Shifts in Climate Foster Exceptional Opportunities for Species Radiation: The Case of South African Geraniums

Climate change is often assumed to be a major driver of biodiversity loss. However, it can also set the stage for novel diversification in lineages with the evolutionary ability to colonize new environments. Here we tested if the extraordinary evolutionary success of the genus Pelargonium was related to the ability of its species to capitalize on the climate niche variation produced by the historical changes in southern Africa. We evaluated the relationship between rates of climate niche evolution and diversification rates in the main Pelargonium lineages and disentangled the roles of deep and recent historical events in the modification of species niches. Pelargonium clades exhibiting higher ecological differentiation along summer precipitation (SPP) gradients also experienced higher diversification rates. Faster rates of niche differentiation in spatially structured variables, along with lower levels of niche overlap among closely related species, suggest recent modification in species niches (e.g. dispersal or range shift) and niche lability. We suggest that highly structured SPP gradients established during the aridification process within southern Africa, in concert with niche lability and low niche overlap, contributed to species divergence. These factors are likely to be responsible for the extensive diversification of other lineages in this diversity hot spot.     

Ecological release exposes genetically based niche variation

The evolutionary trajectories of ecological niches have profound impacts on community, population and speciation dynamics, yet the underlying causes of niche lability vs. stasis are poorly understood. Here, we conducted a field experiment to quantify the effects of competition and, conversely, competitive release on the microevolutionary processes driving microhabitat niche evolution in an annual plant population restricted to California vernal pool wetlands. Removing competitors generated a strong increase in mean fitness, the exposure of genetically based niche variation and directional selection for niche evolution in the experimental population. In contrast, genetic variation in the microhabitat niche and directional selection for niche evolution were not detected in individuals growing with competitors. These results indicate that ecological opportunity (here, the removal of competitors) can trigger the immediate expression of latent, heritable niche variation that is necessary for rapid evolutionary responses; conversely, competitors may restrict niche evolution, contributing to niche conservatism in saturated communities.     

Ecological niche dimensionality and the evolutionary diversification of stick insects

The degree of phenotypic divergence and reproductive isolation between taxon pairs can vary quantitatively, and often increases as evolutionary divergence proceeds through various stages, from polymorphism to population differentiation, ecotype and race formation, speciation, and post-speciational divergence. Although divergent natural selection promotes divergence, it does not always result in strong differentiation. For example, divergent selection can fail to complete speciation, and distinct species pairs sometimes collapse ('speciation in reverse'). Widely-discussed explanations for this variability concern genetic architecture, and the geographic arrangement of populations. A less-explored possibility is that the degree of phenotypic and reproductive divergence between taxon pairs is positively related to the number of ecological niche dimensions (i.e., traits) subject to divergent selection. Some data supporting this idea stem from laboratory experimental evolution studies using Drosophila, but tests from nature are lacking. Here we report results from manipulative field experiments in natural populations of herbivorous Timema stick insects that are consistent with this 'niche dimensionality' hypothesis. In such insects, divergent selection between host plants might occur for cryptic colouration (camouflage to evade visual predation), physiology (to detoxify plant chemicals), or both of these niche dimensions. We show that divergent selection on the single niche dimension of cryptic colouration can result in ecotype formation and intermediate levels of phenotypic and reproductive divergence between populations feeding on different hosts. However, greater divergence between a species pair involved divergent selection on both niche dimensions. Although further replication of the trends reported here is required, the results suggest that dimensionality of selection may complement genetic and geographic explanations for the degree of diversification in nature.     

Dispersal is associated with morphological innovation,but not increased diversification,in Cyphostemma (Vitaceae)

Multiple processes − including dispersal, morphological innovation, and habitat change − are frequently cited as catalysts for increased diversification. We investigate these processes and the causal linkages among them in the genus Cyphostemma (Vitaceae), a clade comprising ∼200 species that is unique in the Vitaceae for its diversity of growth habits. We reconstruct time‐calibrated evolutionary relationships among 64 species in the genus using five nuclear and chloroplast markers and infer the group's morphological and biogeographic history. We test for changes in speciation rate and evaluate the temporal association and sequencing of events with respect to dispersal, habitat change, and morphological evolution using a Monte Carlo simulation approach. In Cyphostemma, neither dispersal nor morphological evolution is associated with shifts in speciation rate, but dispersal is associated with evolutionary shifts in growth form. Evolution of stem succulence, in particular, is associated with adaptation to local, pre‐existing conditions following long‐distance dispersal, not habitat change in situ. We suggest that the pattern of association between dispersal, morphological innovation, and diversification may depend on the particular characters under study. Lineages with evolutionarily labile characters, such as stem succulence, do not necessarily conform to the notion of niche conservatism and instead demonstrate remarkable morphological adaptation to local climate and edaphic conditions following dispersal.     

Evolutionary and ecological causes of the latitudinal diversity gradient in hylid frogs: treefrog trees unearth the roots of high tropical diversity

Why are there more species in the tropics than in temperate regions? In recent years, this long-standing question has been addressed primarily by seeking environmental correlates of diversity. But to understand the ultimate causes of diversity patterns, we must also examine the evolutionary and biogeographic processes that directly change species numbers (i.e., speciation, extinction, and dispersal). With this perspective, we dissect the latitudinal diversity gradient in hylid frogs. We reconstruct a phylogeny for 124 hylid species, estimate divergence times and diversification rates for major clades, reconstruct biogeographic changes, and use ecological niche modeling to identify climatic variables that potentially limit dispersal. We find that hylids originated in tropical South America and spread to temperate regions only recently (leaving limited time for speciation). There is a strong relationship between the species richness of each region and when that region was colonized but not between the latitudinal positions of clades and their rates of diversification. Temperature seasonality seemingly limits dispersal of many tropical clades into temperate regions and shows significant phylogenetic conservatism. Overall, our study illustrates how two general principles (niche conservatism and the time-for-speciation effect) may help explain the latitudinal diversity gradient as well as many other diversity patterns across taxa and regions.     

Diversification and the evolution of dispersal ability in the tribe Brassiceae (Brassicaceae)

MethodsThe phylogenetic analysis included 72 taxa sampled from across the Brassiceae and included both nuclear and chloroplast markers. Dispersal-related fruit characters were scored and climate information for each taxon was retrieved from a database. Correlations between fruit traits, seed characters, habitat, range and climate were determined, together with trait-dependent diversification rates.ConclusionsThis study provides evidence that the evolution of increased dispersal ability and larger seed size, which may increase establishment ability, can also influence macro-evolutionary processes, possibly by increasing the propensity for long-distance dispersal. In particular, it may increase speciation and consequent diversification rates by increasing the likelihood of geographic and thereby reproductive isolation.     

Speciational evolution of coloration in the genus Carduelis

Sexual selection has been hypothesized to promote speciation, but evidence relating sexual selection to differences in speciation rates among taxa is equivocal. We note that evolutionary changes in ornaments are the link connecting sexual selection to speciation, and that ornament evolution is influenced by many factors so that its relationship with the strength of sexual selection may not be linear. We test if the evolution of ornamental coloration in Carduelis finches is related with speciation and if more ornamented lineages speciate more. We found that coloration evolves with a speciational pattern, but we found no evidence that the evolutionary changes associated with speciation are predominantly gains in ornamentation. The speciational pattern was found for both carotenoid- and melanin-based coloration, suggesting that traits putatively under stronger sexual selection by female choice (carotenoid coloration) are not the sole ones facilitating reproductive isolation. We conclude that in the genus Carduelis the evolutionary lability of ornaments influences speciation more than the strength of sexual selection, and we suggest that ornament lability should be considered as a possible causal factor in studies comparing cladogenesis among taxa.     

蜘蛛飞航研究进展

种群扩散能够避免生物类群在同一栖息地中的资源竞争和同类相残, 促进基因流动, 具有重要的进化和生态意义。相比于昆虫的迁飞行为, 某些不具翅的节肢动物也能够借助细丝的作用而实现远距离扩散, 即飞航。蜘蛛为肉食性动物, 是陆地生态系统的重要组成部分, 在生态系统中防控害虫方面作用显著。飞航是蜘蛛的一种重要扩散方式, 在其物种形成、生态位分化及害虫防控等方面具有重要意义, 但有关其飞航扩散的报道较少。本文回顾了蜘蛛飞航的研究历程, 大致可分为三个阶段; 总结了其研究现状, 包括飞航蜘蛛的优势类群及特征、飞航过程中的三个环节(即起飞、空中航行和降落)及影响因子、两种飞航机制及四种相关模型、飞航蜘蛛采集和研究方法。同时基于过往研究, 指出了未来蜘蛛飞航研究的重点领域: 加强飞航蜘蛛类群鉴定工作, 继续深度挖掘飞航数据与环境因子间的关系, 探索蜘蛛飞航的全过程(包括起飞、空中航行和降落), 更加明确蜘蛛飞航作为一种扩散方式的意义。本综述将为蜘蛛飞航的知识普及、资源保护和利用等方面提供重要参考。     

Speciation and the evolution of dispersal along environmental gradients

We analyze the joint evolution of an ecological character and of dispersal distance in asexual and sexual populations inhabiting an environmental gradient. Several interesting phenomena resulting from the evolutionary interplay of these characters are revealed. First, asexual and sexual populations exhibit two analogous evolutionary regimes, in which either speciation in the ecological character occurs in conjunction with evolution of short-range dispersal, or dispersal distance remains high and speciation does not occur. Second, transitions between these two regimes qualitatively differ between asexual and sexual populations, with the former showing speciation with long-range dispersal and the latter showing no speciation with short-range dispersal. Third, a phenotypic gradient following the environmental gradient occurs only in the last case, i.e., for non-speciating sexual populations evolving towards short-range dispersal. Fourth, the transition between the evolutionary regimes of long-range dispersal with no speciation and short-range dispersal with speciation is typically abrupt, mediated by a positive feedback between incipient speciation and the evolution of short-range dispersal. Fifth, even though the model of sexual evolution analyzed here does not permit assortative mating preferences, speciation occurs for a surprisingly wide range of conditions. This illustrates that dispersal evolution is a powerful alternative to preference evolution in enabling spatially distributed sexual populations to respond to frequency-dependent disruptive selection.     

Plant diversity and ecology on the Qinghai–Tibet Plateau

The Qinghai–Tibet Plateau (QTP) comprises a platform (sometimes called the Qinghai–Tibet Plateau sensu stricto), the Himalayas, and Hengduan Mountains (Liang et al.,2018; Mao et al.,2021). The latter two parts and adjacent highlands are also called the Pan-Himalaya. Numerous plants are distributed there with many endemic species, probably because of the high diverse landscapes created by continuous geological and climatic activities (Favre et al.,2015; Mao et al.,2021). As the well known biodiversity hotspot of the alpine plants in the world (Sun et al.,2017), many studies have been conducted on evolutionary origin and ecological adaptation of those species occurring in the QTP (e.g., Wen et al.,2014, 2019; Zhang et al.,2019). In the present special issue, we collected 15 related papers on this topic. Among them, two are invited reviews. Mao et al. (2021) provide a comprehensive review of evolutionary origin of species diversity on the QTP. Especially, they outlined major disputes and likely causes in this research topic, including circumscribing and naming the QTP, the QTP uplifts, dating of molecular phylogenetic trees, non-causal correlations between QTP uplifts and species diversification and the unified ice sheet. The authors also summarized genomic advancements related to high-altitude adaptation of both plants and animals. Tong et al. (2021) reviewed the reproductive strategies of animal-pollinated alpine plants on the QTP, involving pollination system, pollen limitation, self-pollination, and sexual system. In this region, 95.4% of animal-pollinated plants are pollinated by insects (i.e., bees, moths, butterflies, and flies) with only 4% by vertebrates (i.e., bats and birds). Self-pollination through self-compatibility shift from outcrossing has become an effective reproductive strategy to overcome pollen limitation in alpine plants. The other 13 research papers aimed to address origin and adaptation of alpine flora involving three major lines of evidence: genomics, ecology, and paleobotany. We hope that the collection of these papers will increase our understanding of the origin, speciation, and adaptation of alpine species on the QTP.     

Synergistic selection between ecological niche and mate preference primes diversification

The ecological niche and mate preferences have independently been shown to be important for the process of speciation. Here, we articulate a novel mechanism by which ecological niche use and mate preference can be linked to promote speciation. The degree to which individual niches are narrow and clustered affects the strength of divergent natural selection and population splitting. Similarly, the degree to which individual mate preferences are narrow and clustered affects the strength of divergent sexual selection and assortative mating between diverging forms. This novel perspective is inspired by the literature on ecological niches; it also explores mate preferences and how they may contribute to speciation. Unlike much comparative work, we do not search for evolutionary patterns using proxies for adaptation and sexual selection, but rather we elucidate how ideas from niche theory relate to mate preference, and how this relationship can foster speciation. Recognizing that individual and population niches are conceptually and ecologically linked to individual and population mate preference functions will significantly increase our understanding of rapid evolutionary diversification in nature. It has potential to help solve the difficult challenge of testing the role of sexual selection in the speciation process. We also identify ecological factors that are likely to affect individual niche and individual mate preference in synergistic ways and as a consequence to promote speciation. The ecological niche an individual occupies can directly affect its mate preference. Clusters of individuals with narrow, differentiated niches are likely to have narrow, differentiated mate preference functions. Our approach integrates ecological and sexual selection research to further our understanding of diversification processes. Such integration may be necessary for progress because these processes seem inextricably linked in the natural world.     

Summary and perspective on evolutionary ecology of fishes

This special issue of Evolutionary Ecology provides ten papers that have been presented at a conference on Evolutionary Ecology of Fishes in 2009. In addition to briefly summarizing the main content of the papers which is related to adaptive radiations, processes of ecological divergence, and fisheries-induced evolution, we review and synthesize in short the recent advance in studies on evolutionary ecology of fishes. We conclude that fishes are excellent model systems to study evolutionary ecology of animals, and suggest three promising new research avenues; (1) the contribution of behavioural processes to evolution, in particular the consideration of animal personalities and predator–prey interactions, (2) metabolic physiology and parasite-host interactions as new niche dimensions to be considered for adaptive diversification, and (3) the opportunities for mechanistic understanding of adaptation and speciation emerging from new genetic tools.     

Can the tropical conservatism hypothesis explain temperate species richness patterns? An inverse latitudinal biodiversity gradient in the New World snake tribe Lampropeltini

Aim  A latitudinal gradient in species richness, defined as a decrease in biodiversity away from the equator, is one of the oldest known patterns in ecology and evolutionary biology. However, there are also many known cases of increasing poleward diversity, forming inverse latitudinal biodiversity gradients. As only three processes (speciation, extinction and dispersal) can directly affect species richness in areas, similar factors may be responsible for both classical (high tropical diversity) and inverse (high temperate diversity) gradients. Thus, a modified explanation for differential species richness which accounts for both patterns would be preferable to one which only explains high tropical biodiversity.
Location  The New World.
Methods  We test several proposed ecological, temporal, evolutionary and spatial explanations for latitudinal diversity gradients in the New World snake tribe Lampropeltini, which exhibits its highest biodiversity in temperate regions.
Results  We find that an extratropical peak in species richness is not explained by latitudinal variation in diversification rate, the mid-domain effect, or Rapoport's rule. Rather, earlier colonization and longer duration in the temperate zones allowing more time for speciation to increase biodiversity, phylogenetic niche conservatism limiting tropical dispersal and the expansion of the temperate zones in the Tertiary better explain inverse diversity gradients in this group.
Main conclusions  Our conclusions are the inverse of the predictions made by the tropical conservatism hypothesis to explain higher biodiversity near the equator. Therefore, we suggest that the processes invoked are not intrinsic to the tropics but are dependent on historical biogeography to determine the distribution of species richness, which we refer to as the 'biogeographical conservatism hypothesis'.     

Ecological opportunity and diversification in a continental radiation of birds: climbing adaptations and cladogenesis in the Furnariidae

Ecological theories of adaptive radiation predict that ecological opportunity stimulates cladogenesis through its effects on competitive release and niche expansion. Given that key innovations may confer ecological opportunity, we investigated the effect of the acquisition of climbing adaptations on rates of cladogenesis in a major avian radiation, the Neotropical bird family Furnariidae, using a species-level phylogeny. Morphological specializations for vertical climbing originated in the woodcreepers ～23 million years ago, well before that adaptation occurred in woodpeckers (Picidae) or in other potential competitors in South America. This suggests that the acquisition of climbing adaptations conferred ample ecological opportunity to early woodcreepers. Nonetheless, we found that increases in speciation rates in Furnariidae did not coincide with the acquisition of climbing adaptations and that the relationship between the accumulation of climbing adaptations and rates of speciation was negative. In addition, we did not detect a diversity-dependent decline in woodcreeper diversification rates consistent with saturation of the trunk-climbing niche. These findings do not support the hypothesis that ecological opportunity related to trunk foraging stimulated cladogenesis in this radiation. The negative effect of climbing on diversification may be mediated by an indirect positive effect of climbing on dispersal ability, which may reduce speciation rates over evolutionary timescales.     

Diversification and biogeographic patterns in four island radiations of passerine birds

Declining diversification rates over time are a well-established evolutionary pattern, often interpreted as indicating initial rapid radiation with filling of ecological niche space. Here, we test the hypothesis that island radiations may show constant net diversification rates over time, due to continued expansion into new niche space in highly dispersive taxa. We investigate diversification patterns of four passerine bird families originating from the Indo-Pacific archipelagos, and link these to biogeographic patterns to provide independent indications of niche filling. We find a declining diversification rate for only one family, the Paradisaeidae (41 species). These are almost completely restricted to New Guinea, and have on average smaller species ranges and higher levels of species richness within grid cells than the other three families. In contrast, we cannot reject constant diversification rates for Campephagidae (93 species), Oriolidae (35 species), and Pachycephalidae (53 species), groups that have independently colonized neighboring archipelagos and continents. We propose that Paradisaeidae have reached the diversity limit imposed by their restricted distribution, whereas high dispersal and colonization success across the geologically dynamic Indo-Pacific archipelagos may have sustained high speciation rates for the other three families. Alternatively, increasing extinction rates may have obscured declining speciation rates in those three phylogenies.     

Genomic and phenomic analysis of island ant community assembly

Island biodiversity has long fascinated biologists as it typically presents tractable systems for unpicking the eco‐evolutionary processes driving community assembly. In general, two recurring themes are of central theoretical interest. First, immigration, diversification, and extinction typically depend on island geographical properties (e.g., area, isolation, and age). Second, predictable ecological and evolutionary trajectories readily occur after colonization, such as the evolution of adaptive trait syndromes, trends toward specialization, adaptive radiation, and eventual ecological decline. Hypotheses such as the taxon cycle draw on several of these themes to posit particular constraints on colonization and subsequent eco‐evolutionary dynamics. However, it has been challenging to examine these integrated dynamics with traditional methods. Here, we combine phylogenomics, population genomics and phenomics, to unravel community assembly dynamics among Pheidole (Hymenoptera, Formicidae) ants in the isolated Fijian archipelago. We uphold basic island biogeographic predictions that isolated islands accumulate diversity primarily through in situ evolution rather than dispersal, and population genomic support for taxon cycle predictions that endemic species have decreased dispersal ability and demography relative to regionally widespread taxa. However, rather than trending toward island syndromes, ecomorphological diversification in Fiji was intense, filling much of the genus‐level global morphospace. Furthermore, while most endemic species exhibit demographic decline and reduced dispersal, we show that the archipelago is not an evolutionary dead‐end. Rather, several endemic species show signatures of population and range expansion, including a successful colonization to the Cook islands. These results shed light on the processes shaping island biotas and refine our understanding of island biogeographic theory.     

The evolution of conditional dispersal and reproductive isolation along environmental gradients

Dispersal modulates gene flow throughout a population's spatial range. Gene flow affects adaptation at local spatial scales, and consequently impacts the evolution of reproductive isolation. A recent theoretical investigation has demonstrated that local adaptation along an environmental gradient, facilitated by the evolution of limited dispersal, can lead to parapatric speciation even in the absence of assortative mating. This and other studies assumed unconditional dispersal, so individuals start dispersing without regard to local environmental conditions. However, many species disperse conditionally; their propensity to disperse is contingent upon environmental cues, such as the degree of local crowding or the availability of suitable mates. Here, we use an individual-based model in continuous space to investigate by numerical simulation the relationship between the evolution of threshold-based conditional dispersal and parapatric speciation driven by frequency-dependent competition along environmental gradients. We find that, as with unconditional dispersal, parapatric speciation occurs under a broad range of conditions when reproduction is asexual, and under a more restricted range of conditions when reproduction is sexual. In both the asexual and sexual cases, the evolution of conditional dispersal is strongly influenced by the slope of the environmental gradient: shallow environmental gradients result in low dispersal thresholds and high dispersal distances, while steep environmental gradients result in high dispersal thresholds and low dispersal distances. The latter, however, remain higher than under unconditional dispersal, thus undermining isolation by distance, and hindering speciation in sexual populations. Consequently, the speciation of sexual populations under conditional dispersal is triggered by a steeper gradient than under unconditional dispersal. Enhancing the disruptiveness of frequency-dependent selection, more box-shaped competition kernels dramatically lower the speciation-enabling slope of the environmental gradient.     

Increased seed dispersal potential towards geographic range limits in a Pacific coast dune plant

Dispersal may be favoured at geographic range edges by unstable population and metapopulation dynamics. However, dispersal may also evolve in response to geographic variation in other life-history traits, especially the mating system. Here, increased dispersal at range margins was tested for with a range-wide analysis of seed dispersal and mating system traits in Abronia umbellata, a plant endemic to Pacific coastal dunes of North America. Seeds disperse within winged anthocarps. Anthocarps from 34 populations varied widely in wing size (mass-corrected wing index). Wing index correlated negatively with threshold wind velocity for dispersal in wind tunnel tests, suggesting that wings facilitate tumbling over open sandy substrate. As predicted, wing index increased and threshold velocity decreased towards both range limits. Flower size, herkogamy and self-incompatibility declined towards range limits, indicating a shift to self-fertilization, and flower size and wing index correlated negatively. However, the increase in wing index towards range limits remained after statistically controlling flower size. These results are consistent with selection favouring dispersal at range margins. The evolutionary lability of dispersal across the range may affect the interaction between selection and gene flow in the establishment and maintenance of geographic range limits.     

Dispersal has inhibited avian diversification in Australasian archipelagoes

Different models of speciation predict contrasting patterns in the relationship between the dispersal ability of lineages and their diversification rates. This relationship is expected to be negative in isolation-limited models and positive in founder-event models. In addition, the combination of negative and positive effects of dispersal on speciation can result in higher diversification rates at intermediate levels of dispersal ability. Using molecular phylogenies to estimate diversification rates, and wing morphology to estimate dispersal ability, we analysed the influence of dispersal on diversification in the avifauna of Australasian archipelagoes. Contrary to expectations given the fragmented nature of island systems, the relationship between dispersal ability and diversification rate was monotonically negative. While multiple mechanisms could generate this pattern, they all share a phase of range expansion that is decoupled from speciation.     

Mutualism with plants drives primate diversification

Understanding the origin of diversity is a fundamental problem in biology. Evolutionary diversification has been intensely explored during the last years due to the development of molecular tools and the comparative method. However, most studies are conducted using only information from extant species. This approach probably leads to misleading conclusions, especially because of inaccuracy in the estimation of extinction rates. It is critical to integrate the information generated by extant organisms with the information obtained from the fossil record. Unfortunately, this integrative approach has been seldom performed, and thus, our understanding of the factors fueling diversification is still deficient. Ecological interactions are a main factor shaping evolutionary diversification by influencing speciation and extinction rates. Most attention has focused on the effect of antagonistic interactions on evolutionary diversification. In contrast, the role of mutualistic interactions in shaping diversification has been much less explored. In this study, by combining phylogenetic, neontological, and paleontological information, we show that a facultative mutualistic plant-animal interaction emerging from frugivory and seed dispersal has most likely contributed to the diversification of our own lineage, the primates. We compiled diet and seed dispersal ability in 381 extant and 556 extinct primates. Using well-established molecular phylogenies, we demonstrated that mutualistic extant primates had higher speciation rates, lower extinction rates, and thereby higher diversification rates than nonmutualistic ones. Similarly, mutualistic fossil primates had higher geological durations and smaller per capita rates of extinction than nonmutualistic ones. As a mechanism underlying this pattern, we found that mutualistic extinct and extant primates have significantly larger geographic ranges, which promotes diversification by hampering extinction and increasing geographic speciation. All these outcomes together strongly suggest that the establishment of a facultative mutualism with plants has greatly benefited primate evolution and fueled its taxonomic diversification.