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Compared to other regions, the drivers of diversification in Africa are poorly understood. We studied a radiation of insects with over 100 species occurring in a wide range of habitats across the Afrotropics to investigate the fundamental evolutionary processes and geological events that generate and maintain patterns of species richness on the continent. By investigating the evolutionary history of Bicyclus butterflies within a phylogenetic framework, we inferred the group’s origin at the Oligo-Miocene boundary from ancestors in the Congolian rainforests of central Africa. Abrupt climatic fluctuations during the Miocene (ca. 19–17 Ma) likely fragmented ancestral populations, resulting in at least eight early-divergent lineages. Only one of these lineages appears to have diversified during the drastic climate and biome changes of the early Miocene, radiating into the largest group of extant species. The other seven lineages diversified in forest ecosystems during the late Miocene and Pleistocene when climatic conditions were more favorable—warmer and wetter. Our results suggest changing Neogene climate, uplift of eastern African orogens, and biotic interactions have had different effects on the various subclades of Bicyclus, producing one of the most spectacular butterfly radiations in Africa. [Afrotropics; biodiversity; biome; biotic interactions; Court Jester; extinction; grasslands; paleoclimates; Red Queen; refugia forests; dependent-diversification; speciation.]

Aim Rates of colonization, speciation and extinction determine species richness and endemism in insular systems. The general dynamic model of island biogeography (GDM) predicts that speciation and extinction rates depend on island area and elevation via their control on ecological limits to diversification and therefore covary with an island's geological history. Additionally, the colonization rate may increase with area and elevation through the 'target effect', which can be mediated by reduced 'environmental filtering'. Here we test whether the area and depth expansion of an island-like ecosystem, a lake, caused a shift in diversity dynamics. Location Lake Biwa, Japan, whose geological history and biota are well documented. Taxon Fishes. Methods We extended the phylogenetic island biogeography framework DAISIE (Dynamic Assembly of Island biota through Speciation, Immigration and Extinction) to accommodate time-shifts in macroevolutionary rates and in carrying capacity. Using phylogenetic information on colonization and speciation times for the complete Lake Biwa fish community (70 taxa), we tested for a shift in macroevolutionary assembly rates and reconstructed the temporal diversity trajectory in the lake. We assessed the power to identify a shift through a wide range of scenarios and benchmarked against simulated fossil records. Results We detected an increase in colonization rate of fishes at 0.2 million years ago (Ma), with limited support for the existence of ecological limits. The reconstructed diversity trajectory was close to a source-sink equilibrium diversity prior to the shift and remained well below a new shift-driven elevated equilibrium thereafter. We found sufficient power to identify an increase in colonization rate up to 1.5 Ma, whereas extinction concealed the signal of earlier shifts. Main conclusions The fish diversity dynamics of Lake Biwa show a response to changes in area and depth and phylogenies carry a signature of these changes. The detected increase in colonization rate following Lake Biwa's expansion, elevating the fish diversity, is unlikely due to a predicted increase in ecological limits feeding back on colonization rate. We therefore call for (additional) explanations: the target effect, whereby larger islands attract more species, and reduced environmental filtering due to higher habitat diversity associated with increased lake area/depth.

Low-elevation regions harbour the majority of the world's species diversity compared to high-elevation areas. This global gradient suggests that lowland species have had more time to diversify, or that net diversification rates have been higher in the lowlands. However, highlands seem to be cradles of diversity as they contain many young endemics, suggesting that their rates of speciation are exceptionally fast. Here we use a phylogenetic diversification model that accounts for the dispersal of species between different elevations to examine the evolutionary dynamics of the elevational diversity gradient in passerine birds, a group that has radiated globally to occupy almost all elevations and latitudes. We find strong support for a model in which passerines diversify at the same rate in the highlands and the lowlands but in which the per-capita rate of dispersal from high to low elevations is more than twice as fast as that in the reverse direction. This suggests that while there is no consistent trend in diversification across elevations, part of the diversity generated by highland regions migrates into the lowlands, thus setting up the observed gradient in passerine diversity. We find that this process drives tropical regions but for temperate areas, the analysis could be hampered by their lower richness. Despite their lower diversity, highland regions are disproportionally important for maintaining diversity in the adjacent lowlands.

The latitudinal diversity gradient (LDG) is one of Earth’s most iconic biodiversity patterns and still one of the most debated. Explanations for the LDG are often categorized into three broad pathways in which the diversity gradient is created by (1) differential diversification rates, (2) differential carrying capacities (ecological limits), or (3) differential time to accumulate species across latitude. Support for these pathways has, however, been mostly verbally expressed. Here, we present a minimal model to clarify the essential assumptions of the three pathways and explore the sensitivity of diversity dynamics to these pathways. We find that an LDG arises most easily from a gradient in ecological limits compared with a gradient in the time for species accumulation or diversification rate in most modeled scenarios. Differential diversification rates create a stronger LDG than ecological limits only when speciation and dispersal rates are low, but then the predicted LDG seems weaker than the observed LDG. Moreover, range dynamics may reduce an LDG created by a gradient in diversification rates or time for species accumulation, but they cannot reduce an LDG induced by differential ecological limits. We conclude that our simple model provides a null prediction for the effectiveness of the three LDG pathways and can thus aid discussions about the causal mechanisms underlying the LDG or motivate more complex models to confirm or falsify our findings., The American Naturalist, 194 (5), ISSN:0003-0147, ISSN:1537-5323

Incomplete or geographically biased sampling poses significant problems for research in phylogeography, population genetics, phylogenetics, and species delimitation. Despite the power of using genome-wide genetic markers in systematics and related fields, approaches such as the multispecies coalescent remain unable to easily account for unsampled lineages. The Empidonax difficilis / E. occidentalis complex of small tyrannid flycatchers (Aves: Tyrannidae) is a classic example of a widely distributed species with limited phenotypic geographic variation that was broken into two largely cryptic (or sibling) lineages following extensive study. Though the group is well-characterized north of the U.S. Mexico border, the evolutionary distinctiveness and phylogenetic relationships of southern populations remain obscure. In this paper, we use dense genomic and geographic sampling across the majority of the range of the E. difficilis / E . occidentalis complex to assess whether current taxonomy and species limits reflect underlying evolutionary patterns, or whether they are an artifact of historically biased or incomplete sampling. We find that additional samples from Mexico render the widely recognized species-level lineage E. occidentalis paraphyletic, though it retains support in the best-fit species delimitation model from clustering analyses. We further identify a highly divergent unrecognized lineage in a previously unsampled portion of the group's range, which a cline analysis suggests is more reproductively isolated than the currently recognized species E. difficilis and E. occidentalis. Our phylogeny supports a southern origin of these taxa. Our results highlight the pervasive impacts of biased geographic sampling, even in well-studied vertebrate groups like birds, and illustrate what is a common problem when attempting to define species in the face of recent divergence and reticulate evolution.

Biodiversity accumulates hierarchically by means of ecological and evolutionary processes and feedbacks. Within ecological communities drift, dispersal, speciation, and selection operate simultaneously to shape patterns of biodiversity. Reconciling the relative importance of these is hindered by current models and inference methods, which tend to focus on a subset of processes and their resulting predictions. Here we introduce Massive Eco-evolutionary Synthesis Simulations (MESS), a unified mechanistic model of community assembly, rooted in classic island biogeography theory, which makes temporally explicit joint predictions across three biodiversity data axes: i) species richness and abundances; ii) population genetic diversities; and iii) trait variation in a phylogenetic context. Using simulations we demonstrate that each data axis captures information at different timescales, and that integrating these axes enables discriminating among previously unidentifiable community assembly models. MESS is unique in generating predictions of community-scale genetic diversity, and in characterizing joint patterns of genetic diversity, abundance, and trait values. MESS unlocks the full potential for investigation of biodiversity processes using multi-dimensional community data including a genetic component, such as might be produced by contemporary eDNA or metabarcoding studies. We combine with supervised machine learning to fit the parameters of the model to real data and infer processes underlying how biodiversity accumulates, using communities of tropical trees, arthropods, and gastropods as case studies that span a range of data availability scenarios, and spatial and taxonomic scales., Funding was provided by grants from FAPESP (BIOTA, 2013/50297-0 to MJH and AC Carnaval), the Synthesis Centre of iDiv (DFG FZT 118), NASA through the Dimensions of Biodiversity Program (DOB 1343578) and the National Science Foundation (DEB-1253710 to MJH; DEB 1745562 to AC Carnaval; DBI 1927319 to AJR). IO was supported by the Mina Rees Dissertation Fellowship in the Sciences provided by the Graduate Center of the City University of New York. MR was supported by the Bioinformatics and Computational Biology Fellowship through the Institute for Bioinformatics and Evolutionary Studies at the University of Idaho. AJR was supported by the Santa Fe Institute Omidyar Fellowship. JR was supported by fellowships from the Natural Environment Research Council (NERC) (NE/I021179, NE/L011611/1). RSE was supported by an NWO-VICI grant. This work is a contribution to Imperial College’s Grand Challenges in Ecosystems and the Environment initiative, through JR.

Vascular epiphytes form a diverse group of almost 30 000 species, yet theory concerning their community structure is still largely lacking. We therefore employed the simplest models of biodiversity, (near-)neutral models, to generate hypotheses concerning their community structure. With recently developed tools for (near-)neutral models we analyzed species abundance data from many samples in Central and South America which we divided into four metacommunities (Mesoamerica, Central America, Amazonia and Paraná), where for each metacommunity we considered two subsets differing in dispersal syndrome: an animal-dispersed guild and a wind-dispersed guild. We considered three models differing in the underlying speciation mode. Across all metacommunities, we found observed patterns to be indistinguishable from patterns generated by neutral or near-neutral processes. Furthermore, we found that subdivision in different dispersal guilds was often supported, with recruitment limitation being stronger for animal-dispersed species than for wind-dispersed species. This is the first time that (near-)neutral theory has been applied to epiphyte communities. Future efforts with additional data sets and more refined models are expected to further improve our understanding of community structure in epiphytes and will have to test the generality of our findings.

Aim Island biogeography theory describes how island size and isolation determine population colonization success. Large islands sustain larger populations than small ones and experience less demographic stochasticity, thus a lower extinction risk. Nearby islands are more likely to be colonized than distant ones, because they receive more immigrants from the mainland. However, local conditions on islands are often different from those on the mainland; therefore, populations on recently colonized islands also need to adapt. Island size and isolation are known to impact the build-up of genetic variation necessary for adaptation; hence, we integrated island biogeography with evolution experimentally to gain a better understanding of the roles of island size and isolation in biodiversity patterns. Location Laboratory, Ghent University, Belgium. Time period October 2013 to June 2014. Major taxa studied Two-spotted spider mite (Tetranychus urticae). Methods Using experimental evolution, we studied the effects of island size and isolation on colonization, extinction and adaptation of the two-spotted spider mite to new islands. The mainland population consisted of bean plants and the islands of tomato plants (a known challenging condition). Islands differed in their size (number of plants) and in the number of immigrants (females, the dispersive stage) they received from the mainland. Results Island size and dispersal decreased extinction risk and increased colonization success and adaptation. Populations on small islands, which are most affected by extinction, were rescued demographically by an increase in dispersal. However, they were never able to adapt. Main conclusions Evolutionary rescue via dispersal is possible only when populations are sufficiently large; small populations cannot adapt, because they lack the genetic variation necessary for local adaptation. Hence, in addition to the effects of island size and dispersal on the ecological processes of colonization and extinction, our results show that island size and dispersal can jointly affect the evolutionary process of adaptation to novel habitats.

Colonization, speciation and extinction are dynamic processes that influence global patterns of species richness1,2,3,4,5,6. Island biogeography theory predicts that the contribution of these processes to the accumulation of species diversity depends on the area and isolation of the island7,8. Notably, there has been no robust global test of this prediction for islands where speciation cannot be ignored9, because neither the appropriate data nor the analytical tools have been available. Here we address both deficiencies to reveal, for island birds, the empirical shape of the general relationships that determine how colonization, extinction and speciation rates co-vary with the area and isolation of islands. We compiled a global molecular phylogenetic dataset of birds on islands, based on the terrestrial avifaunas of 41 oceanic archipelagos worldwide (including 596 avian taxa), and applied a new analysis method to estimate the sensitivity of island-specific rates of colonization, speciation and extinction to island features (area and isolation). Our model predicts—with high explanatory power—several global relationships. We found a decline in colonization with isolation, a decline in extinction with area and an increase in speciation with area and isolation. Combining the theoretical foundations of island biogeography7,8 with the temporal information contained in molecular phylogenies10 proves a powerful approach to reveal the fundamental relationships that govern variation in biodiversity across the planet.

The unified neutral theory of biodiversity and biogeography has gained the status of a quantitative null model for explaining patterns in ecological (meta)communities. The theory assumes that individuals of trophically similar species are functionally equivalent. We empirically evaluate the relative contribution of neutral and deterministic processes in shaping fruit‐feeding butterfly assemblages in three tropical forests in Africa, using both direct (confronting the neutral model with species abundance data) and indirect approaches (testing the predictions of neutral theory using data other than species abundance distributions). Abundance data were obtained by sampling butterflies using banana baited traps set at the forest canopy and understorey strata. Our results indicate a clear consistency in the kind of species or species groups observed at either the canopy or understorey in the three studied communities. Furthermore, we found significant correlation between some flight‐related morphological traits and species abundance at the forest canopy, but not at the understorey. Neutral theory's contribution to explaining our data lies largely in identifying dispersal limitation as a key process regulating fruit‐feeding butterfly community structure. Our study illustrates that using species abundance data alone in evaluating neutral theory can be informative, but is insufficient. Species‐level information such as habitat preference, host plants, geographical distribution, and phylogeny is essential in elucidating the processes that regulate biodiversity community structures and patterns.

Summary Half a century ago, MacArthur and Wilson proposed that the number of species on islands tends toward a dynamic equilibrium diversity around which species richness fluctuates [1]. The current prevailing view in island biogeography accepts the fundamentals of MacArthur and Wilson's theory [2] but questions whether their prediction of equilibrium can be fulfilled over evolutionary timescales, given the unpredictable and ever-changing nature of island geological and biotic features [3–7]. Here we conduct a complete molecular phylogenetic survey of the terrestrial bird species from four oceanic archipelagos that make up the diverse Macaronesian bioregion—the Azores, the Canary Islands, Cape Verde, and Madeira [8, 9]. We estimate the times at which birds colonized and speciated in the four archipelagos, including many previously unsampled endemic and non-endemic taxa and their closest continental relatives. We develop and fit a new multi-archipelago dynamic stochastic model to these data, explicitly incorporating information from 91 taxa, both extant and extinct. Remarkably, we find that all four archipelagos have independently achieved and maintained a dynamic equilibrium over millions of years. Biogeographical rates are homogeneous across archipelagos, except for the Canary Islands, which exhibit higher speciation and colonization. Our finding that the avian communities of the four Macaronesian archipelagos display an equilibrium diversity pattern indicates that a diversity plateau may be rapidly achieved on islands where rates of in situ radiation are low and extinction is high. This study reveals that equilibrium processes may be more prevalent than recently proposed, supporting MacArthur and Wilson's 50-year-old theory.

Molecular phylogenies have been increasingly recognized as an important source of information on species diversification. For many models of macroevolution, analytical likelihood formulas have been derived to infer macroevolutionary parameters from phylogenies. A few years ago, a general framework to numerically compute such likelihood formulas was proposed, which accommodates models that allow speciation and/or extinction rates to depend on diversity. This framework calculates the likelihood as the probability of the diversification process being consistent with the phylogeny from the root to the tips. However, while some readers found the framework presented in Etienne et al. (Proc R Soc Lond B Biol Sci 279(1732):1300–1309, 2012) convincing, others still questioned it (personal communication), despite numerical evidence that for special cases the framework yields the same (i.e., within double precision) numerical value for the likelihood as analytical formulas do that were independently derived for these special cases. Here we prove analytically that the likelihoods calculated in the new framework are correct for all special cases with known analytical likelihood formula. Our results thus add substantial mathematical support for the overall coherence of the general framework.

The study of biodiversity started as a single unified field that spanned both ecology and evolution and both macro and micro phenomena. But over the 20th century, major trends drove ecology and evolution apart and pushed an emphasis towards the micro perspective in both disciplines. Macroecology and macroevolution re-emerged as self-consciously distinct fields in the 1970s and 1980s, but they remain largely separated from each other. Here, we argue that despite the challenges, it is worth working to combine macroecology and macroevolution. We present 25 fundamental questions about biodiversity that are answerable only with a mixture of the views and tools of both macroecology and macroevolution., This paper is a joint effort of the working group sEcoEvo, kindly supported by sDiv, the Synthesis Centre of the German Centre for Integrative Biodiversity Research (iDiv), Halle‐Jena‐Leipzig, funded by the German Research Foundation (FZT 118). Brian McGill acknowledges US Department of Agriculture Hatch grant to Maine Agricultural and Forestry Experimental Station #1011538 and National Science Foundation Advances in Biological Infrastructure grant #1660000.

This thesis is about Life and how it is unevenly distributed across time, space and hierarchical levels. When comparing the fossil record with the current diversity of life, the extent of the change in composition of flora and fauna over time is enormous, where some groups reigned in the past but have only few delegates surviving to the present. Spatial variation in the number of organisms is also astonishing, with the exuberance of jungles contrasting with plain arid ecosystems, one of many gradients of diversity. With a more detailed examination, one can also find examples of groups of animals or plants which have tremendously flourished producing endless forms while other groups have no more than a handful of members. In this thesis, I developed methods based on simulations and likelihood calculations. They include predictive theoretical models, and methods that are applied to case studies varying in taxonomic scope (songbirds; fish) and geographic scale (worldwide elevational distribution; lake depth gradient). I show how the spatial distribution of species is dynamic over time and has important consequences for patterns of species origination and extinction. I show that species do not have the same rates of diversification, and this can be the result of 1) regional equilibrium or out-of-equilibrium dynamics or 2) spatial distribution across gradients.

Islands are at the frontline of the anthropogenic extinction crisis [1]. A vast number of island birds have gone extinct since human colonization [2], and an important proportion is currently threatened with extinction [3]. While the number of lost or threatened avian species has often been quantified [4], the macroevolutionary consequences of human impact on island biodiversity have rarely been measured [5]. Here, we estimate the amount of evolutionary time that has been lost or is under threat due to anthropogenic activity in a classic example, New Zealand. Half of its bird taxa have gone extinct since humans arrived [6, 7] and many are threatened [8], including lineages forming highly distinct branches in the avian tree of life [9-11]. Using paleontological and ancient DNA information, we compiled a dated phylogenetic dataset for New Zealand's terrestrial avifauna. We extend the method DAISIE developed for island biogeography [12] to allow for the fact that many of New Zealand's birds are evolutionarily isolated and use it to estimate natural rates of speciation, extinction, and colonization. Simulating under a range of human-induced extinction scenarios, we find that it would take approximately 50 million years (Ma) to recover the number of species lost since human colonization of New Zealand and up to 10 Ma to return to today's species numbers if currently threatened species go extinct. This study puts into macroevolutionary perspective the impact of humans in an isolated fauna and reveals how conservation decisions we take today will have repercussions for millions of years.

Aim: Islands are often hotspots of endemism due to their isolation, making colonization a rare event and hence facilitating allopatric speciation. Dispersal usually occurs between nearby locations according to a stepping-stone model. We aimed to reconstruct colonization and speciation processes in an endemic-rich system of land-based islands that does not seem to follow the obvious stepping-stone model of dispersal.Location: Five land-based habitat archipelagos of limestone outcrops in the flood-plain of the Kinabatangan River in Sabah, Malaysian Borneo.Methods: We studied the phylogeography of three species complexes of endemic land snails, using multiple genetic markers. We calculated genetic distances between populations, applied beast2 to reconstruct phylogenies for each taxon and subsequently reconstructed ancestral ranges using 'BioGeoBEARS'.Results: We found spatial-genetic structure among nearby locations to be highly pronounced for each taxon. Genetic correlation was present at small spatial scales only and disappeared at distances of 5 km and above. Most archipelagos have been colonized from within the region multiple times over the past three million years, in 78% of the cases as a result of long-distance dispersal (LDD) or dispersal from non-adjacent limestone outcrops. The flow of the main geographical feature within the region, the Kinabatangan River, did not play a role.Main conclusions: Phylogeographic structure in these Bornean land snails has only partly been determined by small-scale dispersal, where it leads to isolation-by-distance, but mostly by LDD. Our results demonstrate that island endemic taxa only very locally follow a simple stepping-stone model, whilst dispersal to non-adjacent islands and especially LDD, is most important. This leads to the formation of highly localized, isolated " endemic populations" forming the onset of a complex radiation of endemic species.

4 figuras, 1 tabla, The interactions between plants and arbuscular mycorrhizal fungi (AMF) maintain a crucial link between macroscopic organisms and the soil microbial world. These interactions are of extreme importance for the diversity of plant communities and ecosystem functioning. Despite this importance, only recently has the structure of plant-AMF interaction networks been studied. These recent studies, which used genetic data, suggest that these networks are highly structured, very similar to plant-animal mutualistic networks. However, the assembly process of plant- AMF communities is still largely unknown, and an important feature of plant-AMF interactions has not been incorporated: they occur at an extremely localized scale. Studying plant-AMF networks in a spatial context seems therefore a crucial step. This paper studies a plant-AMF spatial co-occurrence network using novel methodology based on information theory and a unique set of spatially explicit species-level data. We apply three null models of which only one accounts for spatial effects. We find that the data show substantial departures from null expectations for the two non-spatial null models. However, for the null model considering spatial effects, there are few significant co-occurrences compared with the other two null models. Thus, plant-AMF spatial co-occurrences seem to be mostly explained by stochasticity, with a small role for other factors related to plant-AMF specialization. Furthermore, we find that the network is not significantly nested or modular. We conclude that this plant-AMF spatial co-occurrence network lacks substantial structure and, therefore, plants and AMF species do not track each other over space. Thus, random encounters seem more important in the first step of the assembly of plant-AMF communities., R.S. Etienne and E.R. Chang share last authorship We would like to thank Roger Guimerá for providing us with the simulated annealing algorithm to estimate network modularity, the Natural Sciences and Engineering Research Council of Canada (NSERC) for a post-doctoral fellowship awarded to ERC, the Spanish "Ministerio de Economía y Competitividad" under the project SITES (CGL2012-39964) to DA and the Netherlands Organization for Scientific Research (NWO) for financial assistance through a VIDI grant awarded to RSE.

The role of ecological limits in regulating the distribution and diversification of species remains controversial. Although such limits must ultimately arise from constraints on local species coexistence, this spatial context is missing from most macroevolutionary models. Here, we develop a stochastic, spatially explicit model of species diversification to explore the phylogenetic and biogeographic patterns expected when local diversity is bounded. We show how local ecological limits, by regulating opportunities for range expansion and thus rates of speciation and extinction, lead to temporal slowdowns in diversification and predictable differences in equilibrium diversity between regions. However, our models also show that even when regions have identical diversity limits, the dynamics of diversification and total number of species supported at equilibrium can vary dramatically depending on the relative size of geographic and local ecological niche space. Our model predicts that small regions with higher local ecological limits support a higher standing diversity and more balanced phylogenetic trees than large geographic areas with more stringent constraints on local coexistence. Our findings highlight how considering the spatial context of diversification can provide new insights into the role of ecological limits in driving variation in biodiversity across space, time, and clades.

This research was funded by the Netherlands Organisation for Scientific Research VENI grant 863.13.003 (to A.L.P.), NASA Biodiversity grant NNX11AP72G and NSF grants NSF DBI 1262600, DBI 0960550 and DEB 1026764 (to W.J.), the Oxford Clarendon Fund and US-UK Fulbright Commission (to C.S.), and the John Fell Fund and NERC grant NE/I028068/1 (to J.A.T.). Ecological communities are assembled from the overlapping of species in geographic space, but the mechanisms facilitating or limiting such overlaps are difficult to resolve. Here, we combine phylogenetic, morphological and environmental data to model how multiple processes regulate the origin and maintenance of geographic range overlap across 1,115 pairs of avian sister species globally. We show that coexistence cannot be adequately predicted by either dispersal-assembly (that is, biogeographic) models or niche-assembly models alone. Instead, our results overwhelmingly support an integrated model with different assembly processes dominating at different stages of coexistence. The initial attainment of narrow geographic overlap is dictated by intrinsic dispersal ability and the time available for dispersal, whereas wider coexistence is largely dependent on niche availability, increasing with ecosystem productivity and divergence in niche-related traits, and apparently declining as communities become saturated with species. Furthermore, although coexistence of any individual pair of species is highly stochastic, we find that integrating assembly processes allows broad variation in the incidence and extent of coexistence to be predicted with reasonable accuracy. Our findings demonstrate how phylogenetic data coupled with environmental factors and functional traits can begin to clarify the multi-layered processes shaping the distribution of biodiversity at large spatial scales. Postprint

International audience; Past global change may have forced animal-dispersed plants with megafaunal fruits to adapt or go extinct, but these processes have remained unexplored at broad spatio-temporal scales. Here, we combine phylogenetic, distributional and fruit size data for more than 2500 palm (Arecaceae) species in a time-slice diversification analysis to quantify how extinction and adaptation have changed over deep time. Our results indicate that extinction rates of palms with megafaunal fruits have increased in the New World since the onset of the Quaternary (2.6 million years ago). In contrast, Old World palms show a Quaternary increase in transition rates towards evolving small fruits from megafaunal fruits. We suggest that Quaternary climate oscillations and concurrent habitat fragmentation and defaunation of megafaunal frugivores in the New World have reduced seed dispersal distances and geographical ranges of palms with megafaunal fruits, resulting in their extinction. The increasing adaptation to smaller fruits in the Old World could reflect selection for seed dispersal by ocean-crossing frugivores (e.g. medium-sized birds and bats) to colonize Indo-Pacific islands against a background of Quaternary sea-level fluctuations. Our macro-evolutionary results suggest that megafaunal fruits are increasingly being lost from tropical ecosystems, either due to extinctions or by adapting to smaller fruit sizes.

A contemporary goal in both ecology and evolutionary biology is to develop theory that transcends the boundary between the two disciplines, to understand phenomena that cannot be explained by either field in isolation. This is challenging because macroevolution typically uses lineage-based models, whereas ecology often focuses on individual organisms. Here, we develop a new parsimonious individual-based theory by adding mild selection to the neutral theory of biodiversity. We show that this model generates realistic phylogenies showing a slowdown in diversification and also improves on the ecological predictions of neutral theory by explaining the occurrence of very common species. Moreover, we find the distribution of individual fitness changes over time, with average fitness increasing at a pace that depends positively on community size. Consequently, large communities tend to produce fitter species than smaller communities. These findings have broad implications beyond biodiversity theory, potentially impacting, for example, invasion biology and paleontology.

Time-calibrated phylogenies of living species have been widely used to study the tempo and mode of species diversification. However, it is increasingly clear that inferences about species diversification - extinction rates in particular - can be unreliable in the absence of paleontological data. We introduce a general framework based on the fossilized birth-death process for studying speciation-extinction dynamics on phylogenies of extant and extinct species. The model assumes that phylogenies can be modeled as a mixture of distinct evolutionary rate regimes and that a hierarchical Poisson process governs the number of such rate regimes across a tree. We implemented the model in BAMM, a computational framework that uses reversible jump Markov chain Monte Carlo to simulate a posterior distribution of macroevolutionary rate regimes conditional on the branching times and topology of a phylogeny. The implementation we describe can be applied to paleontological phylogenies, neontological phylogenies, and to phylogenies that include both extant and extinct taxa. We evaluate performance of the model on datasets simulated under a range of diversification scenarios. We find that speciation rates are reliably inferred in the absence of paleontological data. However, the inclusion of fossil observations substantially increases the accuracy of extinction rate estimates. We demonstrate that inferences are relatively robust to at least some violations of model assumptions, including heterogeneity in preservation rates and misspecification of the number of occurrences in paleontological datasets.

A focus of ecology is to determine drivers of community assembly. Here, we investigate effects of immigration and species sorting (environmental selection) on structuring aquatic bacterial communities in both colonised and previously uncolonised environments. We used nonsterilised and presterilised water from three chemically distinct ponds to establish microcosms, which were opened for 12, 24, 48, 96 or 167h and then closed again to allow airborne bacterial immigration and subsequent succession. Community similarity, richness, evenness and the parameters of a neutral model were investigated after 167h. Immigration appeared to govern the assembly of communities in the presterilised water as there were no significant differences in evenness among microcosm communities containing water from each pond. Statistical estimation of neutral model parameters confirmed these findings, because the estimated immigration rate changed significantly with time of exposure to immigration. Species sorting also occurred because significant differences in community similarity (for presterilised and nonsterilised communities) and evenness (only for nonsterilised communities) were detected among microcosms containing different pond water; the magnitude of these differences was greater for communities in nonsterilised microcosms. Our study provides evidence for both processes being important during the colonisation of aquatic environments and presents a novel way to apply the neutral model.

Islands are ideal systems to model temporal changes in biodiversity and reveal the influence of humans on natural communities.Although theory predicts biodiversity on islands tends towards an equilibrium value, the recent extinction of large proportionsof island biotas complicates testing this model. The well-preserved subfossil record of Caribbean bats—involving multipleinsular radiations—provides a rare opportunity to model diversity dynamics in an insular community. Here, we reconstruct thediversity trajectory in noctilionoid bats of the Greater Antilles by applying a dynamic model of colonization, extinction andspeciation to phylogenetic and palaeontological data including all known extinct and extant species. We show species richnessasymptotes to an equilibrium value, a demonstration of natural equilibrium dynamics across an entire community. However,recent extinctions—many caused by humans—have wiped out nearly a third of island lineages, dragging diversity away fromequilibrium. Using a metric to measure island biodiversity loss, we estimate it will take at least eight million years to regainpre-human diversity levels. Our integrative approach reveals how anthropogenic extinctions can drastically alter the naturaltrajectory of biological communities, resulting in evolutionary disequilibrium.

Quantifying the role of biodiversity in ecosystems not only requires understanding the links between species and the ecological functions and services they provide, but also how these factors relate to measurable indices, such as functional traits and phylogenetic diversity. However, these relationships remain poorly understood, especially for heterotrophic organisms within complex ecological networks. Here, we assemble data on avian traits across a global sample of mutualistic plant–frugivore networks to critically assess how the functional roles of frugivores are associated with their intrinsic traits, as well as their evolutionary and functional distinctiveness. We find strong evidence for niche complementarity, with phenotypically and phylogenetically distinct birds interacting with more unique sets of plants. However, interaction strengths—the number of plant species dependent on a frugivore—were unrelated to evolutionary or functional distinctiveness, largely because distinct frugivores tend to be locally rare, and thus have fewer connections across the network. Instead, interaction strengths were better predicted by intrinsic traits, including body size, gape width and dietary specialization. Our analysis provides general support for the use of traits in quantifying species ecological functions, but also highlights the need to go beyond simple metrics of functional or phylogenetic diversity to consider the multiple pathways through which traits may determine ecological processes.

The association between species richness and ecosystem energy availability is one of the major geographic trends in biodiversity. It is often explained in terms of energetic constraints, such that coexistence among competing species is limited in low productivity environments. However, it has proven challenging to reject alternative views, including the null hypothesis that species richness has simply had more time to accumulate in productive regions, and thus the role of energetic constraints in limiting coexistence remains largely unknown. We use the phylogenetic relationships and geographic ranges of sister species (pairs of lineages who are each other’s closest extant relatives) to examine the association between energy availability and coexistence across an entire vertebrate class (Aves). We show that the incidence of coexistence among sister species increases with overall species richness and is elevated in more productive ecosystems, even when accounting for differences in the evolutionary time available for coexistence to occur. Our results indicate that energy availability promotes species coexistence in closely related lineages, providing a key step toward a more mechanistic understanding of the productivity–richness relationship underlying global gradients in biodiversity., An analysis of phylogenetic and geographic data across the world's bird species reveals that the probability of species coexistence is constrained by the availability of energy in the environment., Author Summary The increase in the number of species with the availability of energy in the environment is one of the most general but least understood patterns in global biodiversity. The finite amount of energy flowing through an ecosystem has long been suspected to place a fundamental constraint on the ability of species to subdivide ecological resources, with greater energy availability—and, thus, ecosystem productivity—in the tropics, potentially facilitating increased levels of coexistence. However, empirical support for this hypothesis has been lacking, raising the possibility that richness is higher in productive ecosystems for largely historical reasons, including the greater geological age and area of tropical biomes, which increases the period of time available for diversity to accumulate. By combining phylogenetic and geographic data from across the world’s bird species, we show that greater ecosystem productivity is associated with an increased probability of coexistence among closely related lineages and that this pattern contributes to the higher species richness in the tropics. Our results confirm that contemporary gradients in species richness are fundamentally shaped by energetic constraints on coexistence.

aeres : ACL; International audience; In this article we propose a new framework for studying adaptive radiations in the context of diversity-dependent diversification. Diversity dependence causes diversification to decelerate at the end of an adaptive radiation but also plays a key role in the initial pulse of diversification. In particular, key innovations (which in our definition include novel traits as well as new environments) may cause decoupling of the diversity-dependent dynamics of the innovative clade from the diversity-dependent dynamics of its ancestral clade. We present a likelihood-based inference method to test for decoupling of diversity dependence using molecular phylogenies. The method, which can handle incomplete phylogenies, identifies when the decoupling took place and which diversification parameters are affected. We illustrate our approach by applying it to the molecular phylogeny of the North American clade of the legume tribe Psoraleeae (47 extant species, of which 4 are missing). Two diversification rate shifts were previously identified for this clade; our analysis shows that the first, positive shift can be associated with decoupling of two Pediomelum subgenera from the other Psoraleeae lineages, while we argue that the second, negative shift can be attributed to speciation being protracted. The latter explanation yields nonzero extinction rates, in contrast to previous findings. Our framework offers a new perspective on macroevolution: new environments and novel traits (ecological opportunity) and diversity dependence (ecological limits) cannot be considered separately.

Adaptive radiation is the rapid diversification of a single lineage into many species that inhabit a variety of environments or use a variety of resources and differ in traits required to exploit these. Why some lineages undergo adaptive radiation is not well-understood, but filling unoccupied ecological space appears to be a common feature. We construct a complete, dated, species-level phylogeny of the endemic Vangidae of Madagascar. This passerine bird radiation represents a classic, but poorly known, avian adaptive radiation. Our results reveal an initial rapid increase in evolutionary lineages and diversification in morphospace after colonizing Madagascar in the late Oligocene some 25 Mya. A subsequent key innovation involving unique bill morphology was associated with a second increase in diversification rates about 10 Mya. The volume of morphospace occupied by contemporary Madagascan vangas is in many aspects as large (shape variation)—or even larger (size variation)—as that of other better-known avian adaptive radiations, including the much younger Galapagos Darwin's finches and Hawaiian honeycreepers. Morphological space bears a close relationship to diet, substrate use, and foraging movements, and thus our results demonstrate the great extent of the evolutionary diversification of the Madagascan vangas.

Phylogenetic trees show a remarkable slowdown in the increase of number of lineages towards the present, a phenomenon which cannot be explained by the standard birth–death model of diversification with constant speciation and extinction rates. The birth–death model instead predicts a constant or accelerating increase in the number of lineages, which has been called the pull of the present. The observed slowdown has been attributed to nonconstancy of the speciation and extinction rates due to some form of diversity dependence (i.e., species-level density dependence), but the mechanisms underlying this are still unclear. Here, we propose an alternative explanation based on the simple concept that speciation takes time to complete. We show that this idea of “protracted” speciation can be incorporated in the standard birth–death model of diversification. The protracted birth–death model predicts a realistic slowdown in the rate of increase of number of lineages in the phylogeny and provides a compelling fit to four bird phylogenies with realistic parameter values. Thus, the effect of recognizing the generally accepted fact that speciation is not an instantaneous event is significant; even if it cannot account for all the observed patterns, it certainly contributes substantially and should therefore be incorporated into future studies.

Aims Neutral theory consists of a suite of models that assume ecological equivalence among individual organisms. They have been most commonly applied to tropical forest tree communities either as null models or as approximations. Neutral models typically only include reproductive adults; therefore, fitting to empirical tree community data requires defining a reproductive-size threshold, which for trees is usually set arbitrarily to a diameter at breast height (DBH) of 100 mm. The inevitable exclusion of some reproductive adults and inclusion of some saplings cause a non-random sampling bias in neutral model fits. Here, we investigate this problem and resolve it by introducing simple age structure into a neutral model. Methods We compared the performance and sensitivity of DBH threshold of three variants of a spatially explicit neutral model: the traditional model, a model incorporating random sampling and a model with two distinct age classes—reproductive adults and saplings. In the age-structured model, saplings are offspring from adults that disperse according to a Gaussian dispersal kernel around the adults. The only extra parameter is the ratio of adults to saplings, which is not a free parameter but directly measurable. We used species–area relationships (SARs) to explore the predicted effect of saplings on the species richness at different scales in our model. We then evaluated the three model variations to find the parameters required to maintain the observed level of species richness in the 50-ha plot on Barro Colorado Island (BCI). We repeated our analysis filtering the data at different minimum tree-size thresholds in order to find the effect this threshold has on our results. Lastly, we used empirical species–individual relationships (SIRs) to test the pre-existing hypothesis that environmental filtering is the primary cause of differences between the assemblage of saplings and that of adults on BCI. Important Findings Our age-structured neutral model was characterized by SARs that were insensitive to the presence of saplings at large scales and highly sensitive to them at small scales. Both models without age structure were highly sensitive to the DBH threshold chosen in a way that could not be explained based on random samplings alone. The age-structured neutral model, which allowed for non-random sampling based on life stage, was consistent with species richness observations. Our analysis of empirical SIRs did not support environmental filtering as a dominant force, but it did show evidence for other differences between age classes. Age can now be easily incorporated into future studies of neutral models whenever there is a concern that a sample is not entirely composed of reproductive adult individuals. More generally, we suggest that modeling studies using tree data subject to a minimum size threshold should consider the sensitivity of their results to that threshold.

International audience; Entropy maximization (EM, also known as MaxEnt) is a general inference procedure that originated in statistical mechanics. It has been applied recently to predict ecological patterns, such as species abundance distributions and species-area relationships. It is well known in physics that the EM result strongly depends on how elementary configurations are described. Here we argue that the same issue is also of crucial importance for EM applications in ecology. To illustrate this, we focus on the EM prediction of species-level spatial abundance distributions. We show that the EM outcome depends on (1) the choice of configuration set, (2) the way constraints are imposed, and (3) the scale on which the EM procedure is applied. By varying these choices in the EM model, we obtain a large range of EM predictions. Interestingly, they correspond to spatial abundance distributions that have been derived previously from mechanistic models. We argue that the appropriate choice of the EM model assumptions is nontrivial and can be determined only by comparison with empirical data.

The Cape region of South Africa is a hotspot of flowering plant biodiversity. However, the reasons why levels of diversity and endemism are so high remain obscure. Here, we reconstructed phylogenetic relationships among species in the genus Protea, which has its center of species richness and endemism in the Cape, but also extends through tropical Africa as far as Eritrea and Angola. Contrary to previous views, the Cape is identified as the ancestral area for the radiation of the extant lineages: most species in subtropical and tropical Africa are derived from a single invasion of that region. Moreover, diversification rates have been similar within and outside the Cape region. Migration out of the Cape has opened up vast areas, but those lineages have not diversified as extensively at fine spatial scales as lineages in the Cape. Therefore, higher net rates of diversification do not explain the high diversity and endemism of Protea in the Cape. Instead, understanding why the Cape is so diverse requires an explanation for how Cape species are able to diverge and persist at such small spatial scales.

The species abundance distribution (SAD) is one of the few universal patterns in ecology. Research on this fundamental distribution has primarily focused on the study of numerical counts, irrespective of the traits of individuals. Here we show that considering a set of Generalized Species Abundance Distributions (GSADs) encompassing several abundance measures, such as numerical abundance, biomass and resource use, can provide novel insights into the structure of ecological communities and the forces that organize them. We use a taxonomically diverse combination of macroecological data sets to investigate the similarities and differences between GSADs. We then use probability theory to explore, under parsimonious assumptions, theoretical linkages among them. Our study suggests that examining different GSADs simultaneously in natural systems may help with assessing determinants of community structure. Broadening SADs to encompass multiple abundance measures opens novel perspectives in biodiversity research and warrants future empirical and theoretical developments.

Understanding the processes that underlie biodiversity requires insight into the evolutionary history of the taxa involved. Accurate estimation of speciation, extinction, and diversification rates is a prerequisite for gaining this insight. Here, we develop a stochastic birth-death model of speciation and extinction that predicts the probability distribution of both extinct and extant numbers of species in a clade. We present two estimation methods based on this model given data on the number of extinct species (from the fossil record) and extant species (from diversity assessments): a multivariate method of moments approach and a maximum-likelihood approach. We show that, except for some special cases, the two estimation methods produce very similar estimates. This is convenient, because the usually preferred maximum-likelihood approach is much more computationally demanding, so the method of moments can serve as a proxy. Furthermore, we introduce a correction for possible bias that can arise by the mere fact that we will normally only consider extant clades. We find that in some cases the bias correction affects the estimates profoundly. Finally, we show how our model can be extended to incorporate incomplete preservation. Preservation rates can, however, not be reliably estimated on the basis of numbers of extant and extinct species alone.

It is widely believed that the neutral theory of biodiversity cannot be used for parameter inference if the assumption of neutrality is not met. The goal of this work is to extend this neutral framework to quantify the intensity of recruitment limitation (limited dispersal plus environmental filtering) in natural species assemblages. We model several local communities as part of a larger metacommunity, and we assume that neutrality holds in each local community, but not in the metacommunity. The immigration rate m does not only reflect dispersal limitation into a given local community, but also the intensity of environmental filtering. We develop a novel statistical method to infer the immigration parameter m in each local community. Using simulated datasets, we show that m indeed depends on both dispersal limitation and on the intensity of environmental filtering. We then apply this method to a network of tropical tree plots in central Panama. Inferred recruitment rates m were positively correlated with the fraction of trees dispersed by mammals, and with annual rainfall, possibly due to a weaker environmental filtering as rainfall increases. Finally, m, as estimated from trees greater than 1 cm trunk diameter, were significantly larger than an estimation based on trees greater than 10 cm trunk diameter. This suggests a cumulative effect of environmental filtering upon trees throughout their ontogeny.

As the utility of the neutral theory of biodiversity is increasingly being recognized, there is also an increasing need for proper tools to evaluate the relative importance of neutral processes (dispersal limitation and stochasticity). One of the key features of neutral theory is its close link to data: sampling formulas, giving the probability of a data set conditional on a set of model parameters, have been developed for parameter estimation and model comparison. However, only single local samples can be handled with the currently available sampling formulas, whereas data are often available for many small spatially separated plots. Here, I present a sampling formula for multiple, spatially separated samples from the same metacommunity, which is a generalization of earlier sampling formulas. I also provide an algorithm to generate data sets with the model and I introduce a general test of neutrality that does not require an alternative model; this test compares the probability of the observed data (calculated using the new sampling formula) with the probability of model-generated data sets. I illustrate this with tree abundance data from three large Panamanian neotropical forest plots. When the test is performed with model parameters estimated from the three plots, the model cannot be rejected; however, when parameter estimates previously reported for BCI are used, the model is strongly rejected. This suggests that neutrality cannot explain the structure of the three Panamanian tree communities on the local (BCI) and regional (Panama Canal Zone) scale simultaneously. One should be aware, however, that aspects of the model other than neutrality may be responsible for its failure. I argue that the spatially implicit character of the model is a potential candidate.

Tropical mountains are hot spots of biodiversity and endemism,but the evolutionary origins of their unique biotas are poorlyunderstood. In varying degrees, local and regional extinction,long-distance colonization, and local recruitment may all contribute to the exceptional character of these communities. Also, it isdebated whether mountain endemics mostly originate from locallowland taxa, or from lineages that reach the mountain by long-range dispersal from cool localities elsewhere. Here we investigatethe evolutionary routes to endemism by sampling an entire tropical mountain biota on the 4,095-metre-high Mount Kinabalu inSabah, East Malaysia. We discover that most of its unique biodiversity is younger than the mountain itself (6 million years), andcomprises a mix of immigrant pre-adapted lineages and descendants from local lowland ancestors, although substantial shiftsfrom lower to higher vegetation zones in this latter group wererare. These insights could improve forecasts of the likelihood ofextinction and ‘evolutionary rescue in montane biodiversity hotspots under climate change scenarios.

International audience; Over the past decade, the neutral theory of biodiversity has stirred up community assembly theory considerably by suggesting that stochasticity in the form of ecological drift is an important factor determining community composition and community turnover. The neutral theory assumes that all species within a community are functionally equivalent (the neutrality assumption), and therefore applies best to communities of trophically similar species. Evidently, trophically similar species may still differ in dispersal ability, and therefore may not be completely functionally equivalent. Here we present a new sampling formula that takes into account the partitioning of a community into two guilds that differ in immigration rate. We show that, using this sampling formula, we can accurately detect a subdivision into guilds from species abundance distributions, given ecological data about dispersal ability. We apply our sampling formula to tropical tree data from Barro Colorado Island, Panama. Tropical trees are divided depending on their dispersal mode, where biotically dispersed trees are grouped as one guild, and abiotically dispersed trees represent another guild. We find that breaking neutrality by adding guild structure to the neutral model significantly improves the fit to data and provides a better understanding of community assembly on BCI. Our findings are thus an important step towards an integration of neutral and niche theory.

There has recently been a surge of literature examining microbial invasions into a variety of environments. These studies often include a component of biological diversity as a major factor determining an invader's fate, yet common results are rarely cross-compared. Since many studies only present a snapshot of the entire invasion process, a bird's eye view is required to piece together the entire continuum, which we find consists of introduction, establishment, spread, and impact phases. We further examine the patterns and mechanisms associated with invasion resistance and create a mechanistic synthesis governed by the species richness, species evenness, and resource availability of resident communities. We conclude by exploring the advantages of using a theoretical invasion framework across different fields.

Neutral community theory explains biodiversity, i.e. the coexistence of several species, as the result of a stochastic balance between immigration and extinction on a local level, and between speciation and extinction on a regional level. The most popular model, presented by Hubbell in 2001, has seen many analytical developments in recent years, which can be used in model analysis, model testing and model comparison. We review these developments here, and present alternative derivations and shine previously unnoticed lights on them.

Ongoing large-scale habitat disturbance requires quick identification of conservation priorities such as targeting sites rich in species and/or endemics. Biodiversity assessments are time consuming and expensive, so surveys often rely on partial sampling. Optimal use should be made of all currently available sources of information, but methodological differences between surveys hamper direct comparison. Because diversity depends on spatial scale, diversity characteristics of different sites are best compared on the basis of species-area relationships. As a result of the incompleteness of sampling, the observed species-area relationship deviates from the "true" species-area relationship.In this paper, we identify five key factors affecting the shape of the species-area relationship due to incomplete sampling: (1) the total spatial extent of the observations, (2) the spatial distribution of the observations, (3) the proportion of the total extent sampled, (4) the proportion of the individuals in the sampled area included in the survey, and (5) the proportion of the included individuals successfully identified. We outline how methodologically different surveys can be combined to optimize the use of existing data in the evaluation of conservation needs, particularly for tropical forests.As an illustration, we analyzed four methodologically different botanical surveys in the same area of old growth lowland forest in South Cameroon with the aim of reconciling these surveys. The four surveys were (1) reconnaissance scale vegetation mapping, (2) detailed botanical assessment (all individuals), (3) incomplete botanical assessment (10% individuals), and (4) herbarium collections.By correcting for the five key factors we were able to match the results of the four different biodiversity surveys. The five key factors affected the recorded number of species and endemics differently; partial sampling of extent (3) and individuals (4) and partial identification of individuals (5) were the three most important factors.We conclude that reconciliation of biodiversity assessments is possible if the differences between methods can be accounted for. We advocate reliable documentation of survey methods, especially the five key factors, because it greatly enhances the potential of combining methodologically different surveys for comparative biodiversity analyses.

Under allopatric speciation models, a key step in the build-up of species richness is population dispersal leading to the co-occurrence of previously geographically isolated forms. Despite its central importance for community assembly, the extent to which the transition from spatial segregation (allopatry or parapatry) to coexistence (sympatry) is a predictable process, or alternatively one governed by chance and the vagaries of biogeographic history, remains poorly understood. Here, we use estimated divergence times and current patterns of geographical range overlap among sister species to explore the evolution of sympatry in vertebrates. We show that rates of transition to sympatry vary predictably according to ecology, being faster in marine or strongly dispersive terrestrial clades. This association with organism vagility is robust to the relative frequency of geographical speciation modes and consistent across taxonomic scales and metrics of dispersal ability. These findings reject neutral models of dispersal assembly based simply on evolutionary age and are not predicted by the main alternative view that range overlap is primarily constrained by biotic interactions. We conclude that species differences in dispersal limitation are fundamental in organizing the assembly of ecological communities and shaping broad-scale patterns of biodiversity over space and time.

22 páginas, 4 tablas, 4 figuras, Species abundance distributions (SAD) are probably ecology’s most well-known empirical pattern, and over the last decades many models have been proposed to explain their shape. There is no concensus over which model is correct, because the degree to which different processes can be discerned from SAD patterns has not yet been rigorously quantified. We present a power calculation to quantify our ability to detect deviations from neutrality using species abundance data. We study non-neutral stochastic community models, and show that the presence of non-neutral processes is detectable if sample size is large enough and/or the amplitude of the effect is strong enough. Our framework can be used for any candidate community model that can be simulated on a computer, and determines both the sampling effort required to distinguish between alternative processes, and a range for the strength of non-neutral processes in communities whose patterns are statistically consistent with neutral theory. We find that even data sets of the scale of the 50 Ha forest plot on Barro Colorado Island, Panama, are unlikely to be large enough to detect deviations from neutrality caused by competitive interactions alone, though the presence of multiple non-neutral processes with contrasting effects on abundance distributions may be detectable., OAH, DA, and SJC were supported by the National Environment Research Council (grant no. NE/H007458/1). DA is supported by grant CGL2012- 39964 and Ramon y Cajal program from the Spanish Government. RSE thanks the Netherlands Organisation for Scientific Research (NWO) for funding through VIDI and VICI grants. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

A major goal of island biogeography is to understand how island communities are assembled over time. However, we know little about the influence of variable area and ecological opportunity on island biotas over geological timescales. Islands have limited life spans, and it has been posited that insular diversity patterns should rise and fall with an island's ontogeny. The potential of phylogenies to inform us of island ontogenetic stage remains unclear, as we lack a phylogenetic framework that focuses on islands rather than clades. Here, we present a parsimonious island-centric model that integrates phylogeny and ontogeny into island biogeography and can incorporate a negative feedback of diversity on species origination. This framework allows us to generate predictions about species richness and phylogenies on islands of different ages. We find that peak richness lags behind peak island area, and that endemic species age increases with island age on volcanic islands. When diversity negatively affects rates of immigration and cladogenesis, our model predicts speciation slowdowns on old islands. Importantly, we find that branching times of in situ radiations can be informative of an island's ontogenetic stage. This novel framework provides a quantitative means of uncovering processes responsible for island biogeography patterns using phylogenies.

Gorgona Island, the major insular area in the Colombian Pacific Ocean, is characterized by a remarkably high biological and ecosystem diversity for this area of the world. Coral reefs are well developed and their fish communities have been described using conventional visual surveys. These methods, however, are known to be biased towards detecting larger and more mobile species, tending to ignore small and cryptobenthic species. The two main objectives of this study were to describe the assemblage structure of the cryptobenthic fish fauna and estimate the extent to which this fauna is underestimated by visual surveys.At the beginning and the end of the warm season, we compared the cryptobenthic fish assemblage recorded using visual surveys against the one recorded using “enclosed anesthetic/rotenone samples” on isolated coral colonies (N=54 beginning of warm season; N=17 end of warm season). The crypthobenthic fish fauna associated to coral colonies was characterized by small body sizes and was composed mainly by species of the families Antennaridae, Blennidae, Gobiidae, Labrisomidae, Muraenidae, Serranidae, Scorpaenidae and Syngnathidae. Conventional visual surveys underestimated overall species richness by 28-36% and number of individuals by 16-35%. Noteworthy, four species recorded during this study using “enclosed anesthetic/rotenone samples” were new records for Gorgona Island. Although both sampling methods can detect a largely overlapping group of species, the “enclosed anesthetic/rotenone samples” method was able to detect more individuals and species, including several species that visual surveys fail to detect. Although this study is the first effort to describe the cryptobenthic fish assemblage associated to coral reefs in the Tropical Eastern Pacific, our results suggest that these assemblages are an important component of the reef fish community in the region in terms of biodiversity and functional roles. Rev. Biol. Trop. 62 (Suppl. 1): 359-371. Epub 2014 February 01. Métodos de muestreo convencionales (censos visuales) usados en el estudio de peces arrecifales han llevado a una constante subestimación de la riqueza de especies, ya que estos tienden a ignorar especies pequeñas y crípticas. En este estudio describimos la estructura del ensamblaje de peces criptobentónicos en Isla Gorgona, Colombia y estimamos el grado en que este ensamblaje es subestimado al usar censos visuales. Al comienzo y al final de la estación cálida comparamos la capacidad de detección de censos visuales vs “muestreos cerrados con anestésico/rotenona” (N=54 comienzo; N=17 final). La fauna criptobentónica se caracterizó por mantener individuos de tamaños corporales pequeños, pertenecientes principalmente a las familias Antennaridae, Blennidae, Gobiidae, Labrisomidae, Muraenidae, Serranidae, Scorpaenidae and Syngnathidae. Los censos visuales subestimaron la riqueza de especies en un 28-36% y el número de individuos en un 16-35%. Aunque un gran número de especies son detectadas usando ambos métodos, los “muestreos cerrados con anestésico/rotenona” detectan un mayor rango de especies. Este estudio, el primer esfuerzo para describir el ensamblaje de peces criptobentónicos asociados a arrecifes coralinos en Colombia, sugiere que la fauna criptobentónica puede ser un componente importante de la comunidad arrecifal en términos de biodiversidad y papel funcional.

International audience; The roles of species richness, resource use, and resource availability are central to many hypotheses explaining the diversityinvasion phenomenon but are generally not investigated together. Here, we created a large diversity gradient of soil microbial communities by either assembling communities of pure bacterial strains or removing the diversity of a natural soil. Using data on the resource-use capacities of the soil communities and an invader that were gathered from 71 carbon sources, we quantified the niches available to both constituents by using the metrics community niche and remaining niche available to the invader. A strong positive relationship between species richness and community niche across both experiments indicated the presence of resource complementarity. Moreover, community niche and the remaining niche available to the invader predicted invader abundance well. This suggested that increased competition in communities of higher diversity limits community invasibility and underscored the importance of resource availability as a key mechanism through which diversity hinders invasions. As a proof of principle, we subjected selected invaded communities to a resource pulse, which progressively uncoupled the link between soil microbial diversity and invasion and allowed the invader to rebound after nearly being eliminated in some communities. Our results thus show that (1) resource competition suppresses invasion, (2) biodiversity increases resource competition and decreases invasion through niche preemption, and (3) resource pulses that cannot be fully used, even by diverse communities, are favorable to invasion.

Plant-pollinator interactions are among the best known and ubiquitous plant-animal mutualisms and are crucial for ecosystem functioning and the maintenance of biodiversity. Most pollinators are insects with several life-stages (e.g. egg, larva, pupa, adult) and the mutualistic interaction depends on the pollinator surviving these different life-stages. However, to our knowledge, pollinator population structure has been ignored in most theoretical models of plant-pollinator dynamics, and we lack understanding of the role of different life-stages in determining the stability of the mutualism. Here we therefore develop a simple plant-pollinator model with a facultative plant and an obligate pollinator with stage-structure. Our model predicts a globally stable equilibrium when pollinator demography is dominated by adults and a locally stable equilibrium when the plants are strongly dependent on pollination and pollinator demography is dominated by the larval stage. In the latter case, the mutualism is vulnerable to fluctuations in the pollinator population size or structure caused by external factors (e.g. pesticides) reducing larval development and increasing adult mortality. This may cause a sudden collapse rather than gradual decrease of the mutualism, after which the pollination service cannot be recovered by reducing these detrimental external factors, but must be accompanied by large increases in pollinator populations. This highlights the importance of considering population structure in plant-pollinator interactions. (C) 2014 Elsevier Ltd. All rights reserved.

A popular line of research in evolutionary biology is the use of time-calibrated phylogenies for the inference of diversification processes. This requires computing the likelihood of a given ultrametric tree as the reconstructed tree produced by a given model of diversification. Etienne & Rosindell (2012) proposed a lineage-based model of diversification, called protracted speciation, where species remain incipient during a random duration before turning good species, and showed that this can explain the slowdown in lineage accumulation observed in real phylogenies. However, they were unable to provide a general likelihood formula. Here, we present a likelihood formula for protracted speciation models, where rates at which species turn good or become extinct can depend both on their age and on time. Our only restrictive assumption is that speciation rate does not depend on species status. Our likelihood formula utilizes a new technique, based on the contour of the phylogenetic tree and first developed in Lambert (2010). We consider the reconstructed trees spanned by all extant species, by all good extant species, or by all representative species, which are either good extant species or incipient species representative of some good extinct species. Specifically, we prove that each of these trees is a coalescent point process, that is, a planar, ultrametric tree where the coalescence times between two consecutive tips are independent, identically distributed random variables. We characterize the common distribution of these coalescence times in some, biologically meaningful, special cases for which the likelihood reduces to an elegant analytical formula or becomes numerically tractable., 27 pages, 5 figures

Ricklefs and Renner (Reports, 27 January 2012, p. 464) showed correlations of species richness and individual abundance within families across continents and claimed that neutral theory predicts no such correlation. However, they did not substantiate this claim quantitatively with a neutral model. Here, we show that neutral theory can be consistent with these correlations and, consequently, that the correlations alone cannot reject neutrality.

Most empirical and theoretical studies have shown that sex increases the rate of evolution, although evidence of sex constraining genomic and epigenetic variation and slowing down evolution also exists. Faster rates with sex have been attributed to new gene combinations, removal of deleterious mutations, and adaptation to heterogeneous environments. Slower rates with sex have been attributed to removal of major genetic rearrangements, the cost of finding a mate, vulnerability to predation, and exposure to sexually transmitted diseases. Whether sex speeds or slows evolution, the connection between reproductive mode, the evolutionary rate, and species diversity remains largely unexplored. Here we present a spatially explicit model of ecological and evolutionary dynamics based on DNA sequence change to study the connection between mutation, speciation, and the resulting biodiversity in sexual and asexual populations. We show that faster speciation can decrease the abundance of newly formed species and thus decrease long-term biodiversity. In this way, sex can reduce diversity relative to asexual populations, because it leads to a higher rate of production of new species, but with lower abundances. Our results show that reproductive mode and the mechanisms underlying it can alter the link between mutation, evolutionary rate, speciation and biodiversity and we suggest that a high rate of evolution may not be required to yield high biodiversity., Author Summary The role of sex in driving genetic variation and the speed at which new species emerge has been debated for over a century. There is experimental and theoretical evidence that sex increases genetic variation and the speed at which new species emerge, although evidence that sex reduces variation and slows the formation of new species also exists. Surprisingly, given the link between sex and genetic variation, little work has been done on the impact of sex on biodiversity. In the present theoretical study we show that a faster evolutionary rate can decrease the abundance of newly formed species and thus decrease long-term biodiversity. This leads to the paradoxical result that sexual reproduction can increase genetic variation but reduce species diversity. These results suggest that reducing the rate of appearance of genetic variation and the speed at which new species emerge may increase biodiversity in the long-term. This unexpected link between reproductive mode, the speed of evolution and biodiversity suggests that a high evolutionary rate may not be required to yield a large number of species in natural ecosystems.