest understoreys

Non-native plant species richness may be either negatively or positively correlated with native species due to differences in resource availability, propagule pressure or the scale of vegetation sampling. We investigated the relationships between these factors and both native and non-native plant species at 12 mainland and island forested sites in southeastern Ontario, Canada. In general, the presence of non-native species was limited: <20% of all species at a site were non-native and non-native species cover was <4% m⁻² at 11 of the 12 sites. Non-native species were always positively correlated with native species, regardless of spatial scale and whether islands were sampled. Additionally, islands had a greater abundance of non-native species. Non-native species richness across mainland sites was significantly negatively correlated with mean shape index, a measure of the ratio of forest edge to area, and positively correlated with the mean distance to the nearest forest patch. Other factors associated with disturbance and propagule pressure in northeastern North America forests, including human land use, white-tailed deer populations, understorey light, and soil nitrogen, did not explain non-native richness nor cover better than the null models. Our results suggest that management strategies for controlling non-native plant invasions should aim to reduce the propagule pressure associated with human activities, and maximize the connectivity of forest habitats to benefit more poorly dispersed native species.     

Testing island biogeography theory with visitation rates of birds to British islands

Aim We consider three hypotheses – MacArthur and Wilson’s island biogeography theory (IBT), Lack’s habitat diversity idea and the ‘target effect’– that explain the pattern of decreased species richness on small and distant islands. Location We evaluate these hypotheses using a detailed dataset on the occurrence and abundance of terrestrial birds on nine islands off the coast of Britain and the Republic of Ireland. Methods  Unlike previous studies, we compile data on species that visit the islands, rather than just those that breed on them. We divided the species into five mutually exclusive categories based upon their migratory status and where they regularly breed: British residents, summer visitors to Britain, winter visitors to Britain, and vagrants from Europe or beyond Europe. For each species group on each island we calculated the average number of species visiting each year. We then regressed the average number of species against island area and distance to the mainland (all variables were log‐transformed). We also compared the average number of species visiting each island with the average number of species breeding on each island. Results  Average number of visiting British residents decreased significantly with increasing island distance, but showed no relationship with island area. There was no significant relationship between island area or island distance and average number of summer or winter visitors. European and non‐European vagrants likewise showed no relationship between numbers of species visiting and island distance. However, the relationship between island area and number of visiting species was significant for both these categories; as island area increases so too does the number of visiting species. Main conclusions  As predicted by IBT, there were fewer visiting species on more distant islands. There were substantially more visitors to each island than breeding species, supporting Lack’s argument that lower bird richness is not a result of varying immigration rates (as predicted by IBT) but rather a result of some other island property, e.g. fewer resources. Birds make a decision to either leave an island or stay and breed. The target effect was also clearly demonstrated by the increase in European and non‐European breeders with increasing island size.