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Incomplete sampling of species’ geographic distributions has challenged biogeographers for many years to precisely quantify global-scale biodiversity patterns. After correcting for the spatial inequality of sample completeness, we generated a global species diversity map for woody angiosperms (82,974 species, 13,959,780 occurrence records). The estimated diversity demonstrated non-linear latitudinal and longitudinal patterns that were potentially related to region-specific biogeographic factors including current climate, paleoclimate, and topographical factors, while energy availability was the most important predictor at a global level. We identified the areas with potentially high species richness and rarity, but poorly explored, unprotected, and threatened by deforestation: they are distributed mostly at low latitudes across central South America, central Africa, subtropical China, and Indomalayan islands. These priority areas for botanical exploration would help to efficiently fill spatial knowledge gaps for better describing the status of biodiversity and improve the effectiveness of the protected area network for global woody plant conservation.TeaserBias-corrected diversity map based on occurrence records sheds new light on global macroecology and conservation of woody angiosperms.

As Earth's climate has varied strongly through geological time, studying the impacts of past climate change on biodiversity helps to understand the risks from future climate change. However, it remains unclear how paleo-climate shapes spatial variation in biodiversity. Here, we assessed the influence of Quaternary climate change on spatial dissimilarity in taxonomic, phylogenetic, and functional composition among neighboring 200-kilometer cells (beta-diversity) for angiosperm trees worldwide. We found that larger glacial-interglacial temperature change was strongly associated with lower spatial turnover (species replacements) and higher nestedness (rich-ness changes) components of beta-diversity across all three biodiversity facets. Moreover, phylogenetic and functional turnover was lower and nestedness higher than random expectations based on taxonomic beta -di-versity in regions that experienced large temperature change, reflecting phylogenetically and functionally se-lective processes in species replacement, extinction, and colonization during glacial-interglacial oscillations. Our results suggest that future human-driven climate change could cause local homogenization and reduction in taxonomic, phylogenetic, and functional diversity of angiosperm trees worldwide. W.-B.X. and A.O. acknowledge support from the Aarhus University Research Foundation (AUFF) Starting Grant (AUFF-F-2018-7-8). J.-C.S. acknowledges support from the Danish Council for Independent Research|Natural Sciences (grant 6108-00078B) to the TREECHANGE Project. J.-C.S. also considers this work a contribution to the Center for Ecological Dynamics in a Novel Biosphere (ECONOVO) funded by Danish National Research Foundation (grant DNRF173) and the VILLUM Investigator project “Biodiversity Dynamics in a Changing World” funded by VILLUM FONDEN (grant 16549). W.L.E. was supported by the VILLUM FONDEN (grant 00025354). M.A. was supported by the Natural Sciences and Engineering Research Council of Canada (NSERC) Discovery grants program. C.B. was supported by the National Research Foundation of Korea (NRF) grant funded by the Korea government (MSIT) (2022R1A2C1003504). A.G.G. was supported by the ANID PIA/BASAL (grant FB210006) and FONDECYT (grant 1200468). R.E.O. was supported by the German Research Foundation to German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig (DFG-FZT 118, 202548816). J.Pe. was supported by the Fundación Ramón Areces project (CIVP20A6621) and Spanish Government project (PID2019-110521GB-I00). J.Pi. was supported by the Estonian Research Council Grants (PUT1355, PRG 1405) and Mobilitas Pluss (MOBERC11). V.D.P. was supported by the National Council of Scientific and Technological Development (CNPq) of Brazil (grant 307689/2014-0).

With 178 species, palm subtribe Dypsidinae is one of the largest plant radiations on Madagascar. A well-resolved species-level phylogeny is required not only to unpick the drivers of this spectacular radiation, but also to define natural and useful generic limits in this taxonomically difficult group. The only recent taxonomic revision of Dypsidinae recognised just four genera, including the large genus Dypsis (currently with 172 species), which was considered impossible to divide further based on morphology alone. Here we describe the first comprehensively sampled species-level phylogeny of Dypsidinae, including 157 described species (88% of the subtribe) as well as 10 samples of uncertain taxonomic identity (potentially new to science). Our tree is based on target sequence capture data for 161 nuclear DNA loci. The phylogenetic relationships recovered render Dypsis s.l. paraphyletic, requiring the resurrection of two previously synonymised genera. In total, we recognise six genera in Dypsidinae (Dypsis, Chrysalidocarpus, Marojejya, Vonitra, Lemurophoenix, Masoala), all of which are monophyletic and consistent with morphology.

Functional traits play a key role in driving plant community effects on ecosystem function. We examined nine functional traits in various palm (Arecaceae) species and their relationships with moisture, tree-fall gaps, slope, and forest type at 29 transects (500×5 m) in the northeastern region of the Colombian Amazon. Redundancy analysis of mean trait values of species within a plot weighted by their abundance and Pearson correlations were used to evaluate the relationships between traits and environmental factors. The community trait composition was correlated with local environmental factors, which explained 23% of the trait variance. We detected functional dominance of the tallest palms in soils with high moisture and in floodplain forests (p ≤0.05). Palms with relatively long leaves were dominant in the flooded forests. Acaulescent and small palms were dominant on high slopes, and interra firmeforests, long-petioled palms were dominant in forest gaps. The number of seeds per fruit was not correlated with any environmental variable. Thus, hydrology is one of the main drivers of the functional composition of neotropical palm communities at the local scale, segregating tall palms with competitive and evasive strategies from small understory palms, which are mainly stress tolerant.

SummaryThe traits of plants determine how they interact with each other and their environment, constituting key knowledge for diverse fields. The lack of comprehensive knowledge of plant traits (the “Raunkiærian shortfall”) poses a major, cross-disciplinary, barrier to scientific advancement. Spatial biases in trait coverage may also lead to erroneous conclusions affecting ecosystem management and conservation planning. Thus, there is an urgent need to assess the spatial completeness of plant trait data, understand drivers of geographic biases, and to identify solutions for filling regional gaps. Here, we leverage a comprehensive set of regional species checklists for vascular plants and trait data for 2,027 traits and 128,929 plant species from the TRY database to assess trait data completeness across the globe. We show that trait data availability in TRY is associated with socioeconomic and biological factors influencing sampling likelihood: trait completeness was positively associated with mean species range size, research expenditure, and human population density and negatively associated with endemism and vascular plant species richness. Integration of a second, regional trait database (AusTraits) more than doubled trait completeness for the continent covered, indicating that the creation and integration of regional databases can rapidly expand trait completeness.Plain Language SummaryThe traits of plants determine how they interact with each other and their environment. Our knowledge of plant traits is incomplete, limiting scientific advancement as well as our ability to manage ecosystems and plan conservation actions. We show that there are large biases in trait data availability which are associated with both biological factors (range size, endemism, species richness) and socioeconomic factors (research expenditure, human population density). We also show how regionally-focused efforts can help rapidly expand trait data availability.

Safeguarding Earth?s tree diversity is a conservation priority due to the importance of trees for biodiversity and ecosystem functions and services such as carbon sequestration. Here, we improve the foundation for effective conservation of global tree diversity by analyzing a recently developed database of tree species covering 46,752 species. We quantify range protection and anthropogenic pressures for each species and develop conservation priorities across taxonomic, phylogenetic, and functional diversity dimensions. We also assess the effectiveness of several influential proposed conservation prioritization frameworks to protect the top 17% and top 50% of tree priority areas. We find that an average of 50.2% of a tree species? range occurs in 110-km grid cells without any protected areas (PAs), with 6,377 small-range tree species fully unprotected, and that 83% of tree species experience nonnegligible human pressure across their range on average. Protecting highpriority areas for the top 17% and 50% priority thresholds would increase the average protected proportion of each tree species? range to 65.5% and 82.6%, respectively, leaving many fewer species (2,151 and 2,010) completely unprotected. The priority areas identified for trees match well to the Global 200 Ecoregions framework, revealing that priority areas for trees would in large part also optimize protection for terrestrial biodiversity overall. Based on range estimates for >46,000 tree species, our findings show that a large proportion of tree species receive limited protection by current PAs and are under substantial human pressure. Improved protection of biodiversity overall would also strongly benefit global tree diversity. Made available in DSpace on 2022-06-21T15:23:55Z (GMT). No. of bitstreams: 1 Guo-et-al.-2022-PNAS-High-exposure-of-global-tree-diversity-to-human-pressure.pdf: 2523289 bytes, checksum: 245be3df3fb487ed152d25bf50f4e984 (MD5) Previous issue date: 2022 Proceedings of the National Academy of Sciences of the United States of America.

Aim: Since Wallace's and Engler's 19th-century biogeographical schemes, biogeographers have sought to classify the world into biogeographical regions according to patterns in biotic distribution. Yet, while most of the world's plant biodiversity can be found in the tropics, basic phytogeographical relationships and boundaries within this zone remain debated. We investigated whether palms, a species-rich, well-studied pantropical family, reflect traditional floristic schemes or support a recently published phylogenetic regionalisation that contests the traditional views on phytogeographical tropical regions. Location: Tropics and subtropics. Taxon: Palm family (Arecaceae). Methods: We use a species-level dated phylogenetic tree of palms along with a dataset of all palm species distributions to calculate pairwise phylogenetic beta diversity among palm assemblages. From these pairwise dissimilarity values, we compute the relationship between assemblages in NMDS ordination space and use hierarchical clustering to define biogeographical units for palms. Results: We found a Neotropical versus Palaeotropical division of palm assemblages as the major biogeographical split. Two Neotropical and four Palaeotropical clusters were identified with the following relationships: ((Southern Neotropical, Northern Neotropical), ((Indian Ocean), ((African), (Eurasian-Australian, Melanesian-Pacific)))). Main Conclusions: Our analysis supports many delineations suggested in traditional floristic schemes, most importantly a clear Neotropical versus Palaeotropical division and units resembling Takhtajan's African, Indo-Malesian, New Caledonian and Polynesian Palaeotropical subkingdoms. However, as recently suggested in phylogenetic regionalisation, our results also show a clustering of Southeast Asian and Australian assemblages, substantiating the suggestion of a combined Asian-Australian region. Biogeographical regionalisation for palms does not support the existence of trans-continental dry tropical or subtropical phytogeographical clusters that have recently been found in dicot trees, despite known Northern Hemisphere connections for palms. Further modern regionalisations based on multiple datasets with complementary strengths are needed to establish a credible set of phytogeographical regions for the tropics.

Six new species of Maesa (Primulaceae-Maesoideae) from Papua New Guinea, namely M. angustibracteolata, M. aurulenta, M. brassii, M. oblanceolatifolia, M. pusilliflora and M. prolatifructa are described and illustrated based on observations from herbarium specimens; the taxonomic affinities of each species are also discussed.

The Darwinian shortfall, i.e. the lack of knowledge of phylogenetic relationships, significantly impedes our understanding of evolutionary drivers of global patterns of biodiversity. Spatial bias in the Darwinian shortfall, where phylogenetic knowledge in some regions is more complete than others, could undermine eco- and biogeographic inferences. Yet, spatial biases in phylogenetic knowledge for major groups – such as plants – remain poorly understood. Using data for 337 023 species (99.7%) of seed plants (Spermatophyta), we produced a global map of phylogenetic knowledge based on regional data and tested several potential drivers of the observed spatial variation. Regional phylogenetic knowledge was defined as the proportion of the regional seed plant flora represented in GenBank's nucleotide database with phylogenetically relevant data. We used simultaneous autoregressive models to explain variation in phylogenetic knowledge based on three biodiversity variables (species richness, range size and endemism) and six socioeconomic variables representing funding and accessibility. We compared observed patterns and relationships to established patterns of the Wallacean shortfall (the lack of knowledge of species distributions). We found that the Darwinian shortfall is strongly and significantly related to the macroecological distribution of species' range sizes. Small-ranged species were significantly less likely to have phylogenetic data, leading to a concentration of the Darwinian shortfall in species-rich, tropical countries where range sizes are small on average. Socioeconomic factors were less important, with significant but quantitatively small effects of accessibility and funding. In conclusion, reducing the Darwinian shortfall and smoothen its spatial bias will require increased efforts to sequence the world's small-ranged (endemic) species.

Species richness varies immensely around the world. Variation in the rate of diversification (speciation minus extinction) is often hypothesized to explain this pattern, while alternative explanations invoke time or ecological carrying capacities as drivers. Focusing on seed plants, the world’s most important engineers of terrestrial ecosystems, we investigated the role of diversification rate as a link between the environment and global species richness patterns. Applying structural equation modeling to a comprehensive distribution dataset and phylogenetic tree covering all circa 332,000 seed plant species and 99.9% of the world’s terrestrial surface (excluding Antarctica), we test five broad hypotheses postulating that diversification serves as a mechanistic link between species richness and climate, climatic stability, seasonality, environmental heterogeneity, or the distribution of biomes. Our results show that the global patterns of species richness and diversification rate are entirely independent. Diversification rates were not highest in warm and wet climates, running counter to the Metabolic Theory of Ecology, one of the dominant explanations for global gradients in species richness. Instead, diversification rates were highest in edaphically diverse, dry areas that have experienced climate change during the Neogene. Meanwhile, we confirmed climate and environmental heterogeneity as the main drivers of species richness, but these effects did not involve diversification rates as a mechanistic link, calling for alternative explanations. We conclude that high species richness is likely driven by the antiquity of wet tropical areas (supporting the “tropical conservatism hypothesis”) or the high ecological carrying capacity of warm, wet, and/or environmentally heterogeneous environments.

The tree of life is the fundamental biological roadmap for navigating the evolution and properties of life on Earth, and yet remains largely unknown. Even angiosperms (flowering plants) are fraught with data gaps, despite their critical role in sustaining terrestrial life. Today, high-throughput sequencing promises to significantly deepen our understanding of evolutionary relationships. Here, we describe a comprehensive phylogenomic platform for exploring the angiosperm tree of life, comprising a set of open tools and data based on the 353 nuclear genes targeted by the universal Angiosperms353 sequence capture probes. The primary goals of this article are to (i) document our methods, (ii) describe our first data release, and (iii) present a novel open data portal, the Kew Tree of Life Explorer (https://treeoflife.kew.org). We aim to generate novel target sequence capture data for all genera of flowering plants, exploiting natural history collections such as herbarium specimens, and augment it with mined public data. Our first data release, described here, is the most extensive nuclear phylogenomic data set for angiosperms to date, comprising 3099 samples validated by DNA barcode and phylogenetic tests, representing all 64 orders, 404 families (96%⁠) and 2333 genera (17%⁠). A “first pass” angiosperm tree of life was inferred from the data, which totaled 824,878 sequences, 489,086,049 base pairs, and 532,260 alignment columns, for interactive presentation in the Kew Tree of Life Explorer. This species tree was generated using methods that were rigorous, yet tractable at our scale of operation. Despite limitations pertaining to taxon and gene sampling, gene recovery, models of sequence evolution and paralogy, the tree strongly supports existing taxonomy, while challenging numerous hypothesized relationships among orders and placing many genera for the first time. The validated data set, species tree and all intermediates are openly accessible via the Kew Tree of Life Explorer and will be updated as further data become available. This major milestone toward a complete tree of life for all flowering plant species opens doors to a highly integrated future for angiosperm phylogenomics through the systematic sequencing of standardized nuclear markers. Our approach has the potential to serve as a much-needed bridge between the growing movement to sequence the genomes of all life on Earth and the vast phylogenomic potential of the world’s natural history collections. [Angiosperms; Angiosperms353; genomics; herbariomics; museomics; nuclear phylogenomics; open access; target sequence capture; tree of life.], This work was supported by grants from the Calleva Foundation and the Sackler Trust to the Plant and Fungal Trees of Life project at the Royal Botanic Gardens, Kew. Additional support was received from the Garfield Weston Foundation, as part of the Global Tree Seed Bank Programme.

The island of Sulawesi formed from the joining of proto-islands roughly three million years ago. Regions of zoological endemism, corresponding to the proto-islands, have been reported. Sulawesi's tree communities, however, remain poorly documented. In better-studied tropical regions, soil types similar to those found in Sulawesi often have distinctive tree communities. To gather data on Sulawesi's tree communities, we established ten (0.25 ha) plots on four soil types across three regions. We documented diversity, endemism, dominance, and species composition. Linear models of species composition showed greater influence of geographic distance rather than soil, and no relationship with climate. This suggests that the legacy of Sulawesi's formation may have influenced tree communities more so than the soil types we sampled. Most of our plots were on stressful soil types making it difficult to conclude on the importance of edaphic specialization in the Sulawesi tree flora. The lack of climatic effects reflects Sulawesi's position within the wet tropics where the small climatic differences are unlikely to have large influence on tree communities. Abstract in Indonesian is available with online material.

The extinction of all Madagascar's megafrugivores ca 1000 years ago, may have left its signature on the current distribution of vertebrate-dispersed plants across the island, due to the loss of effective seed dispersal. In this study, we dissect the roles of extinct and extant frugivore distributions, abiotic variables, human impact and spatial predictors on the compositional turnover, or beta-diversity, of palm (Arecaceae) species and their dispersal-related traits across 40 assemblages in Madagascar. Variation partitioning showed that palm beta-diversity is mostly shaped by the distribution of extant frugivores (eight lemur, three bird, two rodent and one bat species) and the abiotic environment (e.g. forest cover, slope and temperature), and to a lesser extent by the distribution of extinct megafrugivores (several giant lemur and elephant bird species). However, the contribution of these variables differed between dry western assemblages and wet eastern assemblages, with a more prominent role, albeit still small, of extinct megafrugivores in the west. These results suggest that palm distributions in the dry west of Madagascar, where megafrugivores were probably most abundant in the past, still show signatures of past interactions. With a fourth-corner analysis we observed that the distribution of palm species with relatively large fruits and seeds was negatively associated with frugivore richness of both past and present communities and home range sizes of extant mammalian frugivores. This suggests that palm species with relatively large fruits tend to occur in places with fewer, small-ranged mammalian frugivores, which may indicate dysfunctional seed dispersal. Nevertheless, our results also indicate that several wide-ranging bird species with high dispersal ability (large hand-wing index) that also occasionally feed on fruits may compensate for this potential dispersal loss. Our study sheds new light on dispersal anachronisms in Madagascar, and how defaunation and past species interactions may underlie current plant distributions.

Extinction has increased as human activities impact ecosystems, yet relatively few species have conservation assessments. Novel approaches are needed to highlight threatened species that are currently data-deficient. Many Madagascan plant species have extremely narrow ranges, but this may not have always been the case—it is unclear how the island's diverse flora evolved. To assess this, we generated restriction-site associated DNA sequence data for 10 Madagascan plant species, estimated effective population size (Ne) for each species and compared this to census (Nc) sizes. In each case,Newas an order of magnitude larger thanNc—signifying rapid, recent population decline. We then estimated species' demographic history, tracking changes inNeover time. We show that it is possible to predict extinction risk, particularly in the most threatened species. Furthermore, simulations showed that our approach has the power to detect population decline during the Anthropocene. Our analyses reveal that Madagascar's micro-endemics were not always rare, having experienced a rapid decline in their recent history. This casts further uncertainty over the processes that generated Madagascar's exceptional biodiversity. Our approach targets data-deficient species in need of conservation assessment, particularly in regions where human modification of the environment has been rapid.

Aim Palms are an iconic, diverse and often abundant component of tropical ecosystems that provide many ecosystem services. Being monocots, tree palms are evolutionarily, morphologically and physiologically distinct from other trees, and these differences have important consequences for ecosystem services (e.g., carbon sequestration and storage) and in terms of responses to climate change. We quantified global patterns of tree palm relative abundance to help improve understanding of tropical forests and reduce uncertainty about these ecosystems under climate change. Location Tropical and subtropical moist forests. Time period Current. Major taxa studied Palms (Arecaceae). Methods We assembled a pantropical dataset of 2,548 forest plots (covering 1,191ha) and quantified tree palm (i.e., ≥10cm diameter at breast height) abundance relative to co‐occurring non‐palm trees. We compared the relative abundance of tree palms across biogeographical realms and tested for associations with palaeoclimate stability, current climate, edaphic conditions and metrics of forest structure. Results On average, the relative abundance of tree palms was more than five times larger between Neotropical locations and other biogeographical realms. Tree palms were absent in most locations outside the Neotropics but present in >80% of Neotropical locations. The relative abundance of tree palms was more strongly associated with local conditions (e.g., higher mean annual precipitation, lower soil fertility, shallower water table and lower plot mean wood density) than metrics of long‐term climate stability. Life‐form diversity also influenced the patterns; palm assemblages outside the Neotropics comprise many non‐tree (e.g., climbing) palms. Finally, we show that tree palms can influence estimates of above‐ground biomass, but the magnitude and direction of the effect require additional work. Conclusions Tree palms are not only quintessentially tropical, but they are also overwhelmingly Neotropical. Future work to understand the contributions of tree palms to biomass estimates and carbon cycling will be particularly crucial in Neotropical forests.

The date palm, Phoenix dactylifera, has been a cornerstone of Middle Eastern and North African agriculture for millennia. It was first domesticated in the Persian Gulf, and its evolution appears to have been influenced by gene flow from two wild relatives, P. theophrasti, currently restricted to Crete and Turkey, and P. sylvestris, widespread from Bangladesh to the West Himalayas. Genomes of ancient date palm seeds show that gene flow from P. theophrasti to P. dactylifera may have occurred by ∼2,200 years ago, but traces of P. sylvestris could not be detected. We here integrate archeogenomics of a ∼2,100-year-old P. dactylifera leaf from Saqqara (Egypt), molecular-clock dating, and coalescence approaches with population genomic tests, to probe the hybridization between the date palm and its two closest relatives and provide minimum and maximum timestamps for its reticulated evolution. The Saqqara date palm shares a close genetic affinity with North African date palm populations, and we find clear genomic admixture from both P. theophrasti, and P. sylvestris, indicating that both had contributed to the date palm genome by 2,100 years ago. Molecular-clocks placed the divergence of P. theophrasti from P. dactylifera/P. sylvestris and that of P. dactylifera from P. sylvestris in the Upper Miocene, but strongly supported, conflicting topologies point to older gene flow between P. theophrasti and P. dactylifera, and P. sylvestris and P. dactylifera. Our work highlights the ancient hybrid origin of the date palms, and prompts the investigation of the functional significance of genetic material introgressed from both close relatives, which in turn could prove useful for modern date palm breeding., Molecular Biology and Evolution, 38 (10), ISSN:0737-4038, ISSN:1537-1719

Sequencing of target-enriched libraries is an efficient and cost-effective method for obtaining DNA sequence data from hundreds of nuclear loci for phylogeny reconstruction. Much of the cost of developing targeted sequencing approaches is associated with the generation of preliminary data needed for the identification of orthologous loci for probe design. In plants, identifying orthologous loci has proven difficult due to a large number of whole-genome duplication events, especially in the angiosperms (flowering plants). We used multiple sequence alignments from over 600 angiosperms for 353 putatively single-copy protein-coding genes identified by the One Thousand Plant Transcriptomes Initiative to design a set of targeted sequencing probes for phylogenetic studies of any angiosperm group. To maximize the phylogenetic potential of the probes, while minimizing the cost of production, we introduce a k-medoids clustering approach to identify the minimum number of sequences necessary to represent each coding sequence in the final probe set. Using this method, 5–15 representative sequences were selected per orthologous locus, representing the sequence diversity of angiosperms more efficiently than if probes were designed using available sequenced genomes alone. To test our approximately 80,000 probes, we hybridized libraries from 42 species spanning all higher-order groups of angiosperms, with a focus on taxa not present in the sequence alignments used to design the probes. Out of a possible 353 coding sequences, we recovered an average of 283 per species and at least 100 in all species. Differences among taxa in sequence recovery could not be explained by relatedness to the representative taxa selected for probe design, suggesting that there is no phylogenetic bias in the probe set. Our probe set, which targeted 260 kbp of coding sequence, achieved a median recovery of 137 kbp per taxon in coding regions, a maximum recovery of 250 kbp, and an additional median of 212 kbp per taxon in flanking non-coding regions across all species. These results suggest that the Angiosperms353 probe set described here is effective for any group of flowering plants and would be useful for phylogenetic studies from the species level to higher-order groups, including the entire angiosperm clade itself., This work was supported by the Texas Tech University College of Arts and Sciences [to M.G.J.], the National Science Foundation [DEB-1239992, DEB-1342873 to N.J.W.], and by the Calleva Foundation, the Garfield Weston Foundation, and the Sackler Trust to the Royal Botanic Gardens, Kew.

Phylogenetic analyses are increasingly being performed with data sets that incorporate hundreds of loci. Due to incomplete lineage sorting, hybridization, and horizontal gene transfer, the gene trees for these loci may often have topologies that differ from each other and from the species tree. The effect of these topological incongruences on divergence time estimation has not been fully investigated. Using a series of simulation experiments and empirical analyses, we demonstrate that when topological incongruence between gene trees and the species tree is not accounted for, the temporal duration of branches in regions of the species tree that are affected by incongruence is underestimated, whilst the duration of other branches is considerably overestimated. This effect becomes more pronounced with higher levels of topological incongruence. We show that this pattern results from the erroneous estimation of the number of substitutions along branches in the species tree, although the effect is modulated by the assumptions inherent to divergence time estimation, such as those relating to the fossil record or among-branch-substitution-rate variation. By only analyzing loci with gene trees that are topologically congruent with the species tree, or only taking into account the branches from each gene tree that are topologically congruent with the species tree, we demonstrate that the effects of topological incongruence can be ameliorated. Nonetheless, even when topologically congruent gene trees or topologically congruent branches are selected, error in divergence time estimates remains. This stems from temporal incongruences between divergence times in species trees and divergence times in gene trees, and more importantly, the difficulty of incorporating necessary assumptions for divergence time estimation. [Divergence time estimation; gene trees; species tree; topological incongruence.]

Well-supported phylogenies are a prerequisite for the study of the evolution and diversity of life on earth. The subfamily Calamoideae accounts for more than one fifth of the palm family (Arecaceae), occurs in tropical rainforests across the world, and supports a billion-dollar industry in rattan products. It contains ca. 550 species in 17 genera, 10 subtribes and three tribes, but their phylogenetic relationships remain insufficiently understood. Here, we sequenced almost one thousand nuclear genomic regions for 75 systematically selected Calamoideae, representing the taxonomic diversity within all calamoid genera. Our phylogenomic analyses resolved a maximally supported phylogenetic backbone for the Calamoideae, including several higher-level relationships not previously inferred. In-depth analysis revealed low gene tree conflict for the backbone but complex deep evolutionary histories within several subtribes. Overall, our phylogenomic framework sheds new light on the evolution of palms and provides a robust foundation for future comparative studies, such as taxonomy, systematics, biogeography, and macroevolutionary research.

The date palm (Phoenix dactylifera) has been a cornerstone of Middle Eastern and North African agriculture for millennia. It is presumed that date palms were first domesticated in the Persian Gulf and subsequently introduced into North Africa, where their evolution in the latter region appears to have been influenced by gene flow from the wild relative P. theophrasti, which is restricted to Crete and Turkey. However, the timing of gene flow from P. theophrasti to P. dactylifera remains unknown due to the limited archaeobotanical evidence of P. theophrasti and their exclusion from population genomic studies.We addressed this issue by investigating the relatedness and ancestry of a ~2,100-year-old P. dactylifera leaf from Saqqara (Egypt), combining genome sequencing of this ancient specimen with a broad sample of date palm cultivars and closely related species.The ancient Saqqara date palm shares close genetic ancestry with North African date palm populations. We find clear genomic admixture between the Saqqara date palm, P. theophrasti and the closest known relative P. sylvestris.Our study highlights that gene flow from P. theophrasti and P. sylvestris to North African date palms had already occurred at least ~2,100 years ago, providing a minimum timestamp for hybridisation between species.

Maesa brevipedicellata, a new species of Maesa ( Primulaceae-Maesoideae) from Papua New Guinea, is described and illustrated based on herbarium specimen observations. The collections of this species resemble M. rufovillosa and were previously determined as that species. Maesa brevipedicellata is unique with its self- supporting habit, hispid hairs throughout and paniculate inflorescences with very short pedicels. This new species mainly differs from M. rufovillosa by the habit (tree/shrub in M. brevipedicellata vs climber in M. rufovillosa) and the inflorescence structure (panicles in M. brevipedicellata vs simple racemes in M. rufovillosa).

Trees are of vital importance for ecosystem functioning and services at local to global scales, yet we still lack a detailed overview of the global patterns of tree diversity and the underlying drivers, particularly the imprint of paleoclimate. Here, we present the high-resolution (110 km) worldwide mapping of tree species richness, functional and phylogenetic diversities based on ∼7 million quality-assessed occurrences for 46,752 tree species (80.5% of the estimated total number of tree species), and subsequent assessments of the influence of paleo-climate legacies on these patterns. All three tree diversity dimensions exhibited the expected latitudinal decline. Contemporary climate emerged as the strongest driver of all diversity patterns, with Pleistocene and deeper-time (>107 years) paleoclimate as important co-determinants, and, notably, with past cold and drought stress being linked to reduced current diversity. These findings demonstrate that tree diversity is affected by paleoclimate millions of years back in time and highlight the potential for tree diversity losses from future climate change.

Although trees are key to ecosystem functioning, many forests and tree species across the globe face strong threats. Preserving areas of high biodiversity is a core priority for conservation; however, different dimensions of biodiversity and varied conservation targets make it difficult to respond effectively to this challenge. Here, we (i) identify priority areas for global tree conservation using comprehensive coverage of tree diversity based on taxonomy, phylogeny, and functional traits; and (ii) compare these findings to existing protected areas and global biodiversity conservation frameworks. We find that ca. 51% of the top-priority areas for tree biodiversity are located in current protected areas. The remaining half top-priority areas are subject to moderate to high human pressures, indicating conservation actions are needed to mitigate these human impacts. Our findings emphasize the effectiveness of using tree conservation priority areas for future global conservation planning.

Spatial patterns of plant diversity follow the well-known global latitudinal biodiversity gradient, however there is little consensus about the underlying causes for this pattern. Here we present a spatial analysis of a complete checklist of the world’s seed plants, integrated with a comprehensive plant Tree of Life. This combination allows insights into the evolutionary drivers of plant species richness patterns, specifically current plant biodiversity patterns, and the diversification processes that shaped them. Our study provides a comprehensive global species richness map and relates the observed species richness pattern to speciation rates derived from phylogeny, and with environmental variables, which are hypothesized to impact speciation rates. Initial results show that tropical rain forest regions, although being areas that contain among the highest numbers of species, are regions with comparatively low speciation rates, contradicting the widespread notion that rainforests are “cradles” of biodiversity. This finding seems further supported by contrasting association of environmental variables, like precipitation and temperature, with speciation rates and species richness.

Extinction has increased as human activities impact ecosystems. Conservation assessments for the IUCN red list are a fundamental tool in aiding the prevention of further extinction, yet, relatively few species have been thoroughly assessed. To increase the efficiency of assessments, novel approaches are needed to highlight threatened species that are currently data deficient. Many Madagascan plant species currently have extremely narrow ranges, but this may not have always been the case. To assess this, we used high-throughput DNA sequencing for 2-5 individuals of each species - reflecting the paucity of samples available for rare species. We estimated effective population size (Ne) for each species and compared this to census population (Nc) sizes when known. In each case, Ne was an order of magnitude larger than Nc – a signature of rapid, recent population decline. We then estimated the demographic history of each species, tracking changes in Ne over time. Five out of ten species displayed significant population declines towards the present (68–90% decreases). Our results for palm trees indicate that it is possible to predict extinction risk, particularly in the most threatened species. We performed simulations to show that our approach has the power to detect population decline during the Anthropocene, but performs less well when less data is used. Similar declines to those in palms were observed in data deficient species or those assessed as of least concern. These analyses reveal that Madagascar’s narrow endemics were not always rare, having experienced rapid decline in their recent history. Our approach offers the opportunity to target species in need of conservation assessment with little prior information, particularly in regions where human modification of the environment has been rapid.SummaryCurrent IUCN conservation assessment methods are reliant on observed declines in species population and range sizes over the last one hundred years, but for the majority of species this information is not available. We used a population genetic approach to reveal historical demographic decline in the rare endemic flora of Madagascar. These results show that it is possible to predict extinction risk from demographic patterns inferred from genetic data and that destructive human influence is likely to have resulted in the very high frequency of narrow endemics present on the island. Our approach will act as an important tool for rapidly assessing the threatened status of poorly known species in need of further study and conservation, particularly for tropical flora and fauna.

Aims Glacial-interglacial climate stability and contemporary climate could affect bird diversity through their direct effects on diversification and physiological tolerance, and indirect effects on plant species richness, which could provide more food and ecological niches. This study aims to quantitatively assess the direct and indirect effects of these drivers on species richness, phylogenetic and functional structure of bird assemblages.Location Mainland China.Taxon Birds.Methods Ordinary least squares models and simultaneous autoregressive models were used to test the associations between glacial-interglacial climate anomaly, contemporary climate, plant species richness, elevation range and species richness, phylogenetic structure, functional structure of bird assemblages in mainland China. Structural equation models were used to distinguish between direct and indirect effects of these variables on bird diversity.Results High bird species richness with phylogenetically and functionally clustered structure occurred in regions with more plant species and stable glacial-interglacial climate. Notably, the phylogenetically clustered bird assemblages were composed of young lineages. Contemporary precipitation and elevation range affected bird diversity through their indirect effects on plant species richness.Main Conclusions These findings suggest that more plant species and stable glacial-interglacial climate may promote bird diversity through their effects on in situ diversification and species dispersal rate in southwestern China. Therefore, the well-preserved plant and bird diversity through the long-term climate fluctuations should be of great importance of biodiversity conservation in China.

Providing science and society with an integrated, up-to-date, high quality, open, reproducible and sustainable plant tree of life would be a huge service that is now coming within reach. However, synthesizing the growing body of DNA sequence data in the public domain and disseminating the trees to a diverse audience are often not straightforward due to numerous informatics barriers. While big synthetic plant phylogenies are being built, they remain static and become quickly outdated as new data are published and tree-building methods improve. Moreover, the body of existing phylogenetic evidence is hard to navigate and access for non-experts. We propose that our community of botanists, tree builders, and informaticians should converge on a modular framework for data integration and phylogenetic analysis, allowing easy collaboration, updating, data sourcing and flexible analyses. With support from major institutions, this pipeline should be re-run at regular intervals, storing trees and their metadata long-term. Providing the trees to a diverse global audience through user-friendly front ends and application development interfaces should also be a priority. Interactive interfaces could be used to solicit user feedback and thus improve data quality and to coordinate the generation of new data. We conclude by outlining a number of steps that we suggest the scientific community should take to achieve global phylogenetic synthesis.

I. II. III. IV. V. VI. VII. VIII. IX. References SUMMARY: Tropical rainforest (TRF) is the most species-rich terrestrial biome on Earth, harbouring just under half of the world's plant species in c. 7% of the land surface. Phylogenetic trees provide important insights into mechanisms underpinning TRF hyperdiversity that are complementary to those obtained from the fossil record. Phylogenetic studies of TRF plant diversity have mainly focused on whether this biome is an evolutionary 'cradle' or 'museum', emphasizing speciation and extinction rates. However, other explanations, such as biome age, immigration and ecological limits, must also be considered. We present a conceptual framework for addressing the drivers of TRF diversity, and review plant studies that have tested them with phylogenetic data. Although surprisingly few in number, these studies point to old age of TRF, low extinction and high speciation rates as credible drivers of TRF hyperdiversity. There is less evidence for immigration and ecological limits, but these cannot be dismissed owing to the limited number of studies. Rapid methodological developments in DNA sequencing, macroevolutionary analysis and the integration of phylogenetics with other disciplines may improve our grasp of TRF hyperdiversity in the future. However, such advances are critically dependent on fundamental systematic research, yielding numerous, additional, well-sampled phylogenies of TRF lineages.

Four new palm species are described in the genus Dypsis: three tree palms from the Masoala peninsula in north-eastern Madagascar (D. mijoroana sp. nov., D. ovojavavy sp. nov. and D. rabepierrei sp. nov.), and one understorey palm, D. aurantiaca sp. nov. from eastern-central Madagascar. Illustrations, a distribution map and conservation status are provided for all four species.

Several attempts have been made to generate complete species-level phylogenies for large clades, enabling comprehensive analyses of ecological or evolutionary hypotheses at the species level. No such phylogeny has, however, been generated for any major plant group yet, but here we generate such a phylogeny for the palm family (Arecaceae). We do this using a novel Bayesian approach, estimating the validity of intra-generic taxonomic groupings as topological constraints to assist in placing species without genetic or morphological data. From these we implement those that are supported by genetic or morphological data for a given genus or for related genera. The intergeneric relationships in our new phylogeny are surprisingly different from earlier phylogenies in the placement of genera within tribes, but largely identical to previous findings in the deeper branches in the phylogeny, pointing to the need for incorporating phylogenetic uncertainty in analyses based on this phylogeny. Initial analyses of the new phylogeny suggest non-constancy in diversification rates over time within genera, with an apparent increase in diversification rate over time, but no evidence for any geographic variation in the magnitude of this increase. We hope that our study will stimulate further evolutionary or ecological studies using palms as study organisms as well as discussions of the optimal way to place the many species without genetic or morphological data., We are grateful to the Danish Council for Independent Research Natural Sciences (grant 12-125079 to JCS and 4090-00227 to SF) and the European Union’s FP7-PEOPLE programme (grant 327259 to WLE) for financial support.

Earth's climate has experienced strong changes on timescales ranging from decades to millions of years. As biodiversity has evolved under these circumstances, dependence on these climate dynamics is expected. In this review, we assess the current state of knowledge on paleoclimatic legacies in biodiversity and ecosystem patterns. Paleoclimate has had strong impacts on past biodiversity dynamics, driving range shifts and extinctions as well as diversification. We outline theory for how these dynamics may have left legacies in contemporary patterns and review the empirical evidence. We report ample evidence that Quaternary glacial–interglacial climate change affects current patterns of species distributions and diversity across a broad range of organisms and regions. We also report emerging evidence for paleoclimate effects on current patterns in phylogenetic and functional diversity and ecosystem functioning and for legacies of deeper-time paleoclimate conditions. Finally, we discuss implications for Anthropocene ecology and outline an agenda to improve our understanding of paleoclimate's role in shaping contemporary biodiversity and ecosystems.

Aim Using conifers as a model system, we aim to test four hypotheses. H1: the processes that shape the phylogenetic structure of regional species assemblages depend on climate. H2: apparent effects of current climate can be equally well explained by past climate. H3: strong Quaternary climate oscillations have led to phylogenetically non-random assemblages, either with few closely related species because isolated populations do not persist long enough to become new species or with many close relatives due to increased allopatric speciation. H4: strong late Cenozoic aridification has led to assemblages with many close relatives due to extinction and adaptive radiation. Location Global. Methods We used boosted regression trees to relate the net relatedness index (NRI) of regional conifer assemblages to current climate, past climate (0.021, 3 and 7.3-11.6 Ma), and gradual and cyclic late Cenozoic climate change while simultaneously accounting for habitat and biogeographic covariates. Results Climate was the most important predictor of NRI, supporting H1. Current and past climate showed similar relationships with NRI, supporting H2. Conifer NRI was further related to Quaternary climate oscillations and gradual late Cenozoic climate trends, but the shape of the relationships supported neither H3 nor H4. Main conclusions The climate-NRI relationships suggest that late Cenozoic climate consistently influenced the dynamics of conifer speciation, extinction and dispersal, leading to global patterns of phylogenetic assemblage structure. We deduce from the phylogenetic structure that diversification has been highest in warm or dry climates over the last ≥11.6 Myr. The fact that phylogenetic structure is related to climate trends and oscillations indicates that climate change plays an important role in addition to climate per se, but the exact underlying mechanisms remain unclear. Our results suggest that past climate needs to be taken into account when aiming to explain the phylogenetic structure of regional assemblages and other related aspects of biodiversity.

When taxa go extinct, unique evolutionary history is lost. If extinction is selective, and the intrinsic vulnerabilities of taxa show phylogenetic signal, more evolutionary history may be lost than expected under random extinction. Under what conditions this occurs is insufficiently known. We show that late Cenozoic climate change induced phylogenetically selective regional extinction of northern temperate trees because of phylogenetic signal in cold tolerance, leading to significantly and substantially larger than random losses of phylogenetic diversity (PD). The surviving floras in regions that experienced stronger extinction are phylogenetically more clustered, indicating that non-random losses of PD are of increasing concern with increasing extinction severity. Using simulations, we show that a simple threshold model of survival given a physiological trait with phylogenetic signal reproduces our findings. Our results send a strong warning that we may expect future assemblages to be phylogenetically and possibly functionally depauperate if anthropogenic climate change affects taxa similarly.

Accepted manuscript version. Published version available at https://doi.org/10.1016/j.ppees.2017.09.008. Accepted manuscript version, licensed CC BY-NC-ND 4.0. In the face of climate change, populations have two survival options − they can remain in situ and tolerate the new climatic conditions (“stay”), or they can move to track their climatic niches (“go”). For sessile and small-stature organisms like alpine plants, staying requires broad climatic tolerances, realized niche shifts due to changing biotic interactions, acclimation through plasticity, or rapid genetic adaptation. Going, in contrast, requires good dispersal and colonization capacities. Neither the magnitude of climate change experienced locally nor the capacities required for staying/going in response to climate change are constant across landscapes, and both aspects may be strongly affected by local microclimatic variation associated with topographic complexity. We combine ideas from population and community ecology to discuss the effects of topographic complexity in the landscape on the immediate “stay” or “go” opportunities of local populations and communities, and on the selective pressures that may have shaped the stay or go capacities of the species occupying contrasting landscapes. We demonstrate, using example landscapes of different topographical complexity, how species’ thermal niches could be distributed across these landscapes, and how these, in turn, may affect many population and community ecological processes that are related to adaptation or dispersal. Focusing on treeless alpine or Arctic landscapes, where temperature is expected to be a strong determinant, our theorethical framework leads to the hypothesis that populations and communities of topographically complex (rough and patchy) landscapes should be both more resistant and more resilient to climate change than those of topographically simple (flat and homogeneous) landscapes. Our theorethical framework further points to how meta-community dynamics such as mass effects in topographically complex landscapes and extinction lags in simple landscapes, may mask and delay the long-term outcomes of these landscape differences under rapidly changing climates.

Arctic ecosystems face strong changes in snow conditions due to global warming. In contrast to habitat specialists, species occupying a wide range of microhabitats under different snow conditions may better cope with such changes. We studied how growth and reproduction of the dominant dwarf shrub Empetrum hermaphroditum varied among three habitat types differing in winter snow depth and summer irradiation, and whether the observed patterns were consistent along a local climatic gradient (sub-continental vs. sub-oceanic climates) and along a latitudinal gradient (northern Sweden vs. central Norway). Habitat type explained most of the variation in growth and reproduction. Shoots from shallow snow cover and high summer irradiation habitats had higher numbers of flowers and fruits, lower ramet heights, shorter shoot segments, lower numbers of lateral shoots and total biomass but higher leaf density and higher relative leaf allocation than shoots from habitats with higher snow depth and lower summer irradiation. In addition, biomass, leaf allocation and leaf life expectancy were strongly affected by latitude, whereas local climate had strong effects on seed number and seed mass. Empetrum showed high phenotypic trait variation, with a consistent match between local habitat conditions and its growth and reproduction. Although study areas varied strongly with respect to latitude and local climatic conditions, response patterns of growth and reproduction to habitats with different environmental conditions were consistent. Large elasticity of traits suggests that Empetrum may have the potential to cope with changing snow conditions expected in the course of climate change.

The concepts of phylogenetic community structure (PCS) and phylogenetic niche conservatism (PNC) allow ecologists to address the role of species’ evolutionary history in community assembly. It is important to test the role of historical legacies relative to environmental constraints at local scales, where communities are assembled. We studied phylogenetic structure and niche conservatism for palms (Arecaceae) in the 64-km2 Ducke Reserve in the central Amazon, near Manaus. The 72 study plots, each covering 0.1 ha, were distributed regularly in a terra firme forest along a hydro-edaphic gradient. We compared the observed palm PCS with assemblages generated by null models. We also analyzed whether morphological and ecological traits are labile or conserved along the phylogeny and quantified the spatial structure of morphological traits in each plot. We found an overall neutral PCS in combination with low PNC (labile traits), suggesting that evolutionary history poses little constraint on palm community assembly in this Amazonian landscape. Still, there was a tendency towards phylogenetic overdispersion in bottomlands, suggesting competitive exclusion among close relatives or, more likely, environmental filtering acting on convergent traits that affect co-occurrence in flood-prone areas. We conclude that (1) PCS of local communities is random as a whole and morphological traits are overall labile, but that (2) the hydro-edaphic gradient within terra firme forests leads to differences in species co-occurrence so that closely related species occur less often than expected in bottomlands due to diffuse competition among close relatives or environmental filtering on convergent traits.

The extent to which species' ecological and phylogenetic relatedness shape their co-occurrence patterns at large spatial scales remains poorly understood. By quantifying phylogenetic assemblage structure within geographic ranges of >8000 bird species, we show that global co-occurrence patterns are linked - after accounting for regional effects - to key ecological traits reflecting diet, mobility, body size and climatic preference. We found that co-occurrences of carnivorous, migratory and cold-climate species are phylogenetically clustered, whereas nectarivores, herbivores, frugivores and invertebrate eaters tend to be more phylogenetically overdispersed. Preference for open or forested habitats appeared to be independent from the level of phylogenetic clustering. Our results advocate for an extension of the tropical niche conservatism hypothesis to incorporate ecological and life-history traits beyond the climatic niche. They further offer a novel species-oriented perspective on how biogeographic and evolutionary legacies interact with ecological traits to shape global patterns of species coexistence in birds.

Phylogenetic information can provide important insight into the assembly of hyper-diverse tropical rainforest communities. The hierarchical nature and scale dependency of phylogenetic community structure can be used to disentangle complex networks of assembly mechanisms. Here, we outline how particular assembly mechanisms can be identified by analysing phylogenetic community structure on different scales related to space and environment. We present a case study on a large set of species-rich palm communities from the western Amazon basin. We aimed to elucidate the assembly of these communities by separating the roles of broad-scale biogeographic processes, habitat filtering and intra-habitat processes. Overall, we found phylogenetic clustering, which could be caused by a variety of different processes. When we reduced the species pool according to regions, habitats or soil sand content, we found that the structure was consistent with a combination of regional biogeographic differentiation, habitat filtering and within-habitat processes. The habitat filtering effect was strongest, although it varied in strength among communities in different habitats. Some phylogenetic structure could not be accounted for by biogeography or habitat, suggesting that intra-habitat processes such as micro-environmental filtering or biotic interactions also play a role. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ••, ••–••.

Aim It is a central issue in ecology and biogeography to understand what governs community assembly and the maintenance of biodiversity in tropical rain forest ecosystems. A key question is the relative importance of environmental species sorting (niche assembly) and dispersal limitation (dispersal assembly), which we investigate using a large dataset from diverse palm communities. Location Lowland rain forest, western Amazon River Basin, Peru. Methods We inventoried palm communities, registering all palm individuals and recording environmental conditions in 149 transects of 5 m × 500 m. We used ordination, Mantel tests and indicator species analysis (ISA) to assess compositional patterns, species responses to geographical location and environmental factors. Mantel tests were used to assess the relative importance of geographical distance (as a proxy for dispersal limitation) and environmental differences as possible drivers of dissimilarity in palm species composition. We repeated the Mantel tests for subsets of species that differ in traits of likely importance for habitat specialization and dispersal (height and range size). Results We found a strong relationship between compositional dissimilarity and environmental distance and a weaker but also significant relationship between compositional dissimilarity and geographical distance. Consistent with expectations, relationships with environmental and geographical distance were stronger for understorey species than for canopy species. Geographical distance had a higher correlation with compositional dissimilarity for small-ranged species compared with large-ranged species, whereas the opposite was true for environmental distance. The main environmental correlates were inundation and soil nutrient levels. Main conclusions The assembly of palm communities in the western Amazon appears to be driven primarily by species sorting according to hydrology and soil, but with dispersal limitation also playing an important role. The importance of environmental characteristics and geographical distance varies depending on plant height and geographical range size in agreement with functional predictions, increasing our confidence in the inferred assembly mechanisms.

To advance our understanding of the processes that govern the assembly of palm communities and the local coexistence of numerous palm species, we here synthesize available information in the literature on species diversity and growth-form composition in palm communities across the Americas. American palm communities surveyed had 4–48 (median 16) species in study plots covering 0.09–7.2 ha. Climate, soils, hydrology, and topography are the main factors determining palm community species richness. Tropical lowland terra firme rain forests are the most species-rich whereas forests that are inundated or grow on sandy soils or in areas with seasonal climate have much fewer species. Palm communities in the central-western Amazon and in Central America are significantly richer than the average region and those in the Caribbean significantly poorer in species. As for branching, the 789 species of tropical American palms belong to Corner’s model (solitary, 268 species, 33%), Tomlinsons model (cespitose, 521 species, 66%) and Schoute’s model (dichotomous branching, three species

Background The palm family occurs in all tropical and sub-tropical regions of the world. Palms are of high ecological and economical importance, and display complex spatial patterns of species distributions and diversity. Scope This review summarizes empirical evidence for factors that determine palm species distributions, community composition and species richness such as the abiotic environment (climate, soil chemistry, hydrology and topography), the biotic environment (vegetation structure and species interactions) and dispersal. The importance of contemporary vs. historical impacts of these factors and the scale at which they function is discussed. Finally a hierarchical scale framework is developed to guide predictor selection for future studies. Conclusions Determinants of palm distributions, composition and richness vary with spatial scale. For species distributions, climate appears to be important at landscape and broader scales, soil, topography and vegetation at landscape and local scales, hydrology at local scales, and dispersal at all scales. For community composition, soil appears important at regional and finer scales, hydrology, topography and vegetation at landscape and local scales, and dispersal again at all scales. For species richness, climate and dispersal appear to be important at continental to global scales, soil at landscape and broader scales, and topography at landscape and finer scales. Some scale–predictor combinations have not been studied or deserve further attention, e.g. climate on regional to finer scales, and hydrology and topography on landscape and broader scales. The importance of biotic interactions – apart from general vegetation structure effects – for the geographic ecology of palms is generally underexplored. Future studies should target scale–predictor combinations and geographic domains not studied yet. To avoid biased inference, one should ideally include at least all predictors previously found important at the spatial scale of investigation.

Summary 1. Local and regional patterns of plant species richness in tropical rain forests, as well as their possible drivers, remain largely unexplored. The main hypotheses for local species richness (alpha diversity) are (i) local environmental determinism with species-saturated communities, and (ii) regional control, in which the immigration of species from the regional species pool (gamma diversity) determines how many species coexist locally. The species pool hypothesis suggests a combined influence of local and regional drivers on alpha diversity. Differences in gamma diversity may arise from divergent environmental conditions or biogeographic histories. 2. We investigated the cross-scale determinants of palm alpha and gamma diversity across the western Amazon using a large field-based data set: a census of all palm individuals in 312 transects, totalling 98 species. We used regression-based variation partitioning to understand how habitat, topography and region influence alpha diversity, and correlations to assess the importance of the present environment (climate, soil, regional topography) and history (long-term habitat stability) for average regional alpha diversity and gamma diversity, including the link between these two diversity measures (species pool effect). 3. Variation in alpha diversity was primarily explained by region (36%) and habitat (18%), whereas the effect of topography was negligible (1%). Within habitats, region was even more important (up to 69% explained variation). Within regions, habitat and topography covaried and had a variable but an important influence. The pure effect of topography remained of minor importance (up to 13%). 4. Average regional alpha diversity was related to gamma diversity, precipitation seasonality and possibly long-term habitat stability. Gamma diversity was related to long-term habitat stability, and possibly current climate. 5. Synthesis. Gamma diversity strongly influenced alpha diversity, although a clear influence of local environment was also evident, notably habitat type, with a minor, more geographically variable effect of small-scale topography. Apart from gamma diversity, the factor most strongly related to regional alpha diversity was precipitation seasonality, while gamma diversity itself was strongly linked to long-term habitat stability. These results imply that plant species richness is contingent on both contemporary and historical factors with a strong link between local species richness and the regional species pool.

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Manter Laboratory of Parasitology University of Nebraska-Lincoln Robert Bebb Herbarium (OKL) University of Oklahoma Oklahoma Biological Survey Department of Microbiology and Plant Biology Fiocruz Mata Atlântica Fundação Oswaldo Cruz Department of Civil and Environmental Engineering Ehime University Department of Zoology Hungarian Natural History Museum Department of Science Education Hiroshima University Department of Earth Sciences Natural History Museum Kazan Federal University Museum für Naturkunde Leibniz-Institut für Evolutions-und Biodiversitätsforschung Centro de Investigación de la Biodiversidad y Cambio Climático (BioCamb) e Ingeniería en Biodiversidad y Recursos Genéticos Universidad Tecnológica Indoamérica Integrated Taxonomic Information System National Museum of Natural History Smithsonian Institution Natural History Museum of Denmark Department of Botany Szent István University University of Sydney Zoological Museum of Lomonosov Moscow State University Biodiversity Institute University of Kansas CITES Scientific Authority Finnish Museum of Natural History (Botany) University of Helsinki Instituto de Botânica Department of Entomology Kerala Agricultural University University of Michigan Herbarium EEB Chelonian Research Foundation Turtle Conservancy Instituto de Ciencias Marinas y Limnológicas Facultad de Ciencias Universidad Austral de Chile Kansas Biological Survey and the Biodiversity Institute University of Kansas Laboratório de Ecologia e Conservação Universidade Federal do Mato Grosso do Sul Sciences Department Museums Victoria Department of Biology and Museum of Vertebrate Biology Portland State University Department of Biological Sciences Texas Tech University Staatliches Museum für Naturkunde Stuttgart Department of Bioinformatics and Genetics Swedish Museum of Natural History Laboratorio de Sistemática y Biología Comparada de Insectos Instituto de Ciencias Naturales Universidad Nacional de Colombia Global Biodiversity Information Facility Fragile Inheritance Natural History National Focal Point to the Convention on Biological Diversity Royal Belgian Institute of Natural Sciences Department of Biology Pittsburg State University Departamento de Zoologia Instituto de Biociências Universidade de São Paulo Agricultural Research Council Plant Protection Research Instituten National Collection of Insects Section of Mollusks Carnegie Museum of Natural History Department of Zoology Faculty of Science Charles University Museo Argentino de Ciencias Naturales Bernardino Rivadavia National Parks Board Botanic Gardens Department of Ecology and Evolutionary Biology University of Kansas Judicial Commission on Prokaryote Nomenclature Botanische Staatssammlung München and SNSB IT Center Staatliche Naturwissenschaftliche Sammlungen Bayerns Permanent ICN Nomenclature Committee for Fungi International Mycological Association Museo Nacional de Ciencias Naturales (CSIC) Dipartimento di Scienze della Vita e Biologia dei Sistemi Università di Torino Istituto per la Protezione Sostenibile delle Piante sez. di Torino CNR Department of Environmental Sciences University of Basel Allwetterzoo Münster Department of Biology University of Hawai'i at Manoa UNC Herbarium (NCU) North Carolina Botanical Garden University of North Carolina at Chapel Hill Abteilung Morphologie und Systematik der Tiere und Zoologisches Museum Universität Göttingen Unité Mixte de Recherche Peuplements Végétaux et Bioaggresseurs en Milieu Tropical Université de La Réunion Essig Museum of Entomology University of California Berkeley General Committee for Nomenclature [for algae fungi and plants] Royal Botanic Gardens and Domain Trust Smithsonian Marine Station Molecular Ecology and Fisheries Genetics Lab School of Biological Sciences Bangor University Department of Entomology Entomology Research Museum University of California Australian National Insect Collection CSIRO National Research Collections Australian Capital Territory School of Life Sciences Biodiversity Knowledge Integration Center Hasbrouck Insect Collection Arizona State University Landcare Research School of Biological Sciences University of Auckland Institute of Zoology Chinese Academy of Sciences, Chaoyang District Laboratório de Ictiologia Departamento de Zoologia IBB-UNESP Campus de Botucatu Universidade Estadual Paulista (Unesp) Faculdade de Ciências Agrárias e Veterinárias Jaboticabal Departamento de Biologia Aplicada à Agropecuária

Local communities are assembled from larger-scale species pools via dispersal, environmental filtering, biotic interactions and local stochastic demographic processes. The relative importance, scaling and interplay of these assembly processes can be elucidated by comparing local communities to variously circumscribed species pools. Here we present the first study applying this approach to forest tree communities across East Asia, focusing on community phylogenetic structure and using data from a global network of tropical, subtropical and temperate forest plots. We found that Net Relatedness Index (NRI) and Nearest Taxon Index (NTI) values were generally lower with geographically broad species pools (global and Asian species pools) than with an East Asian species pool, except that global species pool produced higher NTI than the East Asian species pool. The lower NRI for the global relative to the East Asian species pool may indicate an important role of intercontinental migration during the Neogene and Quaternary and climatic conservatism in shaping the deeper phylogenetic structure of tree communities in East Asia. In contrast, higher NTI for the global relative to the East Asian species pool is consistent with recent localized diversification determining the shallow phylogenetic structure.