Speciation Rates Research Articles

A diversification relay race from Caribbean-Mesoamerica to the Andes: historical biogeography of Xylophanes hawkmoths.

The regions of the Andes and Caribbean-Mesoamerica are both hypothesized to be the cradle for many Neotropical lineages, but few studies have fully investigated the dynamics and interactions between Neotropical bioregions. The New World hawkmoth genus Xylophanes is the most taxonomically diverse genus in the Sphingidae, with the highest endemism and richness in the Andes and Caribbean-Mesoamerica. We integrated phylogenomic and DNA barcode data and generated the first time-calibrated tree for this genus, covering 93.8% of the species diversity. We used event-based likelihood ancestral area estimation and biogeographic stochastic mapping to examine the speciation and dispersal dynamics of Xylophanes across bioregions. We also used trait-dependent diversification models to compare speciation and extinction rates of lineages associated with different bioregions. Our results indicate that Xylophanes originated in Caribbean-Mesoamerica in the Late Miocene, and immediately diverged into five major clades. The current species diversity and distribution of Xylophanes can be explained by two consecutive phases. In the first phase, the highest Xylophanes speciation and emigration rates occurred in the Caribbean-Mesoamerica, and the highest immigration rates occurred in the Andes, whereas in the second phase the highest immigration rates were found in Amazonia, and the Andes had the highest speciation and emigration rates.

Drivers of diversification in freshwater gastropods vary over deep time.

Unravelling the drivers of species diversification through geological time is of crucial importance for our understanding of long-term evolutionary processes. Numerous studies have proposed different sets of biotic and abiotic controls of speciation and extinction rates, but typically they were inferred for a single, long geological time frame. However, whether the impact of biotic and abiotic controls on diversification changes over time is poorly understood. Here, we use a large fossil dataset, a multivariate birth–death model and a comprehensive set of biotic and abiotic predictors, including a new index to quantify tectonic complexity, to estimate the drivers of diversification for European freshwater gastropods over the past 100 Myr. The effects of these factors on origination and extinction are estimated across the entire time frame as well as within sequential time windows of 20 Myr each. Our results find support for temporal heterogeneity in the factors associated with changes in diversification rates. While the factors impacting speciation and extinction rates vary considerably over time, diversity-dependence and topography are consistently important. Our study highlights that a high level of heterogeneity in diversification rates is best captured by incorporating time-varying effects of biotic and abiotic factors.

The multicausal twilight of\xa0South American native mammalian predators (Metatheria, Sparassodonta)

Sparassodonts were the apex mammalian predators of South America throughout most of the Cenozoic, diversifying into a wide array of niches including fox-like and even saber-toothed forms. Their extinction is still controversial, with different authors suggesting competition with other predators (placental carnivorans, terror birds, and carnivorous opossums), extinction of prey, and climate change as causal explanations. Here, we analyse these hypotheses using a novel approach implicating Bayesian analyses. We find that speciation and extinction rates of sparassodonts can be correlated with (i) intrinsic biotic factors such as changes in body mass and diversity of sparassodonts, (ii) extrinsic biotic factors such as potential prey diversity, and iii) extrinsic abiotic factors like the atmospheric CO2, sea level, temperature, and uplift of the Andes. Thus, sparassodonts are a good example of a multilevel mixed model of evolution, where various factors drove the evolutionary history of this clade in a pluralistic way. There is no evidence for competition between Sparassodonta and others predators, and the effect of competition in the face of extinctions of fossil species should be tested and not assumed. Furthermore, we propose a novel approach for evaluating the fossil record when performing macroevolutionary analyses.

No link between population isolation and speciation rate in squamate reptiles

Rates of species formation vary widely across the tree of life and contribute to massive disparities in species richness among clades. This variation can emerge from differences in metapopulation-level processes that affect the rates at which lineages diverge, persist, and evolve reproductive barriers and ecological differentiation. For example, populations that evolve reproductive barriers quickly should form new species at faster rates than populations that acquire reproductive barriers more slowly. This expectation implicitly links microevolutionary processes (the evolution of populations) and macroevolutionary patterns (the profound disparity in speciation rate across taxa). Here, leveraging extensive field sampling from the Neotropical Cerrado biome in a biogeographically controlled natural experiment, we test the role of an important microevolutionary process-the propensity for population isolation-as a control on speciation rate in lizards and snakes. By quantifying population genomic structure across a set of codistributed taxa with extensive and phylogenetically independent variation in speciation rate, we show that broad-scale patterns of species formation are decoupled from demographic and genetic processes that promote the formation of population isolates. Population isolation is likely a critical stage of speciation for many taxa, but our results suggest that interspecific variability in the propensity for isolation has little influence on speciation rates. These results suggest that other stages of speciation-including the rate at which reproductive barriers evolve and the extent to which newly formed populations persist-are likely to play a larger role than population isolation in controlling speciation rate variation in squamates.

Linking high diversification rates of rapidly growing Amazonian plants to geophysical landscape transformations promoted by Andean uplift

AbstractAmazonia is extremely biodiverse, but the mechanisms for the origin of this diversity are still under debate. We propose a diversification model for Amazonia based on the interplay of intrinsic clade functional traits, habitat associations and past geological events, using as a model group the species-rich Neotropical family Marantaceae. Our results show that the species richness of the lineage is predicted by functional strategy, rather than clade age, and thus the fast vs. slow growth functional trade-off is a major determinant of clade diversification in Marantaceae. Rapidly growing clades were mostly associated with highly productive habitats, and their origin and diversification dynamics matched the expansion of fertile soils mediated by Andean uplift c. 23 Mya. Fast-growth strategies probably led to fast molecular evolution, speeding up speciation rates and species accumulation, resulting in higher numbers of extant species. Our results indicate that pure allopatric-dispersal models disconnected from past geological and ecological forces may be inadequate for explaining the evolutionary and diversity patterns in Amazonian lowlands. We suggest that a coupling of the functional trait-niche framework with diversification dynamics provides insights into the evolutionary history of tropical forests and helps elucidate the mechanisms underlying the origin and evolution of its spectacular biodiversity.

Faster evolution of a premating reproductive barrier is not associated with faster speciation rates in New World passerine birds.

Why are speciation rates so variable across the tree of life? One hypothesis is that this variation is explained by how rapidly reproductive barriers evolve. We tested this hypothesis by conducting a comparative study of the evolution of bird song, a premating barrier to reproduction. Speciation in birds is typically initiated when geographically isolated (allopatric) populations evolve reproductive barriers. We measured the strength of song as a premating barrier between closely related allopatric populations by conducting 2339 field experiments to measure song discrimination for 175 taxon pairs of allopatric or parapatric New World passerine birds, and estimated recent speciation rates from molecular phylogenies. We found evidence that song discrimination is indeed an important reproductive barrier: taxon pairs with high song discrimination in allopatry did not regularly interbreed in parapatry. However, evolutionary rates of song discrimination were not associated with recent speciation rates. Evolutionary rates of song discrimination were also unrelated to latitude or elevation, but species with innate song (suboscines) evolved song discrimination much faster than species with learned song (oscines). We conclude that song is a key premating reproductive barrier in birds, but faster evolution of this reproductive barrier between populations does not consistently result in faster diversification between species.

Species-range-size distributions: Integrating the effects of speciation, transformation, and extinction.

The species‐range size distribution is a product of speciation, transformation of range‐sizes, and extinction. Previous empirical studies showed that it has a left‐skewed lognormal‐like distribution. We developed a new mathematical framework to study species‐range‐size distributions, one in which allopatric speciation, transformation of range size, and the extinction process are explicitly integrated. The approach, which we call the gain‐loss‐allopatric speciation model, allows us to explore the effects of various speciation scenarios. Our model captures key dynamics thought to lead to known range‐size distributions. We also fitted the model to empirical range‐size distributions of birds, mammals, and beetles. Since geographic range dynamics are linked to speciation and extinction, our model provides predictions for the dynamics of species richness. When a species‐range‐size distribution initially evolves away from the range sizes at which the likelihood of speciation is low, it tends to cause diversification slowdown even in the absence of (bio)diversity dependence in speciation rate. Using the mathematical model developed here, we give a potential explanation for how observed range‐size distributions emerge from range‐size dynamics. Although the framework presented is minimalistic, it provides a starting point for examining hypotheses based on more complex mechanisms.

Mountain radiations are not only rapid and recent: Ancient diversification of South American frog and lizard families related to Paleogene Andean orogeny and Cenozoic climate variations

Mountainous areas host a disproportionately large fraction of Earth's biodiversity, suggesting a causal relationship between mountain building and biological diversification. Mountain clade radiations are generally associated with changes in environment, climate, and the increase in heterogeneity therein during mountain building. However, examining the causal relationship between mountain building and diversification is a complex challenge, because isolating the effects of surface uplift from other abiotic (climate) or biotic variables is not straightforward. In this study, we investigate the relative contributions of abiotic climate-driven (temperature) and geology-driven (elevation) drivers on evolutionary rates of ancient groups of organisms in the South American Andes. We present regional curves of Andean elevation based on a recent compilation of paleo-elevational data back to the Late Cretaceous, and analyse the diversification history of six Andean frog and lizard families that originated equally far back in time. For two clades (Aromobatidae and Leptodactylidae), we find that they diversified most rapidly during the early phase of mountain building (Late Cretaceous - Paleogene), when the first high-elevation habitats emerged in South America. The diversification of one clade (Centrolenidae) is correlated with Cenozoic temperature variations, with higher speciation rates during warm periods. The last three clades (Dendrobatidae, Hemiphractidae and Liolaemidae) are best explained by environment-independent diversification, although for Liolaemidae, an almost equally strong positive correlation was found between speciation and Andean elevation since the late Eocene. Our findings imply that throughout the long-lived history of surface uplift in the Andes, mountain building drove the diversification of different clades at different times, while not directly affecting other clades. Our study illustrates the importance of paleogeographic reconstructions that capture the complexity and heterogeneity of mountain building in our understanding of the effects that a changing environment plays in shaping biodiversity patterns observed today.

Nuclear–plastid discordance indicates past introgression inEpidendrumspecies (Laeliinae: Orchidaceae) with highly variable chromosome numbers

AbstractHere we study a challenging group of karyotypically highly variable Epidendrum spp. using phylogenetic methods to help understand how hybridization/introgression contributes to karyotype evolution. We hypothesize that species with great chromosome number variation are a result of past hybridization/introgression. Conflicting topologies in trees constructed using separate plastid and nuclear datasets suggest past hybridization events that occurred most probably at least 3.7 Mya. A basic number x = 14 and substantial karyotype change followed by species divergence are suggested. Descending dysploidy and polyploidy were the most frequent changes estimated across the phylogenetic tree of the group. Two species, Epidendrum secundum and E. xanthinum, have probably experienced unidirectional gene flow involving their ancestors (the pollen recipients) and ancestors of E. puniceoluteum and E. denticulatum/E. flammeum, respectively, the pollen donors. However, it is not possible to say whether hybridization participated in the origin of E. secundum and E. xanthinum or merely contributed to their genomic divergence and karyotype change through introgression as has been observed in modern hybrid zones in Epidendrum. This pattern of introgression causing karyotype disruption and divergence could help explain the enigma of some highly diverse genera, such as Epidendrum. Further studies using a wider sampling of the genus could test if gene flow and karyotype variability are associated with the increase of speciation rates.

Improving phylogenetic resolution of the Lamiales using the complete plastome sequences of six Penstemon species.

The North American endemic genus Penstemon (Mitchell) has a recent geologic origin of ca. 3.6 million years ago (MYA) during the Pliocene/Pleistocene transition and has undergone a rapid adaptive evolutionary radiation with ca. 285 species of perennial forbs and sub-shrubs. Penstemon is divided into six subgenera occupying all North American habitats including the Arctic tundra, Central American tropical forests, alpine meadows, arid deserts, and temperate grasslands. Due to the rapid rate of diversification and speciation, previous phylogenetic studies using individual and concatenated chloroplast sequences have failed to resolve many polytomic clades. We investigated the efficacy of utilizing the plastid genomes (plastomes) of 29 species in the Lamiales order, including five newly sequenced Penstemon plastomes, for analyzing phylogenetic relationships and resolving problematic clades. We compared whole-plastome based phylogenies to phylogenies based on individual gene sequences (matK, ndhF, psaA, psbA, rbcL, rpoC2, and rps2) and concatenated sequences. We also We found that our whole-plastome based phylogeny had higher nodal support than all other phylogenies, which suggests that it provides greater accuracy in describing the hierarchal relationships among taxa as compared to other methods. We found that the genus Penstemon forms a monophyletic clade sister to, but separate from, the Old World taxa of the Plantaginaceae family included in our study. Our whole-plastome based phylogeny also supports the rearrangement of the Scrophulariaceae family and improves resolution of major clades and genera of the Lamiales.

Identifying traits that enable lizard adaptation to different habitats

AbstractAimSpecies adapt differently to contrasting environments, such as open habitats with sparse vegetation and forested habitats with dense forest cover. We investigated colonization patterns in the open and forested environments in the diagonal of open formations and surrounding rain forests (i.e. Amazonia and Atlantic Forest) in Brazil, tested whether the diversification rates were affected by the environmental conditions and identified traits that enabled species to persist in those environments.LocationSouth America, Brazil.TaxonSquamata, Lizards.MethodsWe used phylogenetic information and the current distribution of species in open and forested habitats to estimate ancestral ranges and identify range shifts relative to the current habitats. To evaluate whether these environments influenced species diversification, we tested 12 models using a Hidden Geographic State Speciation and Extinction analysis. Finally, we combined phylogenetic relatedness and species traits in a machine learning framework to identify the traits permitting adaptation in those contrasting environments.ResultsWe identified 41 total transitions between open and forested habitats, of which 80% were from the forested habitats to the open habitats. Widely distributed species had higher speciation, turnover, extinction, and extinction fraction rates than species in forested or open habitats, but had also the lower net diversification rate. Mean body temperature, microhabitat, female snout–vent length and diet were identified as putative traits that enabled adaptation to different environments, and phylogenetic relatedness was an important predictor of species occurrence.Main conclusionsTransitions from forested to open habitats are most common, highlighting the importance of habitat shift in current patterns of biodiversity. The combination of phylogenetic reconstruction of ancestral distributions and the machine learning framework enables us to integrate organismal trait data, environmental data and evolutionary history in a manner that could be applied on a global scale.

Key roles for the freezing line and disturbance in driving the low plant species richness of temperate regions

AbstractAimAt the macroscale, climate strongly correlates with species richness gradients, resulting from differences in in‐situ diversification and dispersal. One historical explanation for the pattern is that regions spanning temperate climates contain few species because past disturbances have generated high extinction rates, and species from tropical regions are unable to easily colonize temperate regions. We test these postulates for Himalayan plants, which span subtropical to temperate climates over steep elevational gradients.LocationHimalaya.Time periodPresent day.Major taxa studiedAngiosperms.MethodsWe use a comprehensive survey of 31 floras to document the elevational and geographical distributions of native Himalayan plants, augmented by field studies of trees in both the east and west Himalaya. We use grade of membership models to cluster species according to locations shared and phylogenetic analysis to evaluate diversification rates.ResultsSpecies fall into four cohesive biotas, organized by climate. Points of turnover between biotas occur where the mean minimum temperature of the coldest month is approximately 0 °C (2,000–2,500 m), and at the point of occasional annual freezing (1,000–1,500 m); these boundaries run the length of the Himalaya. The patterns are retained when we consider whole clades rather than species. All plants (and the subsets trees, herbs and shrubs) belonging to the biota above the 2,000–2,500 m line have higher recent speciation rates than those lower down.Main conclusionsWe attribute the high rate of recent speciation in temperate climates to high rates of turnover, creating ecological and geographical opportunity. The high elevation biota has few species, but spans the largest area, implying species numbers are far from any carrying capacity, at least with respect to accumulation of allopatric forms. This study thus links climatic restrictions of clades to differences in diversification rates, and by inference species numbers.

Global dispersal and diversification in ground beetles of the subfamily Carabinae

The origin and diversification process of lineages of organisms that are currently widely distributed among continents is an interesting subject for exploring the evolutionary history of global species diversity. Ground beetles of the subfamily Carabinae are flightless except for one lineage, but nevertheless occur on all continents except Antarctica. Here, we used sequence data from ultraconserved elements to reconstruct the phylogeny, divergence time, biogeographical history, ancestral state of hind wings and changes in the speciation rate of Carabinae. Our results show that Carabinae originated in the Americas and diversified into four tribes during the period from the late Jurassic to the late Cretaceous, with two in South America (Celoglossini) and Australasia (Pamborini) and two in Laurasia (Cychrini and Carabini). The ancestral Carabinae were inferred to be winged; three of four tribes (Cychrini, Ceglossini and Pamborini) have completely lost their hind wings and flight capability. The remaining tribe, Carabini, diverged into the subtribes Carabina (wingless) and Calosomina (winged) in the Oligocene. Carabina originated in Europe, spread over Eurasia and diversified into approximately 1000 species, accounting for around 60% of all Carabinae species. Calosomina that were flight-capable dispersed from North America or Eurasia to South America, Australia, and Africa, and then flightless lineages evolved on oceanic islands and continental highlands. The speciation rate increased in the Cychrini and Carabini clades in Eurasia. Within Carabini, the speciation rate was higher for wingless than winged states. Our study showed that the global distribution of Carabinae resulted from ancient dispersal before the breakup of Gondwana and more recent dispersal through flight around the world. These patterns consequently illustrate the causal relationships of geographical history, evolution of flightlessness, and the global distribution and species diversity of Carabinae.

Macroevolutionary consequences of mast seeding.

Masting characterizes large, intermittent and highly synchronous seeding events among individual plants and is found throughout the plant Tree of Life (ToL). Although masting can increase plant fitness, little is known about whether it results in evolutionary changes across entire clades, such as by promoting speciation or enhanced trait selection. Here, we tested if masting has macroevolutionary consequences by combining the largest existing dataset of population-level reproductive time series and time-calibrated phylogenetic tree of vascular plants. We found that the coefficient of variation (CVp) of reproductive output for 307 species covaried with evolutionary history, and more so within clades than expected by random. Speciation rates estimated at the species level were highest at intermediate values of CVp and regional-scale synchrony (Sr) in seed production, that is, there were unimodal correlations. There was no support for monotonic correlations between either CVp or Sr and rates of speciation or seed size evolution. These results were robust to different sampling decisions, and we found little bias in our dataset compared with the wider plant ToL. While masting is often adaptive and encompasses a rich diversity of reproductive behaviours, we suggest it may have few consequences beyond the species level.This article is part of the theme issue ‘The ecology and evolution of synchronized seed production in plants’.

Pulled Diversification Rates, Lineages-Through-Time Plots, and Modern Macroevolutionary Modeling.

Estimating time-dependent rates of speciation and extinction from dated phylogenetic trees of extant species (timetrees), and determining how and why they vary, is key to understanding how ecological and evolutionary processes shape biodiversity. Due to an increasing availability of phylogenetic trees, a growing number of process-based methods relying on the birth–death model have been developed in the last decade to address a variety of questions in macroevolution. However, this methodological progress has regularly been criticized such that one may wonder how reliable the estimations of speciation and extinction rates are. In particular, using lineages-through-time (LTT) plots, a recent study has shown that there are an infinite number of equally likely diversification scenarios that can generate any timetree. This has led to questioning whether or not diversification rates should be estimated at all. Here, we summarize, clarify, and highlight technical considerations on recent findings regarding the capacity of models to disentangle diversification histories. Using simulations, we illustrate the characteristics of newly proposed “pulled rates” and their utility. We recognize that the recent findings are a step forward in understanding the behavior of macroevolutionary modeling, but they in no way suggest we should abandon diversification modeling altogether. On the contrary, the study of macroevolution using phylogenetic trees has never been more exciting and promising than today. We still face important limitations in regard to data availability and methods, but by acknowledging them we can better target our joint efforts as a scientific community. [Birth–death models; extinction; phylogenetics; speciation.]

Earth history events shaped the evolution of uneven biodiversity across tropical moist forests

Far from a uniform band, the biodiversity found across Earth's tropical moist forests varies widely between the high diversity of the Neotropics and Indomalaya and the relatively lower diversity of the Afrotropics. Explanations for this variation across different regions, the "pantropical diversity disparity" (PDD), remain contentious, due to difficulty teasing apart the effects of contemporary climate and paleoenvironmental history. Here, we assess the ubiquity of the PDD in over 150,000 species of terrestrial plants and vertebrates and investigate the relationship between the present-day climate and patterns of species richness. We then investigate the consequences of paleoenvironmental dynamics on the emergence of biodiversity gradients using a spatially explicit model of diversification coupled with paleoenvironmental and plate tectonic reconstructions. Contemporary climate is insufficient in explaining the PDD; instead, a simple model of diversification and temperature niche evolution coupled with paleoaridity constraints is successful in reproducing the variation in species richness and phylogenetic diversity seen repeatedly among plant and animal taxa, suggesting a prevalent role of paleoenvironmental dynamics in combination with niche conservatism. The model indicates that high biodiversity in Neotropical and Indomalayan moist forests is driven by complex macroevolutionary dynamics associated with mountain uplift. In contrast, lower diversity in Afrotropical forests is associated with lower speciation rates and higher extinction rates driven by sustained aridification over the Cenozoic. Our analyses provide a mechanistic understanding of the emergence of uneven diversity in tropical moist forests across 110 Ma of Earth's history, highlighting the importance of deep-time paleoenvironmental legacies in determining biodiversity patterns.

Conserved ancestral tropical niche but different continental histories explain the latitudinal diversity gradient in brush-footed butterflies

The global increase in species richness toward the tropics across continents and taxonomic groups, referred to as the latitudinal diversity gradient, stimulated the formulation of many hypotheses to explain the underlying mechanisms of this pattern. We evaluate several of these hypotheses to explain spatial diversity patterns in a butterfly family, the Nymphalidae, by assessing the contributions of speciation, extinction, and dispersal, and also the extent to which these processes differ among regions at the same latitude. We generate a time-calibrated phylogeny containing 2,866 nymphalid species (~45% of extant diversity). Neither speciation nor extinction rate variations consistently explain the latitudinal diversity gradient among regions because temporal diversification dynamics differ greatly across longitude. The Neotropical diversity results from low extinction rates, not high speciation rates, and biotic interchanges with other regions are rare. Southeast Asia is also characterized by a low speciation rate but, unlike the Neotropics, is the main source of dispersal events through time. Our results suggest that global climate change throughout the Cenozoic, combined with tropical niche conservatism, played a major role in generating the modern latitudinal diversity gradient of nymphalid butterflies.

The little fishes that could: smaller fishes demonstrate slow body size evolution but faster speciation in the family Percidae

Abstract Body size affects numerous aspects of organismal biology and many factors have been invoked to explain body size distributions in a macroecological and macroevolutionary context. Body size in the freshwater fish family Percidae is strongly right-skewed (i.e. dominated by small sizes), with small body size potentially being associated with fast water habitats. We constructed a new species-level, multi-locus, time-calibrated phylogeny of Percidae, and used it to test for changes in the rate and pattern of maximum body size evolution. We also tested whether speciation rates varied as a function of body size. We found that Etheostomatinae evolved towards a smaller adaptive optimum in body size compared to the other subfamilies of Percidae, and that this shift was associated with a reduction in the rate of body size evolution. Speciation rates were associated with body size across percids, showing a peak around small to medium body size. Small body size appears to partially, but not fully, explain the diversity of small percids, as many darters fall well below the “optimum” body size. Reinforcement of selection for small body size via selection for novel morphologies or via sexual selection may help to fully explain the remarkable diversity of darter radiation.

The Bamboo Rhizome Evolution in China Is Driven by Geographical Isolation and Trait Differentiation

Plant endemic species are the result of continuous evolution under the combined action of long-term climatic and geological conditions. There are 534 bamboo species in China, and 371 endemic species account for about 70% of all bamboo species. However, little is known about the differences in the rhizome evolution rate between endemic and non-endemic bamboos. Here, we collected morphological traits (height and leaf length) and environmental variables (including climate, space, and soil) of all 534 Chinese bamboo species to determine the relative contribution of environmental factors and traits of bamboo rhizome evolution with different endemism by structural equation modeling. We found that endemic bamboo had a higher speciation rate than non-endemic bamboo. The distribution centers of sympodial bamboos are mainly located in the mountain range of southwest China, while amphipodial and monopodial bamboos are distributed with higher latitude farther east in China. The height of non-endemic sympodial and monopodial bamboos was significantly higher than endemic sympodial and monopodial bamboos. The leaf length of non-endemic sympodial bamboos was significantly higher than endemic sympodial bamboo, while the leaf length of non-endemic amphipodial bamboo was significantly lower than endemic amphipodial bamboo. Environmental factors and traits explain 47% of the evolutionary variation of non-endemic bamboo species, while they explain 17% of that of endemic bamboo species. Longitude, latitude, and leaf length were the critical factors in the rhizome evolution of non-endemic bamboo, while longitude and height were the critical factors in the rhizome evolution of endemic bamboo. Our results imply that for higher species formation rates, endemic bamboo should be more concerned than non-endemic bamboo in the process of bamboo rhizome evolution. It will likely appear that new non-endemic bamboo species have a short leaf length in higher latitude and farther east in China (the lower Yangtze plain), as well as new endemic bamboo species with a low height farther east in China (the Wuyi Mountains). Meanwhile, ancient non-endemic bamboo with a long leaf length in Yunnan Province and ancient endemic bamboo with a high height in the Hengduan Mountains may be more likely to become extinct. Our findings highlight the importance of understanding the difference in the rhizome evolution of endemic and non-endemic bamboos, which provides new insights into the conservation of Chinese bamboo biodiversity.

Adding stage‐structure to a spatial neutral model: implications for explaining local and regional patterns of biodiversity

Theoretical ecologists have analysed a range of neutral models but few including stage structure. Here we introduce a stage‐structured neutral model, by extending the standard spatial neutral model to have two‐stage classes: a juvenile stage and a reproductive stage. We find that formulas for biodiversity patterns (e.g. species–area relationships and species abundance distributions) of reproductives in the stage‐structured model can be obtained from the corresponding standard formulas via a parameter rescaling, which involves calculating an effective speciation rate parameter and an effective dispersal parameter. This is useful because it means existing knowledge about the non‐stage‐structured model can be transferred to the stage‐structured model, providing that applications focus exclusively on the reproductive stage. One surprising implication is that the presence of a juvenile stage can substantially increase the species richness of reproductive individuals: a juvenile stage with a length fraction k that of the reproductive stage increases reproductive species richness by roughly the same factor. We apply our new formulas to a case study of tropical forest trees in Panama and find that while the stage‐structured model makes different predictions than the standard model, it does not fix known problems with cross‐scale predictions. We speculate that some of our results, in particular the result that the presence of a juvenile stage increases community diversity, likely apply to non‐neutral systems as well.