A62 A major likelihood-based approach gives problematic estimates of diversification dynamics and rates

Are polyploids really evolutionary dead-ends (again)? A critical reappraisal of Mayrose etal. ().

Corrigendum for Rojas etal. (2018) DOI: 10.1111/ele.12911.

The Proteaceae is a large angiosperm family displaying the common pattern of uneven distribution of species among genera. Previous studies have shown that this disparity is a result of variation in diversification rates across lineages, but the reasons for this variation are still unclear. Here, we tested the impact of floral symmetry and occurrence in Mediterranean climate regions on speciation and extinction rates in the Proteaceae. A rate shift analysis was conducted on dated genus-level phylogenetic trees of the Proteaceae. Character-dependent analyses were used to test for differences in diversification rates between actinomorphic and zygomorphic lineages and between lineages located within or outside Mediterranean climate regions. The rate shift analysis identified 5-10 major diversification rate shifts in the Proteaceae tree. The character-dependent analyses showed that speciation rates, extinction rates and net diversification rates of the Proteaceae were significantly higher for lineages occurring in Mediterranean hotspots. Higher speciation and extinction rates were also detected for zygomorphic species, but net diversification rates appeared to be similar in actinomorphic and zygomorphic Proteaceae. Presence in Mediterranean hotspots favors Proteaceae diversification. In contrast with observations at the scale of angiosperms, floral symmetry is not a trait that strongly influences their evolutionary success.

At present there are many animal phyla that contain only a few species. The existence of these small phyla can be used to test assumptions about speciation and extinction in multicellular animals.We first model the number of species in a monophyletic clade with a birth and death process that assumes rates of speciation and extinction are constant. If no new phyla have evolved since the Cambrian and speciation and extinction rates for minor phyla are similar to or greater than those estimated from fossils, then our model shows that the probabilities of minor phyla surviving to the present are small. Random variation in extinction and speciation rates does not improve the chances for persistence. If speciation rates exceed extinction rates at the initial radiation of the clade, but before the clade becomes large, speciation rates come to equal extinction rates and both are low, persistence from before the Ordovician up to the present becomes likely. These models show that if speciation and extinction rates are independent of the number of species in a clade, then conditions before the Ordovician strongly influence today's distribution of species among taxa.We also discuss a model in which speciation and extinction rates depend on the number of species in a clade. This alternative model can account for the persistence of phyla with few species to the present and predicts a short duration for phyla that did not exceed a threshold number of species.

Species‐specific diversification rates, or ‘tip rates’, can be computed quickly from phylogenies and are widely used to study diversification rate variation in relation to geography, ecology and phenotypes. These tip rates provide a number of theoretical and practical advantages, such as the relaxation of assumptions of rate homogeneity in trait‐dependent diversification studies. However, there is substantial confusion in the literature regarding whether these metrics estimate speciation or net diversification rates. Additionally, no study has yet compared the relative performance and accuracy of tip rate metrics across simulated diversification scenarios.We compared the statistical performance of three model‐free rate metrics (inverse terminal branch lengths; node density metric; DR statistic) and a model‐based approach (Bayesian analysis of macroevolutionary mixtures [BAMM]). We applied each method to a large set of simulated phylogenies that had been generated under different diversification processes. We summarized performance in relation to the type of rate variation, the magnitude of rate heterogeneity and rate regime size. We also compared the ability of the metrics to estimate both speciation and net diversification rates.We show decisively that model‐free tip rate metrics provide a better estimate of the rate of speciation than of net diversification. Error in net diversification rate estimates increases as a function of the relative extinction rate. In contrast, error in speciation rate estimates is low and relatively insensitive to extinction. Overall, and in particular when relative extinction was high, BAMM inferred the most accurate tip rates and exhibited lower error than non‐model‐based approaches. DR was highly correlated with true speciation rates but exhibited high error variance, and was the best metric for very small rate regimes.We found that, of the metrics tested, DR and BAMM are the most useful metrics for studying speciation rate dynamics and trait‐dependent diversification. Although BAMM was more accurate than DR overall, the two approaches have complementary strengths. Because tip rate metrics are more reliable estimators of speciation rate, we recommend that empirical studies using these metrics exercise caution when drawing biological interpretations in any situation where the distinction between speciation and net diversification is important.

The taxonomic evolution of Jurassic and Cretaceous calcareous nannofossil species is described using the following indices: species diversity, rate of speciation, rate of extinction, rate of diversification, rate of turnover, survivorship, and species accretion. The Jurassic prior to the late Oxfordian is characterized by positive diversification rates, that is, rates of speciation exceeded rates of extinction. Highest rates of diversification occurred in the late Lias and early Oxfordian. During the generally regressive latest Jurassic, diversification rates remained low and rates of extinctions exceed rates of speciation. In the early Cretaceous, rates of diversification are positive and peak in the early Valanginian, early Aptian, and middle Albian, after which time rates of extinction generally exceed rates of speciation. Such peaks in rate of evolution coincide with times of increased accumulation of organic carbon in the ocean (“anoxic events”). Peaks in rates of extinction result in very high rates of turnover during times of major regressions, in particular, in the Tithonian and Maastrichtian. Survivorship analyses for three datum planes (74.5, 144, and 160 Ma) show relatively constant extinction rates with some stepping in the older part; they are best explained by a temporally fluctuating abiotic environment causing changes in the probability of extinction. Species accretion curves are also relatively linear with some indication of changing rates of speciation. The coincidences of major changes in evolutionary rates with major paleoceanographic events are indicative of a predominantly abiotic control of nannoplankton evolution. Relationships of evolutionary rates of calcareous nannoplankton with deep ocean ventilation, sea level, and ocean fertility indicates that global tectonic processes are the ultimate causes of evolutionary change.

One of the most striking patterns observed among animals is that smaller-bodied taxa are generally much more diverse than larger-bodied taxa. This observation seems to be explained by the mere fact that smaller-bodied taxa tend to have an older evolutionary origin and have therefore had more time to diversify. A few studies, based on the prevailing null model of diversification (i.e. the stochastic constant-rate birth–death model), have suggested that this is indeed the correct explanation, and body-size dependence of speciation and extinction rates does not play a role. However, there are several potential shortcomings to these studies: a suboptimal statistical procedure and a relatively narrow range of body sizes in the analysed data. Here, we present a more coherent statistical approach, maximizing the likelihood of the constant-rate birth–death model with allometric scaling of speciation and extinction rates, given data on extant diversity, clade age and average body size in each clade. We applied our method to a dataset compiled from the literature that includes a wide range of Metazoan taxa (range from midges to elephants). We find that the higher diversity among small animals is indeed, partly, caused by higher clade age. However, it is also partly caused by the body-size dependence of speciation and extinction rates. We find that both the speciation rate and extinction rate decrease with body size such that the net diversification rate is close to 0. Even more interestingly, the allometric scaling exponent of speciation and extinction rates is approximately −0.25, which implies that the per generation speciation and extinction rates are independent of body size. This suggests that the observed relationship between diversity and body size pattern can be explained by clade age alone, but only if clade age is measured in generations rather than years. Thus, we argue that the most parsimonious explanation for the observation that smaller-bodied taxa are more diverse is that their evolutionary clock ticks faster.

A key measure of humanity's global impact is by how much it has increased species extinction rates. Familiar statements are that these are 100-1000 times pre-human or background extinction levels. Estimating recent rates is straightforward, but establishing a background rate for comparison is not. Previous researchers chose an approximate benchmark of 1 extinction per million species per year (E/MSY). We explored disparate lines of evidence that suggest a substantially lower estimate. Fossil data yield direct estimates of extinction rates, but they are temporally coarse, mostly limited to marine hard-bodied taxa, and generally involve genera not species. Based on these data, typical background loss is 0.01 genera per million genera per year. Molecular phylogenies are available for more taxa and ecosystems, but it is debated whether they can be used to estimate separately speciation and extinction rates. We selected data to address known concerns and used them to determine median extinction estimates from statistical distributions of probable values for terrestrial plants and animals. We then created simulations to explore effects of violating model assumptions. Finally, we compiled estimates of diversification-the difference between speciation and extinction rates for different taxa. Median estimates of extinction rates ranged from 0.023 to 0.135 E/MSY. Simulation results suggested over- and under-estimation of extinction from individual phylogenies partially canceled each other out when large sets of phylogenies were analyzed. There was no evidence for recent and widespread pre-human overall declines in diversity. This implies that average extinction rates are less than average diversification rates. Median diversification rates were 0.05-0.2 new species per million species per year. On the basis of these results, we concluded that typical rates of background extinction may be closer to 0.1 E/MSY. Thus, current extinction rates are 1,000 times higher than natural background rates of extinction and future rates are likely to be 10,000 times higher.

Quantitative traits have long been hypothesized to affect speciation and extinction rates. For example, smaller body size or increased specialization may be associated with increased rates of diversification. Here, I present a phylogenetic likelihood-based method (quantitative state speciation and extinction [QuaSSE]) that can be used to test such hypotheses using extant character distributions. This approach assumes that diversification follows a birth-death process where speciation and extinction rates may vary with one or more traits that evolve under a diffusion model. Speciation and extinction rates may be arbitrary functions of the character state, allowing much flexibility in testing models of trait-dependent diversification. I test the approach using simulated phylogenies and show that a known relationship between speciation and a quantitative character could be recovered in up to 80% of the cases on large trees (500 species). Consistent with other approaches, detecting shifts in diversification due to differences in extinction rates was harder than when due to differences in speciation rates. Finally, I demonstrate the application of QuaSSE to investigate the correlation between body size and diversification in primates, concluding that clade-specific differences in diversification may be more important than size-dependent diversification in shaping the patterns of diversity within this group.

Body mass is thought to influence diversification rates, but previous studies have produced ambiguous results. We investigated patterns of diversification across 100 trees obtained from a new Bayesian inference of primate phylogeny that sampled trees in proportion to their posterior probabilities. First, we used simulations to assess the validity of previous studies that used linear models to investigate the links between IUCN Red List status and body mass. These analyses support the use of linear models for ordinal ranked data on threat status, and phylogenetic generalized linear models revealed a significant positive correlation between current extinction risk and body mass across our tree block. We then investigated historical patterns of speciation and extinction rates using a recently developed maximum-likelihood method. Specifically, we predicted that body mass correlates positively with extinction rate because larger bodied organisms reproduce more slowly, and body mass correlates negatively with speciation rate because smaller bodied organisms are better able to partition niche space. We failed to find evidence that extinction rates covary with body mass across primate phylogeny. Similarly, the speciation rate was generally unrelated to body mass, except in some tests that indicated an increase in the speciation rate with increasing body mass. Importantly, we discovered that our data violated a key assumption of sample randomness with respect to body mass. After correcting for this bias, we found no association between diversification rates and mass.

The integration of Geographic Information System (GIS) methodology within a phylogenetic and statistical framework provides a background against which to evaluate the relationship between biogeographic changes and evolution in the fossil record. A case study based on patterns in Middle and Late Devonian phyllocarids (Crustacea) illustrates the usefulness of this integrated approach. Using a combined approach enhances determination of rates of biodiversity change and the relationship between biogeographic and evolutionary changes. Because the interaction between speciation and extinction rates fundamentally determines biodiversity dynamics, and speciation and extinction rates are influenced by the geographic ranges of component taxa, the relationship between biogeography and evolution is important. Furthermore, GIS makes it possible to quantify paleobiogeographic ranges.Phylogenetic biogeography resolved patterns of both vicariance and geodispersal and revealed that range expansions were more abundant than range contractions in Devonian phyllocarids. In addition, statistical tests on GIS-constrained species ranges and evolutionary-rate data revealed a relationship between increasing species' ranges and increases in both speciation and extinction rates. Extinction rate, however, increased more rapidly than speciation rate in the phyllocarids. The pattern of extinction rate increasing faster than speciation rate in the phyllocarids may illuminate aspects of the Late Devonian biodiversity crisis in particular, and protracted biodiversity crises in general.

Species diversity patterns are governed by complex interactions among biotic and abiotic factors over time and space, but are essentially the result of the diversification dynamics (differential speciation and extinction rates) over the long‐term evolutionary history of a clade. Previous studies have suggested that temporal variation in global temperature drove long‐term diversity changes in Crocodylia, a monophyletic group of large ectothermic organisms. We use a large database of crocodylian fossil occurrences (192 spp.) and body mass estimations, under a taxic approach, to characterize the global diversification dynamics of crocodylians since the Cretaceous, and their correlation with multiple biotic and abiotic factors in a Bayesian framework. The diversification dynamic of crocodylians, which appears to have originated in the Turonian (c. 92.5 Ma), is characterized by several phases with high extinction and speciation rates within a predominantly low long‐term mean rate. Our results reveal long‐term diversification dynamics of Crocodylia to be a highly complex process driven by a combination of biotic and abiotic factors which influenced the speciation and extinction rates in dissimilar ways. Higher crocodylian extinction rates are related to low body mass disparity, indicating selective extinctions of taxa at both ends of the body mass spectrum. Speciation rate slowdowns are noted when the diversity of the clade is high and the warm temperate climatic belt is reduced. Our finding supports the idea that temporal variations of body mass disparity, self‐diversity, and the warm climate belt size provided more direct mechanistic explanations for crocodylian diversification than do proxies of global temperature.

Author SummaryWhy are there more species in the tropics? This question has fascinated ecologists and evolutionary biologists for decades, generating hundreds of hypotheses, yet basic questions remain: Are rates of speciation higher in the tropics? Are rates of extinction higher in temperate regions? Do the tropics act as a source of diversity for temperate regions? We estimated rates of speciation, extinction, and range expansion associated with mammals living in tropical and temperate regions, using an almost complete mammalian phylogeny. Contrary to what has been suggested before for this class of vertebrates, we found that diversification rates are strikingly consistent with diversity patterns, with latitudinal peaks in species richness being associated with high speciation rates, low extinction rates, or both, depending on the mammalian order (rodents, bats, primates, etc.). We also found evidence for an asymmetry in range expansion, with more expansion “out of” than “into” the tropics. Taken together, these results suggest that tropical regions are not only a reservoir of biodiversity, but also the main place where biodiversity is generated.

Phylogenies are increasingly being used as a basis to provide insight into macroevolutionary history. Here, we use simulation experiments and empirical analyses to evaluate methods that use phylogenies as a basis to make estimates of divergence times and rates of diversification. This is the first study to present a comprehensive assessment of the key variables that underpin analyses in this field-including substitution rates, speciation rates, and extinction, plus character sampling and taxon sampling. We show that in unrealistically simplistic cases (where substitution rates and speciation rates are constant, and where there is no extinction), increased character and taxon sampling lead to more accurate and precise parameter estimates. By contrast, in more complex but realistic cases (where substitution rates, speciation rates, and extinction rates vary), gains in accuracy and precision from increased character and taxon sampling are far more limited. The lack of accuracy and precision even occurs when using methods that are designed to account for more complex cases, such as relaxed clocks, fossil calibrations, and models that allow speciation rates and extinction rates to vary. The problem also persists when analyzing genomic scale data sets. These results suggest two interrelated problems that occur when the processes that generated the data are more complex. First, methodological assumptions are more likely to be violated. Second, limitations in the information content of the data become more important.[Divergence time estimation; diversification rates; macroevolution; phylogeny.].

Net diversification rates were estimated for samples of primarily tribe-to-family-level clades of passerine birds, taking into account extinction as well as speciation. Two samples were used. The first consisted of 37 clades of primarily temperate North American and primarily tropical South American passerines; the second comprised a global set of 90 clades, each distributed within one or more zoogeographic regions. Circumscription and ages of clades were taken from Sibley and Ahlquist's phylogeny based on DNA hybridization, with updates from more recent sequence analysis. Under a homogeneous speciation (rate = lamda) and extinction (rate = mu) process, the expected number of species (N) after t units of time can be described by the expression, N(t)= [exp(lamda(1 - kappa) t - kappa]/(1 - kappa), where kappa = mu/lamda. A nonlinear least-squares regression for the temperate and tropical American clades with more than one species estimated kappa = 0.938 +/- 0.076 (mean +/- SE), suggesting a high rate of turnover of lineages within clades. Because of the broad confidence limits in kappa, I used values ranging from 0.80 to 0.98 to calculate speciation rates in subsequent analyses, assuming that kappa is uniform among clades and does not vary with latitude. Speciation rate among South American clades exceeded that among North American clades for all kappa, whether monophyletic lineages were included or not. The estimated speciation rate was negatively related to clade age, suggesting that proliferation within clades slows with time. In the global data set, rate of speciation decreased with clade age and increased with the area of the region or regions within which a clade is distributed, and for any given value of kappa the speciation rate was significantly higher in tropical than in temperate regions. Relaxing the assumption of latitude independence in kappa, larger clade size in the tropics could be achieved by various combinations of relative speciation and extinction rates that obscure the underlying causes of global biodiversity patterns. Nonetheless, the results of this analysis clearly indicate that a higher rate of diversification in the tropics contributes to the pervasive latitudinal gradient in diversity observed in passerine birds.