Clade age and species richness are decoupled across the eukaryotic tree of life

Explaining the dramatic variation in species richness across the tree of life remains a key challenge in evolutionary biology. At the largest phylogenetic scales, the extreme heterogeneity in species richness observed among different groups of organisms is almost certainly a function of many complex and interdependent factors. However, the most fundamental expectation in macroevolutionary studies is simply that species richness in extant clades should be correlated with clade age: all things being equal, older clades will have had more time for diversity to accumulate than younger clades. Here, we test the relationship between stem clade age and species richness across 1,397 major clades of multicellular eukaryotes that collectively account for more than 1.2 million described species. We find no evidence that clade age predicts species richness at this scale. We demonstrate that this decoupling of age and richness is unlikely to result from variation in net diversification rates among clades. At the largest phylogenetic scales, contemporary patterns of species richness are inconsistent with unbounded diversity increase through time. These results imply that a fundamentally different interpretative paradigm may be needed in the study of phylogenetic diversity patterns in many groups of organisms.     

Rapid diversification and not clade age explains high diversity in neotropical Adelpha butterflies

Latitudinal gradients in species richness are among the most well-known biogeographic patterns in nature, and yet there remains much debate and little consensus over the ecological and evolutionary causes of these gradients. Here, we evaluated whether two prominent alternative hypotheses (namely differences in diversification rate or clade age) could account for the latitudinal diversity gradient in one of the most speciose neotropical butterfly genera (Adelpha) and its close relatives. We generated a multilocus phylogeny of a diverse group of butterflies in the containing tribe Limenitidini, which has both temperate and tropical representatives. Our results suggest there is no relationship between clade age and species richness that could account for the diversity gradient, but that instead it could be explained by a significantly higher diversification rate within the predominantly tropical genus Adelpha. An apparent early larval host-plant shift to Rubiaceae and other plant families suggests that the availability of new potential host plants probably contributed to an increase in diversification of Adelpha in the lowland Neotropics. Collectively, our results support the hypothesis that the equatorial peak in species richness observed within Adelpha is the result of increased diversification rate in the last 10-15 Myr rather than a function of clade age, perhaps reflecting adaptive divergence in response to the dramatic host-plant diversity found within neotropical ecosystems.     

Ecological limits and diversification rate: alternative paradigms to explain the variation in species richness among clades and regions

Diversification rate is one of the most important metrics in macroecological and macroevolutionary studies. Here I demonstrate that diversification analyses can be misleading when researchers assume that diversity increases unbounded through time, as is typical in molecular phylogenetic studies. If clade diversity is regulated by ecological factors, then species richness may be independent of clade age and it may not be possible to infer the rate at which diversity arose. This has substantial consequences for the interpretation of many studies that have contrasted rates of diversification among clades and regions. Often, it is possible to estimate the total diversification experienced by a clade but not diversification rate itself. I show that the evidence for ecological limits on diversity in higher taxa is widespread. Finally, I explore the implications of ecological limits for a variety of ecological and evolutionary questions that involve inferences about speciation and extinction rates from phylogenetic data.     

Cryptic animal species are homogeneously distributed among taxa and biogeographical regions

Background  

Cryptic species are two or more distinct but morphologically similar species that were classified as a single species. During the past two decades we observed an exponential growth of publications on cryptic species. Recently published reviews have demonstrated cryptic species have profound consequences on many biological disciplines. It has been proposed that their distribution is non-random across taxa and biomes.     

净多样化速率和进化时间对虎耳草目科间物种多样性差异的影响

不同生物类群包含的物种数目常存在巨大差异,这是生态学和生物学研究中普遍观察到的现象。然而,这一现象产生的原因仍然是未解之谜。从宏观进化的角度,进化时间假说和多样化速率假说是两个比较流行的假说。进化时间假说认为类群的演化时间越长,积累的物种丰富度越高;而多样化速率假说认为类群的净多样化速率越快,则其物种丰富度越高。为验证这两个假说,该文以一棵包含1 539个物种化石定年的虎耳草目系统发育树为基础,通过宏观进化分析获取了虎耳草目内15个科的物种形成和灭绝速率,并计算了每个科的平均多样化速率。结果表明：(1)虎耳草目的物种多样化速率有着增加的趋势,并且多样化速率的增加主要出现在温带和高山类群,如茶藨子科、景天科和芍药科等。(2)采用系统发育广义最小二乘模型(PGLS)和线性回归模型(LM)结果表明,虎耳草目15个科的物种丰富度与科的分化时间和科内物种的最近共同祖先年龄都没有显著相关关系,而与净多样化速率显著正相关(R² =0.380,P<0.05)。该研究支持了多样化速率假说,认为不同科的净多样化速率的差异是导致虎耳草目科间物种数目差异的主要原因之一。全球气候变冷...     

Concordance of species richness patterns among multiple freshwater taxa: a regional perspective

Geographical gradients in species richness and the degree to which different taxa show congruent patterns remain unknown for many taxonomic groups. Here, I examined broad-scale species richness patterns in five groups of freshwater organisms; macrophytes, dragonflies, stoneflies, aquatic beetles and fishes. The analyses were based on provincial distribution records in Denmark, Norway, Sweden and Finland. In general, variation in species richness across provinces was concordant among the groups, but stoneflies showed weaker negative relationships with the other taxonomic groups. Species richness in most groups decreased with increasing latitude and altitude, and a considerable part of the variation was explained by mean July temperature. However, stoneflies showed a reversed pattern, with species richness correlating positively, albeit more weakly, with mean provincial altitude. Nevertheless, combined species richness of all five taxa showed a strong relationship with mean July temperature, accounting for 74% of variation in provincial species richness alone. Such temperature-controlled patterns suggest that regional freshwater biodiversity will strongly respond to climate change, with repercussions for local community organization in freshwater ecosystems in Fennoscandia.     

Primary controls on species richness in higher taxa

The disparity in species richness across the tree of life is one of the most striking and pervasive features of biological diversity. Some groups are exceptionally diverse, whereas many other groups are species poor. Differences in diversity among groups are frequently assumed to result from primary control by differential rates of net diversification. However, a major alternative explanation is that ecological and other factors exert primary control on clade diversity, such that apparent variation in net diversification rates is a secondary consequence of ecological limits on clade growth. Here, I consider a likelihood framework for distinguishing between these competing hypotheses. I incorporate hierarchical modeling to explicitly relax assumptions about the constancy of diversification rates across clades, and I propose several statistics for a posteriori evaluation of model adequacy. I apply the framework to a recent dated phylogeny of ants. My results reject the hypothesis that net diversification rates exert primary control on species richness in this group and demonstrate that clade diversity is better explained by total time-integrated speciation. These results further suggest that it may not possible to estimate meaningful speciation and extinction rates from higher-level phylogenies of extant taxa only.     

Global patterns of diversification and species richness in amphibians

Geographic patterns of species richness ultimately arise through the processes of speciation, extinction, and dispersal, but relatively few studies consider evolutionary and biogeographic processes in explaining these diversity patterns. One explanation for high tropical species richness is that many species-rich clades originated in tropical regions and spread to temperate regions infrequently and more recently, leaving little time for species richness to accumulate there (assuming similar rates of diversification in temperate and tropical regions). However, the major clades of anurans (frogs) and salamanders may offer a compelling counterexample. Most salamander families are predominately temperate in distribution, but the one primarily tropical clade (Bolitoglossinae) contains nearly half of all salamander species. Similarly, most basal clades of anurans are predominately temperate, but one largely tropical clade (Neobatrachia) contains approximately 96% of anurans. In this article, I examine patterns of diversification in frogs and salamanders and their relationship to large-scale patterns of species richness in amphibians. I find that diversification rates in both frogs and salamanders increase significantly with decreasing latitude. These results may shed light on both the evolutionary causes of the latitudinal diversity gradient and the dramatic but poorly explained disparities in the diversity of living amphibian clades.     

Body size does not predict species richness among the metazoan phyla

We present a comparative study of the relationship between body size and described taxonomic diversity in the Metazoa. We find no pattern between body size and taxonomic diversity; neither the smallest organisms nor organisms at an intermediate body size are consistently more diverse than their closest relatives. This conclusion holds for both nonphylogenetic analysis, in which phyla are treated as independent points, and analysis of independent contrasts using several recent hypotheses of metazoan phylogeny. These results appear surprising in the context of existing models of body size distributions. However, such models are built around the prevalence of right‐skewed distributions and we find no evidence for such a distribution.     

Ecosystem process rate increases with animal species richness: evidence from leaf-eating, aquatic insects

Effects of species number and identity on the breakdown rate of leaf litter were estimated in a laboratory experiment using leaf-eating insects, three species of Plecoptera, as detritivores. We found significant differences between the different species on this process in single-species experiment, but not when animal biomass was accounted for. When species were combined the effect of species identity was strongly reduced and rendered insignificant, whereas the number of species had a significant effect. This shows that rates of ecosystem processes may benefit from species richness even when all species belong to the same guild, which is in contrast to hypotheses predicting redundancy within guilds. Facilitation between species and negative interactions, where intraspecific interactions are greater than interspecific interactions, are two potential mechanisms which could explain increasing decomposition rates with species richness.     

Independence of total species richness and richness difference are not the same thing

Leprieur and Oikonomou (2014) criticized that a replacement index β_− 3, a partitioned component of Jaccard index β_jac, was not richness independent and should not be used in biogeographic and ecological studies. However, Leprieur and Oikonomou failed to recognize the difference between richness and richness difference. Independence of total species richness is not equal to independence of richness difference. Theoretically and ideally, it is true that β_− 3 (and β_jac as well) is not independent of richness difference while Simpson index (β_sim) is fully independent of richness difference. However, all these indices actually are independent of total species richness. At last, it is worth mentioning that the ideal independence studied here is easily violated in computational simulation and real-world settings.     

Biogeography of species richness gradients: linking adaptive traits,demography and diversification

Here we review how adaptive traits contribute to the emergence and maintenance of species richness gradients through their influence on demographic and diversification processes. We start by reviewing how demographic dynamics change along species richness gradients. Empirical studies show that geographical clines in population parameters and measures of demographic variability are frequent along latitudinal and altitudinal gradients. Demographic variability often increases at the extremes of regional species richness gradients and contributes to shape these gradients. Available studies suggest that adaptive traits significantly influence demographic dynamics, and set the limits of species distributions. Traits related to thermal tolerance, resource use, phenology and dispersal seem to play a significant role. For many traits affecting demography and/or diversification processes, complex mechanistic approaches linking genotype, phenotype and fitness are becoming progressively available. In several taxa, species can be distributed along adaptive trait continuums, i.e. a main axis accounting for the bulk of inter‐specific variation in some correlated adaptive traits. It is shown that adaptive trait continuums can provide useful mechanistic frameworks to explain demographic dynamics and diversification in species richness gradients. Finally, we review the existence of sequences of adaptive traits in phylogenies, the interactions of adaptive traits and community context, the clinal variation of traits across geographical gradients, and the role of adaptive traits in determining the history of dispersal and diversification of clades. Overall, we show that the study of demographic and evolutionary mechanisms that shape species richness gradients clearly requires the explicit consideration of adaptive traits. To conclude, future research lines and trends in the field are briefly outlined.     

Functional group dominance and not productivity drives species richness

Background: There is a lack of consensus about the productivity–richness relationship, with several recent studies suggesting that it is not productivity but other factors that are the important drivers that determine species richness.

Aims: Here, we examine the relationship between productivity, functional group dominance and plant species richness at the plot scale in Tibetan Plateau meadows. These alpine meadows are ideal to examine the species productivity-richness relationship because they have a very high species richness, a large gradient in productivity, and can be dominated by either graminoids (grasses and sedges) or forbs.

Methods: We measured plant species richness and above-ground biomass along a natural gradient of functional group abundance in 44 plots distributed across five natural, winter-grazed but otherwise undisturbed sites in the eastern part of the Qing-Hai Tibetan Plateau, in Gansu province, China in 2008.

Results: Graminoid abundance (i.e. graminoid biomass as percent of the total above-ground biomass) explained 39% of plot differences in species richness while neither productivity nor the biomass of the three most abundant plant species, either individually or combined, were a significant predictor of species richness.

Conclusions: Our results show that within these alpine meadows, a shift from graminoid to forb dominance, rather than the individual dominant species or productivity itself, is strongly correlated with species richness. Thus, differences in functional group abundance can be a strong driver of observed plant species richness patterns.     

Distribution of alien animal species richness in the Czech Republic

Biogeographical barriers formed by natural forces over billions of years have been substantially disrupted by human activity, particularly in recent centuries. In response to these anthropogenic changes, global homogenization of biota is observed at an ever‐increasing rate, causing environmental and economic losses as well as emerging health risks. Identifying factors underlying alien species richness is essential for prevention of future introductions and subsequent spread. In this study, we examined the effects of environmental and human‐related factors on distribution of alien animal species richness in the Czech Republic (Central Europe). We compiled a set of maps showing the level of invasion of six categories of alien animal species in each of 628 grid cells (ca. 12.0 × 11.1 km) covering the Czech Republic. Relationships between alien species richness and 12 variables characterizing climatic conditions, topography, land cover, and human population size were calculated using the generalized least squares method. Species richness of all alien species, of invertebrates, and of terrestrial species showed the strongest positive relationship with mean annual temperature, while the number of black and grey (proposed prominent invaders) and aquatic species was most closely related to the presence of large rivers. Alien vertebrates showed a strong negative relationship with annual precipitation. The highest alien animal species richness was found in and near large population centers and in agricultural landscapes in warm and dry lowlands. The gateways for alien aquatic species are rather large rivers over sport fishing and aquaculture import. Compiled maps create a powerful visual communication tool, useful in development of programs to prevent future introductions.     

代谢速率调控物种丰富度格局的研究进展

提出生物多样性分布格局的普适性理论和探索其内在形成机制一直是生态学家们研究的焦点之一.到目前为止,已有很多假说被用来解释生物多样性分布规律,但是这些假说的普适性均受到学者们的质疑.最新理论--代谢速率假说以能量相当法则和代谢分形分配网络模型为基础,定量预测了个体及种群生态进化动态过程与群落生物多样性分布格局之间的关系,以及物种丰富度和环境因子之间的关系.代谢速率假说解释了生物多样性的起源问题,也回答了生物多样性如何维持的问题.该文重点综述了代谢生物多样性理论的发展及其相关研究进展.通过和其他假说比较、分析,我们认为随着代谢理论假说的不断发展和完善,代谢生物多样性理论将更具有普适性.同时我们也提出了进一步完善该假说需要解决的一些科学问题.