Integrating the effects of area, isolation, and habitat heterogeneity on species diversity: a unification of island biogeography and niche theory

We present an analytical model that unifies two of the most influential theories in community ecology, namely, island biogeography and niche theory. Our model captures the main elements of both theories by incorporating the combined effects of area, isolation, stochastic colonization and extinction processes, habitat heterogeneity, and niche partitioning in a unified, demographically based framework. While classical niche theory predicts a positive relationship between species richness and habitat heterogeneity, our unified model demonstrates that area limitation and dispersal limitation (the main elements of island biogeography) may create unimodal and even negative relationships between species richness and habitat heterogeneity. We attribute this finding to the fact that increasing heterogeneity increases the potential number of species that may exist in a given area (as predicted by niche theory) but simultaneously reduces the amount of suitable area available for each species and, thus, increases the likelihood of stochastic extinction. Area limitation, dispersal limitation, and low reproduction rates intensify the latter effect by increasing the likelihood of stochastic extinction. These analytical results demonstrate that the integration of island biogeography and niche theory provides new insights about the mechanisms that regulate the diversity of ecological communities and generates unexpected predictions that could not be attained from any single theory.     

Insular carnivore biogeography: island area and mammalian optimal body size

Patterns of size variation in insular mammals have been used to support the claim that mammals have a single optimal body size. This hypothesis enjoys wide support, despite having been questioned on both theoretical and empirical grounds. It is claimed that species of optimal size maintain the highest population densities. Therefore these species are thought to inhabit the smallest islands, where larger and smaller species are generally absent. We sought such a pattern by testing how area affects the body sizes of the largest and smallest carnivore species on islands. Using data on carnivores from 322 islands, we found that the sizes of carnivores on small islands tend to be close to the order's mode. Furthermore, we found that the size distribution of carnivore species that inhabit islands resembles that of those whose range is entirely continental. We conclude that insular carnivores provide no support for theories proposing a single optimal size, and we suspect such theories are also flawed on theoretical grounds.     

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.     

Small island biogeography in the Gulf of California: lizards, the subsidized island biogeography hypothesis, and the small island effect

Aim We used insular lizard communities to test the predictions of two hypotheses that attempt to explain patterns of species richness on small islands. We first address the subsidized island biogeography (SIB) hypothesis, which predicts that spatial subsidies may cause insular species richness to deviate from species–area predictions, especially on small islands. Next, we examine the small island effect (SIE), which suggests small islands may not fit the traditional log‐linear species–area curve. Location Islands with arthropodivorous lizard communities throughout the Gulf of California. Methods To evaluate the SIB hypothesis, we first identified subsidized and unsubsidized islands based on surrogate measures of allochthonous productivity (i.e. island size and bird presence). Subsequently, we created species–area curves from previously published lizard species richness and island area data. We used the residuals and slopes from these analyses to compare species richness on subsidized and unsubsidized islands. To test for an SIE, we used breakpoint regression to model the relationship between lizard species richness and island area. We compared results from this model to results from the log‐linear regression model. Results Subsidized islands had a lower slope than unsubsidized islands, and the difference between these groups was significant when small islands were defined as < 1 km². In addition to comparing slopes, we tested for differences in the magnitude of the residuals (from the species–area regression of all islands) for subsidized vs. unsubsidized islands. We found no significant patterns in the residual values for small vs. large islands, or between islands with and without seabirds. The SIE was found to be a slightly better predictor of lizard species richness than the traditional log‐linear model. Main conclusions Predictions of the SIB hypothesis were partially supported by the data. The absence of a significant SIE may be a result of spatial subsidies as explained by the SIB hypothesis and data presented here. We conclude by suggesting potential scenarios to test for interactions between these two small island hypotheses. Future studies considering factors affecting species richness should examine the possible role of spatial subsidies, an SIE, or a synergistic effect of the two in data sets with small islands.     

Fungi,leaves, and the theory of island biogeography

Species dynamics of fungi (filamentous fungi and yeasts) on apple leaves were studied within the framework of the theory of island biogeography by following immigration and extinction patterns on individual apple leaf islands over time. Total fungi were censused on unmanipulated leaves collected throughout two seasons; filamentous fungi only were monitored additionally for several weeks in one season on newly created, axenic, model (seedling) islands introduced to the orchard, and on surface-sterilized, preexisting leaves. Analyses based on both the natural and the surface-sterilized systems showed that an equilibrium in species number was reached and turnover in species composition occurred in both. Immigration and extinction events were strongly related to number of species present on each island. The balance between immigration and extinction implies that species number on leaves and real (oceanic) islands is determined by a common mechanism, and emphasizes the need to regard leaf microbial communities as dynamic.     

Trophic theory of island biogeography

MacArthur and Wilson's Theory of Island Biogeography (TIB) is among the most well-known process-based explanations for the distribution of species richness. It helps understand the species-area relationship, a fundamental pattern in ecology and an essential tool for conservation. The classic TIB does not, however, account for the complex structure of ecological systems. We extend the TIB to take into account trophic interactions and derive a species-specific model for occurrence probability. We find that the properties of the regional food web influence the species-area relationship, and that, in return, immigration and extinction dynamics affect local food web properties. We compare the accuracy of the classic TIB to our trophic TIB to predict community composition of real food webs and find strong support for our trophic extension of the TIB. Our approach provides a parsimonious explanation to species distributions and open new perspectives to integrate the complexity of ecological interactions into simple species distribution models.     

The island biogeography of languages

Aim To examine the degree to which area, isolation, environmental conditions and time since first settlement explain variation in language richness among islands. Location Pacific islands ranging east–west from Rapa Nui to Indonesia and north–south from Hawaii to New Zealand. Methods We constructed a dataset of 264 Pacific islands that support 1640 languages (c. 24% of the world's languages). We examined possible predictors of language richness using three different types of models: linear regression models, linear mixed models that included random effects for language phylogeny and simultaneous autoregressive models. We tested whether the following variables, alone or in combination, predict language richness: island area and isolation, climate (rainfall, temperature), mean growing season, soil fertility, habitat heterogeneity (elevation, number of ecoregions), time since first human settlement. Results We identified two optimal models (delta Akaike information criterion < 2). One (R²= 0.52) included area, with 86% of remaining variation accounted for by random effects for phylogeny. The other (R²= 0.56) included a spatial component, area and a suite of other variables (of which isolation and settlement scale were significant). Of the hypotheses tested (mean growing season, ecological risk, habitat heterogeneity, climate, time since settlement, area–isolation theory), area–isolation performed best, alone explaining 44% of variation in language richness. Main conclusions Language diversity relates strongly to island area, and, after controlling for area, with variables linked to isolation (e.g. distance to continent, time since settlement). The influence of environmental productivity may be scale and context dependent. Although environmental productivity may shape language diversity patterns at a global scale, it plays little role on Pacific islands. Approximately half the variance in language richness remains unexplained. Unlike other taxa, for which area, isolation and environmental conditions explain up to 90% of variation in richness, human diversity patterns appear to also be influenced by other variables (e.g. economic, political and social factors).     

A generalized model of island biogeography

MacArthur and Wilson’s equilibrium theory is one of the most influential theories in ecology. Although evolution on islands is to be important to island biodiversity, speciation has not been well integrated into island biogeography models. By incorporating speciation and factors influencing it into the MacArthur-Wilson model, we propose a generalized model unifying ecological and evolutionary processes and island features. Intra-island speciation may play an important role in both island species richness and endemism, and the contribution of speciation to local species diversity may eventually be greater than that of immigration under certain conditions. Those conditions are related to the per species speciation rate, per species extinction rate, and island features, and they are independent of immigration rate. The model predicts that large islands will have a high, though not the highest, proportional endemism when other parameters are fixed. Based on the generalized model, changes in species richness and endemism on an oceanic island over time were predicted to be similar to empirical observations. Our model provides an ideal starting point for re-evaluating the role of speciation and re-analyzing available data on island species diversity, especially those biased by the MacArthur-Wilson model.     

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.     

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.     

Freshwater snail diversity: effects of pond area,habitat heterogeneity and isolation

Summary A large number of eutrophic ponds were surveyed for the presence of freshwater gastropods. Factors thought to influence the distribution of the snails were evaluated. As the investigated area has a homogeneous geological background physicochemical factors probably have a low effect on the local distribution of snails. There was a significant, positive regression between pond area and the number of gastropod species in the pond, but the regression only explained a minor part of the variation in species numbers. Multiple regression of an extended number of variables, associated with habitat complexity and dispersal, indicated that, in addition to area, macrophyte diversity and the mean number of gastropods in the 5 closest ponds (S₅) were important in explaining the distribution of gastropods. An increase in the number of macrophyte species increases the number of available microhabitats and refuges from predators. S₅ probably influences the dispersal rates between ponds. The gastropods in this area are thought to have additional dispersal modes, besides aerial dispersal with birds, and this probably increases the immigration rates and/or decreases the extinction rate.     

A system dynamics model of island biogeography

The MacArthur-Wilson equilibrium theory of island biogeography has been one of the more influential concepts in modern biogeography and ecology. In this paper, we synthesize the theory and examine effects of different immigration/extinction rate-species diversity curves on original predictions from the theory by using the System Dynamics simulation modeling approach. Moreover, we develop a comprehensive and generic System Dynamics model to incorporate a variety of recent modifications and extensions of the theory, including area effect, distance effect, competition effect, habitat diversity effect, target effect, and rescue effect. Through computer simulation with STELLA, a more profound understanding of the theory of island biogeography can be gained. The System Dynamics modeling approach is especially appropriate for such a study because it maximizes the utilization of the ecological data by incorporating qualitative information so that a complex, imprecisely-defined ecological system can be studied quantitatively, effectively, and comprehensively. Our simulation results show that different monotonic rate-species diversity curves do not affect the essence of the theory of island biogeography, while the magnitude of equilibrium species diversity may be greatly affected. Non-monotonic rate-species diversity curves may result in potential multiple equilibria of species diversity. In addition, our model suggests that a non-monotonic relationship may exist between the equilibrium turnover rate and island area and between the equilibrium turnover rate and distance.     

The island biogeography of exotic bird species

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

Pollination,biogeography and phylogeny of oceanic island bellflowers (Campanulaceae)

We studied the pollination biology of nine island Campanulaceae species: Azorina vidalii, Musschia aurea, M. wollastonii, Canarina canariensis, Campanula jacobaea, Nesocodon mauritianus, and three species of Heterochaenia. In addition, we compared C. canariensis to its two African mainland relatives C. eminii and C. abyssinica. We asked to what extent related species converge in their floral biology and pollination in related habitats, i.e. oceanic islands. Study islands were the Azores, Madeira, Canary Islands, Cape Verde, Mauritius, and Réunion. Information about phylogenetic relationships of these species and their relatives were gathered from atpB, matK, rbcL and trnL-F regions, building the most complete phylogeny of Campanulaceae to date. Six of the island bellflower species were bird-pollinated and two (A. vidalii and M. aurea) were lizard-pollinated. Insects also visited some of the species, and at least C. jacobaea had both insect- and self-pollination. Several morphological traits were interpreted as adaptations to bird and lizard pollination, e.g. all had a robust flower morphology and, in addition, bird-pollinated species were scentless, whereas lizard-pollinated species had a weak scent. These examples of vertebrate pollination evolved independently on each island or archipelago. We discuss if these pollination systems have an island or mainland origin and when they may have evolved, and finally, we attempt to reconstruct the pollinator-interaction history of each species.     

Concluding remarks: historical perspective and the future of island biogeography theory

MacArthur and Wilson’s equilibrium theory revolutionized the field of island biogeography and, to a large degree, ecology as well. The theory, which quickly became the ruling paradigm of island biogeography, has changed little over the past three decades. It has not kept pace with relevant theory and our growing appreciation for the complexity of nature, especially with empirical findings that species diversity on many islands: 1) is not in equilibrium; 2) is influenced by differences in speciation, colonization, and extinction among taxa; and 3) is influenced by differences among islands in characteristics other than area and isolation. The discipline of biogeography, itself, is in a state of disequilibrium. We may again be about to witness another paradigm shift, which will see the replacement of MacArthur and Wilson’s theory. Wherever this shift may take us, we are confident that the next generation of biogeographers will still look to islands for insights into the forces that shape biological diversity.     

The lizard fauna of Guam's fringing islets: island biogeography, phylogenetic history, and conservation implications

We sampled the lizard fauna of twenty-two small islets fringing the Pacific island of Guam and used these data to shed light on the processes responsible for present-day diversity. Habitat diversity, measured by islet area and vegetation complexity, was significantly correlated with the number of species found on an islet. However, islet distance and elevation were not significant predictors of diversity. Distribution patterns were slightly different for the two major families in our sample, Scincidae and Gekkonidae: skinks needed larger islets to maintain a population than did geckos. Presence/absence patterns were highly and significantly nested, and population density was correlated with the number of islets on which a species was found. An area cladogram was poorly supported and showed no faunal similarity between nearby islands. These patterns indicate that extinctions on most islets were due mostly to non-catastrophic, long-acting biological causes. The presence on the islets of species extirpated on Guam and the lack of significant nestedness on islands with greater maximum elevation highlight the impact that predators (primarily brown treesnakes) can have. Our findings also show that small reserves will not suffice to protect endangered lizard faunas, and that the islets may serve as a short-term repository of such species until snake-free areas can be established on Guam