Contrasting patterns of diversification in a bird family (Aves: Gruiformes: Rallidae) are revealed by analysis of geospatial distribution of species and phylogenetic diversity

Geospatial patterns in the distribution of regional biodiversity reflect the composite processes that underpin evolution: speciation, dispersal and extinction. The spatial distribution and phylogeny of a globally widespread and species rich bird family (Rallidae) were used to help assess the role of large‐scale biogeographical processes in diversity and diversification. Here, we examine how different geostatistical diversity metrics enhance our understanding of species distribution by linking occurrence records of rail species to corresponding species level phylogeny. Tropical regions and temperate zones contained a large proportion of rail species richness and phylogenetic diversity whilst small islands in Australian, Oceanian and Oriental regions held the highest weighted and phylogenetic endemism. Our results suggest that habitat connectivity and dispersal were important ecological features in rail evolution and distribution. Spatial isolation was a significant driver of diversification where islands in Oceania were centres of neo‐endemism with recent multiple and independent speciation events and could be considered as nurseries of biodiversity. Palaeo‐endemism was mostly associated with older stable regions, so despite extensive long distance range shifting these areas retain their own ancient and distinct character. Madagascar was the major area of palaeo‐endemism associated with the oldest rail lineages and could be considered a museum of rail diversity. This implies a mixture of processes determine the current distribution and diversity of rail clades with some areas dominated by recent ‘in situ’ speciation while others harbour old diversity with ecological traits that have stood the test of time.     

Does the colonization of new biogeographic regions influence the diversification and accumulation of clade richness among the Corvides (Aves: Passeriformes)?

Regional variation in clade richness can be vast, reflecting differences in the dynamics of historical dispersal and diversification among lineages. Although it has been proposed that dispersal into new biogeographic regions may facilitate diversification, to date there has been limited assessment of the importance of this process in the generation, and maintenance, of broad‐scale biodiversity gradients. To address this issue, we analytically derive biogeographic regions for a global radiation of passerine birds (the Corvides, c. 790 species) that are highly variable in the geographic and taxonomic distribution of species. Subsequently, we determine rates of historical dispersal between regions, the dynamics of diversification following regional colonization, and spatial variation in the distribution of species that differ in their rates of lineage diversification. The results of these analyses reveal spatiotemporal differences in the build‐up of lineages across regions. The number of regions occupied and the rate of transition between regions both predict family richness well, indicating that the accumulation of high clade richness is associated with repeated expansion into new geographic areas. However, only the largest family (the Corvidae) had significantly heightened rates of both speciation and regional transition, implying that repeated regional colonization is not a general mechanism promoting lineage diversification among the Corvides.     

DO FRESHWATER FISHES DIVERSIFY FASTER THAN MARINE FISHES? A TEST USING STATE‐DEPENDENT DIVERSIFICATION ANALYSES AND MOLECULAR PHYLOGENETICS OF NEW WORLD SILVERSIDES (ATHERINOPSIDAE)

Freshwater habitats make up only ～0.01% of available aquatic habitat and yet harbor 40% of all fish species, whereas marine habitats comprise >99% of available aquatic habitat and have only 60% of fish species. One possible explanation for this pattern is that diversification rates are higher in freshwater habitats than in marine habitats. We investigated diversification in marine and freshwater lineages in the New World silverside fish clade Menidiinae (Teleostei, Atherinopsidae). Using a time‐calibrated phylogeny and a state‐dependent speciation–extinction framework, we determined the frequency and timing of habitat transitions in Menidiinae and tested for differences in diversification parameters between marine and freshwater lineages. We found that Menidiinae is an ancestrally marine lineage that independently colonized freshwater habitats four times followed by three reversals to the marine environment. Our state‐dependent diversification analyses showed that freshwater lineages have higher speciation and extinction rates than marine lineages. Net diversification rates were higher (but not significant) in freshwater than marine environments. The marine lineage‐through time (LTT) plot shows constant accumulation, suggesting that ecological limits to clade growth have not slowed diversification in marine lineages. Freshwater lineages exhibited an upturn near the recent in their LTT plot, which is consistent with our estimates of high background extinction rates. All sequence data are currently being archived on Genbank and phylogenetic trees archived on Treebase.     

Time,space and ecology: why some clades have more species than others

Aim We seek biotic and abiotic explanations for differences in lineage sizes of Afromontane sedges (Cyperaceae, Carex) and buttercups (Ranunculaceae, Ranunculus). Location Mountains of sub‐Saharan Africa and Madagascar. Methods We investigated differences in the species richness and diversification rates of 18 lineages of the pan‐temperate plant groups Carex and Ranunculus, established by long‐distance dispersal on African sky islands. We built generalized linear models to test the individual and the cumulative power of biotic and abiotic factors for predicting variation in the size of lineages. Tested variables were: ages of the lineages, their geographic distributions, number of mountain systems occupied, isolation/distance from ancestral areas, elevation range, number of vegetation zones and habitat types in which lineages are found, light requirement and water availability for each lineage, and the sum of the habitat factors, representing habitat heterogeneity. Habitat conservatism was measured by the overlap in habitats among the species within each lineage. Diversification rate changes were investigated using ‘laser’ in R. Results The number of Carex and Ranunculus lineages on the African mountains accumulated gradually through time. The size of these lineages could best be explained by a model combining age and distribution together with a measure of environmental heterogeneity (either elevation and water availability or habitat heterogeneity). Extensive overlap in environmental characteristics and distribution ranges among the species indicates a relatively high degree of conservatism of these characters. Main conclusions Lineages that are species‐rich are those that have the ability and time to occupy many mountain regions and a wide range of habitats. If allopatric or ecological speciation plays a role, then secondary dispersal and/or niche expansion soon obscures the patterns that may have existed at the point of speciation.     

Speciation and radiations track climate transitions since the Miocene Climatic Optimum: a case study of southern African chameleons

Aim The high amount of species diversity concentrated in southern Africa has been attributed to palaeoclimatic factors, and the timing of radiations in some taxa corresponds to global palaeoclimatic trends. Using dwarf chameleons (Bradypodion: Chamaeleonidae) as a model system, we explored the relationship between palaeoclimatic fluctuations and cladogenesis with respect to both temporal and spatial patterns in an effort to understand the process of speciation in southern Africa. Location South Africa, with particular emphasis on the Cape Floristic Region and the Maputaland–Pondoland–Albany hotspot. Methods Mitochondrial sequence data (ND2 and 16S) were used to estimate the timing of major radiations and to examine the number of lineages through time. A dated phylogeny was constructed using Bayesian phylogenetic reconstruction, and a Bayesian relaxed molecular clock was used to estimate divergence times. Spatial data and lineage‐through‐time plots were used to identify geographic regions that underwent diversification in connection with major climatic events. Both parsimony and likelihood optimizations of habitat type on the phylogeny were used to determine whether major habitat shifts have occurred. On a coarse scale (half‐degree grid cells), phylogenetic diversity (sum of the branch lengths linking terminals) was compared with species richness (absolute number of species) to identify areas of conservation importance. Results The complete species phylogeny of dwarf chameleons shows that the timing and mode of diversification exhibit spatio‐temporal patterns that link to phases in the evolution of southern Africa’s climate over the last 14 Myr. Optimizations of habitat on the phylogenetic tree show a progression from closed to open habitats since the Mid‐Miocene, corresponding to the shift from C₃ to C₄ environments, and later with the development of south‐western Africa’s winter‐rainfall regime. These shifts are not simultaneous across the region, with different geographic centres of diversity generated during different time periods. Main conclusions Regions that are prominent centres of chameleon diversification are encompassed by the current biodiversity hotspots as shown by chameleon species richness and phylogenetic diversity. Diversity within the Cape Floristic Region appears to be the result of a Late Pliocene radiation, whereas the diversity encompassed within the Maputaland–Pondoland–Albany hotspot is an aggregate of asynchronous radiation events, probably influenced by lineage losses. Overall, dwarf chameleons have experienced a shift in habitat types, with recent radiations occupying open habitats, and older lineages persisting in relictual forested habitats, corresponding to the continental shift of vegetation types since the Miocene Climatic Optimum.     

The effect of environmental diversification on species diversification in New Caledonian caddisflies (Insecta: Trichoptera: Hydropsychidae)

Aim To test whether environmental diversification played a role in the diversification of the New Caledonian Hydropsychinae caddisflies. Location New Caledonia, south‐west Pacific. Methods The phylogeny of the New Caledonian Hydropsychinae caddisflies was hypothesized using parsimony and Bayesian methods on molecular characters. The Bayesian analysis was the basis for a comparative analysis of the correlation between phylogeny and three environmental factors: geological substrate (ultrabasic, non‐ultrabasic), elevation and precipitation. Phylogenetic divergence times were estimated using a relaxed clock method, and environmental factors were mapped onto a lineage‐through‐time plot to investigate the timing of environmental diversification in relation to species radiation. The correlation between rainfall and elevation was tested using independent contrasts, and the gamma statistic was calculated to infer the diversification pattern of the group. Results The diversification of extant Orthopsyche–Caledopsyche species began in the Middle–Late Oligocene, when much of the island of New Caledonia was covered by ultrabasic substrate and mountain forming was prevalent. Most lineages originated in the Middle–Late Miocene, a period associated with long‐term climate oscillation. Optimization of environmental factors on the phylogeny demonstrated that the New Caledonian Hydropsychinae group adapted to ultrabasic substrate early in its evolutionary history. The clade living mostly on ultrabasic substrate was far more species‐rich than the clade living mostly on non‐ultrabasic substrate. Elevation and rainfall were significantly correlated with each other. The lineage‐through‐time plot revealed that the main environmental diversification preceded species diversification. A constant speciation through time was rejected, and the negative gamma indicates that most of the diversification occurred early in the history of the clade. According to the inferred phylogeny, the genus Orthopsyche McFarlane is a synonym under Caledopsyche Kimmins, and Abacaria caledona Oláh & Barnard should also be included in Caledopsyche. Main conclusions The age of the radiation does not support a vicariance origin of New Caledonian Hydropsychinae caddisflies. Environmental diversification pre‐dates lineage diversification, and thus environmental heterogeneity potentially played a role in the diversification of the group, by providing a variety of fragmented habitats to disperse into, promoting speciation. The negative gamma indicates that the speciation rate slowed as niches started to fill.     

Hidden biodiversity: total evidence phylogenetics and evolution of morphological traits in a highly diverse lineage of endogean ground beetles,Typhlocharis Dieck, 1869 (Carabidae,Trechinae, Anillini)

Typhlocharis is the most diverse eyeless endogean ground beetle genus known to date, with 62 species all endemic to the West Mediterranean region. The lineage is characterized by a conservative and singular body plan within Carabidae that contrasts with a high morphological diversity in many traits. We provide an exhaustive phylogeny of the lineage through the study of 92 morphological characters from all 62 described species and 45 potential new species from 70 additional populations, and the combination of morphological and available molecular data, in the first total evidence phylogenetic approach for a highly diverse endogean lineage. We tracked the evolution of morphological traits over the obtained phylogenies. Results suggest eight morphologically distinct clades, which do not correspond to the species groups proposed formerly. Ancestral state reconstructions and phylogenetic signal analyses of morphological traits revealed that some of the previously key characters to the classification of Typhlocharis, such as the umbilicate series or the apical denticles of elytra, are highly homoplasic, whereas other characters show stronger phylogenetic signal, including structures in the antennae, gula, pronotum and last abdominal ventrite. This evidence supports the split of Typhlocharis into three genera: Lusotyphlus gen. nov. ; Typhlocharis Dieck, 1869 and Microcharidius Coiffait, 1969 (revalidated), forming the subtribe Typhlocharina Jeanne, 1973.     

Niche evolution and diversification in a Neotropical radiation of birds (Aves: Furnariidae)

Rapid diversification may be caused by ecological adaptive radiation via niche divergence. In this model, speciation is coupled with niche divergence and lineage diversification is predicted to be correlated with rates of niche evolution. Studies of the role of niche evolution in diversification have generally focused on ecomorphological diversification but climatic‐niche evolution may also be important. We tested these alternatives using a phylogeny of 298 species of ovenbirds (Aves: Furnariidae). We found that within Furnariidae, variation in species richness and diversification rates of subclades were best predicted by rate of climatic‐niche evolution than ecomorphological evolution. Although both are clearly important, univariate regression and multivariate model averaging more consistently supported the climatic‐niche as the best predictor of lineage diversification. Our study adds to the growing body of evidence, suggesting that climatic‐niche divergence may be an important driver of rapid diversification in addition to ecomorphological evolution. However, this pattern may depend on the phylogenetic scale at which rate heterogeneity is examined.     

Does migration promote or inhibit diversification? A case study involving the dominant radiation of temperate Southern Hemisphere freshwater fishes

Although theory predicts that dispersal has a pivotal influence on speciation and extinction rates, it can have contradictory effects on each, such that empirical quantification of its role is required. In many studies, dispersal reduction appears to promote diversification, although some comparisons of migratory and nonmigratory species suggest otherwise. We tested for a relationship between migratory status and diversification rate within the dominant radiation of temperate Southern Hemisphere freshwater fishes, the Galaxiidae. We reconstructed a molecular phylogeny comprising >95% of extant taxa, and applied State-dependent Speciation Extinction models to estimate speciation, extinction, and diversification rates. In contrast to some previous studies, we revealed higher diversification rates in nonmigratory lineages. The reduced gene flow experienced by nonmigratory galaxiids appears to have increased diversification under conditions of allopatry or local adaptation. Migratory galaxiid lineages, by contrast, are genetically homogeneous within landmasses, but may also be rarely able to diversify by colonizing other landmasses in the temperate Southern Hemisphere. Apparent contradictions among studies of dispersal-diversification relationships may be explained by the spatial context of study systems relative to species dispersal abilities, by means of the “intermediate dispersal” model; the accurate quantification of dispersal abilities will aid in the understanding of these proposed interactions.     

Local above‐ground persistence of vascular plants: Life‐history trade‐offs and environmental constraints

Questions: 1. Which plant traits and habitat characteristics best explain local above‐ground persistence of vascular plant species and 2. Is there a trade‐off between local above‐ground persistence and the ability for seed dispersal and below‐ground persistence in the soil seed bank? Locations: 845 long‐term permanent plots in terrestrial habitats across the Netherlands. Methods: We analysed the local above‐ground persistence of vascular plants in permanent plots (monitored once a year for ca. 16 year) with respect to functional traits and habitat preferences using survival statistics (Kaplan‐Meier analysis and Cox’ regression). These methods account for censored data and are rarely used in vegetation ecology. Results: Local above‐ground persistence is determined by both functional traits (especially the ability to form long‐lived clonal connections) and habitat preferences (especially nutrient requirements). Above‐ground persistence is negatively related to the ability for dispersal by wind and to the ability to accumulate a long‐term persistent soil seed bank (‘dispersal through time’) and is positively related to the ability for dispersal by water. Conclusions: Most species have a half‐life expectation over 15 years, which may contribute to time lags after changes in habitat quality or ‐configuration (‘extinction debt’). There is evidence for a trade‐off relationship between local above‐ground persistence and below‐ground seed persistence, while the relationship with dispersal in space is vector specific. The rate of species turnover increases with productivity.     

Detecting local diversity‐dependence in diversification

Whether there are ecological limits to species diversification is a hotly debated topic. Molecular phylogenies show slowdowns in lineage accumulation, suggesting that speciation rates decline with increasing diversity. A maximum‐likelihood (ML) method to detect diversity‐dependent (DD) diversification from phylogenetic branching times exists, but it assumes that diversity‐dependence is a global phenomenon and therefore ignores that the underlying species interactions are mostly local, and not all species in the phylogeny co‐occur locally. Here, we explore whether this ML method based on the nonspatial diversity‐dependence model can detect local diversity‐dependence, by applying it to phylogenies, simulated with a spatial stochastic model of local DD speciation, extinction, and dispersal between two local communities. We find that type I errors (falsely detecting diversity‐dependence) are low, and the power to detect diversity‐dependence is high when dispersal rates are not too low. Interestingly, when dispersal is high the power to detect diversity‐dependence is even higher than in the nonspatial model. Moreover, estimates of intrinsic speciation rate, extinction rate, and ecological limit strongly depend on dispersal rate. We conclude that the nonspatial DD approach can be used to detect diversity‐dependence in clades of species that live in not too disconnected areas, but parameter estimates must be interpreted cautiously.     

BOOM AND BUST: ANCIENT AND RECENT DIVERSIFICATION IN BICHIRS (POLYPTERIDAE: ACTINOPTERYGII), A RELICTUAL LINEAGE OF RAY‐FINNED FISHES

Understanding the history that underlies patterns of species richness across the Tree of Life requires an investigation of the mechanisms that not only generate young species‐rich clades, but also those that maintain species‐poor lineages over long stretches of evolutionary time. However, diversification dynamics that underlie ancient species‐poor lineages are often hidden due to a lack of fossil evidence. Using information from the fossil record and time calibrated molecular phylogenies, we investigate the history of lineage diversification in Polypteridae, which is the sister lineage of all other ray‐finned fishes (Actinopterygii). Despite originating at least 390 million years (Myr) ago, molecular timetrees support a Neogene origin for the living polypterid species. Our analyses demonstrate polypterids are exceptionally species depauperate with a stem lineage duration that exceeds 380 million years (Ma) and is significantly longer than the stem lineage durations observed in other ray‐finned fish lineages. Analyses of the fossil record show an early Late Cretaceous (100.5–83.6 Ma) peak in polypterid genus richness, followed by 60 Ma of low richness. The Neogene species radiation and evidence for high‐diversity intervals in the geological past suggest a “boom and bust” pattern of diversification that contrasts with common perceptions of relative evolutionary stasis in so‐called “living fossils.”     

Not going with the flow: a comprehensive time‐calibrated phylogeny of dragonflies (Anisoptera: Odonata: Insecta) provides evidence for the role of lentic habitats on diversification

Ecological diversification of aquatic insects has long been suspected to have been driven by differences in freshwater habitats, which can be classified into flowing (lotic) waters and standing (lentic) waters. The contrasting characteristics of lotic and lentic freshwater systems imply different ecological constraints on their inhabitants. The ephemeral and discontinuous character of most lentic water bodies may encourage dispersal by lentic species in turn reducing geographical isolation among populations. Hence, speciation probability would be lower in lentic species. Here, we assess the impact of habitat use on diversification patterns in dragonflies (Anisoptera: Odonata). Based on the eight nuclear and mitochondrial genes, we inferred species diversification with a model‐based evolutionary framework, to account for rate variation through time and among lineages and to estimate the impact of larval habitat on the potentially nonrandom diversification among anisopteran groups. Ancestral state reconstruction revealed lotic fresh water systems as their original primary habitat, while lentic waters have been colonized independently in Aeshnidae, Corduliidae and Libellulidae. Furthermore, our results indicate a positive correlation of speciation and lentic habitat colonization by dragonflies: speciation rates increased in lentic Aeshnidae and Libellulidae, whereas they remain mostly uniform among lotic groups. This contradicts the hypothesis of inherently lower speciation in lentic groups and suggests species with larger ranges are more likely to diversify, perhaps due to higher probability of larger areas being dissected by geographical barriers. Furthermore, larger range sizes may comprise more habitat types, which could also promote speciation by providing additional niches, allowing the coexistence of emerging species.     

Insights into the historical construction of species-rich Mesoamerican seasonally dry tropical forests: the diversification of Bursera (Burseraceae, Sapindales)

? Mesoamerican arid biomes epitomize neotropical rich and complex biodiversity. To document some of the macroevolutionary processes underlying the vast species richness of Mesoamerican seasonally dry tropical forests (SDTFs), and to evaluate specific predictions about the age, geographical structure and niche conservatism of SDTF-centered woody plant lineages, the diversification of Bursera is reconstructed. ? Using a nearly complete Bursera species-level phylogeny from nuclear and plastid genomic markers, we estimate divergence times, test for phylogenetic and temporal diversification heterogeneity, test for geographical structure, and reconstruct habitat shifts. ? Bursera became differentiated in the earliest Eocene, but diversified during independent early Miocene consecutive radiations that took place in SDTFs. The late Miocene average age of Bursera species, the presence of phylogenetic geographical structure, and its strong conservatism to SDTFs conform to expectations derived from South American SDTF-centered lineages. ? The diversification of Bursera suggests that Mesoamerican SDTF richness derives from high speciation from the Miocene onwards uncoupled from habitat shifts, during a period of enhanced aridity resulting mainly from global cooling and regional rain shadows.     

A Dated Phylogeny Complements Macroecological Analysis to Explain the Diversity Patterns in Geonoma (Arecaceae)

Integrating phylogenetic data into macroecological studies of biodiversity patterns may complement the information provided by present‐day spatial patterns. In the present study, we used range map data for all Geonoma (Arecaceae) species to assess whether Geonoma species composition forms spatially coherent floristic clusters. We then evaluated the extent to which the spatial variation in species composition reflects present‐day environmental variation vs. nonenvironmental spatial effects, as expected if the pattern reflects historical biogeography. We also examined the degree of geographic structure in the Geonoma phylogeny. Finally, we used a dated phylogeny to assess whether species richness within the floristic clusters was constrained by a specific historical biogeographic driver, namely time‐for‐diversification. A cluster analysis identified six spatially coherent floristic clusters, four of which were used to reveal a significant geographic phylogenetic structure. Variation partitioning analysis showed that 56 percent of the variation in species composition could be explained by spatial variables alone, consistent with historical factors having played a major role in generating the Geonoma diversity pattern. To test for a time‐for‐diversification effect, we correlated four different species richness measures with the diversification time of the earliest large lineage that is characteristic of each cluster. In support of this hypothesis, we found that geographic areas with higher richness contained older radiations. We conclude that current geographic diversity patterns in Geonoma reflect the present‐day climate, but to a larger extent are related to nonenvironmental spatial constraints linked to colonization time, dispersal limitation, and geological history, followed by within‐area evolutionary diversification. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Patterns of population structure and complex haplotype sharing among field isolates of the green alga Chlamydomonas reinhardtii

The nature of population structure in microbial eukaryotes has long been debated. Competing models have argued that microbial species are either ubiquitous, with high dispersal and low rates of speciation, or that for many species gene flow between populations is limited, resulting in evolutionary histories similar to those of macroorganisms. However, population genomic approaches have seldom been applied to this question. Here, we analyse whole‐genome resequencing data for all 36 confirmed field isolates of the green alga Chlamydomonas reinhardtii. At a continental scale, we report evidence for putative allopatric divergence, between both North American and Japanese isolates, and two highly differentiated lineages within N. America. Conversely, at a local scale within the most densely sampled lineage, we find little evidence for either spatial or temporal structure. Taken together with evidence for ongoing admixture between the two N. American lineages, this lack of structure supports a role for substantial dispersal in C. reinhardtii and implies that between‐lineage differentiation may be maintained by reproductive isolation and/or local adaptation. Our results therefore support a role for allopatric divergence in microbial eukaryotes, while also indicating that species may be ubiquitous at local scales. Despite the high genetic diversity observed within the most well‐sampled lineage, we find that pairs of isolates share on average ~9% of their genomes in long haplotypes, even when isolates were sampled decades apart and from different locations. This proportion is several orders of magnitude higher than the Wright–Fisher expectation, raising many further questions concerning the evolutionary genetics of C. reinhardtii and microbial eukaryotes generally.     

Relationships of diversity,disparity, and their evolutionary rates in squirrels (Sciuridae)

Several theories predict that rapidly diversifying clades will also rapidly diverge phenotypically; yet, there are also reasons for suspecting that diversification and divergence might not be correlated. In the widely distributed squirrel clade (Sciuridae), we test for correlations between per lineage speciation rates, species richness, disparity, and a time‐invariant measure of disparity that allows for comparing rates when evolutionary modes differ, as they do in squirrels. We find that species richness and speciation rates are not correlated with clade age or with each other. Disparity appears to be positively correlated with clade age because young, rapidly diversifying Nearctic grassland clades are strongly pulled to a single stable optimum but older, slowly diversifying Paleotropical forest clades contain lineages that diverge along multiple ecological and morphological lines. That contrast is likely due to both the environments they inhabit and their phylogenetic community structure. Our results argue against a shared explanation for diversity and disparity in favor of geographically mediated modes of speciation and ecologically mediated modes of phenotypic evolution.     

Phylogenetic diversity as a window into the evolutionary and biogeographic histories of present-day richness gradients for mammals

Phylogenetic diversity (PD) captures the shared ancestry of species, and is increasingly being recognized as a valuable conservation currency. Regionally, PD frequently covaries closely with species richness; however, variation in speciation and extinction rates and/or the biogeographic history of lineages can result in significant deviation. Locally, these differences may be pronounced. Rapid recent speciation or high temporal turnover of lineages can result in low PD but high richness. In contrast, rare dispersal events, for example, between biomes, can elevate PD but have only small impact on richness. To date, environmental predictors of species richness have been well studied but global models explaining variation in PD are lacking. Here, we contrast the global distribution of PD versus species richness for terrestrial mammals. We show that an environmental model of lineage diversification can predict well the discrepancy in the distribution of these two variables in some places, for example, South America and Africa but not others, such as Southeast Asia. When we have information on multiple diversity indices, conservation efforts directed towards maximizing one currency or another (e.g. species richness versus PD) should also consider the underlying processes that have shaped their distributions.     

High tropical net diversification drives the New World latitudinal gradient in palm (Arecaceae) species richness

Aim Species richness exhibits striking geographical variation, but the processes that drive this variation are unresolved. We investigated the relative importance of two hypothesized evolutionary causes for the variation in palm species richness across the New World: time for diversification and evolutionary (net diversification) rate. Palms have a long history in the region, with the major clades diversifying during the Tertiary (65–2 Ma). Location Tropical and subtropical America (34° N–34° S; 33–120° W). Methods Using range maps, palm species richness was estimated in a 1° × 1° grid. Mean lineage net diversification was estimated by the mean phylogenetic root distance (MRD), the average number of nodes separating a species from the base of the palm phylogeny for the species in each grid cell. If evolutionary rate limits richness, then richness should increase with MRD. If time limits richness, then old, relict species (with low root distance) should predominantly occur in long‐inhabited and therefore species‐rich areas. Hence, richness should decrease with MRD. To determine the influence of net diversification across different time frames, MRD was computed for subtribe, genus and species levels within the phylogeny, and supplemented with the purely tip‐level measure, mean number of species per genus (MS/G). Correlations and regressions, in combination with eigenvector‐based spatial filtering, were used to assess the relationship between species richness, the net diversification measures, and potential environmental and geographical drivers. Results Species richness increased with all net diversification measures. The regression models showed that richness and the net diversification measures increased with decreasing (absolute) latitude and, less strongly, with increasing energy/temperature and water availability. These patterns therefore reflect net diversification at both deep and shallow levels in the phylogeny. Richness also increased with range in elevation, but this was only reflected in the MS/G pattern and therefore reflects recent diversification. Main conclusions The geographical patterns in palm species richness appear to be predominantly the result of elevated net diversification rates towards the equator and in warm, wet climates, sustained throughout most of the Tertiary. Late‐Tertiary orogeny has caused localized increases in net diversification rates that have also made a mark on the richness pattern.     

Pliocene forest dynamics as a primary driver of African bird speciation

Aim Montane tropics are areas of high endemism, and mechanisms driving this endemism have been receiving increasing attention at a global scale. A general trend is that climatic factors do not explain the species richness of species with small to medium‐sized geographic ranges, suggesting that geological and evolutionary processes must be considered. On the African continent, several hypotheses including both refugial and geographic uplift models have been advanced to explain avian speciation and diversity in the lowland forest and montane regions of central and eastern Africa; montane regions in particular are recognized as hotspots of vertebrate endemism. Here, we examine the possible role of these models in driving speciation in a clade of African forest robins. Location Africa. Methods We constructed the first robustly supported molecular phylogenetic hypothesis of forest robins. On this phylogeny, we reconstructed habitat‐based distributions and geographic distributions relative to the Albertine Rift. We also estimated the timing of lineage divergences via a molecular clock. Results Robust estimates of phylogenetic relationships and clock‐based divergences reject Miocene tectonic uplift and Pleistocene forest refugia as primary drivers of speciation in forest robins. Instead, our data suggest that most forest robin speciation took place in the Late Pliocene, from 3.2 to 2.2 Ma. Distributional patterns are complex, with the Albertine Rift region serving as a general east–west break across the group. Montane distributions are inferred to have evolved four times. Main conclusions Phylogenetic divergence dates coincide with a single period of lowland forest retraction in the late Pliocene, suggesting that most montane speciation resulted from the rapid isolation of populations in montane areas, rather than montane areas themselves being drivers of speciation. This conclusion provides additional evidence that Pliocene climate change was a major driver of speciation in broadly distributed African animal lineages. We further show that lowland forest robins are no older than their montane relatives, suggesting that lowland areas are not museums which house ‘ancient’ taxa; rather, for forest robins, montane areas should be viewed as living museums of a late Pliocene diversification event. A forest refugial pattern is operating in Africa, but it is not constrained to the Pleistocene.