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bioRxiv2018

Evolution of the latitudinal diversity gradient in the hyperdiverse ant genus Pheidole

AimThe latitudinal diversity gradient is the dominant pattern of life on Earth, but a consensus understanding of its origins has remained elusive. The analysis of recently diverged, hyper-rich invertebrate groups provides an opportunity to investigate latitudinal patterns with the statistical power of large trees while minimizing potentially confounding variation in ecology and history. Here, we synthesize global phylogenetic and macroecological data on a hyperdiverse (>1100 species) ant radiation, Pheidole, and evaluate the roles of three general explanations for the latitudinal gradient: variation in diversification rate, tropical conservatism, and ecological regulation.\n\nLocationGlobal.\n\nTime PeriodThe past 35 million years.\n\nMajor taxa studiedThe hyperdiverse ant genus Pheidole Westwood.\n\nMethodsWe assembled geographic data for 1499 species and morphospecies, and inferred a dated phylogeny of Pheidole of 449 species, including 150 species newly sequenced for this study. We tested correlations between diversification rate and latitude with BAMM, HiSSE, GeoSSE, and FiSSE, and examined patterns of diversification as Pheidole spread around the globe.\n\nResultsWe found that Pheidole diversification occurred in series of bursts when new continents were colonized, followed by a slowdown in each region. There was no evidence of systematic variation of net diversification rates with latitude across any of the methods. Additionally, we found latitudinal affinity is moderately conserved with a Neotropical ancestor and phylogenetic inertia alone is sufficient to produce the gradient pattern.\n\nMain ConclusionsOverall our results are consistent with tropical conservatism explaining the diversity gradient, while providing no evidence that diversification rate varies systematically with latitude. There is evidence of ecological regulation on continental scales through the pattern of diversification after colonization. These results shed light on the mechanisms underlying the diversity gradient, while contributing toward a much-needed invertebrate perspective on global biodiversity dynamics.

evolutionary biology↗

bioRxiv2018

Applying generalised allometric regressions to predict live body mass of tropical and temperate arthropods

1. The ecological implications of body size extend from the biology of individual organisms to ecosystem-level processes. Measuring body mass for high numbers of invertebrates can be logistically challenging, making length-mass regressions useful for predicting body mass with minimal effort. However, standardised sets of scaling relationships covering a large range in body length, taxonomic groups, and multiple geographical regions are scarce.\n\n2. We collected 6293 arthropods from 19 higher-level taxa in both temperate and tropical locations to compile a comprehensive set of linear models relating live body mass to a range of predictor variables. For each individual, we measured live weight (hereafter, body mass), body length and width, and conducted linear regressions to predict body mass using body length, body width, taxonomic group and geographic region. Additionally, we quantified prediction error when using parameters from arthropods of a different geographic region.\n\n3. Incorporating body width into taxon- and region-specific length-mass regressions yielded the highest prediction accuracy for body mass. Using regression parameters from a different geographic location increased prediction error, causing over- or underestimation of body mass depending on geographical origin and whether body width was included.\n\n4. We present a comprehensive range of parameters for predicting arthropod body mass and provide guidance for selecting optimal scaling relationships. Given the importance of body mass for functional invertebrate ecology and a paucity of adequate regressions to predict arthropod body mass from different geographical regions, our study provides a long-needed resource for quantifying live body mass in invertebrate ecology research.

zoology↗

bioRxiv2018

Taxa-driven functional shifts associated with stormflow in an urban stream microbial community

Urban streams are susceptible to stormwater and sewage inputs that can impact their ecological health and water quality. Microbial communities in streams play important functional roles and their composition and metabolic potential can help assess ecological state and water quality. Although these environments are highly heterogenous, little is known about the influence of isolated perturbations, such as those resulting from rain events on urban stream microbiota. Here, we examined the microbial community composition and diversity in an urban stream during dry and wet weather conditions with both 16S rRNA gene sequencing across multiple years and shotgun metagenomics to more deeply analyze a single stormflow event. Metagenomics was used to assess population-level dynamics as well as shifts in the microbial community taxonomic profile and functional potential before and after a substantial rainfall. Results demonstrated general trends present in the stream under stormflow vs. baseflow conditions across years and seasons and also highlighted the significant influence of increased effluent flow following rain in shifting the stream microbial community from abundant freshwater taxa to those more associated with urban/anthropogenic settings. Shifts in the taxonomic composition were also linked to changes in functional gene content, particularly for transmembrane transport and organic substance biosynthesis. We also observed an increase in relative abundance of genes encoding degradation of organic pollutants and antibiotic resistance after rain. Overall, this study provided evidence of stormflow impacts on an urban stream microbiome from an environmental and public health perspective.\n\nImportanceUrban streams in various parts of the world are facing increased anthropogenic pressure on their water quality, and stormflow events represent one such source of complex physical, chemical and biological perturbations. Microorganisms are important components of these streams from both ecological and public-health perspectives, and analyzing the effect of such perturbations on the stream microbial community can help improve current knowledge on the impact such chronic disturbances can have on these water resources. This study examines microbial community dynamics during rain-induced stormflow conditions in an urban stream of the Chicago Area Waterway System. Additionally, using shotgun metagenomics we identified significant shifts in the microbial community composition and functional gene content following a high rainfall event, with potential environment and public health implications. Previous work in this area has been limited to specific genes/organisms or has not assessed immediate stormflow impact.

microbiology↗

bioRxiv2018

Fresh Snowfall Microbiology and Chemistry are Driven by Geography in Storm-Tracked Events

Snowfall is a global phenomenon highly integrated with hydrology and ecology. Forays into studying bioaerosols and their dependence on aeolian movement are largely constrained to either precipitation-independent analyses or in-silico models. Though snowpack and glacial microbiological studies have been conducted, little is known about the biological component of meteoric snow. Through culture-independent phylogenetic and geochemical analyses, we show that the geographical location at which snow precipitates determines snowfalls geochemical and microbiological composition. Storm-tracking, furthermore, can be used as a valuable environmental indicator to trace down what factors are influencing bioaerosols. We estimate annual deposits of up to ~10 kg of bacterial / archaeal biomass per hectare along our study area of the eastern Front Range in Colorado. The dominant kinds of microbiota captured in an analysis of seven snow events at two different locations, one urban, one rural, across the winter of 2016/2017 included phyla Proteobacteria, Bacteroidetes, Firmicutes and Acidobacteria, though a multitude of different kinds of organisms were found in both. Taxonomically, Bacteroidetes were more abundant in Golden (urban plain) snow while Proteobacteria were more common in Sunshine (rural mountain) samples. Chemically, Golden snowfall was positively correlated with some metals and anions. The work also hints at better informing the everything is everywhere hypotheses of the microbial world and that atmospheric transport of microbiota is not only common, but is capable of disseminating vast amounts of microbiota of different physiologies and genetics that then affect ecosystems globally. Snowfall, we conclude, is a significant repository of microbiological material with strong implications for both ecosystem genetic flux and general bio-aerosol theory.\n\nImportanceSnowfall is commonplace to the temperate and polar regions of the world. As an interface between the atmosphere, hydrosphere and earth, snow is responsible for high annual deposits of moisture globally, and, can serve as a water bank in the form of both permanent snow fields and glaciers. Essential to general ecosystem function, snow can also be considered a transporter of aerosolized material. Given the magnitude of microbiota deposited by snowfall, which we report, it is likely that biological material within snowfall, with its geochemical underpinning-- and the associated genetic banks--have significant downstream ecological effects.\n\nUnderstanding what is contained in snowfall becomes especially urgent in a warming climate where high-impact meteorological and ecological changes are imminent and likely. With climate-induced changes to snowfall patterns, surface ecosystems are likely to be impacted by ensuing changes in microbiota deposition. Thus, the ecosystem function of soils, rock and surface waters are also likely to be impacted; these changes, in turn, greatly influence agriculture, weathering and infrastructure.

microbiology↗

bioRxiv2018

DivNet: Estimating diversity in networked communities

Diversity is a marker of ecosystem health in ecology, microbiology and immunology, with implications for disease diagnosis and infection resistance. However, accurately comparing diversity across environmental gradients is challenging, especially when number of different taxonomic groups in the community is large. Furthermore, existing approaches to estimating diversity do not perform well when the taxonomic groups in the community interact via an ecological network, such as by competing within their niche, or with mutualistic relationships. To address this, we propose DivNet, a method for estimating within- and between-community diversity in ecosystems where taxa interact via an ecological network. In particular, accounting for network structure permits more accurate estimates of alpha- and beta-diversity, even in settings with a large number of taxa and a small number of samples. DivNet is fast, accurate, precise, performs well with large numbers of taxa, and is robust to both weakly and strongly networked communities. We show that the advantages of incorporating taxon interactions into diversity estimation are especially clear in analyzing microbiomes and other high-diversity, strongly networked ecosystems. Therefore, to illustrate the method, we analyze the microbiome of seafloor basalts based on a 16S amplicon sequencing dataset with 1490 taxa and 13 samples.

bioinformatics↗

bioRxiv2018

Diversity of active viral infections within the Sphagnum microbiome

Sphagnum-dominated peatlands play an important role in global carbon storage and represent significant sources of economic and ecological value. While recent efforts to describe microbial diversity and metabolic potential of the Sphagnum microbiome have demonstrated the importance of its microbial community, little is known about the viral constituents. We used metatranscriptomics to describe the diversity and activity of viruses infecting microbes within the Sphagnum peat bog. The vegetative portions of 6 Sphagnum plants were obtained from a peatland in northern Minnesota and total RNA extracted and sequenced. Metatranscriptomes were assembled and contigs screened for the presence of conserved virus marker genes. Using bacteriophage capsid protein, gp23, as a marker for phage diversity, we identified 33 contigs representing undocumented phage s that were active in the community at the time of sampling. Similarly, RNA-dependent RNA polymerase and the Nucleo-Cytoplasmic Large DNA Virus (NCLDV) major capsid protein were used as markers for ssRNA viruses and NCLDV, respectively. In total 114 contigs were identified as originating from undescribed ssRNA viruses, 22 of which represent near-complete genomes. An additional 64 contigs were identified as being from NCLDVs. Finally, 7 contigs were identified as putative virophage or polinto-like viruses. We developed co-occurrence networks with these markers in relation to the expression of potential-host housekeeping gene rpb1 to predict virus-host relationships, identifying 13 groups. Together, our approach offers new tools for the identification of virus diversity and interactions in understudied clades, and suggest viruses may play a considerable role in the ecology of the Sphagnum microbiome.\n\nSignificanceSphagnum-dominated peatlands play an important role in maintaining atmospheric carbon dioxide levels by modifying conditions in the surrounding soil to favor its own growth over other plant species. This slows rates of decomposition and facilitates the accumulation of fixed carbon in the form of partially decomposed biomass. The unique environment produced by Sphagnum enriches for the growth of a diverse microbial consortia that benefit from and support the mosss growth, while also maintaining the hostile soil conditions. While a growing body of research has begun to characterize the microbial groups that colonize Sphagnum, little is currently known about the ecological factors that constrain community structure and define ecosystem function. Top-down population control by viruses is almost completely undescribed. This study provides insight into the significant viral influence on the Sphagnum microbiome, and identifying new potential model systems to study virus-host interactions in the peatland ecosystem.

microbiology↗

bioRxiv2018

Local adaptation and sexual selection drive intra-island diversification in a songbird lineage: differential divergence in autosomes and sex chromosomes.

Recently diverged taxa showing marked phenotypic and ecological diversity are optimal systems to test the relative importance of two major evolutionary mechanisms, adaptation to local ecological conditions by natural selection, or mechanisms of reproductive isolation such as assortative mating mediated by sexually selected mating signals or post-zygotic incompatibilities. Whereas local adaptation is expected to affect many loci throughout the genome, traits acting as mating signals are expected to be located on sex chromosomes and have a simple genetic basis. We used genome-wide markers to test these predictions in Reunion Islands gray-white eye (Zosterops borbonicus), which has recently diversified into five distinct plumage forms. Two of them correspond to a polymorphic highland population that is separated by a steep ecological gradient from three distinct lowland forms that show narrow contact zones in plumage color traits, yet no association with environmental variables. An analysis of population structure using genome-wide SNP loci revealed two major clades corresponding to highland and lowland forms, respectively, with the latter separated further into three independent lineages corresponding to plumage forms. Coalescent tests of alternative demographic scenarios provided support for divergence of highland and lowland lineages with an intensification of gene flow in the last 60,000 years. Landscapes of genomic variation revealed that signatures of selection associated with elevation are found at multiple regions across the genome, whereas most loci associated with the lowland forms are located on the Z sex chromosome. A gene ontology analysis identified TYRP1, a Z-linked color gene, as a likely candidate locus underlying color variation among lowland forms. Our results are consistent with the role of natural selection in driving the divergence of locally adapted highland populations, and the role of sexual selection in differentiating lowland forms through reproductive isolation mechanisms, showing that both modes of lineage divergence can take place at very small geographic scales in birds.

evolutionary biology↗

bioRxiv2018

Environmental disturbances compromise synthetic symbiosis

By virtue of complex interactions, the behaviour of mutualistic systems is difficult to study and nearly impossible to predict. We have developed a theoretical model of a modifiable experimental yeast system that is amenable to exploring self-organised cooperation while considering the production and use of specific metabolites. Leveraging the simplicity of an artificial yeast system, a simple model of mutualism, we develop and test the assumptions and stability of this theoretical model. We examine how one-off, recurring and permanent changes to an ecological niche affect a cooperative interaction and identify an ecological \"Goldilocks zone\" in which the mutualism can survive. Moreover, we explore how a factor like the cost of mutualism - the cellular burden of cooperating - influences the stability of mutualism and how environmental changes shape this stability. Our results highlight the fragility of mutualisms and suggest the use of synthetic biology to stave off an ecological collapse.

evolutionary biology↗

bioRxiv2018

Interacting effects of the abiotic and biotic environment on fitness of rainforest Drosophila

Competition within and between species can have large effects on fitness and may therefore drive local adaptation. However, these effects are rarely tested systematically, or considered when predicting species responses to environmental change. We used a field transplant experiment to test the effects of intra and interspecific competition on fitness across the ecological niches of two rainforest Drosophila species that replace each other along an elevation gradient. For the species with the broader elevational range, we also tested for adaptation to the local abiotic and biotic environment. In both species, intraspecific competition reduced productivity more than interspecific competition at the centre of its elevational range, while interspecific competition had a stronger effect at the range edge, where the competing species is more abundant. Local adaptation was detected in the centre of the range of the more widespread species, but only in the presence of intraspecific competition. This study is the first to demonstrate that fitness effects of inter-specific competition increase at ecological margins, while intra-specific competition has more pervasive effects at range centres. This is a key assumption of "tangled bank" models of community evolution and has important implications for predicting the resilience of ecological networks to global change.

evolutionary biology↗

bioRxiv2018

Thermal selection drives biodiversity origination across the Atlantic/Indian Ocean boundary

Intraspecific genetic structure in widely distributed marine species often mirrors the boundaries between temperature-defined bioregions. This suggests that the same thermal gradients that maintain distinct species assemblages also drive the evolution of new biodiversity. Ecological speciation scenarios are often invoked to explain such patterns, but the fact that adaptation is usually only identified when phylogenetic splits are already evident makes it impossible to rule out the alternative scenario of allopatric speciation with subsequent adaptation. We integrated large-scale genomic and environmental datasets along one of the worlds best defined marine thermal gradients (the South African coastline) to test the hypothesis that incipient speciation in the sea is due to divergence linked to the thermal environment. We identified temperature-associated gene regions in a coastal fish species that is spatially homogeneous throughout several temperature-defined biogeographical regions on the basis of selectively neutral markers. Based on these gene regions, the species is divided into geographically distinct regional populations. Importantly, the ranges of these populations are delimited by the same ecological boundaries that define distinct infraspecific genetic lineages in co-distributed marine the species, and biogeographical disjunctions in species assemblages. Our results indicate that ecologically-mediated selection represents an early stage of marine speciation in coastal regions that lack physical dispersal barriers.

evolutionary biology↗

bioRxiv2018

Deeply conserved susceptibility in a multi-host, multi-parasite system

Variation in susceptibility is ubiquitous in multi-host, multi-parasite assemblages, and can have profound implications for ecology and evolution. The extent to which susceptibility is phylogenetically conserved among hosts is poorly understood and has rarely been appropriately tested. We screened for haemosporidian parasites in 3983 birds representing 40 families and 523 species, spanning ~4500 meters elevation in the tropical Andes. To quantify the influence of host phylogeny on infection status, we applied Bayesian phylogenetic multilevel models that included a suite of environmental, spatial, temporal, life history, and ecological predictors. We found evidence of deeply-conserved susceptibility across the avian tree; host phylogeny explained substantial variation in infection rate, and results were robust to phylogenetic uncertainty. Our study suggests that susceptibility is governed, in part, by conserved, latent aspects of anti-parasite defense. This demonstrates the importance of deep phylogeny for understanding the outcomes of present-day ecological interactions.\n\nStatement of authorshipLNB, SMM, NM, and CCW designed the study; SMM, SCG, HLL, HS, TV, JDW, and CCW collected the data; LNB and NM analyzed the data; LNB, NM, and CCW wrote the paper with input from all authors.\n\nData accessibility statementSpecimen information is available from the Arctos database (arctosdb.org) and in supplementary tables (Appendix S1). Files used for analysis will be archived in Dryad. DOI: XXX.

evolutionary biology↗

bioRxiv2018

Time calibrated morpho-molecular classification of Nassellaria (Radiolaria)

Nassellaria are marine radiolarian protists belonging to the Rhizaria lineage. Their skeleton, made of opaline silica, exhibit an excellent fossil record, extremely valuable in micro-paleontological studies for paleo-environmental reconstruction. Yet, to date very little is known about the extant diversity and ecology of Nassellaria in contemporary oceans, and most of it is inferred from their fossil record. Here we present an integrative classification of Nassellaria based on taxonomical marker genes (18S and 28S ribosomal DNA) and morphological characteristics obtained by optical and scanning electron microscopy imaging. Our phylogenetic analyses distinguished 11 main morpho-molecular clades relying essentially on the overall morphology of the skeleton and not on internal structures as previously considered. Using fossil calibrated molecular clock we estimated the origin of Nassellaria among radiolarians primitive forms in the Devonian (ca. 420 Ma), that gave rise to living nassellarian groups in the Triassic (ca. 250 Ma), during the biggest diversification event over their evolutionary history. This morpho-molecular framework provides both a new morphological classification easier to identify under light microscopy and the basis for future molecular ecology surveys. Altogether, it brings a new standpoint to improve our scarce understanding of the ecology and worldwide distribution of extant nassellarians.

molecular biology↗

bioRxiv2018

The influence of fluctuating population densities on evolutionary dynamics

The causes and consequences of fluctuating population densities are an important topic in ecological literature. Yet, the effects of such fluctuations on maintenance of variation in spatially structured populations have received little analytic treatment. We analyze what happens when two habitats coupled by migration not only differ in their trade-offs in selection but also in their demographic stability - and show that equilibrium allele frequencies can change significantly due to ecological feedback arising from locally fluctuating population sizes. When an ecological niche exhibits such fluctuations, these drive an asymmetry in the relative impact of gene flow, and therefore, the equilibrium frequency of the locally adapted type decreases. Our results extend the classic conditions on maintenance of diversity under selection and migration by including the effect of fluctuating population densities. We find simple analytic conditions in terms of the strength of selection, immigration, and the extent of fluctuations between generations in a continent-island model. Whereas weak fluctuations hardly affect coexistence, strong recurrent fluctuations lead to extinction of the type better adapted to the fluctuating niche - even if the invader is locally maladapted. There is a disadvantage to specialization to an unstable habitat, as it makes the population vulnerable to swamping from more stable habitats.

evolutionary biology↗

bioRxiv2013

Climate change triggers morphological and life-history evolution in response to predators

Although climate change is expected to reorganize entire communities, this restructuring might reflect either direct ecological or evolutionary responses to abiotic conditions or indirect effects mediated through altered species interactions. We tested the hypothesis that changes in trophic interaction strength due to altered predator abundance have a cascading evolutionary response in a prey species (Daphnia pulex). Using a multiyear / multigenerational field experiment, we manipulated 12 open aquatic mesocosms to simulate hydrological conditions under climate change. After a three-year press manipulation, we collected Daphnia pulex from each pond and raised them in a common garden. Using quantitative genetic methods, we measured a series of quantitative traits every other day on 108 individuals for eight weeks. There was a significant decrease in tail spine length and population growth rate in groups exposed to the most extreme future climate scenarios. Structural equation models demonstrated that trait changes were best explained as an indirect effect of climate change treatments mediated through changes in predator abundance. Our results suggest climate change can trigger a cascade of ecological and evolutionary forces by reducing predator density, which in turn acts as a selective force leading to evolutionary change in prey morphology and life history.

Ecology↗

bioRxiv2014

A novel inference of the fundamental biodiversity number for multiple immigration-limited communities

Neutral community theory postulates a fundamental quantity, {theta}, which reflects the species diversity on a regional scale. While the recent genealogical formulation of community dynamics has considerably enhanced quantitative neutral ecology, its inferential aspects have remained computationally prohibitive. Here, we make use of a generalized version of the original two-level hierarchical framework in order to define a novel estimator for{theta} ;, which proves to be computationally efficient and robust when tested on a wide range of simulated neutral communities. Estimating{theta} ; from field data is also illustrated using two tropical forest datasets consisting of spatially separated permanent field plots. Preliminary results also reveal that our inferred regional diversity parameter based on community dynamics may be linked to widely used ordination techniques in ecology. This paper essentially paves the way for future work dealing with the parameter inference of neutral communities with respect to their spatial scale and structure.

Ecology↗

bioRxiv2014

Building a better frog trap: The benefits of mal-adaptive habitat choice for metapopulations with different life history strategies

By spatially distributing offspring among several habitat patches in varying environments, an organism can \"hedge its bets\" to protect against bad conditions in any single patch. This strategy can maintain populations even when some or even all locations are, on average, population sinks. However, species may not have evolved this bet-hedging mechanism, especially when sink environments are anthropogenic \"traps\" - locations where traditional habitat cues have been altered. Using a model based on the life history of the Cascades frog (Rana cascadae), we examine the conditions that maximize growth in an environment with ecological traps created by the introduction of predators. In a temporally stochastic environment, maximum growth rates occur when some juveniles disperse to the trap. We then examine how different life histories and predation regimes affect the ability of an organism gain an advantage by bet-hedging, and find that bet-hedging can be less useful when the ecological trap drives adult, rather than juvenile, mortality.

Ecology↗

bioRxiv2014

A continuum of specialists and generalists in empirical communities

Understanding the persistence of specialists and generalists within ecological communities is a topical research question, with far-reaching consequences for the maintenance of functional diversity. Although theoretical studies indicate that restricted conditions may be necessary to achieve co-occurrence of specialists and generalists, analyses of larger empirical (and species-rich) communities reveal the pervasiveness of coexistence. In this paper, we analyze 175 ecological bipartite networks of three interaction types (animal hosts-parasite, plant-herbivore and plant-pollinator), and measure the extent to which these communities are composed of species with different levels of specificity in their biotic interactions. We find a continuum from specialism to generalism. Furthermore, we demonstrate that diversity tends to be greatest in networks with intermediate connectance, and argue this is because of physical constraints in the filling of networks.

Ecology↗

bioRxiv2015

Multi-trait eco-evolutionary dynamics explain niche diversity and evolved neutrality in forests

An enduring challenge in ecology is to understand how diverse plant species coexist when competing for the same basic resources1-4. Two candidate frameworks for meeting this challenge currently exist in forest ecology. Niche-based approaches succeed in describing successional dynamics in response to recurrent disturbances5-9. Approaches based on Hubbells neutral theory describe abundance patterns in highly diverse communities10. Yet both theories are unsuccessful where their competitor is strong, highlighting an enduring need for reconciliation11-13. In fact, the perception that niche and neutral processes are incompatible may have arisen because both types of models have lacked some critical features of real forests. Here we report a productive reconciliation that arises from extending niche models to include three ubiquitous features of vegetation: (1) plant growth under light competition, (2) multiple trait-mediated tradeoffs in plant function, and (3) evolutionary community assembly. We show that fitness equivalence - which neutral theory controversially takes as an assumption4,1213-14 - naturally arises as a mechanistic outcome of niche differentiation, but only within a specific region of trait space, corresponding to shade-tolerant later-successional strategies. In this way, niche diversification can lead, via evolved neutrality, to a proliferation of shade-tolerant strategies, one of the phenomena that motivated the development of neutral theory12. Earlier niche models focussed on a single growth-mortality trait and found no evolved neutrality. In contrast, the model introduced here includes a second trait, height at maturation, in addition to a growth-mortality trait in the form of leaf-construction cost. We show that fitness equivalence only emerges when diversifications include this second trait. We also demonstrate that our extended niche model generates markedly different forest types under different environmental conditions, recovering known biogeographic patterns15-18. By evolving forests from first principles, our study constructively resolves the protracted debate about niche versus neutral dynamics for forest communities, and provides a platform for a synthetic theory of forest structure and diversity.

Ecology↗