Synthesis and Selection: Wynne-Edwards' Challenge to David Lack

David Lack of Oxford Universityand V.C. Wynne-Edwards of Aberdeen Universitywere renowned ornithologists with contrastingviews of the modern synthesis which deeplyinfluenced their interpretation and explanationof bird behavior. In the 1950's and 60's Lackbecame the chief advocate of neo-Darwinism withrespect to avian ecology, while Wynne-Edwardsdeveloped his theory of group selection. Lack's position was consistent with thedeveloping focus on individual leveladaptation, which was a core concept of themodern synthesis. Alternatively, Wynne-Edwardsviewed the emphasis on populations as the mostimportant development provided by the modernsynthesis. In this paper, I present thedevelopment of these two positions and tracetheir roots in the literature of the synthesis.Through an analysis of Lack's 1966 critique ofWynne-Edwards I conclude that Wynne-Edwardswas, in many ways, justified in his pursuit ofgroup level explanations.     

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.     

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.     

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.     

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.     

Bird assemblages on Amazonian river islands: Patterns of species diversity and composition

The principles of island biogeography are rarely applied to the animal assemblages of Amazonian river islands. Here, we compare bird assemblages of Amazonian river islands with a variety of mainland habitats. We also examine how bird species diversity and composition are related to island physical attributes. Birds were sampled with mist nets and qualitative censuses on 11 river islands and 24 mainland sites on the lower reaches of the Rio Negro in the Brazilian Amazon. Island bird assemblages were characterized by lower species richness and a higher abundance of a few dominant species. Additionally, the species composition of the islands was distinct from that of the mainland, including the nearby floodplain habitats. The number of bird species increased with island size and habitat diversity, and decreased with degree of isolation. In addition, small islands tended to harbor an impoverished subset of the species present on larger ones. Bird species diversity and composition on Amazonian river islands are likely influenced by the ecological succession and historical events affecting island formation. Considering their small total area across the Amazon basin, these insular fluvial communities could be disproportionately threatened by river channel disturbances related to climate change or hydroelectric dam development. Abstract in Portughese is available with online material.     

Breeding birds on small islands: island biogeography or optimal foraging?

1. We test MacArthur and Wilson's theory about the biogeography of communities on isolated habitat patches using bird breeding records from 16 small islands off the coasts of Britain and Ireland. 2. A traditional examination of patterns of species richness on these islands suggests that area and habitat diversity are important predictors, but that isolation and latitude have a negligible impact in this system. 3. Unlike traditional studies, we directly examine the fundamental processes of colonization and local extinction (cessation of breeding), rather than higher-order phenomena such as species richness. 4. We find that many of MacArthur and Wilson's predictions hold: colonization probability is lower on more isolated islands, and extinction probability is lower on larger islands and those with a greater diversity of habitats. 5. We also find an unexpected pattern: extinction probability is much lower on more isolated islands. This is the strongest relationship in these data, and isolation is the best single predictor of colonization and extinction. 6. Our results show that examination of species richness alone is misleading. Isolation has a strong effect on both of the dynamic processes that underlie richness, and in this system, the reductions in both colonization and extinction probability seen on more distant islands have opposing influences on species richness, and largely cancel each other out. 7. We suggest that an appropriate model for this system might be optimal foraging theory, which predicts that organisms will stay longer in a resource patch if the distance to a neighbouring patch is large. If nest sites and food are the resources in this system, then optimal foraging theory predicts the pattern we observe. 8. We advance the hypothesis that there is a class of spatial systems, defined by their scale and by the taxon under consideration, at which decision-making processes are a key driver of the spatiotemporal dynamics. The appropriate theory for such systems will be a hybrid of concepts from biogeography/metapopulation theory and behavioural ecology.     

Spatial patterns in species–area relationships and species distribution in a West African forest–savanna mosaic

Aim To investigate the relationship between the slope z of the species–area relationship (SAR) and the intensity of spatial patterns in species number and dissimilarity for woody plants with different modes of seed dispersal. According to island theory we expect, for any given archipelago, steeper slopes and more pronounced spatial patterns for groups of less dispersive species. Location Ivory Coast, West Africa. Methods In a West African forest–savanna mosaic we collected presence–absence data for woody plant species in 49 forest islands. The parameters of the SARs were fitted by nonlinear regressions and then compared for plant species aggregated according to their mode of seed dispersal. We used the Mantel test to calculate the intensity of spatial patterns in species number, i.e. residual deviation from SAR, and species dissimilarity. Results The z‐value for bird‐dispersed species was lower (0.11) than that for wind‐dispersed species (0.27), with mammal‐dispersed species taking an intermediate value (0.16). This result suggests that, as a group, bird‐dispersed species are better colonizers. The spatial pattern in species number as well as species similarity was more pronounced for bird‐ compared with wind‐dispersed species. Main conclusions The standard interpretation of the theory of island biogeography claims that shallow slopes in the SAR imply low isolation of islands, i.e. good dispersal abilities of species. The results of our study appear to contradict this statement. The contradiction can eventually be resolved by a more detailed account of the colonization process, i.e. by distinguishing between dispersal and consecutive establishment of populations.     

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.     

The biogeography of avian extinctions on oceanic islands

Aim To test the influences of island area, island isolation, and human‐introduced mammalian predators on avian extinctions that have occurred on oceanic islands worldwide. Location The oceanic islands of the world. Methods We augmented and re‐examined an existing data set for 218 oceanic islands by means of causal modelling using path analysis (a form of structural equation modelling) and a null model. Results The number of extinctions was not a simple function of the number of bird species on the various islands. Whereas introduced mammalian predators had an influence on the number of extinctions, island area (via indirect influences) and isolation (via direct influences) were equally or more important. Main conclusions The multiple influences of physical and biotic factors on past extinctions can be revealed through modelling the causal influences of physical attributes of islands on biological characteristics, and the causal influences of both physical and biological characteristics on extinctions.     

Macro‐scale bird species richness patterns of the East Asian mainland and islands: energy,area and isolation

Aim To create a map of bird species richness (BSR) in East Asia and to examine the effect of area, isolation, primary productivity, topographic heterogeneity, and human population density on BSR. Location East Asia (from 70° E to 180° E longitude), including the eastern half of the Palaearctic Region, the entire Oriental Region, and the entire Wallacea Subregion. Methods The breeding ranges of 2406 terrestrial bird species were mapped and overlaid to create a species richness map. The BSR map was transformed into a 100 × 100 km quadrat system, and BSR was analysed in relation to land area, average normalized difference vegetation index (NDVI), elevation range, and average population density. Results In general, BSR declined from the Tropics to the Arctic. In mainland East Asia, however, BSR was highest around the Tropic of Cancer, and fluctuated between 30° and 50° N. Islands had lower BSR than adjacent mainland areas. The NDVI was strongly positively correlated with BSR in mainland areas and on islands. For mainland areas, NDVI explained 65% of the BSR variation, and topographic heterogeneity explained an additional 6% in ordinary least‐squares regression. On islands, NDVI explained 66% of BSR variation, island area explained 13%, and distance to mainland accounted for 1%. Main conclusions In East Asia, we suggest that primary productivity is the key factor underpinning patterns of BSR. Primary productivity sets the upper limits of the capacity of habitats to support bird species. In isolated areas such as islands and peninsulas, however, BSR might not reach the richness limits set by primary productivity because the degree of isolation and area size also can affect species richness. Other factors, such as spatial heterogeneity, biotic interactions, and perturbations, may also affect species richness. However, their effects are secondary and are not as strong as primary productivity, isolation, and area size.     

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

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

Patterns of bird species richness and composition on islands off Arnhem Land,Northern Territory,Australia

The bird faunas of the adjacent Wessel and English Company island chains were sampled at two scales (0.25 ha quadrats and entire islands). Ninety‐six species were recorded from 226 quadrats, with the most frequently recorded species being mistletoebird Dicaeum hirundinaceum, brown honeyeater Lichmera indistincta, silver‐crowned friarbird Philemon argenticeps, bar‐shouldered dove Geopelia humeralis, northern fantail Rhipidura rufiventris and yellow white‐eye Zosterops lutea. At the quadrat scale, vegetation type was a major determinant of the abundance of individual species (and hence species composition), species richness and total bird abundance. Bird species composition and richness at the quadrat scale was also significantly affected by island isolation (particularly the amount of land within 20 km of the island perimeter). Island size had no effect on quadrat‐scale richness or total abundance. However, the abundance of 10 of the 38 most frequently recorded individual species was significantly related to island size, in most cases even when the comparison was restricted to similar habitats. The most striking cases were rufous fantail Rhipidura rufifrons, mangrove golden whistler Pachycephala melanura, brown honeyeater and yellow white‐eye, which were all significantly more abundant on smaller islands. One hundred and seventy‐one species were recorded from the 62 islands sampled. There was a very tight relationship between island size and the number of terrestrial species (73% of deviance explained) and of all species (84% of deviance explained). This relationship was improved (marginally) when isolation was included in the model. Ordination of islands by their terrestrial bird species composition was related to island size and isolation, and suggested an erratic species composition on small islands.     

Historical bird and terrestrial mammal extinction rates and causes

Aim Conservation of species is an ongoing concern. Location Worldwide. Methods We examined historical extinction rates for birds and mammals and contrasted island and continental extinctions. Australia was included as an island because of its isolation. Results Only six continental birds and three continental mammals were recorded in standard databases as going extinct since 1500 compared to 123 bird species and 58 mammal species on islands. Of the extinctions, 95% were on islands. On a per unit area basis, the extinction rate on islands was 177 times higher for mammals and 187 times higher for birds than on continents. The continental mammal extinction rate was between 0.89 and 7.4 times the background rate, whereas the island mammal extinction rate was between 82 and 702 times background. The continental bird extinction rate was between 0.69 and 5.9 times the background rate, whereas for islands it was between 98 and 844 times the background rate. Undocumented prehistoric extinctions, particularly on islands, amplify these trends. Island extinction rates are much higher than continental rates largely because of introductions of alien predators (including man) and diseases. Main conclusions Our analysis suggests that conservation strategies for birds and mammals on continents should not be based on island extinction rates and that on islands the key factor to enhance conservation is to alleviate pressures from uncontrolled hunting and predation.     

Terrestrial bird community patterns on the coralline islands of the Dahlak Archipelago, Red Sea, Eritrea

Aim This study aims to explain the patterns of species richness and nestedness of a terrestrial bird community in a poorly studied region. Location Twenty‐six islands in the Dahlak Archipelago, Southern Red Sea, Eritrea. Methods The islands and five mainland areas were censused in summer 1999 and winter 2001. To study the importance of island size, isolation from the mainland and inter‐island distance, I used constrained null models for the nestedness temperature calculator and a cluster analysis. Results Species richness depended on island area and isolation from the mainland. Nestedness was detected, even when passive sampling was accounted for. The nested rank of islands was correlated with area and species richness, but not with isolation. Idiosyncrasies appeared among species‐poor and species‐rich islands, and among common and rare species. Cluster analysis showed differences among species‐rich islands, close similarity among species‐poor and idiosyncratic islands, and that the compositional similarity among islands decreased with increasing inter‐island distance. Thus, faunas of species‐poor, smaller islands were more likely to be subsets of faunas of species‐rich, larger islands if the distance between the islands was short. Main conclusions Species richness and nestedness were related to island area, and nestedness also to inter‐island distances but not to isolation from the mainland. Thus, nestedness and species richness are not affected in the same way by area and distance. Moreover, idiosyncrasies may have been the outcome of species distributions among islands being influenced also by non‐nested distributions of habitats, inter–specific interactions, and differences in species distributions across the mainland. Idiosyncrasies in nested patterns may be as important as the nested pattern itself for conservation – and conservation strategies based on nestedness and strong area effects (e.g. protection of only larger islands) may fail to preserve idiosyncratic species/habitats.     

Revealing Beta-Diversity Patterns of Breeding Bird and Lizard Communities on Inundated Land-Bridge Islands by Separating the Turnover and Nestedness Components

Beta diversity describes changes in species composition among sites in a region and has particular relevance for explaining ecological patterns in fragmented habitats. However, it is difficult to reveal the mechanisms if broad sense beta-diversity indices (i.e. yielding identical values under nestedness and species replacement) are used. Partitioning beta diversity into turnover (caused by species replacement from site to site) and nestedness-resultant components (caused by nested species losses) could provide a unique way to understand the variation of species composition in fragmented habitats. Here, we collected occupancy data of breeding birds and lizards on land-bridge islands in an inundated lake in eastern China. We decomposed beta diversity of breeding bird and lizard communities into spatial turnover and nestedness-resultant components to assess their relative contributions and respective relationships to differences in island area, isolation, and habitat richness. Our results showed that spatial turnover contributed more to beta diversity than the nestedness-resultant component. The degree of isolation had no significant effect on overall beta diversity or its components, neither for breeding birds nor for lizards. In turn, in both groups the nestedness-resultant component increased with larger differences in island area and habitat richness, respectively, while turnover component decreased with them. The major difference among birds and lizards was a higher relevance of nestedness-resultant dissimilarity in lizards, suggesting that they are more prone to local extinctions derived from habitat fragmentation. The dominance of the spatial turnover component of beta diversity suggests that all islands have potential conservation value for breeding bird and lizard communities.     

The Assembly of an Island Fauna by Natural Invasion: Sources and Temporal Patterns in the Avian Colonization of Barbados

By virtue of their isolation and depauperate faunas, oceanic islands offer unique opportunities to characterize the historical development of ecological communities derived from both natural and anthropogenic invasions. Barbados, an outlying island in the Lesser Antilles, was formed approximately 700,000 YBP by tectonic uplift and was then colonized by birds via natural invasion from the much older volcanic islands in the main Lesser Antillean arc. We investigated the timing and sources of the avian invasion of Barbados by determining levels of mitochondrial DNA (mtDNA) divergence between populations of eight bird species from Barbados and those on the nearby putative source islands of St. Lucia and St. Vincent. Although all Barbados populations appeared to be young relative to the geological age of the island, we found differences among species in their inferred times of colonization and we identified at least two sources of immigrants to Barbados. In contrast to these historical differences across species and populations, our characterization of the mitochondrial genotypes of 231 individual birds suggests that each island population represents the descendants of a single founding maternal lineage. Considered in concert, the results of this molecular survey indicate that the Barbados bird community is composed of species with different invasion histories, which in turn suggests that the island's community composition has changed repeatedly over its 700,000 year history.     

Island biogeography theory explains the genetic diversity of a fragmented rock ptarmigan (Lagopus muta) population

The island biogeography theory is one of the major theories in ecology, and its applicability to natural systems is well documented. The core model of the theory, the equilibrium model of island biogeography, predicts that species diversity on an island is positively related to the size of the island, but negatively related by the island's distance to the mainland. In recent years, ecologists have begun to apply this model when investigating genetic diversity, arguing that genetic and species diversity might be influenced by similar ecological processes. However, most studies have focused on oceanic islands, but knowledge on how the theory applies to islands located on the mainland (e.g., mountain islands, forest islands) is scarce. In this study, we examined how the size and degree of isolation of mountain islands would affect the genetic diversity of an alpine bird, the rock ptarmigan (Lagopus muta). Within our study area, we defined the largest contiguous mountain area as the mainland, while smaller mountains surrounding the mainland were defined as islands. We found that the observed heterozygosity (H_o) was significantly higher, and the inbreeding coefficient (F_is) significantly lower, on the mainland compared to islands. There was a positive significant relationship between the unbiased expected heterozygosity (H_n.b.) and island size (log km²), but a negative significant relationship between H_o and the cost distance to the mainland. Our results are consistent with the equilibrium model of island biogeography and show that the model is well suited for investigating genetic diversity among islands, but also on islands located on the mainland.     

Bird species numbers in an archipelago of reeds at Lake Velence, Hungary

1 Bird species numbers were studied on 109 reed islands at Lake Velence, Hungary, in the 1993 and 1994 breeding seasons. The aim was to describe and account for the abundance and distribution patterns of the bird species. 2 It was expected that an exponential model would fit the calculated species–area curves. However, for the 1993 data, both the power function (LogS ~ LogArea) and the exponential (S ~ LogArea) models did so, while the power function, exponential and linear (S ~ A) models fitted the curves for the 1994 data. 3 The results showed that the pattern was not random: a collection of small islands held more species than a few large islands with the same total area. 4 The relative species richness of small islands is a result of the preference of most common passerine bird species for the edges of reed islands. Most individuals were found in the first 5 m of the reedbed, and no edge avoidance was detected on a local spatial scale. Large, rarer species (e.g. Great White Egret), however, were found to be dependent on large reed islands. 5 Comparison of results with two other studies on bird communities of reed islands revealed that the type of landscape matrix (e.g. deep water, shallow water or agricultural lands) among reed patches significantly influences bird communities. Deep water was dominated by grebes and coot, shallow water by reed‐nesting passerines, and farmed areas by reed‐ and bush‐nesting passerines.     

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.