A global model of island biogeography

Aim The goal of our study was to build a global model of island biogeography explaining bird species richness that combines MacArthur and Wilson's area–isolation theory with the species–energy theory. Location Global. Methods We assembled a global data set of 346 marine islands representing all types of climate, topography and degree of isolation on our planet, ranging in size from 10 ha to 800,000 km². We built a multiple regression model with the number of non‐marine breeding bird species as the dependent variable. Results We found that about 85–90% of the global variance in insular bird species richness can be explained by simple, contemporary abiotic factors. On a global scale, the three major predictors — area, average annual temperature and the distance separating the islands from the nearest continent — all have constraining (i.e. triangular rather than linear) relationships with insular bird species richness. We found that the slope of the species–area curve depends on both average annual temperature and total annual precipitation, but not on isolation. Insular isolation depends not only on the distance of an island from the continent, but also on the presence or absence of other neighbouring islands. Range in elevation — a surrogate for diversity of habitats — showed a positive correlation with bird diversity in warmer regions of the world, while its effect was negative in colder regions. We also propose a global statistical model to quantify the isolation‐reducing effect of neighbouring islands. Main conclusions The variation in avian richness among islands worldwide can be statistically explained by contemporary environmental variables. The equilibrium theory of island biogeography of MacArthur and Wilson and the species–energy theory are both only partly correct. Global variation in richness depends about equally upon area, climate (temperature and precipitation) and isolation. The slope of the species richness–area curve depends upon climate, but not on isolation, in contrast to MacArthur and Wilson's theory.     

A null model of exotic plant diversity tested with exotic and native species-area relationships

At large spatial scales, exotic and native plant diversity exhibit a strong positive relationship. This may occur because exotic and native species respond similarly to processes that influence diversity over large geographical areas. To test this hypothesis, we compared exotic and native species–area relationships within six North American ecoregions. We predicted and found that within ecoregions the ratio of exotic to native species richness remains constant with increasing area. Furthermore, we predicted that areas with more native species than predicted by the species–area relationship would have proportionally more exotics as well. We did find that these exotic and native deviations were highly correlated, but areas that were good (or bad) for native plants were even better (or worse) for exotics. Similar processes appear to influence exotic and native plant diversity but the degree of this influence may differ with site quality.     

Causes of exotic bird establishment across oceanic islands

The probability that exotic species will successfully establish viable populations varies between regions, for reasons that are currently unknown. Here, we use data for exotic bird introductions to 41 oceanic islands and archipelagos around the globe to test five hypotheses for this variation: the effects of introduction effort, competition, predation, human disturbance and habitat diversity (island biogeography). Our analyses demonstrate the primary importance of introduction effort for avian establishment success across regions, in concordance with previous analyses within regions. However, they also reveal a strong negative interaction across regions between establishment success and predation; exotic birds are more likely to fail on islands with species-rich mammalian predator assemblages.     

Exotic species richness and native species endemism increase the impact of exotic species on islands

Aim Exotic species pose one of the most significant threats to biodiversity, especially on islands. The impacts of exotic species vary in severity among islands, yet little is known about what makes some islands more susceptible than others. Here we determine which characteristics of an island influence how severely exotic species affect its native biota. Location We studied 65 islands and archipelagos from around the world, ranging from latitude 65° N to 54° S. Methods We compiled a global database of 10 island characteristics for 65 islands and determined the relative importance of each characteristic in predicting the impact of exotic species using multivariate modelling and hierarchical partitioning. We defined the impact of exotic species as the number of bird, amphibian and mammal (BAM) species listed by the International Union for Conservation of Nature (IUCN) as threatened by exotics, relative to the total number of BAM species on that island. Results We found that the impact of exotic species is more severe on islands with more exotic species and a greater proportion of native species that are endemic. Unexpectedly, the level of anthropogenic disturbance did not influence an island's susceptibility to the impacts of exotic species. Main conclusions By coupling our results with studies on the introduction and establishment of exotic species, we conclude that colonization pressure, or invasion opportunities, influences all stages of the invasion process. However, species endemism, the other important factor determining the impact of exotic species, is not known to contribute to introduction and establishment success on islands. This demonstrates that different factors correlate with the initial stages of the invasion process and the subsequent impacts of those invaders, highlighting the importance of studying the impacts of exotic species directly. Our study helps identify islands that are at risk of impact by exotics and where investment should focus on preventing further invasions.     

Invasive species and land bird diversity on remote South Atlantic islands

Invasive species pose significant threats to biodiversity, especially on islands. They cause extinctions and population declines, yet little is known about their consequences on the emergent, metacommunity-level patterns of native species in island assemblages. We investigated differences in species–area relationships, nestedness, and occupancy of 9 species of native land birds between island assemblages with and without invasive Norway rats (Rattus norvegicus) in the Falkland Archipelago. We found that species–area curves, nestedness, and individual species’ occurrences differed between island assemblages with and without rats. Rat-free islands had, on average, 2.1 more land bird species than rat-infested islands of similar size. Passerine bird communities on islands with and without rats were significantly nested, but nestedness was significantly higher on rat-free islands than on rat-infested islands. The presence of rats was associated with differences in the incidence of many, but not all bird species. On rat free islands the occurrence of all species increased with island area. The occurrence of most, albeit not all, bird species was lower on islands with than on islands without rats. Two species of conservation concern, Troglodytes aedon cobbi and Cinclodes antarcticus, were abundant on rat-free islands, but absent or found at very low frequencies on islands with rats. The occurrence of three species was not associated with the presence of rats. The patterns presented here can be used to evaluate the consequences of ongoing rat eradications for passerine diversity, distribution, and abundance.     

Remarkable new evidence for island radiation in birds

If island biogeography has a sweet spot, it's where islands generate their own species diversity rather than merely taking on mainland immigrants. In birds and other highly dispersive taxa, however, this 'zone of radiation', may be vanishingly small. Darwin's finches and Hawaiian Honeycreepers are among only a handful of examples of island radiation in birds (Price 2008), suggesting that winged powers of dispersal make sufficient isolation from mainland colonists unlikely, while also hindering speciation within and among isolated islands. Nevertheless, two studies in this issue of Molecular Ecology join a string of other recent analyses suggesting that island radiation in birds remains under-appreciated (see also Moyle et al. 2009; Kisel & Barraclough 2010; Rosindell & Phillimore 2011). Melo et al. (2011) use a phylogenetic analysis of white-eyes on islands in the Gulf of Guinea to identify two previously overlooked island radiations, and reveal replicated adaptive divergence on islands where species occur in pairs. Sly et al. (2011), meanwhile, consider possible explanations for speciation and geographic differentiation within a large island, and find the same type of oceanic barriers that are critical to bird speciation across archipelagos may also contribute to divergence that appears to have occurred within a single island.     

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.     

Human activities alter biogeographical patterns of reptiles on Mediterranean islands

Aim The theory of island biogeography predicts species richness based on geographical factors that influence the extinction–colonization balance, such as area and isolation. However, human influence is the major cause of present biotic changes, and may therefore modify biogeographical patterns by increasing extinctions and colonizations. Our aim was to evaluate the effect of human activities on the species richness of reptiles on islands. Location Islands in the Mediterranean Sea and Macaronesia. Methods Using a large data set (n = 212 islands) compiled from the literature, we built spatial regression models to compare the effect of geographical (area, isolation, topography) and human (population, airports) factors on native and alien species. We also used piecewise regression to evaluate whether human activities cause deviation of the species–area relationship from the linear (on log–log axes) pattern, and path analysis to reveal the relationships among multiple potential predictors. Results The richness of both native and alien species was best explained by models combining geographical and human factors. The richness of native species was negatively related to human influence, while that of alien species was positively related, with the overall balance being negative. In models that did not take into account human factors, the relationship between island area and species richness was not linear. Large islands hosted fewer native species than expected from a linear (on log–log axes) species–area relationship, because they were more strongly affected by human influence than were small islands. Path analysis showed that island size has a direct positive effect on reptile richness. However, area also had a positive relationship with human impact, which in turn mediated a negative effect on richness. Main conclusion Anthropogenic factors can strongly modify the biogeographical pattern of islands, probably because they are major drivers of present‐day extinctions and colonizations and can displace island biodiversity from the equilibrium points expected by theory on the basis of geographical features.     

'Natural experiment' Demonstrates Top-Down Control of Spiders by Birds on a Landscape Level

The combination of small-scale manipulative experiments and large-scale natural experiments provides a powerful approach for demonstrating the importance of top-down trophic control on the ecosystem scale. The most compelling natural experiments have come from studies examining the landscape-scale loss of apex predators like sea otters, wolves, fish and land crabs. Birds are dominant apex predators in terrestrial systems around the world, yet all studies on their role as predators have come from small-scale experiments; the top-down impact of bird loss on their arthropod prey has yet to be examined at a landscape scale. Here, we use a unique natural experiment, the extirpation of insectivorous birds from nearly all forests on the island of Guam by the invasive brown tree snake, to produce the first assessment of the impacts of bird loss on their prey. We focused on spiders because experimental studies showed a consistent top-down effect of birds on spiders. We conducted spider web surveys in native forest on Guam and three nearby islands with healthy bird populations. Spider web densities on the island of Guam were 40 times greater than densities on islands with birds during the wet season, and 2.3 times greater during the dry season. These results confirm the general trend from manipulative experiments conducted in other systems however, the effect size was much greater in this natural experiment than in most manipulative experiments. In addition, bird loss appears to have removed the seasonality of spider webs and led to larger webs in at least one spider species in the forests of Guam than on nearby islands with birds. We discuss several possible mechanisms for the observed changes. Overall, our results suggest that effect sizes from smaller-scale experimental studies may significantly underestimate the impact of bird loss on spider density as demonstrated by this large-scale natural experiment.     

Richness and composition of plants and birds on land‐bridge islands: effects of island attributes and differential responses of species groups

Aim To test relationships between the richness and composition of vascular plants and birds and attributes of habitat fragments using a model land‐bridge island system, and to investigate whether the effects of fragmentation differ depending on species natural history traits. Location Thousand Island Lake, China. Methods We compiled presence/absence data of vascular plant and bird species through exhaustive surveys of 41 islands. Plant species were assigned to two categories: shade‐intolerant and shade‐tolerant species; bird species were assigned to three categories: edge, interior, and generalist species. We analysed the relationships between island attributes (area, isolation, elevation, shape complexity, and perimeter to area ratio) and species richness using generalized linear models (GLMs). We also investigated patterns of composition in relation to island attributes using ordination (redundancy analysis). Results We found that island area explained a high degree of variation in the species richness of all species groups. The slope of the species–area relationship (z) was 0.16 for all plant species and 0.11 for all bird species. The lowest z‐value was for generalist birds (0.04). The species richness of the three plant species groups was associated with island area per se, while that of all, generalist, and interior birds was explained mainly by elevation, and that of edge bird species was associated primarily with island shape. Patterns of species composition were most strongly related to elevation, island shape complexity, and perimeter to area ratio rather than to island area per se. Species richness had no significant relationship with isolation, but species composition did. We also found differential responses among the species groups to changes in island attributes. Main conclusions Within the Thousand Island Lake system, the effects of fragmentation on both bird and plant species appear to be scale‐dependent and taxon‐specific. The number of plant species occurring on an island is strongly correlated with island area, and the richness of birds and the species composition of plants and birds are associated with variables related to habitat heterogeneity. We conclude that the effects of fragmentation on species diversity and composition depend not only on the degree of habitat loss but also on the specific patterns of habitat fragmentation.     

The value of georeferenced collection records for predicting patterns of mosquito species richness and endemism in the Neotropics

Abstract.  1. Determining large-scale distribution patterns for mosquitoes could advance knowledge of global mosquito biogeography and inform decisions about where mosquito inventory needs are greatest.
2. Over 43 000 georeferenced records are presented of identified and vouchered mosquitoes from collections undertaken between 1899 and 1982, from 1853 locations in 42 countries throughout the Neotropics. Of 492 species in the data set, 23% were only recorded from one location, and Anopheles albimanus Wiedemann is the most common species.
3. A linear log–log species–area relationship was found for mosquito species number and country area. Chile had the lowest relative density of species and Trinidad-Tobago the highest, followed by Panama and French Guiana.
4. The potential distribution of species was predicted using an Ecological Niche Modelling (ENM) approach. Anopheles species had the largest predicted species ranges, whereas species of Deinocerites and Wyeomyia had the smallest.
5. Species richness was estimated for 1° grids and by summing predicted presence of species from ENM. These methods both showed areas of high species richness in French Guiana, Panama, Trinidad-Tobago, and Colombia. Potential hotspots in endemicity included unsampled areas in Panama, French Guiana, Colombia, Belize, Venezuela, and Brazil.
6. Argentina, The Bahamas, Bermuda, Bolivia, Cuba, and Peru were the most under-represented countries in the database compared with known country species occurrence data. Analysis of species accumulation curves suggested patchiness in the distribution of data points, which may affect estimates of species richness.
7. The data set is a first step towards the development of a global-scale repository of georeferenced mosquito collection records.     

Rapid evolution of great kiskadees on Bermuda: an assessment of the ability of the island rule to predict the direction of contemporary evolution in exotic vertebrates

Aim To determine whether an exotic bird species, the great kiskadee (Pitangus sulphuratus), has diverged in morphology from its native source population, and, if so, has done so in a manner predicted by the island rule. The island rule predicts that insular vertebrates will tend towards dwarfism or gigantism when isolated on islands, depending on their body size. For birds, the island rule predicts that species with body sizes below 70–120 g should increase in size. The great kiskadee has a mean mass of c. 60 g in its native range, therefore we predicted that it would increase in size within the exotic, and more insular, Bermudan range. Location The islands of Bermuda (exotic population) and Trinidad (native source population). Methods We took eight morphological measurements on 84 individuals captured in the exotic (Bermudan) population and 62 individuals captured in the native source (Trinidadian) population. We compared morphological metrics between populations using univariate and principal components analyses. We assessed whether the effects of genetic drift could explain observed differences in morphology. We calculated divergence rates in haldanes and darwins for comparison with published examples of contemporary evolution. Finally, we used mark–recapture analysis to determine the effects of the measured morphological characters on survivorship within the exotic Bermudan population. Results Individuals in the exotic Bermudan population have larger morphological dimensions than individuals in the native source population on Trinidad. The degree of divergence in body mass (g) and bill width (mm) is probably not due to genetic drift. This rate of divergence is nearly equal to that observed amongst well‐documented examples of contemporary bird evolution, and is within the mid‐range of rates reported across taxa. There is no clear effect of body size on survivorship as only one character (bill width) was found to have an influence on individual survivorship. Main conclusions Exotic species provide useful systems for examining evolutionary predictions over contemporary time‐scales. We found that divergence between the exotic and native populations of this bird species occurred over c. 17 generations, and was in the direction predicted by the island rule, a principle based on the study of native species.     

Island biogeography of bats in Baja California, Mexico: patterns of bat species richness in a near-shore archipelago

Aim We studied the relationship between the size and isolation of islands and bat species richness in a near‐shore archipelago to determine whether communities of vagile mammals conform to predictions of island biogeography theory. We compared patterns of species richness in two subarchipelagos to determine whether area per se or differences in habitat diversity explain variations in bat species richness. Location Islands in the Gulf of California and adjacent coastal habitats on the Baja California peninsula in northwest Mexico. Methods Presence–absence surveys for bats were conducted on 32 islands in the Gulf of California using acoustic and mist‐net surveys. We sampled for bats in coastal habitats of four regions of the Baja peninsula to characterize the source pool of potential colonizing species. We fitted a semi‐log model of species richness and multiple linear regression and used Akaike information criterion model selection to assess the possible influence of log₁₀ area, isolation, and island group (two subarchipelagos) on the species richness of bats. We compared the species richness of bats on islands with greater vegetation densities in the southern gulf (n = 20) with that on drier islands with less vegetation in the northern gulf (n = 12) to investigate the relationship between habitat diversity and the species richness of bats. Results Twelve species of bats were detected on islands in the Gulf of California, and 15 species were detected in coastal habitats on the Baja peninsula. Bat species richness was related to both area and isolation of islands, and was higher in the southern subarchipelago, which has denser vegetation. Log₁₀ area was positively related to bat species richness, which increased by one species for every 5.4‐fold increase in island area. On average, richness declined by one species per 6.25 km increase in isolation from the Baja peninsula. Main conclusions Our results demonstrate that patterns of bat species richness in a near‐shore archipelago are consistent with patterns predicted by the equilibrium theory of island biogeography. Despite their vagility, bats may be more sensitive to moderate levels of isolation than previously expected in near‐shore archipelagos. Differences in vegetation and habitat xericity appear to be associated with richness of bat communities in this desert ecosystem. Although observed patterns of species richness were consistent with those predicted by the equilibrium theory, similar relationships between species richness and size and isolation of islands may arise from patch‐use decision making by individuals (optimal foraging strategies).     

A global review of island endemic birds

Although fewer than one-fifth of the world's bird species are restricted to islands, over 90% of bird extinctions during historic times have occurred on islands. The major identified cause has been the effects of exotic animal species introduced by man; the largest number of documented extinctions has occurred on islands of the Pacific Ocean.
Some 39% (402) of threatened bird species are restricted to islands and more than 907; of these are endemic to a single geopolitical unit. The largest numbers occurring in such units are in Indonesia (91) and the Philippines (34). As a region, the Pacific holds more threatened species (110) than any other, including almost half of those considered Endangered and over 40% of the Vulnerable species.
Most threatened island species are forest-dwelling. A high proportion of the Endangered species use seasonal/temperate forest. While habitat destruction now poses the greatest overall threat to island birds (affecting over half the species restricted to islands), the presence of introduced species threatens 30 of the 66 Endangered species.
Although immediate extinctions of island species can best be averted by mitigating the effects of introductions, the removal of native forests will be a more severe problem in the longer term. There is an urgent need for ecologists to provide detailed information on the habitats of both threatened and endemic species so that more appropriate and effective conservation programmes can be developed.     

A strong species-area relationship for eukaryotic soil microbes: island size matters for ectomycorrhizal fungi

While the effects of habitat size and isolation have been successfully studied for macro-organisms, there is currently debate about their relative importance in explaining patterns of microbial species richness. In this study, we examine the species richness of a dominant group of eukaryotic soil microbes, ectomycorrhizal fungi, on 'tree islands' of constant age and host composition that range in size from < 10 to > 10 000 m². Our results show that ectomycorrhizal species richness is significantly reduced on smaller and more isolated tree islands, and the species–area slope that we observe (0.20–0.23) is similar to average slopes reported for macro-organisms. Additionally, species' occurrence patterns across tree islands and investment trends in fungal fruit bodies suggest that a trade-off between competition and dispersal could play an important role in structuring ectomycorrhizal assemblages.     

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 bird species richness,endemism, and island network roles in Wallacea and the West Indies: is geography sufficient or does current and historical climate matter?

Island biogeography has greatly contributed to our understanding of the processes determining species' distributions. Previous research has focused on the effects of island geography (i.e., island area, elevation, and isolation) and current climate as drivers of island species richness and endemism. Here, we evaluate the potential additional effects of historical climate on breeding land bird richness and endemism in Wallacea and the West Indies. Furthermore, on the basis of species distributions, we identify island biogeographical network roles and examine their association with geography, current and historical climate, and bird richness/endemism. We found that island geography, especially island area but also isolation and elevation, largely explained the variation in island species richness and endemism. Current and historical climate only added marginally to our understanding of the distribution of species on islands, and this was idiosyncratic to each archipelago. In the West Indies, endemic richness was slightly reduced on islands with historically unstable climates; weak support for the opposite was found in Wallacea. In both archipelagos, large islands with many endemics and situated far from other large islands had high importance for the linkage within modules, indicating that these islands potentially act as speciation pumps and source islands for surrounding smaller islands within the module and, thus, define the biogeographical modules. Large islands situated far from the mainland and/or with a high number of nonendemics acted as links between modules. Additionally, in Wallacea, but not in the West Indies, climatically unstable islands tended to interlink biogeographical modules. The weak and idiosyncratic effect of historical climate on island richness, endemism, and network roles indicates that historical climate had little effects on extinction‐immigration dynamics. This is in contrast to the strong effect of historical climate observed on the mainland, possibly because surrounding oceans buffer against strong climate oscillations and because geography is a strong determinant of island richness, endemism and network roles.     

Differential effects of exotic predator-control on nest success of native and introduced birds in New Zealand

The introduction of mammalian predators has been detrimental to many native birds in New Zealand. One solution to this problem has been the creation of “mainland islands” in which exotic predators are systematically removed. Although mainland islands have been effective in increasing some native bird populations, few studies have measured the effect of predator-control on nest success nor what effect control measures have on sympatric populations of introduced birds. We measured the effect of predator-control on nest survival rates in both native and introduced passerines in a mainland island near Kaikoura, New Zealand. Nest survival was significantly higher in Waimangarara Bush (the site with experimental predator-control) than in Kowhai Bush (the site with no predator-control) and this pattern was found in both groups of birds. However, mammalian predator-control increased nest success of native species significantly more than nest success of introduced species. This suggests that native birds benefit disproportionately from control of introduced predators, most likely because they lack behavioural defences against mammalian predators that are present among the introduced birds.     

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.