Do forest plants conform to the theory of island biogeography: the case study of bog islands

Biodiversity conservation is confronted with increasing risk of extinction in isolated small-area remnants and the limitation of species to colonize recently formed habitats. We hypothesized that the equilibrium pattern of forest herb layer in long-term fragmented landscape should comply with the theory of island biogeography. Forests on mineral soil islands located in large mires of western Estonia were considered as dispersal target habitats, and forests on mainland and peninsulas in mires as sources. Species richness was the lowest in mainland forests and the effect was confounded by habitat structure, suggesting a negative effect of silvicultural management in easily accessible forests. We observed the ‘small island effect’, i.e. greater overall species richness in small-area habitats, which was determined by the habitat preference of shade tolerant generalists. The average species richness of common mainland forest specialists varied little, but capitalizing on the traditional approach and analyzing only island data, weak effects of distance and habitat quality were detected. At single species level, unexpectedly, many habitat specialists were observed to have successfully dispersed to islands, indicating insufficient knowledge of the long-distance dispersal mechanisms of forest-dwelling plants. In fragmented forest landscapes the theory of island biogeography can be applied to habitat specialist plant species, but only regarding the effect of isolation and in conditions of persistent forest structural quality. In the light of global changes, optimized conservation planning should primarily target on (i) the conservation of ancient habitat fragments independent of their current area, and (ii) the promotion of diversity of potential dispersal vectors in the landscape.     

Direct and indirect effects of area,energy and habitat heterogeneity on breeding bird communities

Aim To compare the ability of island biogeography theory, niche theory and species–energy theory to explain patterns of species richness and density for breeding bird communities across islands with contrasting characteristics. Location Thirty forested islands in two freshwater lakes in the boreal forest zone of northern Sweden (65°55′ N to 66°09′ N; 17°43′ E to 17°55′ E). Methods We performed bird censuses on 30 lake islands that have each previously been well characterized in terms of size, isolation, habitat heterogeneity (plant diversity and forest age), net primary productivity (NPP), and invertebrate prey abundance. To test the relative abilities of island biogeography theory, niche theory and species–energy theory to describe bird community patterns, we used both traditional statistical approaches (linear and multiple regressions) and structural equation modelling (SEM; in which both direct and indirect influences can be quantified). Results Using regression‐based approaches, area and bird abundance were the two most important predictors of bird species richness. However, when the data were analysed by SEM, area was not found to exert a direct effect on bird species richness. Instead, terrestrial prey abundance was the strongest predictor of bird abundance, and bird abundance in combination with NPP was the best predictor of bird species richness. Area was only of indirect importance through its positive effect on terrestrial prey abundance, but habitat heterogeneity and spatial subsidies (emerging aquatic insects) also showed important indirect influences. Thus, our results provided the strongest support for species–energy theory. Main conclusions Our results suggest that, by using statistical approaches that allow for analyses of both direct and indirect influences, a seemingly direct influence of area on species richness can be explained by greater energy availability on larger islands. As such, animal community patterns that seem to be in line with island biogeography theory may be primarily driven by energy availability. Our results also point to the need to consider several aspects of habitat quality (e.g. heterogeneity, NPP, prey availability and spatial subsidies) for successful management of breeding bird diversity at local spatial scales and in fragmented or insular habitats.     

Species-area relationships are controlled by species traits

The species-area relationship (SAR) is one of the most thoroughly investigated empirical relationships in ecology. Two theories have been proposed to explain SARs: classical island biogeography theory and niche theory. Classical island biogeography theory considers the processes of persistence, extinction, and colonization, whereas niche theory focuses on species requirements, such as habitat and resource use. Recent studies have called for the unification of these two theories to better explain the underlying mechanisms that generates SARs. In this context, species traits that can be related to each theory seem promising. Here we analyzed the SARs of butterfly and moth assemblages on islands differing in size and isolation. We tested whether species traits modify the SAR and the response to isolation. In addition to the expected overall effects on the area, traits related to each of the two theories increased the model fit, from 69% up to 90%. Steeper slopes have been shown to have a particularly higher sensitivity to area, which was indicated by species with restricted range (slope?=?0.82), narrow dietary niche (slope=?0.59), low abundance (slope=?0.52), and low reproductive potential (slope?=?0.51). We concluded that considering species traits by analyzing SARs yields considerable potential for unifying island biogeography theory and niche theory, and that the systematic and predictable effects observed when considering traits can help to guide conservation and management actions.     

Patterns of invertebrate density and taxonomic richness across gradients of area,isolation, and vegetation diversity in a lake‐island system

Over the past half century, ecologists have tried to unravel the factors that drive species richness patterns in ecological communities. One influential theory is island biogeography theory (IBT), which predicts that island or habitat area and isolation are drivers of species richness. However, relatively few studies testing IBT have considered invertebrate or belowground communities, and it is unclear as to whether the predictions made by IBT hold for these communities. Other theories predict that habitat characteristics such as vegetation diversity may be important drivers of invertebrate species richness. To investigate patterns of invertebrate density and species richness across gradients of area, isolation, and vegetation diversity, we used a system of 30 lake islands in the boreal zone of northern Sweden. We assessed density and taxonomic richness of ground‐dwelling spiders, web‐building spiders, beetles, collembolans, mites, and nematodes, for all islands during two consecutive summers. For all invertebrate groups, both density and taxonomic richness were either neutrally or negatively related to island size, and either neutrally or positively related to island isolation. Meanwhile the density and taxonomic richness for several groups was positively related to vegetation diversity (i.e. habitat heterogeneity). In multiple regression analyses, island size was often the single best predictor for both invertebrate density and taxonomic richness, but in some cases island size and isolation in combination explained more variation than each factor considered singly. Contrary to IBT predictions, invertebrate density and richness was never positively related to island size or negatively related to island isolation. Instead, our results suggest that plant diversity (and thus habitat heterogeneity) was the main driver of the patterns that we found, although other factors could have some influence. We conclude that several factors, but not necessarily those predicted as important by IBT, are important in determining invertebrate abundance and species richness in island systems.     

Spatial processes that maintain biodiversity in plant communities

The composition of communities of sessile organisms, and the change in species diversity with time, is a spatially explicit phenomenon. Three spatial factors clearly affect diversity: (1) the structure and heterogeneity of the landscape that limits species immigration and ultimate community size; (2) neighborhood interactions that determine colonization and extinction rates and influence residence times of local populations; and (3) disturbances that open spatially contiguous areas for recolonization by less abundant species. The importance of these three factors was first reviewed and then examined with a spatially explicit, multi-species model of plant dispersal, competition and establishment, with an assumption of neutrality (all species had equivalent life histories) that reduced the initial dimensionality of the problem. The simulations assumed that the probability of immigration was a linear function of mainland abundance and distance to islands, similar to the equilibrium theory of island biogeography and the unified neutral theory of biodiversity. The rate of increase in species richness was not constant across island sizes, declining as island area became very large. This pattern was explained by the spatial dynamics of colonization and establishment, a non-random process that cannot be explained by passive sampling alone. Simulations showed that population establishment depended critically on rare long-distance dispersal events while population persistence was achieved by the formation of aggregated species distributions that developed through restricted dispersal and local competitive interactions. Nevertheless, species richness always declined to a single species in the absence of disturbances, while up to 40 species could persist to 10,000 years when spatially dependent mortality was added. Further explorations with spatially explicit models will be required to fully appreciate the consequence of land use change and altered disturbance regimes on patterns of species distribution and the maintenance of diversity.     

island biogeography of Day Geckos (Phelsuma) in the Indian Ocean

Summary Step-wise multiple regression was employed to probe the determinants of species diversity of day geckos (Phelsuma) in the Indian Ocean. Independent variables were area, elevation, and two measures of isolation. Distance from Madagascar and island height (an indicator of habitat diversity) were the two most important predictors of species richness. Similar studies on other taxa rarely find isolation to be a major factor. The relatively poor dispersal abilities of reptiles may explain why isolation, rather than attributes of the islands, are more important in this case. The regressions also indicate that habitat diversity (assumed to correlate with maximum island elevation) is more important than area per se in determining species diversity. These results agree with predictions of the equilibrium theory of island biogeography, but historical processes have also greatly influenced species richness.     

Mountaintop island age determines species richness of boreal mammals in the American Southwest

Models that describe the mechanisms responsible for insular patterns of species richness include the equilibrium theory of island biogeography and the nonequilibrium vicariance model. The relative importance of dispersal or vicariance in structuring insular distribution patterns can be inferred from these models. Predictions of the alternative models were tested for boreal mammals in the American Southwest. Age of mountaintop islands of boreal habitat was determined by constructing a geographic cladogram based on characteristics of intervening valley barriers. Other independent variables included area and isolation of mountaintop islands. Island age was the most important predictor of species richness. In contrast with previous studies of species richness patterns in this system, these results supported the nonequilibrium vicariance model, which indicates that vicariance has been the primary determinant of species distribution patterns in this system.     

Environmental gradients and the structure of freshwater snail communities

A fundamental goal of ecology is to understand the factors that influence community structure and, consequently, generate heterogeneity in species richness across habitats. While niche‐assembly (e.g. species‐sorting) and dispersal‐assembly mechanisms are widely recognized as factors structuring communities, there remains substantial debate concerning the relative importance of each of these mechanisms. Using freshwater snails as a model system, we explore how abiotic and biotic factors interact with dispersal to structure local communities and generate regional patterns in species richness. Our data set consisted of 24 snail species from 43 ponds and lakes surveyed for seven years on the Univ. of Michigan's E. S. George Reserve and Pinckney State Recreation Area near Ann Arbor, Michigan. We found that heterogeneity in habitat conditions mediated species‐sorting mechanism to drive patterns in snail species richness across sites. In particular, physical environmental variables (i.e. habitat area, hydroperiod, and canopy cover), pH, and fish presence accounted for the majority of variation in the species richness across sites. We also found evidence of Gleasonian structure (i.e. significant species turnover with stochastic species loss) in the metacommunity. Turnover in snail species distributions was driven by the replacement of several pulmonate species with prosobranch species at the pond permanence transition. Turnover appeared to be driven by physiological constraints associated with differences in respiration mode between the snail orders and shell characteristics that deter molluscivorous fish. In contrast to these niche‐assembly mechanisms, there was no evidence that dispersal‐assembly mechanisms were structuring the communities. This suggests that niche‐assembly mechanisms are more important than dispersal‐assembly mechanisms for structuring local snail communities.     

A neglected facet of island biogeography: The role of internal spatial dynamics in area effects

The classical theory of island biogeography has as its basic variable the presence or absence of species on entire islands, and as its basic processes colonization and extinction rates on entire islands as functions of island area, distance, and so forth. Yet for many organisms with limited dispersal abilities, it may be more reasonable to consider larger islands as comprised of an ensemble of local populations coupled by within-island dispersal. Conceptual arguments and a simple patch occupancy model are used to examine the potential relevance of such internal spatial dynamics in explaining area effects, expressed via the probability that a species is present per unit area as a function of total island area. The model suggests that strong area effects depend on a rather fine balance between local colonization and extinction rates. A fruitful direction of future research should be the application of patch dynamic theory to classic island biogeographic questions and systems.     

Separating the effect of mechanisms shaping species‐abundance distributions at multiple scales in a subtropical forest

Species abundance distributions (SADs) play an important role in the current dispute over mechanisms shaping community assembly. Niche theory assumes differential occurrence of species in different habitats while neutral theory emphasizes stochastic events and dispersal. The previous tests of niche and neutral models shaping SADs lead to the claim that SADs are not informative for inferring underlying processes. Using spatial statistical models in a fully mapped 24‐ha subtropical forest in China, we first demonstrate that one can not distinguish between the effect of habitat heterogeneity and dispersal limitation on SADs by inspecting whether the observed SADs fall within 95% confidence intervals of the simulated SADs. Subsequently, we demonstrate that SADs can be used to detect mechanisms shaping SADS by comparing alternative process‐based models using model selection techniques. We found that dispersal limitation explain SADs at smaller spatial scales, while the combination of niche and dispersal limitation explain SADs at larger scales. These processes are linked with the degree of conspecific aggregation, informing further attempts to refine and parameterize the statistical theory of sampling SADs.     

Evidence for community assembly constraints during succession in dune slack plant communities

Community assembly during succession can be constrained by both local and regional factors. Despite an increasing regional species pool size during succession, we found a limit on the number of species in 1 × 1 m plots in dune slacks. Three alternative hypotheses (habitat heterogeneity, dispersal limitation and niche limitation) explaining this community saturation were tested. A null model analysis showed that species richness in the plots had an unusually low variance suggesting that beta habitat diversity was not likely to explain the limitation on species richness. Because we did not find a correlation between the distribution of species over the slack and their dispersal capacity, we also excluded the dispersal limitation hypothesis. Finally, a guild proportionality analysis revealed that the abundances of forb, graminoid and ruderal species showed low an unusually low variance over all age classes involved. This provides evidence for nonrandom community assembly during succession, likely to be determined by competitive exclusion among species of the same guild.     

Properties of similarity indices under niche-based and dispersal-based processes in communities

Little is known about the combined impact of habitat filtering and dispersal limitation on species turnover patterns. To gain new insights, we constructed a spatially explicit community model wherein we controlled dispersal distances, the strength of habitat filtering, and the grain of habitat heterogeneity to study the distance decay of several (dis)similarity indices. The impact of habitat filtering is dependent on the ratio between the grain of habitats and the mean dispersal distance. The behavior of (dis)similarity indices varies. First, incidence-based measures of species overlap are less affected by habitat filtering than are abundance-based indices. Second, species identity-based indices, derived from population genetics' F(ST), show interesting capacities to infer dispersal processes under neutrality but are also highly sensitive to habitat filtering. All indices except F(ST)-related indices under neutrality are very sensitive to overall species richness. Hence, community patterns showing contrasted diversity levels should be compared with caution. Partitioning similarity indices within and between habitats appears to be an efficient approach to assess the strength of habitat filtering, and we show that a torus-translation test is powerful for this purpose. We finally highlight the need for further analytical studies to achieve theoretical expectations of similarity decay under dispersal and niche processes.     

陆地生境岛屿藓类植物小岛屿效应驱动因素分析——以太行山脉中段山顶为例

小岛屿效应描述了种-面积关系的一种特殊现象,是当前生物地理学和生物多样性研究理论框架的重要组成部分。随着气候变暖,山顶物种的生存受到威胁,然而以山顶生境岛屿为载体对小岛屿效应的研究还十分缺乏。该研究以太行山脉中段19个面积0.06–801.58km²的山顶生境岛屿为研究区,在2019–2021年的夏秋季对藓类进行调查。共记录到藓类131种,隶属于23科68属。采用6种种-面积关系回归模型,分别检测了所有藓和6个常见藓科是否存在小岛屿效应。根据小岛屿效应形成机制的生境多样性假说、灭亡假说和营养补给假说,选择了岛屿高度、温度年变化范围和单位面积净初级生产力作为变量,对小岛屿效应的驱动因素进行分析。在各类群组中,使用多元线性回归和变差分解分别评估上述3个变量对物种丰富度变化的线性影响。首先使用5个面积最小的岛屿进行分析,计算出3个变量对物种丰富度变化的贡献,然后以迭代的方式逐次加入面积更大的1个岛屿,并再次进行变差分解分析。最后使用广义线性回归分析了3个变量对物种丰富度变化的贡献在迭代过程中的变化趋势。结果显示,所有藓和6个常见藓科均存在小岛屿效应,其面积阈值分布在0....     

Habitat diversity associated to island size and environmental filtering control the species richness of rock‐savanna plants in neotropical inselbergs

Disentangling the multiple factors controlling species diversity is a major challenge in ecology. Island biogeography and environmental filtering are two influential theories emphasizing respectively island size and isolation, and the abiotic environment, as key drivers of species richness. However, few attempts have been made to quantify their relative importance and investigate their mechanistic basis. Here, we applied structural equation modelling, a powerful method allowing test of complex hypotheses involving multiple and indirect effects, on an island‐like system of 22 French Guianan neotropical inselbergs covered with rock‐savanna. We separated the effects of size (rock‐savanna area), isolation (density of surrounding inselbergs), environmental filtering (rainfall, altitude) and dispersal filtering (forest‐matrix openness) on the species richness of all plants and of various ecological groups (terrestrial versus epiphytic, small‐scale versus large‐scale dispersal species). We showed that the species richness of all plants and terrestrial species was mainly explained by the size of rock‐savanna vegetation patches, with increasing richness associated with higher rock‐savanna area, while inselberg isolation and forest‐matrix openness had no measurable effect. This size effect was mediated by an increase in terrestrial‐habitat diversity, even after accounting for increased sampling effort. The richness of epiphytic species was mainly explained by environmental filtering, with a positive effect of rainfall and altitude, but also by a positive size effect mediated by enhanced woody‐plant species richness. Inselberg size and environmental filtering both explained the richness of small‐scale and large‐scale dispersal species, but these ecological groups responded in opposite directions to altitude and rainfall, that is positively for large‐scale and negatively for small‐scale dispersal species. Our study revealed both habitat diversity associated with island size and environmental filtering as major drivers of neotropical inselberg plant diversity and showed the importance of plant species growth form and dispersal ability to explain the relative importance of each driver.     

Increase of island endemism with altitude – speciation processes on oceanic islands

Understanding speciation on oceanic islands is a major topic in current research on island biogeography. Within this context, it is not an easy task to differentiate between the influence of elevation as an indicator for habitat diversity and island age as an indicator for the time available for diversification. One reason for this is that erosion processes reduce the elevation of islands over time. In addition, the geographic distance to source ecosystems might differ among habitats, which could lead to habitat‐specific reduction of species immigration, niche occupation and diversification. We used the percentage of single island endemic species (pSIE) in five different zonal ecosystems (distributed in altitude) on the Canary Islands as an indicator for diversification. We tested whether diversification increases with altitude due to a greater ecological isolation of high elevation ecosystems on oceanic islands under the assumption of a low elevation source region on the mainland. In addition we tested whether the ‘hump‐shaped’ (unimodal) relationship between pSIE and island age as well as the linear relationship between species richness and pSIE is consistent across spatial scales. We also analyse a potential influence of island area and habitat area. We found that pSIE increases with elevation. The relations between species richness as well as age with pSIE are consistent across scales. We conclude that high elevation ecosystems are ecologically isolated. Surprisingly, the altitudinal belt with the strongest human influences has the highest values of pSIE. We successfully transfer the ‘general dynamic theory of island biogeography’ to the ecosystem scale, which provides multiple opportunities for future studies. With this approach we find that the effects of elevation on diversification can be separated from those of island age.     

Environmental heterogeneity,species diversity and co‐existence at different spatial scales

The positive relationship between spatial environmental heterogeneity and species diversity is a widely accepted concept, generally associated with niche limitation. However, niche limitation cannot account for negative heterogeneity–diversity relationships (HDR) revealed in several case studies. Here we explore how HDR varies at different spatial scales and provide novel theories for small‐scale species co‐existence that explain both positive and negative HDR. At large spatial scales of heterogeneity (e.g. landscape level), different communities co‐exist, promoting large regional species pool size and resulting in positive HDR. At smaller scales within communities, species co‐existence can be enhanced by increasing the number of different patches, as predicted by the niche limitation theory, or alternatively, restrained by heterogeneity. We conducted meta‐regressions for experimental and observational HDR studies, and found that negative HDRs are significantly more common at smaller spatial scales. We propose three theories to account for niche limitation at small spatial scales. (1) Microfragmentation theory: with increasing spatial heterogeneity, large homogeneous patches lose area and become isolated, which in turn restrains the establishment of new plant individuals and populations, thus reducing species richness. (2) Heterogeneity confounded by mean: when heterogeneity occurs at spatial scales smaller than the size of individual plants, which forage through the patches, species diversity can be either positively or negatively affected by a change in the mean of an environmental factor. (3) Heterogeneity as a separate niche axis: the ability of species to tolerate heterogeneity at spatial scales smaller than plant size varies, affecting HDR. We conclude that processes other than niche limitation can affect the relationship between heterogeneity and diversity.     

Patch quality and context, but not patch number, drive multi-scale colonization dynamics in experimental aquatic landscapes

Colonization and extinction are primary drivers of local population dynamics, community structure, and spatial patterns of biological diversity. Existing paradigms of island biogeography, metapopulation biology, and metacommunity ecology, as well as habitat management and conservation biology based on those paradigms, emphasize patch size, number, and isolation as primary characteristics influencing colonization and extinction. Habitat selection theory suggests that patch quality could rival size, number, and isolation in determining rates of colonization and resulting community structure. We used naturally colonized experimental landscapes to address four issues: (a) how do colonizing aquatic beetles respond to variation in patch number, (b) how do they respond to variation in patch quality, (c) does patch context affect colonization dynamics, and (d) at what spatial scales do beetles respond to habitat variation? Increasing patch number had no effect on per patch colonization rates, while patch quality and context were critical in determining colonization rates and resulting patterns of abundance and species richness at multiple spatial scales. We graphically illustrate how variation in immigration rates driven by perceived predation risk (habitat quality) can further modify dynamics of the equilibrium theory of island biogeography beyond predator-driven effects on extinction rates. Our data support the importance of patch quality and context as primary determinants of colonization rate, occupancy, abundance, and resulting patterns of species richness, and reinforce the idea that management of metapopulations for species preservation, and metacommunities for local and regional diversity, should incorporate habitat quality into the predictive equation.     

HOW DOES POLLEN VERSUS SEED DISPERSAL AFFECT NICHE EVOLUTION?

In heterogeneous landscapes, the genetic and demographic consequences of dispersal influence the evolution of niche width. Unless pollen is limiting, pollen dispersal does not contribute directly to population growth. However, by disrupting local adaptation, it indirectly affects population dynamics. We compare the effect of pollen versus seed dispersal on the evolution of niche width in heterogeneous habitats, explicitly considering the feedback between maladaptation and demography. We consider two scenarios: the secondary contact of two subpopulations, in distinct, formerly isolated habitats, and the colonization of an empty habitat with dispersal between the new and ancestral habitat. With an analytical model, we identify critical levels of genetic variance leading to niche contraction (secondary contact scenario), or expansion (new habitat scenario). We confront these predictions with simulations where the genetic variance freely evolves. Niche contraction occurs when habitats are very different. It is faster as total gene flow increases or as pollen predominates in overall gene flow. Niche expansion occurs when habitat heterogeneity is not too high. Seed dispersal accelerates it, whereas pollen dispersal tends to retard it. In both scenarios very high seed dispersal leads to extinction. Overall, our results predict a wider niche for species dispersing seeds more than pollen.     

Effects of aerial dispersal, habitat specialisation, and landscape structure on spider distribution across fragmented grey dunes

Species distribution patterns have been explained by Hutchinson's niche theory, metapopulation theory and source-sink theory. Empirical verification of this framework, however, remains surprisingly scant. In this paper, we test the hypothesis that landscape characteristics (patch size and connectivity), aerial dispersal ability and niche breadth interact in explaining distribution patterns of 29 spider species inhabiting fragmented grey dunes. Distribution patterns only depended on aerial dispersal potential, and the interaction between patch connectivity and area. Niche breadth, measured as the degree of habitat specialisation in the total coastal dune system, did not contribute to the observed distribution patterns. Additional variation in patch occupancy frequency was strongly species-dependent and was determined by different responses to the degree of patch connectivity for ballooning dispersal. Results from this study suggest that dispersal ability largely affects our perception of a species "fundamental niche", and that source-sink and metapopulation dynamics may have a major impact on the distribution of species. From a conservation point of view, specialised (and hence intrinsically rare) species can be predicted to become rarer if fragmentation increases and connectivity decreases. This study is, to our knowledge, one of the few linking species distribution (and not patch occupancy, species diversity or richness) to landscape ecological (patch connectivity and area) and auto-ecological (niche breadth, dispersal potential) features.     

Biogeographic and anthropogenic correlates of Aleutian Islands plant diversity: A machine‐learning approach

This is the first comprehensive analysis of vascular plant diversity patterns in the Aleutian Islands to identify and quantify the impact of Aleutian Island distance dispersal barriers, geographical, ecological and anthropogenic factors. Data from public Open Access databases, printed floristic accounts, and from collections made by the primary author were used to develop an Aleutian floristic database. The most common plant distribution pattern was “an eastern origin community”, though it compared similarly to the “Western” and “Widespread” distribution pattern. We established an ecological plant community composition class for each island, based on clustering species assemblage dissimilarity measurements (Jaccard Index), and a measurement of phylogenetic dissimilarity (UniFrac). We modelled these composition classes and species richness values in non‐parametric algorithmic models and concepts (data cloning using machine learning, stochastic boosting‐ TreeNet) based on classic and Aleutians‐specific island biogeography hypotheses. Plant species richness is strongly associated with the equilibrium model variables of area and island isolation, as well as distance to the Alaska Peninsula, and island total stream length. Species composition is strongly associated with the landmass groups during the last glacial maximum, maximum island elevation, island isolation and island area. Phylogenetic composition is associated with island area, distance from the islands to the Chukotka Peninsula, maximum island elevation, island geologic age, and island isolation. This study extends the equilibrium theory of island biogeography by including additional drivers of diversity during the Anthropocene, such as the landmass during the LGM, as well as factors that may be related to anthropogenic extinction rate.