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Spatial phylogenetics of Fagales: Investigating drivers of temperate forest distributions
Abstract
Aim
Quantifying the phylogenetic diversity of temperate trees is essential for understanding the processes that have shaped the modern distribution of temperate broadleaf forest and other major forest biomes. Here, we focus on Fagales, an iconic member of forests worldwide, to uncover global diversity and endemism patterns and investigate the distribution of root nodule symbiosis (RNS), an important morphological specialisation in this clade, as a key factor behind these patterns.
Location
Global.
Taxon
Fagales.
Methods
We combined phylogenetic data covering 60.2% of living species, fine‐scale distribution models covering 90% of species, and nodulation data covering all species to investigate the distribution of species richness and phylogenetic diversity at fine spatial scales compared to the distribution of RNS. We identify abiotic environmental factors associated with RNS and with Fagales diversity in general.
Results
We find the highest species richness in temperate east Asia, eastern North America, and equatorial montane regions of Asia and Central America. By contrast, relative phylogenetic diversity (RPD) is highest at higher latitudes, where RNS also predominates. We found a strong spatial structuring of regionalisations of Fagales floras, reflecting distinct Northern and Southern Hemisphere floras (except a unique Afro‐Boreal region), each with distinct RNS‐environment relationships.
Main Conclusions
Although species richness and phylogenetic regionalisation for Fagales accord well with traditional biogeographic concepts for temperate forests, this is not the case for RPD. RNS is almost universal in the highest RPD regions, which may reflect ecological filtering promoting RNS in these regions. Our results highlight the utility of global‐scale, clade‐specific spatial phylogenetics and its utility for understanding drivers of diversity in species‐rich clades.
Astragalus (Fabaceae) is astoundingly diverse in temperate, cold arid regions of Earth, positioning this group as a model clade for investigating the distribution of plant diversity in the face of environmental challenges. Here, we identify the spatial distribution of diversity and endemism in Astragalus using species distribution models for 752 species and a phylogenetic tree comprising 847 species. We integrated these to map centers of species richness (SR) and relative phylogenetic diversity (RPD) and used randomization approaches to investigate centers of endemism. We also used clustering methods to identify phylogenetic regionalizations. We then assembled predictor variables of current climate conditions to test environmental factors predicting these phylogenetic diversity results, especially temperature and precipitation seasonality. We find that SR centers are distributed globally at temperate middle latitudes in arid regions, but the Mediterranean Basin is the most important center of RPD. Endemism centers also occur globally, but Iran represents a key endemic area with a concentration of both paleo‐ and neoendemism. Phylogenetic regionalization recovered an east‐west gradient in Eurasia and an amphitropical disjunction across North and South America; American phyloregions are overall most closely related to east and central Asia. SR, RPD, and lineage turnover are driven mostly by precipitation and seasonality, but endemism is driven primarily by diurnal temperature variation. Endemism and regionalization results point to western Asia and especially Iran as a biogeographic gateway between Europe and Asia. RPD and endemism highlight the importance of temperature and drought stress in determining plant diversity and endemism centers.
Root nodule symbiosis (RNS) is a complex trait that enables plants to access atmospheric nitrogen converted into usable forms through a mutualistic relationship with soil bacteria. Pinpointing the evolutionary origins of RNS is critical for understanding its genetic basis, but building this evolutionary context is complicated by data limitations and the intermittent presence of RNS in a single clade of ca. 30,000 species of flowering plants, i.e., the nitrogen-fixing clade (NFC). We developed the most extensive de novo phylogeny for the NFC and an RNS trait database to reconstruct the evolution of RNS. Our analysis identifies evolutionary rate heterogeneity associated with a two-step process: An ancestral precursor state transitioned to a more labile state from which RNS was rapidly gained at multiple points in the NFC. We illustrate how a two-step process could explain multiple independent gains and losses of RNS, contrary to recent hypotheses suggesting one gain and numerous losses, and suggest a broader phylogenetic and genetic scope may be required for genome-phenome mapping.
Rarely have studies assessed Odonata diversity for the entire Nearctic realm by including
Canada, the United States, and Mexico. For the first time, we explored Odonata diversity in this region according to a definition of natural community assemblages and generated species distribution models (SDMs). Species occurrence data were assembled by reviewing databases of specimens held by significant Odonata repositories and through an extensive search of literature references. Species were categorized as forest-dependent or non-forest-dependent, as lentic or lotic-dependent, and according to conservation status. Predicted distributions were stacked for all species across their entire ranges, including areas outside of the Nearctic. Species richness and corrected weighted endemism (CWE) were then calculated for each grid cell. We found a pattern of greater species richness in the eastern portion of the Nearctic, which can be explained by the higher aquatic habitat diversity at micro and macroscales east of the Rocky Mountains, promoting niche partitioning and specialization. In the Nearctic region, the southeastern US has the highest number of endemic species of dragonflies and damselflies; this degree of endemism is likely due to glacial refuges providing a foundation for the evolution of a rich and unique biota.
Aim
Nitrogen (N)‐fixing plants are an important component of global plant communities, but the drivers of N‐fixing plant diversity, especially in temperate regions, remain underexplored. Here, we examined broad‐scale patterns of N‐fixing and non‐fixing plant phylogenetic diversity (PD) and species richness (SR) across a wide portion of temperate North America, focusing on relationships with soil N and aridity. We also tested whether exotic species, with and without N‐fixing symbiosis, have fewer abiotic limitations compared with native species.
Location
USA and Puerto Rico.
Time period
Current.
Major taxa studied
Vascular plants, focusing on N‐fixing groups (orders Fabales, Fagales, Rosales and Cucurbitales).
Methods
We subset National Ecological Observatory Network (NEON) plant plot data from all sites along two axes (N fixing–non‐N fixing and native–exotic), calculating plot‐level SR, PD and mean pairwise phylogenetic distance (MPD). We then used linear mixed models to investigate relationships between diversity values and key soil measurements, along with aridity, temperature and fire frequency.
Results
Aridity was the sole predictor of proportional phylogenetic diversity of N fixers. The SR of N fixers still decreased marginally in arid regions, whereas native N‐fixer MPD increased with aridity, indicative of unique lineages of N fixers in the driest conditions, in contrast to native non‐N fixers. The SR of both native N fixers and non‐N fixers increased in low‐N soils. Aridity did not affect SR of exotic non‐N fixers, unlike other groups, whereas exotic N fixers showed lower MPD in increasingly high‐N soils, suggesting filtering, contrary what was found for native N fixers.
Main conclusions
Our results suggest that it is not nitrogen, or any soil nutrient, that has the strongest effect on the relative success of N fixers in plant communities. Rather, aridity is the key driver, at least for native species, in line with empirical results from other biomes and increased understanding of N fixation as a key mechanism to avoid water loss.
The documentation of biodiversity distribution through species range identification is crucial for macroecology, biogeography, conservation, and restoration. However, for plants, species range maps remain scarce and often inaccurate.
We present a novel approach to map species ranges at a global scale, integrating polygon mapping and species distribution modelling (SDM). We develop a polygon mapping algorithm by considering distances and nestedness of occurrences. We further apply an SDM approach considering multiple modelling algorithms, complexity levels, and pseudo‐absence selections to map the species at a high spatial resolution and intersect it with the generated polygons.
We use this approach to construct range maps for all 1957 species of Fagales and Pinales with data compilated from multiple sources. We construct high‐resolution global species richness maps of these important plant clades, and document diversity hotspots for both clades in southern and south‐western China, Central America, and Borneo. We validate the approach with two representative genera, Quercus and Pinus, using previously published coarser range maps, and find good agreement.
By efficiently producing high‐resolution range maps, our mapping approach offers a new tool in the field of macroecology for studying global species distribution patterns and supporting ongoing conservation efforts.
Existing global regionalization schemes for plants consider the compositional affinities among biotas, but these have not explicitly considered phylogenetic information. Here, we present for the first time, a phytogeographical delineation of the global vascular flora based on species‐level evolutionary relationships.
We analysed 8737 820 geographical occurrence records for vascular plants together with a time‐calibrated phylogeny including 67 269 species. We constructed a global phylogenetic regionalization by estimating species composition and phylogenetic beta diversity among 200 km × 200 km grid cells across the world.
We identified de novo 16 phytogeographical units that are deeply split into two clusters: Laurasian and Gondwanan. Our regionalization broadly matches previous schemes, but also highlights the separation of the Gondwanan biota into an Holotropical cluster and an Australian–Neozealandic–Patagonian cluster. In contrast, no clear split among Laurasian and Gondwanan biotas was retrieved when omitting phylogenetic information.
The integration of phylogenetic and geographical information provides new insights into the delineation of phytogeographical areas and their historical relationships, enabling the identification of three large, clearly differentiated biotas, here referred to as kingdoms: Holarctic, Holotropical, and Austral. Our results provide further evidence for delineating transition zones and show a clear latitudinal pattern of increasing evolutionary distinctiveness towards the poles.
Premise:
Large phylogenetic data sets have often been restricted to small numbers of loci from GenBank, and a vetted sampling-to-sequencing phylogenomic protocol scaling to thousands of species is not yet available. Here, we report a high-throughput collections-based approach that empowers researchers to explore more branches of the tree of life with numerous loci.
Methods:
We developed an integrated Specimen-to-Laboratory Information Management System (SLIMS), connecting sampling and wet lab efforts with progress tracking at each stage. Using unique identifiers encoded in QR codes and a taxonomic database, a research team can sample herbarium specimens, efficiently record the sampling event, and capture specimen images. After sampling in herbaria, images are uploaded to a citizen science platform for metadata generation, and tissue samples are moved through a simple, high-throughput, plate-based herbarium DNA extraction and sequencing protocol.
Results:
We applied this sampling-to-sequencing workflow to ~15,000 species, producing for the first time a data set with ~50% taxonomic representation of the "nitrogen-fixing clade" of angiosperms.
Discussion:
The approach we present is appropriate at any taxonomic scale and is extensible to other collection types. The widespread use of large-scale sampling strategies repositions herbaria as accessible but largely untapped resources for broad taxonomic sampling with thousands of species.
Aim
It is well known that the distribution of species diversity is spatially heterogeneous, but understanding the factors contributing to this heterogeneity and to the formation of biodiversity hotspots remains a challenge. Here, we seek to improve our understanding of how historical, ecological and evolutionary processes contribute to current patterns of global fern diversity.
Location
Worldwide.
Taxon
Ferns.
Methods
To evaluate the drivers of global fern diversity, we integrate over 800,000 georeferenced species occurrence records of nearly 8000 species, a time‐calibrated phylogeny and seven climate and environmental layers. We use these data to summarize diversity and evolutionary patterns at a resolution of 100 × 100 km, and identify hotspots of fern species richness and endemism. We compare these hotspots to neighbouring non‐hotspot regions to provide insight into the factors controlling global patterns of fern diversity.
Results
Tropical and subtropical mountains harbour a disproportionate amount of species relative to the land area they occupy; 58% of global species richness occur in eight principally montane hotspots together comprising just 7% of Earth’s land area. We identify hotspots of fern species richness and endemism that are universally characterized by disproportionately high ecological variation. We demonstrate that total fern species richness scales linearly with available climate space at regional and global scales.
Main Conclusions
Areas of high environmental heterogeneity harbour a disproportionate amount of fern species, and global patterns of extant fern diversity reflect the distribution of these areas, especially in mountains at lower latitudes. Persistence of ancient lineages in areas with long‐term climatic stability helps explain exceptional endemism in regions such as Malesia.
The symbiosis between plants and nitrogen‐fixing bacteria is widespread among legumes and actinorhizal plants within the nitrogen‐fixing root nodule (NFN) clade. However, there are major differences, as well as similarities, in the symbioses between actinorhizal plants and Frankia and those of legumes and their associated rhizobia.
This review provides an overview of NFN symbioses. We outline the evolution and biogeography of actinorhizal plants and legumes and compare and contrast their microsymbionts and symbiotic processes.
Within the NFN clade, a far greater number of nodulated legumes exists, compared with actinorhizal plants, and legumes have a much wider biogeographical distribution. There are genetic and physiological differences between free‐living diazotrophic Frankia and the phylogenetically diverse rhizobia, most strains of which are unable to fix N2 ex planta. Actinorhizal nodules are modified lateral roots with a central vascular system, whereas legume nodules are stem‐like organs with peripheral vascular systems. Most legumes contain their microsymbionts within symbiosomes, rather than the infection threads found in actinorhizal nodule cells. Legumes have greater control of their microsymbionts, and those within the Inverted Repeat Lacking Clade impose terminal differentiation on their bacteroids. Legumes also have effective processes for autoregulation of nodulation and downregulation of N2 fixation in response to high levels of soil N. These features of the legume‐rhizobia symbiosis have led to increased efficiencies in N2 fixation.
Synthesis. We suggest that these characteristic features of the legume‐rhizobia symbiosis, specifically legumes' greater flexibility in the choice of microsymbiont partner and the evolution of increased efficiencies in N2 fixation, are factors that can explain why the majority of species within the Leguminosae have retained the ability to nodulate and how this has contributed to their evolutionary success.
Conflicting relationships have been found between diversification rate and temperature across disparate clades of life. Here, we use a supermatrix comprising nearly 20,000 species of rosids-a clade of~25% of all angiosperm species-to understand global patterns of diversification and its climatic association. Our approach incorporates historical global temperature, assessment of species' temperature niche, and two broad-scale characterizations of tropical versus non-tropical niche occupancy. We find the diversification rates of most subclades dramatically increased over the last 15 million years (Myr) during cooling associated with global expansion of temperate habitats. Climatic niche is negatively associated with diversification rates, with tropical rosids forming older communities and experiencing speciation rates~2-fold below rosids in cooler climates. Our results suggest long-term cooling had a disproportionate effect on non-tropical diversification rates, leading to dynamic young communities outside of the tropics, while relative stability in tropical climes led to older, slower-evolving but still species-rich communities.
To better understand how ecosystems are changing, a multifaceted approach to measuring biodiversity that considers species richness (SR) and evolutionary history across spatial scales is needed. Here, we compiled 162 datasets for fish, bird and plant assemblages across the globe and measured how taxonomic and phylogenetic diversity changed at different spatial scales (within site α diversity and between sites spatial β diversity). Biodiversity change is measured from these datasets in three ways: across land use gradients, from species lists, and through sampling of the same locations across two time periods. We found that local SR and phylogenetic α diversity (Faith's PD (phylogenetic diversity)) increased for all taxonomic groups. However, when measured with a metric that is independent of SR (phylogenetic species variation, PSV), phylogenetic α diversity declined for all taxonomic groups. Land use datasets showed declines in SR, Faith's PD and PSV. For all taxonomic groups and data types, spatial taxonomic and phylogenetic β diversity decreased when measured with Sorensen dissimilarity and phylogenetic Sorensen dissimilarity, respectively, providing strong evidence of global biotic homogenization. The decoupling of α and β diversity, as well as taxonomic and phylogenetic diversity, highlights the need for a broader perspective on contemporary biodiversity changes. Conservation and environmental policy decisions thus need to consider biodiversity beyond local SR to protect biodiversity and ecosystem services.
The historical course of evolutionary diversification shapes the current distribution of biodiversity, but the main forces constraining diversification are still a subject of debate. We unveil the evolutionary structure of tree species assemblages across the Americas to assess whether an inability to move or an inability to evolve is the predominant constraint in plant diversification and biogeography. We find a fundamental divide in tree lineage composition between tropical and extratropical environments, defined by the absence versus presence of freezing temperatures. Within the Neotropics, we uncover a further evolutionary split between moist and dry forests. Our results demonstrate that American tree lineages tend to retain their ancestral environmental relationships and that phylogenetic niche conservatism is the primary force structuring the distribution of tree biodiversity. Our study establishes the pervasive importance of niche conservatism to community assembly even at intercontinental scales.
North America is a large continent with extensive climatic, geological, soil, and biological diversity. That biota is under threat from habitat destruction and climate change, making a quantitative assessment of biodiversity of critical importance. Rapid digitization of plant specimen records and accumulation of DNA sequence data enable a much‐needed broad synthesis of species occurrences with phylogenetic data. Here we attempted the first such synthesis of a flora from such a large and diverse part of the world: all seed plants for the North American continent (here defined to include Canada, United States, and Mexico) with a focus on examining phylogenetic diversity and endemism. We collected digitized plant specimen records and chose a coarse grain for analysis, recognizing that this grain is currently necessary for reasonable completeness per sampling unit. We found that raw richness and endemism patterns largely support previous hypotheses of biodiversity hotspots. Application of phylogenetic metrics and a randomization test revealed novel results, including significant phylogenetic clustering across the continent, a striking east‐west geographic difference in the distribution of branch lengths, and the discovery of centers of neo‐ and paleo‐endemism in Mexico, the southwestern USA, and the southeastern USA. Finally, our examination of phylogenetic beta‐diversity provides a new approach to comparing centers of endemism. We discuss the empirical challenges of working at the continental scale, and the need for more sampling across large parts of the continent, for both DNA data for terminal taxa and spatial data for poorly understood regions, to confirm and extend these results.
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Should we build our own phylogenetic trees based on gene sequence data, or can we simply use available synthesis phylogenies? This is a fundamental question that any study involving a phylogenetic framework must face at the beginning of the project. Building a phylogeny from gene sequence data (purpose‐built phylogeny) requires more effort, expertise, and cost than subsetting an already available phylogeny (synthesis‐based phylogeny). However, we still lack a comparison of how these two approaches to building phylogenetic trees influence common community phylogenetic analyses such as comparing community phylogenetic diversity and estimating trait phylogenetic signal. Here, we generated three purpose‐built phylogenies and their corresponding synthesis‐based trees (two from Phylomatic and one from the Open Tree of Life, OTL). We simulated 1,000 communities and 12,000 continuous traits along each purpose‐built phylogeny. We then compared the effects of different trees on estimates of phylogenetic diversity (alpha and beta) and phylogenetic signal (Pagel's λ and Blomberg's K). Synthesis‐based phylogenies generally yielded higher estimates of phylogenetic diversity when compared to purpose‐built phylogenies. However, resulting measures of phylogenetic diversity from both types of phylogenies were highly correlated (Spearman's ρ > 0.8 in most cases). Mean pairwise distance (both alpha and beta) is the index that is most robust to the differences in tree construction that we tested. Measures of phylogenetic diversity based on the OTL showed the highest correlation with measures based on the purpose‐built phylogenies. Trait phylogenetic signal estimated with synthesis‐based phylogenies, especially from the OTL, was also highly correlated with estimates of Blomberg's K or close to Pagel's λ from purpose‐built phylogenies when traits were simulated under Brownian motion. For commonly employed community phylogenetic analyses, our results justify taking advantage of recently developed and continuously improving synthesis trees, especially the Open Tree of Life.
Motivation:
Phylogenies are important for fundamental biological research, but also have numerous applications in biotechnology, agriculture, and medicine. Finding the optimal tree under the popular maximum likelihood (ML) criterion is known to be NP-hard. Thus, highly optimized and scalable codes are needed to analyze constantly growing empirical datasets.
Results:
We present RAxML-NG, a from scratch re-implementation of the established greedy tree search algorithm of RAxML/ExaML. RAxML-NG offers improved accuracy, flexibility, speed, scalability, and usability compared to RAxML/ExaML. On taxon-rich datasets, RAxML-NG typically finds higher-scoring trees than IQTree, an increasingly popular recent tool for ML-based phylogenetic inference (although IQ-Tree shows better stability). Finally, RAxML-NG introduces several new features, such as the detection of terraces in tree space and a the recently introduced transfer bootstrap support metric.
Availability:
The code is available under GNU GPL at https://github.com/amkozlov/raxml-ng. RAxML-NG web service (maintained by Vital-IT) is available at https://raxml-ng.vital-it.ch/.
Supplementary information:
Supplementary data are available at Bioinformatics online.
Symbiotic nitrogen (N)‐fixing trees can provide large quantities of new N to ecosystems, but only if they are sufficiently abundant. The overall abundance and latitudinal abundance distributions of N‐fixing trees are well characterised in the Americas, but less well outside the Americas.
Here, we characterised the abundance of N‐fixing trees in a network of forest plots spanning five continents, ~5,000 tree species and ~4 million trees. The majority of the plots (86%) were in America or Asia. In addition, we examined whether the observed pattern of abundance of N‐fixing trees was correlated with mean annual temperature and precipitation.
Outside the tropics, N‐fixing trees were consistently rare in the forest plots we examined. Within the tropics, N‐fixing trees were abundant in American but not Asian forest plots (~7% versus ~1% of basal area and stems). This disparity was not explained by mean annual temperature or precipitation. Our finding of low N‐fixing tree abundance in the Asian tropics casts some doubt on recent high estimates of N fixation rates in this region, which do not account for disparities in N‐fixing tree abundance between the Asian and American tropics.
Synthesis. Inputs of nitrogen to forests depend on symbiotic nitrogen fixation, which is constrained by the abundance of N‐fixing trees. By analysing a large dataset of ~4 million trees, we found that N‐fixing trees were consistently rare in the Asian tropics as well as across higher latitudes in Asia, America and Europe. The rarity of N‐fixing trees in the Asian tropics compared with the American tropics might stem from lower intrinsic N limitation in Asian tropical forests, although direct support for any mechanism is lacking. The paucity of N‐fixing trees throughout Asian forests suggests that N inputs to the Asian tropics might be lower than previously thought.
The grass subfamily Pooideae dominates the grass floras in cold temperate regions, and has evolved complex physiological adaptations to cope with extreme environmental conditions like frost, winter and seasonality. One such adaptation is cold acclimation, wherein plants increase their frost tolerance in response to gradually falling temperatures and shorter days in the autumn. However, understanding how complex traits like cold acclimation evolve remains a major challenge in evolutionary biology. Here, we investigated the evolution of cold acclimation in Pooideae, and found that a phylogenetically diverse set of Pooideae species displayed cold acclimation capacity. However, comparing differential gene expression after cold treatment in transcriptomes of five phylogenetically diverse species revealed widespread species-specific responses of genes with conserved sequences. Furthermore, we studied the correlation between gene family size and number of cold responsive genes, as well as between selection pressure on coding sequences of genes and their cold responsiveness. We saw evidence of protein-coding and regulatory sequence evolution, as well as origin of novel genes and functions contributing towards evolution of a cold response in Pooideae. Our results reflect that selection pressure resulting from global cooling must have acted on already diverged lineages. Nevertheless, conservation of cold induced gene expression of certain genes indicates that the Pooideae ancestor may have possessed some molecular machinery to mitigate cold stress. Evolution of adaptations to seasonally cold climates is regarded particularly difficult. How Pooideae evolved to transition from tropical to temperate biomes sheds light on how complex traits evolve in the light of climate changes.
Species occurrence records from online databases are an indispensable resource in ecological, biogeographical and palaeontological research. However, issues with data quality, especially incorrect geo‐referencing or dating, can diminish their usefulness. Manual cleaning is time‐consuming, error prone, difficult to reproduce and limited to known geographical areas and taxonomic groups, making it impractical for datasets with thousands or millions of records.
Here, we present CoordinateCleaner , an r ‐package to scan datasets of species occurrence records for geo‐referencing and dating imprecisions and data entry errors in a standardized and reproducible way. CoordinateCleaner is tailored to problems common in biological and palaeontological databases and can handle datasets with millions of records. The software includes (a) functions to flag potentially problematic coordinate records based on geographical gazetteers, (b) a global database of 9,691 geo‐referenced biodiversity institutions to identify records that are likely from horticulture or captivity, (c) novel algorithms to identify datasets with rasterized data, conversion errors and strong decimal rounding and (d) spatio‐temporal tests for fossils.
We describe the individual functions available in CoordinateCleaner and demonstrate them on more than 90 million occurrences of flowering plants from the Global Biodiversity Information Facility (GBIF) and 19,000 fossil occurrences from the Palaeobiology Database (PBDB). We find that in GBIF more than 3.4 million records (3.7%) are potentially problematic and that 179 of the tested contributing datasets (18.5%) might be biased by rasterized coordinates. In PBDB, 1205 records (6.3%) are potentially problematic.
All cleaning functions and the biodiversity institution database are open‐source and available within the CoordinateCleaner r ‐package.
Recent availability of biodiversity data resources has enabled an unprecedented ability to estimate phylogenetically based biodiversity metrics over broad scales. Such approaches elucidate ecological and evolutionary processes yielding a biota and help guide conservation efforts. However, the choice of appropriate phylogenetic resources and underlying input data uncertainties may affect interpretation. Here, we address how differences among phylogenetic source trees and levels of phylogenetic uncertainty affect these metrics and test existing hypotheses regarding geographic biodiversity patterns across the diverse vascular plant flora of Florida, US. Ecological niche models for 1,490 Florida species were combined with a “purpose-built” phylogenetic tree (phylogram and chronogram), as well as with trees derived from community resources (Phylomatic and Open Tree of Life). There were only modest differences in phylodiversity metrics given the phylogenetic source tree and taking into account the level of phylogenetic uncertainty; we identify similar areas of conservation interest across Florida regardless of the method used.
Contents
Summary
669
I.
Model clades for the study and integration of ecology and evolution
670
II.
Oaks: an important model clade
671
III.
Insights from the history of the American oaks for understanding community assembly and ecosystem dominance
673
IV.
Bridging the gap between micro‐ and macroevolutionary processes relevant to ecology
679
V.
How do we reconcile evidence for adaptive evolution with niche conservatism and long‐term stasis?
682
VI.
High plasticity and within‐population genetic variation contribute to population persistence
683
VII.
Emerging technologies for tracking functional change
685
VIII.
Conclusions
685
Acknowledgements
686
References
686
Summary
Ecologists and evolutionary biologists are concerned with explaining the diversity and composition of the natural world and are aware of the inextricable linkages between ecological and evolutionary processes that maintain the Earth's life support systems. Yet examination of these linkages remains challenging due to the contrasting nature of focal systems and research approaches. Model clades provide a critical means to integrate ecology and evolution, as illustrated by the oaks (genus Quercus), an important model clade, given their ecological dominance, remarkable diversity, and growing phylogenetic, genomic, and ecological data resources. Studies of the clade reveal that their history of sympatric parallel adaptive radiation continues to influence community assembly today, highlighting questions on the nature and extent of coexistence mechanisms. Flexible phenology and hydraulic traits, despite evolutionary stasis, may have enabled adaptation to a wide range of environments within and across species, contributing to their high abundance and diversity. The oaks offer fundamental insights at the intersection of ecology and evolution on the role of diversification in community assembly processes, on the importance of flexibility in key functional traits in adapting to new environments, on factors contributing to persistence of long‐lived organisms, and on evolutionary legacies that influence ecosystem function.
The latitudinal diversity gradient—the tendency for more species to occur toward the equator—is the dominant pattern of life on Earth, yet the mechanisms responsible for it remain largely unexplained. Recently, the analysis of global data has led to advances in understanding, but these advances have been mostly limited to vertebrates and trees and have not provided consensus answers. Here we synthesize large-scale geographic, phylogenetic, and fossil data for an exemplar invertebrate group—ants—and investigate whether the latitudinal diversity gradient arose due to higher rates of net diversification in the tropics, or due to a longer time period to accumulate diversity due to Earth’s climatic history. We find that latitudinal affinity is highly conserved, temperate clades are young and clustered within tropical clades, and diversification rate shows no systematic variation with latitude. These results indicate that diversification time—and not rate—is the main driver of the diversity gradient in ants.
Topographic variation underpins a myriad of patterns and processes in hydrology, climatology, geography and ecology and is key to understanding the variation of life on the planet. A fully standardized and global multivariate product of different terrain features has the potential to support many large-scale research applications, however to date, such datasets are unavailable. Here we used the digital elevation model products of global 250 m GMTED2010 and near-global 90 m SRTM4.1dev to derive a suite of topographic variables: elevation, slope, aspect, eastness, northness, roughness, terrain roughness index, topographic position index, vector ruggedness measure, profile/tangential curvature, first/second order partial derivative, and 10 geomorphological landform classes. We aggregated each variable to 1, 5, 10, 50 and 100 km spatial grains using several aggregation approaches. While a cross-correlation underlines the high similarity of many variables, a more detailed view in four mountain regions reveals local differences, as well as scale variations in the aggregated variables at different spatial grains. All newly-developed variables are available for download at Data Citation 1 and for download and visualization at http://www.earthenv.org/topography.
High species diversity may result from recent rapid speciation in a ‘cradle’ and/or the gradual accumulation and preservation of species over time in a ‘museum’1,2. China harbours nearly 10% of angiosperm species worldwide and has long been considered as both a museum, owing to the presence of many species with hypothesized ancient origins3,4, and a cradle, as many lineages have originated as recent topographic changes and climatic shifts—such as the formation of the Qinghai–Tibetan Plateau and the development of the monsoon—provided new habitats that promoted remarkable radiation⁵. However, no detailed phylogenetic study has addressed when and how the major components of the Chinese angiosperm flora assembled to form the present-day vegetation. Here we investigate the spatio-temporal divergence patterns of the Chinese flora using a dated phylogeny of 92% of the angiosperm genera for the region, a nearly complete species-level tree comprising 26,978 species and detailed spatial distribution data. We found that 66% of the angiosperm genera in China did not originate until early in the Miocene epoch (23 million years ago (Mya)). The flora of eastern China bears a signature of older divergence (mean divergence times of 22.04–25.39 Mya), phylogenetic overdispersion (spatial co-occurrence of distant relatives) and higher phylogenetic diversity. In western China, the flora shows more recent divergence (mean divergence times of 15.29–18.86 Mya), pronounced phylogenetic clustering (co-occurrence of close relatives) and lower phylogenetic diversity. Analyses of species-level phylogenetic diversity using simulated branch lengths yielded results similar to genus-level patterns. Our analyses indicate that eastern China represents a floristic museum, and western China an evolutionary cradle, for herbaceous genera; eastern China has served as both a museum and a cradle for woody genera. These results identify areas of high species richness and phylogenetic diversity, and provide a foundation on which to build conservation efforts in China.
Background
California is a world floristic biodiversity hotspot where the terms neo- and paleo-endemism were first applied. Using spatial phylogenetics, it is now possible to evaluate biodiversity from an evolutionary standpoint, including discovering significant areas of neo- and paleo-endemism, by combining spatial information from museum collections and DNA-based phylogenies. Here we used a distributional dataset of 1.39 million herbarium specimens, a phylogeny of 1083 operational taxonomic units (OTUs) and 9 genes, and a spatial randomization test to identify regions of significant phylogenetic diversity, relative phylogenetic diversity, and phylogenetic endemism (PE), as well as to conduct a categorical analysis of neo- and paleo-endemism (CANAPE).
Results
We found (1) extensive phylogenetic clustering in the South Coast Ranges, southern Great Valley, and deserts of California; (2) significant concentrations of short branches in the Mojave and Great Basin Deserts and the South Coast Ranges and long branches in the northern Great Valley, Sierra Nevada foothills, and the northwestern and southwestern parts of the state; (3) significant concentrations of paleo-endemism in Northwestern California, the northern Great Valley, and western Sonoran Desert, and neo-endemism in the White-Inyo Range, northern Mojave Desert, and southern Channel Islands. Multiple analyses were run to observe the effects on significance patterns of using different phylogenetic tree topologies (uncalibrated trees versus time-calibrated ultrametric trees) and using different representations of OTU ranges (herbarium specimen locations versus species distribution models).
Conclusions
These analyses showed that examining the geographic distributions of branch lengths in a statistical framework adds a new dimension to California floristics that, in comparison with climatic data, helps to illuminate causes of endemism. In particular, the concentration of significant PE in more arid regions of California extends previous ideas about aridity as an evolutionary stimulus. The patterns seen are largely robust to phylogenetic uncertainty and time calibration but are sensitive to the use of occurrence data versus modeled ranges, indicating that special attention toward improving geographic distributional data should be top priority in the future for advancing understanding of spatial patterns of biodiversity.
Electronic supplementary material
The online version of this article (doi:10.1186/s12915-017-0435-x) contains supplementary material, which is available to authorized users.
We created a new dataset of spatially interpolated monthly climate data for global land areas at a very high spatial resolution (approximately 1 km 2). We included monthly temperature (minimum, maximum and average), precipitation, solar radiation, vapour pressure and wind speed, aggregated across a target temporal range of 1970–2000, using data from between 9000 and 60 000 weather stations. Weather station data were interpolated using thin-plate splines with covariates including elevation, distance to the coast and three satellite-derived covariates: maximum and minimum land surface temperature as well as cloud cover, obtained with the MODIS satellite platform. Interpolation was done for 23 regions of varying size depending on station density. Satellite data improved prediction accuracy for temperature variables 5–15% (0.07–0.17 ∘ C), particularly for areas with a low station density, although prediction error remained high in such regions for all climate variables. Contributions of satellite covariates were mostly negligible for the other variables, although their importance varied by region. In contrast to the common approach to use a single model formulation for the entire world, we constructed the final product by selecting the best performing model for each region and variable. Global cross-validation correlations were ≥ 0.99 for temperature and humidity, 0.86 for precipitation and 0.76 for wind speed. The fact that most of our climate surface estimates were only marginally improved by use of satellite covariates highlights the importance having a dense, high-quality network of climate station data.
This software note announces a new open-source release of the Maxent software for modeling species distributions from occurrence records and environmental data, and describes a new R package for fitting such models. The new release (Version 3.4.0) will be hosted online by the American Museum of Natural History, along with future versions. It contains small functional changes, most notably use of a complementary log-log (cloglog) transform to produce an estimate of occurrence probability. The cloglog transform derives from the recently-published interpretation of Maxent as an inhomogeneous Poisson process (IPP), giving it a stronger theoretical justification than the logistic transform which it replaces by default. In addition, the new R package, maxnet, fits Maxent models using the glmnet package for regularized generalized linear models. We discuss the implications of the IPP formulation in terms of model inputs and outputs, treating occurrence records as points rather than grid cells and interpreting the exponential Maxent model (raw output) as as an estimate of relative abundance. With these two open-source developments, we invite others to freely use and contribute to the software. This article is protected by copyright. All rights reserved.
http://www.earth-syst-sci-data.net/9/1/2017/
The aim of the World Soil Information Service (WoSIS) is to serve quality-assessed, georeferenced soil data (point, polygon, and grid) to the international community upon their standardisation and harmonisation. So far, the focus has been on developing procedures for legacy point data with special attention to the selection of soil analytical and physical properties considered in the GlobalSoilMap specifications (e.g. organic carbon, soil pH, soil texture (sand, silt, and clay), coarse fragments ( > 2 mm), cation exchange capacity, electrical conductivity, bulk density, and water holding capacity). Profile data managed in WoSIS were contributed by a wide range of soil data providers; the data have been described, sampled, and analysed according to methods and standards in use in the originating countries. Hence, special attention was paid to measures for soil data quality and the standardisation of soil property definitions, soil property values, and soil analytical method descriptions. At the time of writing, the full WoSIS database contained some 118 400 unique “shared” soil profiles, of which some 96 000 are georeferenced within defined limits. In total, this corresponds with over 31 million soil records, of which some 20 % have so far been quality-assessed and standardised using the sequential procedure discussed in this paper. The number of measured data for each property varies between profiles and with depth, generally depending on the purpose of the initial studies. Overall, the data lineage strongly determined which data could be standardised with acceptable confidence in accord with WoSIS procedures, corresponding to over 4 million records for 94 441 profiles. The publicly available data – WoSIS snapshot of July 2016 – are persistently accessible from ISRIC WDC-Soils through doi:10.17027/isric-wdcsoils.20160003.
Tropical savannas are hypothesized to be hot spots of nitrogen fixer diversity and activity
because of the high disturbance and low nitrogen characteristic of savanna landscapes. Here we
compare the abundances of nitrogen-fixing and non-fixing trees in both tropical savannas and
tropical forests under climatically equivalent conditions, using plant inventory studies across 566
plots in South America and Africa. A single factor – aridity – explained 19-54% of the variance
in fixer abundance, and unexpectedly was more important than fire frequency, biome, and
continent. Nitrogen fixers were more abundant in arid environments; as a result, African
savannas, which tend to be drier, were richer in nitrogen fixers than South American savannas.
Fixer abundance converged on similar levels in forests in both continents. We conclude that
climate plays a greater role than fire in determining the distribution of nitrogen fixers across
tropical savanna and forest biomes.
Batesian mimicry, in which harmless species (mimics) deter predators by deceitfully imitating the warning signals of noxious species (models), generates striking cases of phenotypic convergence that are classic examples of evolution by natural selection. However, mimicry of venomous coral snakes has remained controversial because of unresolved conflict between the predictions of mimicry theory and empirical patterns in the distribution and abundance of snakes. Here we integrate distributional, phenotypic and phylogenetic data across all New World snake species to demonstrate that shifts to mimetic coloration in nonvenomous snakes are highly correlated with coral snakes in both space and time, providing overwhelming support for Batesian mimicry. We also find that bidirectional transitions between mimetic and cryptic coloration are unexpectedly frequent over both long- and short-time scales, challenging traditional views of mimicry as a stable evolutionary 'end point' and suggesting that insect and snake mimicry may have different evolutionary dynamics.
Aim
We used fossil and phylogenetic evidence to reconstruct climatic niche evolution in Nothofagus , a Gondwana genus distributed in tropical and temperate latitudes. To assess whether the modern distribution of the genus can be explained by the tropical conservatism hypothesis, we tested three predictions: (1) species from all Nothofagus subgenera coexisted under mesothermal climates during the early Eocene; (2) tolerance to microthermal climates evolved during the Eocene–Oligocene cooling from an ancestor that grew under mesothermal conditions; and (3) the climatic niche in Nothofagus is phylogenetically conserved.
Location
Australia, New Zealand, New Caledonia, Papua‐New Guinea and South America.
Methods
We estimated the palaeoclimate of the Early Eocene, fossil‐bearing Ligorio Marquez Formation ( LMF , Chile), using coexistence and leaf physiognomic analysis. We reconstructed ancestral climatic niches of Nothofagus using extant species distributions and a time‐calibrated phylogeny. Finally, we used the morphological disparity index and phylogenetic generalized least squares to assess whether climatic variables follow a Brownian motion ( BM ) or an Ornstein–Uhlenbeck ( OU ) model of evolution.
Results
Our palaeoclimatic estimates suggest mesothermal conditions for the LMF , where macrofossils associated with subgenera Lophozonia and possibly Fuscospora, and fossil pollen of Brassospora and Fuscospora / Nothofagus were recorded. These results are not supported by our phylogenetic analysis, which instead suggests that the ancestor of Nothofagus lived under microthermal to marginally mesothermal conditions, with tolerance to mesothermal conditions evolving only in the subgenus Brassospor a. Precipitation and temperature dimensions of the realized climatic niche fit with a gradual BM or constrained OU model of evolution.
Main Conclusions
Our results suggest that the use of phylogenetic reconstruction methods based only on present distributions of extant taxa to infer ancestral climatic niches is likely to lead to erroneous results when climatic requirements of ancestors differ from their extant descendants, or when much extinction has occurred.
Comparisons of Tertiary floras of North America with those of Europe and Asia document a long history of floristic interchange. The stratigraphic and geographic ranges of selected conifer and angiosperm genera that are easily recognized in the fossil record provide a basis for discerning patterns in the routes and timings of intercontinental dispersals through the Tertiary.
Reconstructing the biogeographical history and timing of the diversification of temperate forests is essential for understanding their history and resolving uncertainties about how flowering plants emerged from their deep tropical origins to dominate in today's freezing terrestrial environments. The angiosperm order Fagales, comprising iconic components of temperate forests worldwide with an extensive fossil record, are an excellent plant system in which to apply a fossil-aware paradigm, such as the fossilized birth-death (FBD) process, for investigating the macroevolution of temperate forest biomes. Here, we improve upon previous efforts to resolve phylogeny and incorporate fossils in Fagales using low-copy nuclear loci and an expanded morphological matrix to reevaluate the Fagales fossil record and: (1) infer the phylogenetic relationships and the time of origin of the clade using the FBD model as implemented in RevBayes, (2) provide a framework for evaluating the climatic and biogeographic history of Fagales, and (3) investigate how the inclusion of fossils via the FBD method influences ancestral reconstruction and diversification estimation. The phylogenetic relationships we recovered are conventional except for the position of Nothofagaceae, while our inferred ages support older timelines than previously proposed, with a mid-Cretaceous date for the most recent common ancestor of the order. Biogeographical analysis shows an origin of Fagales consistent with an ancestral circumboreal temperate distribution corroborated by ancestral niche reconstructions. While distributions today largely reflect the general conservatism of temperate forests, we identified two episodes of high diversification, one at the mid-Cretaceous origin of the clade and the other continuing from the Miocene to the present. Removing fossil taxa from the tree reveals a different story, shifting the origin of extant families from North America to East Asia, reflecting refugial distributions in this biodiversity "museum" and implying a general bias towards low extinction areas in biogeographic reconstruction. Likewise, without fossil data, diversification estimates were higher and unable to detect an early diversification burst. Based on our analyses, we close with recommendations regarding the interpretation of estimates of diversification and ancestral state reconstruction using phylogenetic trees with only extant species as tips.
- While the importance of root nodular symbioses (RNS) in plants has long been recognized, the ecological and evolutionary factors maintaining RNS remain obscure. RNS is associated with environmental stressors such as aridity and nitrogen-poor soils; the ability to tolerate harsh environments may provide ecological opportunities for diversification, yet, nodulators are also diverse outside these environments.
- We test several environmental determinants of increased survival and enhanced diversification of RNS species, using an explicitly phylogenetic approach for the first time. We assembled the largest phylogeny of the nitrogen-fixing clade to date and a comprehensive set of abiotic niche estimates and nodulation data. We used comparative phylogenetic tools to test environmental and diversification associations.
- We found that RNS is associated with warm, arid, and nitrogen-poor habitats. However, RNS was gained long before lineages entered these habitats. RNS is associated with accelerated diversification, but diversification rates are heterogeneous among nodulators, and non-legume nodulators do not show elevated diversification.
- Our findings undermine the interpretation that RNS directly drove the invasion of challenging habitats, and do not support a direct relationship between soil or climate and the diversity of nodulators. Still, RNS may have been an important exaptation allowing further niche evolution.
Aim
Digitization of herbarium specimens and DNA sequencing efforts in the past decade have enabled integrative analyses of patterns of diversity and endemism in a phylogenetic context. Here, we compare the best available floristic databases to a comprehensive specimen database to examine spatial patterns of moss phylogenetic assembly. We test the hypotheses that (1) mosses exhibit phylogenetic regionalization, (2) islands contain significantly high phylogenetic diversity and (3) that moss phylogenetic endemism is low on a global scale.
Location
Global.
Taxon
Mosses.
Methods
We developed a phylogeny of 3654 moss species using 25 markers and compiled a global specimen database from online repositories. We calculated floristic and phylogenetic measures of diversity and endemism and performed randomizations to test for significant deviations from expectations. We use rarefaction and extrapolation to alleviate substantial differences in sampling effort across the globe. We used both phylogenetic and floristic methods to test for spatial regionalization. We compare our specimen‐based results to those obtained using a floristic dataset.
Results
Phylogenetic diversity is more robust to missing data than species richness. Mean phylogenetic distance was significantly higher than expected in areas with high species richness, indicating that reported richness in these areas is likely a product of repeated colonization. Phylogenetic endemism is low globally. Phylogenetic regionalizations cluster into a Holarctic/Holantarctic temperate region, a pantropical region, and a region composed of Australia, New Zealand and South Africa.
Main Conclusions
Future efforts for collecting, sequencing and databasing moss species should focus on the tropics, particularly Africa and Southeast Asia. We provide further evidence to support several important theories developed in moss biogeography, including the role of long‐distance dispersal in shaping floristic patterns, the dominance of anagenesis in driving patterns of island diversity, and the role of climatic instability in driving patterns of assembly in the Holarctic.
Plastid phylogenomic analyses have shed light on many recalcitrant relationships across the angiosperm Tree of Life and continue to play an important role in plant phylogenetics alongside nuclear data sets given the utility of plastomes for revealing ancient and recent introgression. Here we conduct a plastid phylogenomic study of Fagales, aimed at exploring contentious relationships (e.g., the placement of Myricaceae and some intergeneric relationships in Betulaceae, Juglandaceae, and Fagaceae) and dissecting conflicting phylogenetic signals across the plastome. Combining 102 newly sequenced samples with publically available plastomes, we analyzed a dataset including 256 species and 32 of the 34 total genera of Fagales, representing the largest plastome-based study of the order to date. We find strong support for a sister relationship between Myricaceae and Juglandaceae, as well as strongly supported conflicting signal for alternative generic relationships in Betulaceae and Juglandaceae. These conflicts highlight the sensitivity of plastid phylogenomic analyses to genic composition, perhaps due to the prevalence of uninformative loci and heterogeneity in signal across different regions of the plastome. Phylogenetic relationships were geographically structure in subfamily Quercoideae, with Quercus being non-monophyletic and its sections forming clades with co-distributed Old World or New World genera of Quercoideae. Compared against studies based on nuclear genes, these results suggest extensive introgression and chloroplast capture in the early diversification of Quercus and Quercoideae. This study provides a critical plastome perspective on Fagales phylogeny, setting the stage for future studies employing more extensive data from the nuclear genome.
Abstract
Aim
Plants that host root‐symbiotic nitrogen‐fixing bacteria have an important role in driving terrestrial ecosystem processes, but N‐fixing ability is unequally distributed among plant taxa and ecosystems. Here we explore the large‐scale distribution of N‐fixing plant species worldwide.
Location
Global.
Time period
Present.
Major taxa studied
Vascular plants.
Methods
We estimated root‐symbiotic N‐fixing plant species diversity (as Shannon entropy) and relative richness (log‐ratio of N‐fixing to non‐fixing plant species) for c. 7,800 km2 hexagonal grid cells using the NodDB and Global Biodiversity Information Facility (GBIF) databases. Additionally, we explored the distributions of plant species associated with rhizobia, actinobacteria or cyanobacteria (relative to other plant species), and the relative richness of N‐fixing trees (log‐ratio of N‐fixing to non‐fixing trees). We related N‐fixing plant species distribution to environmental (climate, soil) and biogeographical (biome, realm) variables using multiple linear regression.
Results
N‐fixing plant diversity and relative richness showed unimodal relationships with latitude. Diversity of N‐fixing plants was highest in warm and wet climates, but in dry biomes and in Australasia. The relative richness of N‐fixing plants was highest in warm and dry climates, in tropical and temperate grasslands and in Eurasia. Plants associated with cyanobacteria were more widely distributed near the equator, while those associated with rhizobia were more prevalent at the edge of the tropics, and those associated with actinobacteria at higher latitudes (especially in boreal forests). The relative richness of N‐fixing tree species was highest in cold and dry areas and in boreal forests, with contrasting peaks in the Northern and Southern Hemispheres.
Main conclusions
The distribution of N‐fixing plant species exhibits regional hotspots and coldspots related to both environmental conditions and biogeographical history. Global N‐fixing plant distributions are different for the key root‐symbiotic bacterial groups. Information about N‐fixing plant distribution can improve global models of ecosystem functions and contribute to understanding how plants respond to global change.
A hallmark of flowering plants is their ability to invade some of the most extreme and dynamic habitats, including cold and dry biomes, to a far greater extent than other land plants. Recent work has provided insight to the phylogenetic distribution and evolutionary mechanisms which have enabled this success, yet needed is a synthesis of evolutionary perspectives with plant physiological traits, morphology, and genomic diversity. Linking these disparate components will not only lead to better understand the evolutionary parallelism and diversification of plants with these two strategies, but also to provide the framework needed for directing future research. We summarize the primary physiological and structural traits involved in response to cold- and drought stress, outline the phylogenetic distribution of these adaptations, and describe the recurring association of these changes with rapid diversification events that occurred in multiple lineages over the past 15 million years. Across these three-fold facets of dry-cold correlation (traits, phylogeny, and time) we stress the contrast between (1) the amazing diversity of solutions flowering plants have developed in the face of extreme environments and (2) a broad correlation between cold and dry adaptations that in some cases may hint at deep common origins. This article is protected by copyright. All rights reserved.
Biogeographical regionalization schemes have traditionally been constructed based on taxonomic endemism of families, genera, and/or species, and rarely incorporated the phylogenetic relationships between taxa. However, phylogenetic relationships are important for understanding historical connections within and among biogeographical regions. Phylogeny-based delineation of biota is a burgeoning and fruitful field that is expected to provide novel insights into the conservation of regional diversity and the evolutionary history of biota. Using the Chinese flora as an example, we compared regionalization schemes that were based on: (1) taxonomic endemism, (2) taxonomic dissimilarity, and (3) phylogenetic dissimilarity. Our results revealed general consistency among different regionalization schemes and demonstrated that the phylogenetic dissimilarity approach is preferable for biogeographical regionalization studies. Using the phylogenetic dissimilarity approach, we identified five phytogeographical regions within China: the Paleotropic, Holarctic, East Asiatic, Tethyan, and Qinghai–Tibet Plateau Regions. The relationship of these regions was inferred to be: (Paleotropic, ((East Asiatic + Holarctic) + (Tethyan + Qinghai–Tibet Plateau)).
A key step in understanding the distribution of biodiversity is the grouping of regions based on their shared elements. Historically, regionalization schemes have been largely species centric. Recently, there has been interest in incorporating phylogenetic information into regionalization schemes. Phylogenetic regionalization can provide novel insights into the mechanisms that generate, distribute, and maintain biodiversity. We argue that four processes (dispersal limitation, extinction, speciation, and niche conservatism) underlie the formation of species assemblages into phylogenetically distinct biogeographic units. We outline how it can be possible to distinguish among these processes, and identify centers of evolutionary radiation, museums of diversity, and extinction hotspots. We suggest that phylogenetic regionalization provides a rigorous and objective classification of regional diversity and enhances our knowledge of biodiversity patterns.
The deciduous habit of northern temperate trees and shrubs provides one of the most obvious examples of convergent evolution, but how did it evolve? Hypotheses based on the fossil record posit that deciduousness evolved first in response to drought or darkness and preadapted certain lineages as cold climates spread. An alternative is that evergreens first established in freezing environments and later evolved the deciduous habit. We monitored phenological patterns of 20 species of Viburnum spanning tropical, lucidophyllous (subtropical montane and warm temperate), and cool temperate Asian forests. In lucidophyllous forests, all viburnums were evergreen plants that exhibited coordinated leaf flushes with the onset of the rainy season but varied greatly in the timing of leaf senescence. In contrast, deciduous species exhibited tight coordination of both flushing and senescence, and we found a perfect correlation between the deciduous habit and prolonged annual freezing. In contrast to previous stepwise hypotheses, a consilience of independent lines of evidence supports a lockstep model in which deciduousness evolved in situ, in parallel, and concurrent with a gradual cooling climate. A pervasive selective force combined with the elevated evolutionary accessibility of a particular response may explain the massive convergence of adaptive strategies that characterizes the world’s biomes.
An update of the Angiosperm Phylogeny Group (APG) classification of the orders and families of angiosperms is presented. Several new orders are recognized: Boraginales, Dilleniales, Icacinales, Metteniusiales and Vahliales. This brings the total number of orders and families recognized in the APG system to 64 and 416, respectively. We propose two additional informal major clades, superrosids and superasterids, that each comprise the additional orders that are included in the larger clades dominated by the rosids and asterids. Families that made up potentially monofamilial orders, Dasypogonaceae and Sabiaceae, are instead referred to Arecales and Proteales, respectively. Two parasitic families formerly of uncertain positions are now placed: Cynomoriaceae in Saxifragales and Apodanthaceae in Cucurbitales. Although there is evidence that some families recognized in APG III are not monophyletic, we make no changes in Dioscoreales and Santalales relative to APG III and leave some genera in Lamiales unplaced (e.g. Peltanthera). These changes in familial circumscription and recognition have all resulted from new results published since APG III, except for some changes simply due to nomenclatural issues, which include substituting Asphodelaceae for Xanthorrhoeaceae (Asparagales) and Francoaceae for Melianthaceae (Geraniales); however, in Francoaceae we also include Bersamaceae, Ledocarpaceae, Rhynchothecaceae and Vivianiaceae. Other changes to family limits are not drastic or numerous and are mostly focused on some members of the lamiids, especially the former Icacinaceae that have long been problematic with several genera moved to the formerly monogeneric Metteniusaceae, but minor changes in circumscription include Aristolochiaceae (now including Lactoridaceae and Hydnoraceae; Aristolochiales), Maundiaceae (removed from Juncaginaceae; Alismatales), Restionaceae (now re-including Anarthriaceae and Centrolepidaceae; Poales), Buxaceae (now including Haptanthaceae; Buxales), Peraceae (split from Euphorbiaceae; Malpighiales), recognition of Petenaeaceae (Huerteales), Kewaceae, Limeaceae, Macarthuriaceae and Microteaceae (all Caryophyllales), Petiveriaceae split from Phytolaccaceae (Caryophyllales), changes to the generic composition of Ixonanthaceae and Irvingiaceae (with transfer of Allantospermum from the former to the latter; Malpighiales), transfer of Pakaraimaea (formerly Dipterocarpaceae) to Cistaceae (Malvales), transfer of Borthwickia, Forchhammeria, Stixis and Tirania (formerly all Capparaceae) to Resedaceae (Brassicales), Nyssaceae split from Cornaceae (Cornales), Pteleocarpa moved to Gelsemiaceae (Gentianales), changes to the generic composition of Gesneriaceae (Sanango moved from Loganiaceae) and Orobanchaceae (now including Lindenbergiaceae and Rehmanniaceae) and recognition of Mazaceae distinct from Phrymaceae (all Lamiales).
As a primary determinant of spatial structure in angiosperm populations, fruit dispersal may impact large-scale ecological and evolutionary processes. Essential to understanding these mechanisms is an accurate reconstruction of dispersal mode over the entire history of an angiosperm lineage. A total-evidence phylogeny is presented for most fossil fruit and all extant genera in Fagales over its c. 95 million yr history. This phylogeny - the largest of its kind to include plant fossils - was used to reconstruct an evolutionary history directly informed by fossil morphologies and to assess relationships among dispersal mode, biogeographic range size, and diversification rate. Reconstructions indicate four transitions to wind dispersal and seven to biotic dispersal, with the phylogenetic integration of fossils crucial to understanding these patterns. Complexity further increased when more specialized behaviors were considered, with fluttering, gliding, autorotating, and scatter-hoarding evolving multiple times across the order. Preliminary biogeographic analyses suggest larger range sizes in biotically dispersed lineages, especially when pollination mode was held constant. Biotically dispersed lineages had significantly higher diversification rates than abiotically dispersed lineages, although transitions in dispersal mode alone cannot explain all detected diversification rate shifts across Fagales.
© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.
Deciduousness in mesic, broad-leaved plants occurred in disturbed, middle-latitude environments during the Late Cretaceous. Only in polar environments in the Late Cretaceous was the deciduous element dominant, although of low diversity. The terminal Cretaceous event resulted in wide-spread selection for plants of deciduous habit and diversification of deciduous taxa, thus leaving a lasting imprint on Northern Hemisphere vegetation. Various environmental factors have played important roles in subsequent diversification of mesic, broad-leaved deciduous taxa and in origination and decline of broad-leaved deciduous forests. Low diversity and rarity of mesic deciduous plants in the post-Cretaceous of the Southern Hemisphere indicate that the inferred 'impact winter' of the terminal Cretaceous event had little effect on Southern Hemisphere vegetation and climate. -Author
Numerous allied trees, shrubs, and herbs inhabit the discontinuous mixed deciduous hardwood and conifer-hardwood forests of Holarctica. Their close ancestors occupied middle and high latitudes into the Middle Miocene (15-16 m.y.a.). As the forests retreated southward and were disrupted by spreading colder and drier climates following Antarctic glaciation (13 m.y.a.), numerous paired or allied species (or sections, series) originated and have survived in the disjunct segregate forests across Holarctica. In addition, all major climatic-topographic provinces served as centers of evolution from which taxa have not ranged widely. It is this latter pattern to which oaks conform most closely. Only species of allied subsects. Prinoideae and Diversipiloseae had an Holarctic occurrence, occupying lands above latitude 60⚬N during the late Cretaceous and Tertiary. Although climate was favorable for numerous deciduous hardwoods and conifers at high latitudes, taxa of most Quercus sections and subsections have been confined to one continent or to parts of them where they ranged somewhat more widely during the Tertiary. The rarity of Tertiary oaks linking Eurasia-North America may owe to the light factor (long day) at the north; fully four-fifths of all oak species occur below latitude 30-35⚬N and only six or seven reach latitude 50⚬N. Proliferation into 280 to 300 species was chiefly in response to spreading drier and colder climate that commenced in the middle Eocene, to increasing topographic diversity in the later Cenozoic, and to fluctuating Quaternary climate. At times of major climatic shifts or increased relief, oaks responded opportunistically, probably by quantum steps (punctuated equilibria) that account for the numerous series.