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Canopy gaps are widely recognized as being crucial for maintaining the diversity of forest tree communities. But empirical studies have found mixed results because the differences in diversity between individual gaps and non‐gaps are often small and statistically undetectable. One overlooked factor, however, is how small individual gap versus non‐gap differences may accumulate across sites and potentially have a large effect on forest diversity at the plot scale. Our study investigated sapling richness, density, and composition in 124 treefall gaps, and 200 non‐gap sites in the 50‐ha tropical forest plot at Barro Colorado Island (BCI), Panama. Additionally, we analyzed species accumulation curves to understand how species richness increases with increasing stem numbers. We observed that sapling richness and density were only slightly higher in gaps 7 years after formation and statistically indistinguishable from non‐gaps after 12 years. However, species accumulation curves across multiple gaps were substantially higher than those across non‐gaps. Species composition showed small differences between individual gaps and non‐gaps but differed significantly between collections of gaps and non‐gaps. Specifically, 55 species specialized in 7‐year‐old gaps compared with 24 in non‐gaps; of these, 23 gap‐specialized species and zero non‐gap species were pioneers. Our results indicate that tree species richness is higher in gaps because of both higher stem density and the presence of gap‐specialized species. Our study has finally provided compelling evidence to support the idea that gaps enhance the overall diversity of tropical forest tree communities. [ABSTRACT FROM AUTHOR]

Hyperdiverse tropical forest tree communities illustrate a fundamental problem in ecology: How can many species coexist given relatively few limiting resources? Neutral theory provides a solution by positing that species have equal fitness and hence drift to extinction slowly. However, neutral theory seriously under‐predicts temporal changes in species abundances. This can be remedied by breaking neutrality and adding temporal environmental stochasticity (TES), but the mechanisms mediating the effects of TES on species richness remain unclear. Here, we make progress by analysing a local community model with species competing for a common resource under TES, to derive formulae partitioning species richness according to different mechanisms.By applying our formulae to generic parameter sets for tropical forest tree communities, we found that when the autocorrelation time of TES was short, the dominant mechanism driving species richness was non‐linear averaging of the interspecific competition term over time, which reduced the typical strength of interspecific competition and boosted richness relative to the neutral case. However, greater immigration to the community resulted in more species and hence weaker non‐linear averaging due to the law of large numbers. In contrast, when the autocorrelation time of TES was long, the dominant mechanism driving species richness was strong selection between changes in environmental conditions, which increased the typical strength of interspecific competition and reduced richness relative to the neutral case.By applying our formulae to a specific parameter set for a tropical forest tree community in Panama, we found that TES had minor effects on species richness because (i) the immigration rate was sufficiently large for non‐linear averaging of the interspecific competition term to be weak and (ii) the autocorrelation time was sufficiently short to suppress the effects of selection.Synthesis. We provide a novel mechanistic explanation of how TES affects tree species richness in tropical forests, in particular how TES often has minor effects on richness despite having substantial effects on temporal changes in species abundances. This provides a deeper insight into why a neutral model with added TES can accurately capture static and dynamic aspects of tree community diversity in the tropics. [ABSTRACT FROM AUTHOR]

Aim: Artificial island habitats such as human‐made wetlands are emerging novel ecosystems. Understanding the drivers of diversity in such artificial systems is essential for balancing the goals of biodiversity conservation and human socio‐economic needs. Location: Telangana state, India. Methods: We surveyed water birds in a network of 57 artificial wetlands and assessed four macroecological biodiversity patterns: spatial betadiversity, temporal betadiversity, species‐abundance distributions (SADs), and the species–area relationship (SAR). We employed a mix of phenomenological and mechanistic models to examine the four macroecological patterns. We hypothesized that the wetland bird communities are primarily structured by immigration–extinction dynamics and thus that spatial and temporal betadiversity would be high, the within‐wetland SADs would exhibit a large number of rare species and a monotonically declining overall shape, and that the SAR across wetlands would be strongly increasing. Results: Spatial and temporal betadiversity were both high and mostly attributable to turnover rather than nestedness. While the pooled SAD exhibited an interior mode, the SAD for individual wetlands was generally log‐series distributed, consistent with a model in which immigration among wetlands is high. The SAR exhibited an increasing trend, with the 'small‐island effect', which reflects constraints on immigration and is often observed for true island archipelagos, being absent. Main Conclusions: We tentatively conclude that bird diversity in this network of artificial wetlands is mainly structured by immigration–extinction dynamics, although we acknowledge that some of the patterns are also consistent with niche dynamics and future research should measure relevant biotic and abiotic variables in these wetlands. We encourage future work in which our rich dataset is used to fit dynamic models that permit more‐detailed quantitative inferences about mechanisms structuring diversity in this novel ecosystem, which can ultimately also inform conservation management. [ABSTRACT FROM AUTHOR]

Neutral models in ecology assume that all species are demographically equivalent, such that their abundances differ ultimately because of demographic stochasticity rather than selection. In spite of their simplicity, neutral models have been found to accurately reproduce static patterns of biodiversity for diverse communities. However, the same neutral models have been found to exhibit species abundance dynamics that are far too slow compared to reality, resulting in poor fits to temporally dynamic patterns of biodiversity. Here, we show that one of the root causes of these slow dynamics is the additional assumption that a community has reached an equilibrium with a fixed community size, with species that have a net growth rate close to zero. We removed this additional assumption by constructing and analyzing a neutral model with an expected community size that can change over time and is not necessarily at equilibrium, which thus allows the historical formation of a community to be represented explicitly. Our analysis demonstrated that for the general scenario where a small community rapidly grows in size to a carrying capacity, representing recovery from ecological disturbance or assembly of a new community, the model produced much larger changes in species abundances and much shorter species ages than a neutral model at an equilibrium with fixed community size. In addition, the species abundance distribution was biphasic with a subset of abundant species arising from a founder effect. We confirmed these new results in applications of the new model to the specific scenario of recovery of the Amazon tree community after the end-Cretaceous bolide impact, which involved periods of increasing and decreasing community size. We conclude that incorporating transient dynamics in neutral models improves realism by allowing explicit consideration of how a community is formed over realistic time-scales. • Transient dynamics of a neutral model with varying community size are analyzed. • Formulae are derived specifying expected species richness and species age. • Formulae are derived specifying the species-abundance distribution. • Model gives a biphasic SAD with common species arising from a founder effect. • Model also gives realistic species ages for a tree community at an Amazon scale. [ABSTRACT FROM AUTHOR]

Estimating the total number of species on Earth has been a longstanding pursuit. Models project anywhere between 2 and 10 million species, and discovery of new species continues to the present day. Despite this, we hypothesized that our current knowledge of phylogenetic diversity (PD) may be almost complete because new discoveries may be less phylogenetically distinct than past discoveries. Focusing on birds, which are well studied, we generated a robust phylogenetic tree for most extant species by combining existing published trees and calculated each discovery's marginal contribution to known PD since the first formal species descriptions in 1758. We found that PD contributions began to plateau in the early 1900s, about half a century earlier than species richness. Relative contributions of each phylogenetic order to known PD shifted over the first 150 years, with a growing contribution of the hyper-diverse perching birds (Passeriformes) in particular, but after the early 1900s this has remained relatively stable. Altogether, this suggests that our knowledge of the evolutionary history of extant birds is mostly complete, with few discoveries of high evolutionary novelty left to be made, and that conclusions of studies using avian phylogenies are likely to be robust to future species discoveries. [ABSTRACT FROM AUTHOR]

A long‐standing puzzle in ecology is coexistence of many species despite relatively few limiting resources. Studies using competitive community models have found that temporal environmental stochasticity (TES) can provide a solution by providing a rare‐species advantage, for example by creating temporal niches. However, this appears to contradict studies using population models, which have found that TES‐induced temporal fluctuations in species abundances increase the chance of species becoming rare and hence increase their extinction risk. Here, we clarified how TES affects species richness by analysing a competitive community model using a novel mean‐field approach. We found that when temporal correlation in TES was weak, the dominant effect of TES was to create temporal niches by decreasing the strength of interspecific competition via a temporal averaging effect, thereby increasing species richness relative to the neutral case without TES. In contrast, when temporal correlation was strong, the dominant effect of TES was to increase the strength of selection and hence interspecific competition, thereby decreasing species richness. Compared with these indirect community effects of TES, the direct effects of TES on species richness via temporal fluctuations in abundances of species populations were relatively minor. [ABSTRACT FROM AUTHOR]

Theoretical and empirical studies suggest that as immigration increases, ecological communities transition from a niche-structured regime to an immigration-structured regime. The niche-structured regime is the domain of classic niche models; the immigration-structured regime is the domain of island biogeography and related theories. A recent unified model predicted a biphasic species–area relationship (SAR) arising from the transition between the two regimes, but the generality and scope of this relationship remain unclear. Here we study the transition further to address three key questions: (1) Can MacArthur and Wilson's classic graphical paradigm of intersecting immigration and extinction curves be adapted to capture the niche-structured regime that occurs at low-immigration rates? (2) Do different ecological models predict a similar biphasic SAR? (3) Can the biphasic island SAR be reconciled with the classic triphasic SAR observed in mainland biogeography? On the first point, we find that the transition can be incorporated into MacArthur and Wilson's graphical paradigm by forcing the extinction curves sharply downwards at low species richness, reflecting the stabilizing effect of niche processes. On the second point, we confirm that a variety of simple ecological models produce qualitatively similar biphasic SARs. On the third point, we find that a unified model predicts an overall four-phase SAR with the second phase being a shallow niche-structured phase that is rarely observed in mainland SARs, which we hypothesize is because local communities on mainlands are usually in the immigration-structured regime. [ABSTRACT FROM AUTHOR]

Aim: Research in island biogeography has long focused mainly on present‐day island configurations. Recently, there has been an increasing focus on islands' past histories of land connection, shape and size. Moreover, continental islands (=shelf islands) have received less attention than oceanic islands, and species inventories from extremely small islands are lacking in many datasets. We examine the effects of sea‐level rise since the Last Glacial Maximum (LGM) on bird species diversity and composition of tropical shelf islands in Southeast Asia. Location: Sundaland. Taxon: Birds. Methods: We compiled avifaunal island inventories for 94 islands using an exhaustive literature review of historic surveys of larger islands combined with our own comprehensive island surveys from both small and large islands. Using generalised least‐squares models with spatial autocorrelation, we assessed the importance of traditional biogeographical parameters including area, maximum elevation, distance from mainland and geographical isolation, along with post‐LGM effects of change in island area and duration since isolation. We also compared the species composition on similar‐sized shelf islands from two categories—recently submerged and unsubmerged—using non‐metric multidimensional scaling. Results: Post‐LGM effects on species diversity are minimal and insular diversity is instead well explained by present‐day island characteristics, such as area, distance to mainland and proportion of land surrounding an island within a 10 km radius (Cox and Snell Pseudo‐R2 = 0.803). Avifaunal diversity is similar across recently submerged and unsubmerged small shelf islands. Main conclusion: Avifaunal diversity on tropical shelf islands equilibrates rapidly after isolation, indicating that both extinction and immigration rates are high. In particular, a high immigration rate of dispersive species maintains diversity, especially on small islands. Over‐water dispersal is generally restricted to short distances among Sundaic birds. Consequently, the diversity of an island can be maintained by the presence of large or stepping‐stone islands near it. [ABSTRACT FROM AUTHOR]

Theoretical ecologists have analysed a range of neutral models but few including stage structure. Here we introduce a stage‐structured neutral model, by extending the standard spatial neutral model to have two‐stage classes: a juvenile stage and a reproductive stage. We find that formulas for biodiversity patterns (e.g. species–area relationships and species abundance distributions) of reproductives in the stage‐structured model can be obtained from the corresponding standard formulas via a parameter rescaling, which involves calculating an effective speciation rate parameter and an effective dispersal parameter. This is useful because it means existing knowledge about the non‐stage‐structured model can be transferred to the stage‐structured model, providing that applications focus exclusively on the reproductive stage. One surprising implication is that the presence of a juvenile stage can substantially increase the species richness of reproductive individuals: a juvenile stage with a length fraction k that of the reproductive stage increases reproductive species richness by roughly the same factor. We apply our new formulas to a case study of tropical forest trees in Panama and find that while the stage‐structured model makes different predictions than the standard model, it does not fix known problems with cross‐scale predictions. We speculate that some of our results, in particular the result that the presence of a juvenile stage increases community diversity, likely apply to non‐neutral systems as well. [ABSTRACT FROM AUTHOR]

Half a century ago, Janzen and Connell hypothesized that the high tree species diversity in tropical forests is maintained by specialized natural enemies. Along with other mechanisms, these can cause conspecific negative density dependence (CNDD) and thus maintain species diversity. Numerous studies have measured proxies of CNDD worldwide, but doubt about its relative importance remains. We find ample evidence for CNDD in local populations, but methodological limitations make it difficult to assess if CNDD scales up to control community diversity and thereby local and global biodiversity patterns. A combination of more robust statistical methods, new study designs, and eco-evolutionary models are needed to provide a more definite evaluation of the importance of CNDD for geographic variation in plant species diversity. Stabilizing conspecific negative density dependence (CNDD) is a classical explanation for diversity maintenance in species-rich ecosystems, particularly in tropical forests. Fifty years after Janzen and Connell popularized this idea, a plethora of empirical evidence suggests that CNDD can be found at all latitudes and for many tree species. On closer inspection, however, it is still unclear if locally measured density effects are indeed stabilizing community dynamics and, moreover, if they have a causal effect on large-scale diversity and abundance patterns, such as the latitudinal diversity gradient. More robust and comparable CNDD estimates are needed, coupled with a theoretical research program that aims at understanding the role of CNDD for coexistence in stochastic multispecies communities as well as macroecological and macroevolutionary diversity patterns. [ABSTRACT FROM AUTHOR]

A goal of ecology is to identify the stabilizing mechanisms that maintain species diversity in the face of competitive exclusion and drift. For tropical forest tree communities, it has been hypothesized that high diversity is maintained via Janzen-Connell effects, whereby host-specific natural enemies prevent any one species from becoming too abundant. Here we explore the plausibility of this hypothesis with theoretical models. We confirm a previous result that when added to a model with drift but no competitive exclusion—that is, a neutral model where intrinsic fitnesses are perfectly equalized across species—Janzen-Connell effects maintain very high species richness that scales strongly with community size. However, when competitive exclusion is introduced—that is, when intrinsic fitnesses vary across species—the number of species maintained by Janzen-Connell effects is substantially reduced and scales much less strongly with community size. Because fitness variation is pervasive in nature, we conclude that the potential of Janzen-Connell effects to maintain diversity is probably weak and that the mechanism does not yet provide a sufficient explanation for the observed high diversity of tropical forest tree communities. We also show that, surprisingly, dispersal limitation can further reduce the ability of Janzen-Connell effects to maintain diversity. [ABSTRACT FROM AUTHOR]

The relationship between species richness and productivity changes with spatial scale, but the way in which it changes and the underlying mechanisms remain unclear. We address this critical knowledge gap using a new mechanistic model of the spatial scaling of species richness–productivity (SP) relationships for a local community. Our model is neutral and hence assumes that species dynamics are driven by dispersal limitation and demographic stochasticity. We showed analytically that SP relationships predicted by our model are typically unimodal. Consistent with previous simulation-based studies, the positive phase of our unimodal SP relationship was driven by a sampling effect ("more-individuals effect") whereas the negative phase was driven by relatively more propagules being of local as opposed to external origin ("dilution effect"). Our main novel finding related to the spatial scaling of the unimodal SP relationship: the peak shifted to the left with increasing spatial scale, such that the decreasing phase covered a greater range of productivity. This was driven by an increase in the strength of the dilution effect relative to the more-individuals effect, reflecting higher area/perimeter ratios at larger spatial scales. Our theoretical predictions are qualitatively consistent with the spatial scaling of SP relationships documented for trees in localized forest communities across the world. [ABSTRACT FROM AUTHOR]

Among the local processes that determine species diversity in ecological communities, fluctuation‐dependent mechanisms that are mediated by temporal variability in the abundances of species populations have received significant attention. Higher temporal variability in the abundances of species populations can increase the strength of temporal niche partitioning but can also increase the risk of species extinctions, such that the net effect on species coexistence is not clear. We quantified this temporal population variability for tree species in 21 large forest plots and found much greater variability for higher latitude plots with fewer tree species. A fitted mechanistic model showed that among the forest plots, the net effect of temporal population variability on tree species coexistence was usually negative, but sometimes positive or negligible. Therefore, our results suggest that temporal variability in the abundances of species populations has no clear negative or positive contribution to the latitudinal gradient in tree species richness. [ABSTRACT FROM AUTHOR]

Abstract: To estimate species loss from habitat destruction, ecologists typically use species–area relationships, but this approach neglects the spatial pattern of habitat fragmentation. Here, we provide new, easily applied, analytical methods that place upper and lower bounds on immediate species loss at any spatial scale and for any spatial pattern of habitat loss. Our formulas are expressed in terms of what we name the ‘Preston function’, which describes triphasic species–area relationships for contiguous regions. We apply our method to case studies of deforestation and tropical tree species loss at three different scales: a 50 ha forest plot in Panama, the tropical city‐state of Singapore and the Brazilian Amazon. Our results show that immediate species loss is somewhat insensitive to fragmentation pattern at small scales but highly sensitive at larger scales: predicted species loss in the Amazon varies by a factor of 16 across different spatial structures of habitat loss. [ABSTRACT FROM AUTHOR]

Species populations are subjected to fluctuations in their surrounding environment, and the strength of these fluctuations has been hypothesized to be a major determinant of the extinction risk of these populations. Therefore, a key question is: How does temporal environmental variance affect the extinction risk of species populations? Previous theory based on the dynamics of single populations typically predicts an increased risk of extinction from the effects of environmental variance. However, previous studies have focused mainly on the case where environmental effects are temporally uncorrelated (white environmental noise), whereas such effects are typically correlated (colored environmental noise) in nature. Thus, further work on the case of colored environmental noise is required, but this has been hindered by the analytical intractability of corresponding stochastic models. In our study, we address this limitation by developing a new discrete-time Markov chain model of a species population fluctuating under colored environmental noise, with the simplification that the effects of demographic variance are manifested indirectly as an extinction threshold. This simplifying assumption allows us to derive analytical solutions, which show that the expected extinction time of model species declines with the strength of environmental variance under a variety of different scenarios, reflecting greater extinction risk. Our study thus clarifies the situations under which environmental variance tends to increase extinction risk, and provides a novel analytically tractable framework for modeling temporal environmental variance. We also discuss the possible implications of our results for species richness in ecological communities. [ABSTRACT FROM AUTHOR]

The compositional complexity of an assemblage is not expressible as a single number; standard measures such as diversities (Hill numbers) and entropies (Rényi entropies and Tsallis entropies) vary in their order q which determines the measures' emphasis on rare or common species. Ranking and comparing assemblages depend on the choice of q. Rather than selecting one or a few measures to describe an assemblage, it is preferable to convey the complete story by presenting a continuous profile, a plot of diversity or entropy as a function of q ≥ 0. This makes it easy to visually compare the compositional complexities of multiple assemblages and to judge the evenness of the relative abundance distributions of the assemblages. In practice, the profile is plotted for all values of q from 0 to q = 3 or 4 (beyond which it generally changes little)., These profiles are usually generated by substituting species sample proportions into the complexity measures. However, this empirical approach typically underestimates the true profile for low values of q, because samples usually miss some of the assemblage's species due to under-sampling. Although bias-reduction methods exist for individual measures of order q = 0, 1 and 2, there has been no analytic method that unifies these bias-corrected estimates into a continuous profile., For incomplete sampling data, this work proposes a novel analytic method to obtain accurate, continuous, low-bias diversity and entropy profiles with focus on low orders of q (0 ≤ q ≤ 3). Our approach is based on reformulating the diversity and entropy of any order q in terms of the successive discovery rates of new species with respect to sample size, that is the successive slopes of the species accumulation curve. A bootstrap method is applied to obtain approximate variances of our proposed profiles and to construct the associated confidence intervals., Extensive simulations from theoretical models and real surveys show that the proposed profiles greatly reduce under-sampling bias and have substantially lower bias and mean-squared error than the empirical profile, especially for q ≤ 1. Our method is also extended to deal with incidence data. [ABSTRACT FROM AUTHOR]

Many questions in ecology and evolutionary biology consider response variables that are functions (e.g. species-abundance distributions) rather than a single scalar value (e.g. species richness). Although methods for analysing function-valued data have been available for several decades, ecological and evolutionary applications are rare., We outline methods for regression when the response variable is a function ('function regression') and introduce the r package FREE, which focuses on straightforward implementation and interpretation of function regression analyses. Several computational methods are implemented, including machine learning and several Bayesian methods. We compare different methods using simulated data and real ecological data on individual-size distributions ( ISDs) of fish and trees., No single method performed best overall, with several performing equally well for a given data set. Which method to use depends on sample sizes and the questions being considered; in many cases, a consensus approach should be used to combine or compare fitted models., Function regression allows the direct modelling of many function-valued data (e.g. species-abundance distributions) rather than having to reduce those functions to a single scalar response variable (e.g. species diversity or functional diversity indices). Our ecological examples using ISD data show that larger rivers support more-even fish-size distributions than smaller rivers and that low initial planting densities lead to more-even tree-size distributions than high initial planting densities. Function regression provided more informative and intuitive interpretations of these data than conventional non-function-valued approaches. [ABSTRACT FROM AUTHOR]

Individual species are distributed inhomogeneously over space and time, yet, within large communities of species, aggregated patterns of biodiversity seem to display nearly universal behaviour. Neutral models assume that an individual's demographic prospects are independent of its species identity. They have successfully predicted certain static, time-independent patterns. But they have generally failed to predict temporal patterns, such as species ages or population dynamics. We construct a new, multispecies framework incorporating competitive differences between species, and assess the impact of this competition on static and dynamic patterns of biodiversity. We solve this model exactly for the special case of a Red Queen hypothesis, where fitter species are continually arising. The model predicts more realistic species ages than neutral models, without greatly changing predictions for static species abundance distributions. Our modelling approach may allow users to incorporate a broad range of ecological mechanisms. [ABSTRACT FROM AUTHOR]

Abstract: Biogeography seeks to understand the mechanisms that drive biodiversity across long temporal and large spatial scales. Theoretical models of biogeography can be tested by comparing their predictions of quantities such as species ages against empirical estimates. It has previously been claimed that the neutral theory of biodiversity and biogeography predicts species ages that are unrealistically long. Any improved theory of biodiversity must rectify this problem, but first it is necessary to quantify the problem precisely. Here we provide analytical expressions for species ages in neutral biodiversity communities. We analyse a spatially implicit metacommunity model and solve for both the zero-sum and non-zero-sum cases. We explain why our new expressions are, in the context of biodiversity, usually more appropriate than those previously imported from neutral molecular evolution. Because of the time symmetry of the spatially implicit neutral model, our expressions also lead directly to formulas for species persistence times and species lifetimes. We use our new expressions to estimate species ages of forest trees under a neutral model and find that they are about an order of magnitude shorter than those predicted previously but still unrealistically long. In light of our results, we discuss different models of biogeography that may solve the problem of species ages. [Copyright &y& Elsevier]

The relationship between species richness and ecosystem function, as measured by productivity or biomass, is of long-standing theoretical and practical interest in ecology. This is especially true for forests, which represent a majority of global biomass, productivity and biodiversity., Here, we conduct an analysis of relationships between tree species richness, biomass and productivity in 25 forest plots of area 8-50 ha from across the world. The data were collected using standardized protocols, obviating the need to correct for methodological differences that plague many studies on this topic., We found that at very small spatial grains (0.04 ha) species richness was generally positively related to productivity and biomass within plots, with a doubling of species richness corresponding to an average 48% increase in productivity and 53% increase in biomass. At larger spatial grains (0.25 ha, 1 ha), results were mixed, with negative relationships becoming more common. The results were qualitatively similar but much weaker when we controlled for stem density: at the 0.04 ha spatial grain, a doubling of species richness corresponded to a 5% increase in productivity and 7% increase in biomass. Productivity and biomass were themselves almost always positively related at all spatial grains., Synthesis. This is the first cross-site study of the effect of tree species richness on forest biomass and productivity that systematically varies spatial grain within a controlled methodology. The scale-dependent results are consistent with theoretical models in which sampling effects and niche complementarity dominate at small scales, while environmental gradients drive patterns at large scales. Our study shows that the relationship of tree species richness with biomass and productivity changes qualitatively when moving from scales typical of forest surveys (0.04 ha) to slightly larger scales (0.25 and 1 ha). This needs to be recognized in forest conservation policy and management. [ABSTRACT FROM AUTHOR]

Isolating the effects of fragmentation per se (i.e., spatial configuration of habitat patches) on species richness is an ongoing challenge as habitat configuration often covaries with the amount of habitat. Consequently, there is a lack of experimental evidence for configurational effects on species richness in the whole landscape. Here, we developed a novel experimental system for testing the independent and interactive effects of habitat area and configuration on tropical intertidal species richness. Our results confirmed the expectation that average species richness would increase monotonically with habitat area. More intriguingly, we found mixed evidence for a non‐monotonic relationship between species richness and fragmentation per se, with the highest richness at intermediate fragmentation configuration, that is, when habitat tiles were placed in a "several‐small" configuration. The effect of habitat configuration was not due to passive sampling (since area was controlled for), variation in total individual abundance, or niche specialization of species to different landscape configurations. We postulate that a combination of processes, including local negative density dependence and dispersal limitation, could give rise to the observed pattern. We emphasize the importance of considering configurational effects on biodiversity at broader spatial scales and for more experimental research to delve into the mechanisms driving the patterns seen here. [ABSTRACT FROM AUTHOR]

• Probability distribution of species' extinction time derived for a neutral model. • The distribution nearly always gives overestimates of tree species ages. • Probability distributions of species richness derived for two neutral models. • The distributions show that species richness has low variation in large communities. • Formulae derived for immigration and extinction curves from a neutral model. In community ecology, neutral models make the assumption that species are equivalent, such that species abundances differ only because of demographic stochasticity. Despite their ecological simplicity, neutral models have been found to give reasonable descriptions of expected patterns of biodiversity in communities with many species. Such patterns include the expected total number of species and species-abundance distributions describing the expected number of species in different abundance classes. However, the expected patterns represent only the central tendencies of the full distributions of possible outcomes. Thus, ecological inferences and conclusions based only on expected patterns are incomplete, and may be misleading. Here, we address this issue for the spatially implicit neutral model, by using classic results from birth–death processes to derive (1) the probability distribution of extinction time of a species with given abundance for the metacommunity; (2) the probability distributions of total species richness and number of species with given abundance for both the metacommunity and local community; and (3) the probability distributions of the average immigration and extinction rates in the local community, across different values of total species richness. We illustrate the utility of these probability distributions in providing greater ecological insight via statistical inference. Firstly, we show that under the neutral metacommunity model, there is only 2.65 × 10 − 9 probability that the age of a common tree species in the Amazon is ≤ 3 × 108 yr, which is approximately the oldest estimated age of the first angiosperm. Thus, species ages from the model are unrealistically high. Secondly, for a tree community in a 50 ha plot at Barro Colorado Island in Panama, we show that the spatially implicit model can be fitted to observed species richness and an independent estimate of the immigration parameter, with the fitted model predicting a species-abundance distribution close to the observed distribution. Our results complement those using sampling formulae that specify the multivariate probability distribution of species abundances from neutral models. [ABSTRACT FROM AUTHOR]