The return of the Red Queen, or the law of the growth in the mean duration of the existence of genera during evolution

Phanerozoic marine genera apparently do not become less extinction-prone with age. Higher extinction probability in "young" cohorts of genera is better explained by initially different levels of extinction-tolerance of genera in the cohort. This fact agrees with one of the two basic statements of the "Red Queen" hypothesis (Van Valen, 1973). In the second statement (the idea that the increase in fitness lowers extinction probability) the term "fitness" should be changed to "adaptability". The increase of extinction-tolerance, that can be interpreted as the increase of adaptability to unpredictable changes of environment, is found in succession of "generations" of genera that replace one another through time. This increase reveals itself, firstly, in the growth of mean duration of genera, as well as in the decrease of extinction/origination rates, gradual accumulation of long-lived genera and origination of genera with higher duration. The increase of adaptability may be caused by selective extinction of stenotopic, ecologically specialized forms; Cope's law; evolution of ecosystems that involves development of more effective mechanisms of sustaining homeostasis which may stimulate the recovery of a genus after partial extinction.     

Diversification rates and the latitudinal gradient of diversity in mammals

The latitudinal gradient of species richness has frequently been attributed to higher diversification rates of tropical groups. In order to test this hypothesis for mammals, we used a set of 232 genera taken from a mammalian supertree and, additionally, we reconstructed dated Bayesian phylogenetic trees of 100 genera. For each genus, diversification rate was estimated taking incomplete species sampling into account and latitude was assigned considering the heterogeneity in species distribution ranges. For both datasets, we found that the average diversification rate was similar among all latitudinal bands. Furthermore, when we used phylogenetically independent contrasts, we did not find any significant correlation between latitude and diversification parameters, including different estimates of speciation and extinction rates. Thus, other factors, such as the dynamics of dispersal through time, may be required to explain the latitudinal gradient of diversity in mammals.     

Nonstochastic variation of species-level diversification rates within angiosperms

Variations in the origination and extinction rates of species over geological time often are linked with a range of factors, including the evolution of key innovations, changes in ecosystem structure, and environmental factors such as shifts in climate and physical geography. Before hypothesizing causality of a single factor, it is critical to demonstrate that the observed variation in diversification is significantly greater than one would expect due to natural stochasticity in the evolutionary branching process. Here, we use a likelihood-ratio test to compare taxonomic rate heterogeneity to a neutral birth-death model, using data on well-supported sister pairs of taxa and their species richness. We test the likelihood that the distribution of extant species among angiosperm genera and families could be the result of constant diversification rates. Results strongly support the conclusion that there is significantly more heterogeneity in diversity at the species level within angiosperms than would be expected due to stochastic processes. This result is consistent in datasets of genus pairs and family pairs and is not affected significantly by degrading pairs to simulate inaccuracy in the assumption of simultaneous origin of sister taxa. When we parse taxon pairs among higher groups of angiosperms, results indicate that a constant rates model is not rejected by rosid and basal eudicot pairs but is rejected by asterid and eumagnoliid pairs. These results provide strong support for the hypothesis that species-level rates of origination and/or extinction have varied nonrandomly within angiosperms and that the magnitude of heterogeneity varies among major groups within angiosperms.     

Testing reduced evolutionary rates during the Late Palaeozoic Ice Age using the crinoid fossil record

In 2003, Stanley & Powell reported depressed rates of origination and extinction in marine invertebrates during the Late Palaeozoic Ice Age (LPIA). Using a database of crinoid genera, rates of origination, extinction and genus duration were calculated at the stage level from the Early Devonian to the Late Permian. This 165 m.y. time span includes non‐glacial intervals before and after the LPIA, which spanned the Serpukhovian to Sakmarian, providing background rates for comparison. Data generated on crinoid evolutionary rates during the Middle to Late Palaeozoic were analysed and compared to Stanley & Powell's data to determine whether crinoid evolutionary patterns support their findings or suggest an alternative hypothesis. Rates of origination and extinction in all crinoid clades were reduced during the LPIA compared to the combined background intervals before and after the LPIA. However, crinoid diversity was higher during the LPIA than the surrounding time intervals. The difference in diversity trends between crinoids and other marine invertebrates is due to the advanced cladids clade. Unstable, fluctuating environmental conditions during the LPIA may have created habitats suitable for opportunistic crinoid genera that reduced both the probability of origination and extinction. The increased diversity of the advanced cladids is likely due to their unique adaptation of muscular arm articulations, which allowed them to thrive in marine settings with increased siliciclastic influx brought on by the Alleghenian orogeny. Despite the advanced cladids’ departure from the expected diversity count, the results of analyses performed on the updated crinoid database provide independent confirmation of Stanley & Powell's original hypothesis of depressed evolutionary rates in marine invertebrates during the LPIA.     

Macroevolution of hoverflies (Diptera: Syrphidae): the effect of using higher‐level taxa in studies of biodiversity,and correlates of species richness

We test a near‐complete genus level phylogeny of hoverflies (Diptera: Syrphidae) for consistency with a null model of clade growth having uniform probabilities of speciation and extinction among contemporaneous species. The phylogeny is too unbalanced for this null model. Importantly, the degree of imbalance in the phylogeny depends on whether the phylogeny is analysed at the genus level or species level, suggesting that genera ought not to be used uncritically as surrogates for species in large‐scale evolutionary analyses. Tests for a range of morphological, life‐history and ecological correlates of diversity give equivocal results, but suggest that high species‐richness may be associated with sexual selection and diet breadth. We find no correlation between species‐richness and either body size or reproductive rate.     

The impact of gene-tree/species-tree discordance on diversification-rate estimation

Molecular phylogenies are often used to test hypotheses about the tempo and mode of speciation and extinction. One commonly used statistic is Pybus and Harvey's γ, which measures the density of ordered internode distances on an ultrametric tree to infer earlier (negative γ) or later (positive γ) bursts of diversification. However, coalescent theory predicts that γ might be biased toward negative values (inferring early bursts of diversification) when using gene trees rather than species trees. Gene divergences predate species divergences, increasingly so at higher effective population sizes (N(e)), and proportionally more so toward the tips of the tree. Thus, gene trees will have a higher density of older nodes in many cases (particularly at higher N(e)), due to the disproportionate lengthening of terminal branches. This will yield an artifactual signature of early bursts of diversification when estimating γ from gene trees. We simulate gene trees within species trees under both Yule (pure-birth) and birth-death processes, and demonstrate support for these predictions. However, for most realistic estimates of θ in natural populations, gene trees provide relatively good estimates of γ, despite the disproportionate overestimation of younger node ages. This is corroborated with an empirical dataset of North American fence lizards (Sceloporus).     

Global diversity of mayflies (Ephemeroptera, Insecta) in freshwater

The extant global Ephemeroptera fauna is represented by over 3,000 described species in 42 families and more than 400 genera. The highest generic diversity occurs in the Neotropics, with a correspondingly high species diversity, while the Palaearctic has the lowest generic diversity, but a high species diversity. Such distribution patterns may relate to how long evolutionary processes have been carrying on in isolation in a bioregion. Over an extended period, there may be extinction of species, but evolution of more genera. Dramatic extinction events such as the K-T mass extinction have affected current mayfly diversity and distribution. Climatic history plays an important role in the rate of speciation in an area, with regions which have been climatically stable over long periods having fewer species per genus, when compared to regions subjected to climatic stresses, such as glaciation. A total of 13 families are endemic to specific bioregions, with eight among them being monospecific. Most of these have restricted distributions which may be the result of them being the relict of a previously more diverse, but presently almost completely extinct family, or may be the consequence of vicariance events, resulting from evolution due to long-term isolation. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

CALIBRATED DIVERSITY,TREE TOPOLOGY AND THE MOTHER OF MASS EXTINCTIONS: THE LESSON OF TEMNOSPONDYLS

Abstract: Three family‐level cladistic analyses of temnospondyl amphibians are used to evaluate the impact of taxonomic rank, tree topology, and sample size on diversity profiles, origination and extinction rates, and faunal turnover. Temnospondyls are used as a case study for investigating replacement of families across the Permo‐Triassic boundary and modality of recovery in the aftermath of the end‐Permian mass extinction. Both observed and inferred (i.e. tree topology‐dependent) values of family diversity have a negligible effect on the shape of the diversity curve. However, inferred values produce both a flattening of the curve throughout the Cisuralian and a less pronounced increase in family diversity from Tatarian through to Induan than do observed values. Diversity curves based upon counts of genera and species display a clearer distinction between peaks and troughs. We use rarefaction techniques (specifically, rarefaction of the number of genera and species within families) to evaluate the effect of sampling size on the curve of estimated family‐level diversity during five time bins (Carboniferous; Cisuralian; Guadalupian–Lopingian; Early Triassic; Middle Triassic–Cretaceous). After applying rarefaction, we note that Cisuralian and Early Triassic diversity values are closer to one another than they are when the observed number of families is used; both values are also slightly higher than the Carboniferous estimated diversity. The Guadalupian–Lopingian value is lower than raw data indicate, reflecting in part the depauperate land vertebrate diversity from the late Cisuralian to the middle Guadalupian (Olson’s gap). The time‐calibrated origination and extinction rate trajectories plot out close to one another and show a peak in the Induan, regardless of the tree used to construct them. Origination and extinction trajectories are disjunct in at least some Palaeozoic intervals, and background extinctions exert a significant role in shaping temnospondyl diversity in the lowermost Triassic. Finally, species‐, genus‐, and family trajectories consistently reveal a rapid increase in temnospondyl diversity from latest Permian to earliest Triassic as well as a decline near the end of the Cisuralian. However, during the rest of the Cisuralian family diversity increases slightly and there is no evidence for a steady decline, contrary to previous reports.     

Ecological interactions on macroevolutionary time scales: clams and brachiopods are more than ships that pass in the night

Competition among organisms has ecological and evolutionary consequences. However, whether the consequences of competition are manifested and measureable on macroevolutionary time scales is equivocal. Marine bivalves and brachiopods have overlapping niches such that competition for food and space may occur. Moreover, there is a long‐standing debate over whether bivalves outcompeted brachiopods evolutionarily, because brachiopod diversity declined through time while bivalve diversity increased. To answer this question, we estimate the origination and extinction dynamics of fossil marine bivalve and brachiopod genera from the Ordovician through to the Recent while simultaneously accounting for incomplete sampling. Then, using stochastic differential equations, we assess statistical relationships among diversification and sampling dynamics of brachiopods and bivalves and five paleoenvironmental proxies. None of these potential environmental drivers had any detectable influence on brachiopod or bivalve diversification. In contrast, elevated bivalve extinction rates causally increased brachiopod origination rates, suggesting that bivalves have suppressed brachiopod evolution.     

Geographical distribution and extinction risk: lessons from Triassic–Jurassic marine benthic organisms

Aim To evaluate the influence of geographical distribution on the extinction risk of benthic marine invertebrates using data from the fossil record, both during times of background extinction and across a mass‐extinction episode. Total geographical range is contrasted with proxies of global abundance to assess the relationships between the two essential components of geographical distribution and extinction risk. Location A global occurrence data base of fossil benthic macro‐organisms from the Triassic and Jurassic periods was used for this study. Methods Geographical distributions and biodiversity dynamics were assessed for each genus (all taxa) or species (bivalves) based on a sample‐standardized data set and palaeogeographical reconstructions. Geographical ranges were measured by the maximum great circle distance of a taxon within a stratigraphic interval. Global abundance was assessed by the number of localities at which a taxon was recorded. Widespread and rare taxa were separated using median and percentile values of the frequency distributions of occurrences. Results The frequency distribution of geographical ranges is very similar to that for modern taxa. Although no significant correlation could be established between local abundance and geographical range, proxies of global abundance are strongly correlated with geographical range. Taxon longevities are correlated with both mean geographical range and mean global abundance, but range size appears to be more critical than abundance in determining extinction risk. These results are valid when geographical distribution is treated as a trait of taxa and when assessed for individual geological stages. Main conclusions Geographical distribution is a key predictor of extinction risk of Triassic and Jurassic benthic marine invertebrates. An important exception is in the end‐Triassic mass extinction, which equally affected geographically restricted and widespread genera, as well as common and rare genera. This suggests that global diversity crises may curtail the role of geographical distribution in determining extinction risk.     

Diversity,extinction, and threat status in Lagomorphs

A quarter of all lagomorphs (pikas, rabbits, hares and jackrabbits) are threatened with extinction, including several genera that contain only one species. The number of species in a genus correlates with extinction risk in lagomorphs, but not in other mammal groups, and this is concerning because the non‐random extinction of small clades disproportionately threatens genetic diversity and phylogenetic history. Here, we use phylogenetic analyses to explore the properties of the lagomorph phylogeny and test if variation in evolution, biogeography and ecology between taxa explains current patterns of diversity and extinction risk. Threat status was not related to body size (and, by inference, its biological correlates), and there was no phylogenetic signal in extinction risk. We show that the lagomorph phylogeny has a similar clade‐size distribution to other mammals, and found that genus size was unrelated to present climate, topography, or geographic range size. Extinction risk was greater in areas of higher human population density and negatively correlated with anthropogenically modified habitat. Consistent with this, habitat generalists were less likely to be threatened. Our models did not predict threat status accurately for taxa that experience region‐specific threats. We suggest that pressure from human populations is so severe and widespread that it overrides ecological, biological, and geographic variation in extant lagomorphs.     

Genus age,provincial area and the taxonomic structure of marine faunas

Species are unevenly distributed among genera within clades and regions, with most genera species-poor and few species-rich. At regional scales, this structure to taxonomic diversity is generated via speciation, extinction and geographical range dynamics. Here, we use a global database of extant marine bivalves to characterize the taxonomic structure of climate zones and provinces. Our analyses reveal a general, Zipf–Mandelbrot form to the distribution of species among genera, with faunas from similar climate zones exhibiting similar taxonomic structure. Provinces that contain older taxa and/or encompass larger areas are expected to be more species-rich. Although both median genus age and provincial area correlate with measures of taxonomic structure, these relationships are interdependent, nonlinear and driven primarily by contrasts between tropical and extra-tropical faunas. Provincial area and taxonomic structure are largely decoupled within climate zones. Counter to the expectation that genus age and species richness should positively covary, diverse and highly structured provincial faunas are dominated by young genera. The marked differences between tropical and temperate faunas suggest strong spatial variation in evolutionary rates and invasion frequencies. Such variation contradicts biogeographic models that scale taxonomic diversity to geographical area.     

Species-genus ratios reflect a global history of diversification and range expansion in marine bivalves

The distribution of marine bivalve species among genera and higher taxa takes the form of the classic hollow curve, wherein few lineages are species rich and many are species poor. The distribution of species among genera (S/G ratio) varies with latitude, with temperate S/G's falling within the null expectation, and tropical and polar S/G's exceeding it. Here, we test several hypotheses for this polar overdominance in the species richness of small numbers of genera. We find a significant positive correlation between the latitudinal range of a genus and its species richness, both globally and within regions. Genus age and species richness are also positively related, but this relationship breaks down when the analysis is limited to genera endemic to climate zones or with narrow latitudinal ranges. The data suggest a link between speciation and range-expansion, with genera expanding out of the tropical latitudinal bins tending to speciate more prolifically, both globally and regionally. These genera contain more species within climate zones than taxa endemic to that zone. Range expansion thus appears to be fundamentally coupled with speciation, producing the skewed distribution of species among genera, both globally and regionally, whereas clade longevity is achieved through extinction -- resistance conferred by broad geographical ranges.     

楝科（Meliaceae）的地理分布

楝科为泛热带分布科,全世界有５１属,约５５０—６００种,分布于旧世界热带地区有４６属,热带美洲有８属．热带亚洲和热带非洲为楝科两大现代分布中心．中国楝科共１５属,６１种,占世界属总数的２９％,种总数的１０％。中国楝科的分布是在全球楝科分布区的边缘,主要分布于中国西南部及南部诸省,种类由西南向东南递减。中国楝科属的分布区类型可归为５类：１．热带亚洲、非洲和中南美洲间断分布（１属）;２．旧世界热带分布（３属）;３,热带亚洲至热带大洋洲分布（２属）;４．热带亚洲至热带非洲分布（１属）;５．热带亚洲分布（８属）。中国楝科种的分布区类型仅有２类：１．热带亚洲分布（３１种）;２．中国特有分布（３０种）。楝科植物的起源推断在早白垩纪。中国楝科植物由印度—马来西亚成分及特有成分组成。热带亚洲的楝科植物主要是通过中南半岛和中国云南。广西和海南等地发生联系,而菲律宾和台湾之间可有直接的联系。     

A new approach to modeling the diversity dynamics of Phanerozoic marine biota

Modeling of fossil diversity dynamics is usually done with the help of the models borrowed from the population dynamics theory. However there are principal differences between organisms and taxa, reproduction and divergence, mortality and extinction that make this approach doubtful. Another model is presented here, in which absolute origination rate does not depend on diversity, the ability of new genera to sustain unpredictable environmental changes increases three times abruptly at Cambrian/Ordovician, Permian/Triassic and Cretaceous/Tertiary boundaries. In this model the diversity increases due to accumulation of long-lived genera. The computer simulation showed that the model agrees with empirical data by 15 major criteria. The laws of community evolution apparently can explain the general pattern of punctuated equilibrium in the evolution of marine biota.     

Marine phytoplankton: how many species in the world ocean?

Towards the end of the 1980s, living plankton flora of the worldocean amounted to 474-504 genera and 3444–4375 speciesif one neglects the increase rate of taxa during the latestyears In the above figures, the lower estimate is that of the‘reliable’ taxa (of practical value for identificationtasks), whereas the higher estimate includes the insufficientlyknown or doubtful organisms, and synonyms are excluded. Thefrequency distribution of the numbers of species per genus confirmsthe general hyperbolic law, which implies that a relativelylarge number of genera are uni- or paucispecific.     

The role of frugivory in the diversification of bats in the Neotropics

Aim Ecological interactions are among the most important biotic factors influencing the processes of speciation and extinction. Our aim was to test whether diversification rates of New World Noctilionoidea bats are associated with specialization for frugivory, and how this pattern differs between the mainland and the West Indies. Location The New World. Methods We reconstructed a time‐calibrated molecular phylogenetic hypothesis for the New World genera of the superfamily Noctilionoidea. We compiled data on diet, morphology, geographical distribution and number of ecoregions in which each genus occurs. Then, using the phylogenetic tree constructed, we tested whether diversification was driven by diet (animalivorous and sanguinivorous versus nectarivorous and frugivorous) and specialization for frugivory. Afterwards, we conducted phylogenetic comparative analyses to identify correlates of species richness and net diversification rates. Results The diversification rate was higher in mutualistic than in antagonistic clades in mainland and Antillean biogeographical scenarios, but only strictly frugivorous clades showed a markedly higher diversification rate than the rest of the genera. Geographical range and number of ecoregions were positively associated with species richness and diversification rate in continental and insular lineages. Lower body mass, lower forearm length and specialization for frugivory were significantly positively correlated with higher diversification rates in continental lineages, whereas these parameters were negatively correlated in Antillean lineages. Main conclusions The direction of the relationship of intrinsic factors (specialization for frugivory and body size) with diversification of noctilionoid bats depends on the biogeographical context, whereas the direction of the relationship of extrinsic factors (geographical range and number of ecoregions) with diversification is consistent in both mainland and the West Indian lineages.     

中国杜鹃花科的(纟戾)木亚科白珠树亚科北极果亚科地理分布及其与其他地区的关系

<正> 杜鹃花科(Ericaceae)为中国植物的第七大科。因此,它在中国植物地理是中国植物区系上都占有比较重要的地位。杜鹃花科共分五个亚科:杜鹃花亚科、綟木亚科,白珠树亚科、北极果亚科和乌饭树亚科。綟木亚科、白珠树亚科和北极果亚科,虽然在本科中其种类不算太多,但所含10个属,故其多样性在本科中是首屈一指的。无疑,对这三个亚科在中国的地理分布及其与其他地区的关系的讨论是有意义的。     

Amniotes through major biological crises: faunal turnover among Parareptiles and the end‐Permian mass extinction

Abstract: The Parareptilia are a small but ecologically and morphologically diverse clade of Permian and Triassic crown amniotes generally considered to be phylogenetically more proximal to eureptiles (diapsids and their kin) than to synapsids (mammals and their kin). A recent supertree provides impetus for an analysis of parareptile diversity through time and for examining the influence of the end‐Permian mass extinction on the clade’s origination and extinction rates. Phylogeny‐corrected measures of diversity have a significant impact on both rates and the distribution of origination and extinction intensities. Time calibration generally results in a closer correspondence between origination and extinction rate values than in the case of no time correction. Near the end‐Permian event, extinction levels are not significantly higher than origination levels, particularly when time calibration is introduced. Finally, regardless of time calibration and/or phylogenetic correction, the distribution of rates does not differ significantly from unimodal. The curves of rate values are discussed in the light of the numbers and distributions of both range extensions and ghost lineages. The disjoint time distributions of major parareptile clades (e.g. procolophonoids and nycteroleterids‐pareiasaurs) are mostly responsible for the occurrence of long‐range extensions throughout the Permian. Available data are not consistent with a model of sudden decline at the end‐Permian but rather suggest a rapid alternation of originations and extinctions in a number of parareptile groups, both before and after the Permian/Triassic boundary.     

Hyperbolic growth of marine and continental biodiversity through the phanerozoic and community evolution

Among diverse models that are used to describe and interpret the changes in global biodiversity through the Phanerozoic, the exponential and logistic models (traditionally used in population biology) are the most popular. As we have recently demonstrated (Markov, Korotayev, 2007), the growth of the Phanerozoic marine biodiversity at genus level correlates better with the hyperbolic model (widely used in demography and macrosociology). Here we show that the hyperbolic model is also applicable to the Phanerozoic continental biota at genus and family levels, and to the marine biota at species, genus, and family levels. There are many common features in the evolutionary dynamics of the marine and continental biotas that imply similarity and common nature of the factors and mechanisms underlying the hyperbolic growth. Both marine and continental biotas are characterized by continuous growth of the mean longevity of taxa, by decreasing extinction and origination rates, by similar pattern of replacement of dominant groups, by stepwise accumulation of evolutionary stable, adaptable and "physiologically buffered" taxa with effective mechanisms of parental care, protection of early developmental stages, etc. At the beginning of the development of continental biota, the observed taxonomic diversity was substantially lower than that predicted by the hyperbolic model. We suggest that this is due, firstly, to the fact that, during the earliest stages of the continental biota evolution, the groups that are not preserved in the fossil record (such as soil bacteria, unicellular algae, lichens, etc.) played a fundamental role, and secondly, to the fact that the continental biota initially formed as a marginal portion of the marine biota, rather than a separate system. The hyperbolic dynamics is most prominent when both marine and continental biotas are considered together. This fact can be interpreted as a proof of the integrated nature of the biosphere. In the macrosociological models, the hyperbolic pattern of the world population growth arises from a non-linear second-order positive feedback between the demographic growth and technological development (more people - more potential inventors - faster technological growth - the carrying capacity of the Earth grows faster - faster population growth - more people - more potential inventors, and so on). Based on the analogy with macrosociological models and diverse paleontological data, we suggest that the hyperbolic character of biodiversity growth can be similarly accounted for by a non-linear second-order positive feedback between the diversity growth and community structure complexity. The feedback can work via two parallel mechanisms: 1) decreasing extinction rate (more taxa- higher alpha diversity, or mean number of taxa in a community - communities become more complex and stable - extinction rate decreases - more taxa, and so on) and 2) increasing origination rate (new taxa facilitate niche construction; newly formed niches can be occupied by the next "generation" of taxa). The latter possibility makes the mechanisms underlying the hyperbolic growth of biodiversity and human population even more similar, because the total ecospace of the biota is analogous to the "carrying capacity of the Earth" in demography. As far as new species can increase ecospace and facilitate opportunities for additional species entering the community, they are analogous to the "inventors" of the demographic models whose inventions increase the carrying capacity of the Earth. The hyperbolic growth of the Phanerozoic biodiverstiy suggests that "cooperative" interactions between taxa can play an important role in evolution, along with generally accepted competitive interactions. Due to this "cooperation", the evolution of biodiversity acquires some features of a self-accelerating process. Macroevolutionary "cooperation" reveals itself in: 1) increasing stability of communities that arises from alpha diversity growth; 2) ability of species to facilitate opportunities for additional species entering the community.