Gene Hunt

The relative importance of directional change, random walks, and stasis in the evolution of fossil lineages

The nature of evolutionary changes recorded by the fossil record has long been controversial, with particular disagreement concerning the relative frequency of gradual change versus stasis within lineages. Here, I present a large-scale, statistical survey of evolutionary mode in fossil lineages. Over 250 sequences of evolving traits were fit by using maximum likelihood to three evolutionary models: directional change, random walk, and stasis. Evolution in these traits was rarely directional; in only 5% of fossil sequences was directional evolution the most strongly supported of the three modes of change. The remaining 95% of sequences were divided nearly equally between random walks and stasis. Variables related to body size were significantly less likely than shape traits to experience stasis. This finding is in accord with previous suggestions that size may be more evolutionarily labile than shape and is consistent with some but not all of the mechanisms proposed to explain evolutionary stasis. In general, similar evolutionary patterns are observed across other variables, such as clade membership and temporal resolution, but there is some evidence that directional change in planktonic organisms is more frequent than in benthic organisms. The rarity with which directional evolution was observed in this study corroborates a key claim of punctuated equilibria and suggests that truly directional evolution is infrequent or, perhaps more importantly, of short enough duration so as to rarely register in paleontological sampling.

Climate change, body size evolution, and Cope's Rule in deep-sea ostracodes

Causes of macroevolutionary trends in body size, such as Copes Rule, the tendency of body size to increase over time, remain poorly understood. We used size measurements from Cenozoic populations of the ostracode genus Poseidonamicus, in conjunction with phylogeny and paleotemperature estimates, to show that climatic cooling leads to significant increases in body size, both overall and within individual lineages. The magnitude of size increase due to Cenozoic cooling is consistent with temperature-size relationships in geographically separated modern populations (Bergmanns Rule). Thus population-level phenotypic evolution in response to climate change can be an important determinant of macroevolutionary trends in body size.

Larval Ecology, Geographic Range, and Species Survivorship in Cretaceous Mollusks: Organismic versus Species‐Level Explanations

The observation that geographic range size in Cretaceous mollusks is correlated with species survivorship and is heritable at the species level has figured repeatedly in discussions of species selection over the past two decades. However, some authors have suggested that the relationship between mode of larval development and geographic range supports the reduction of this example to selection on organismic properties. Our reanalysis of Jablonskis work on heritability at the species level finds that geographic range is significantly heritable (using a randomization test) in both bivalves and gastropods, even within a single larval mode. Further, generalized linear models show that geographic range size is more important than larval mode in predicting extinction probability in both gastropods and bivalves. These results reaffirm the role and heritability of geographic range as a species‐level property that can promote species selection; the model‐based approach applied here may help to operationalize “screening off ” and related approaches to evaluating hierarchical explanations in evolution.

Evolution toward a new adaptive optimum: phenotypic evolution in a fossil stickleback lineage.

Abstract Natural selection has almost certainly shaped many evolutionary trajectories documented in fossil lineages, but it has proven difficult to demonstrate this claim by analyzing sequences of evolutionary changes. In a recently published and particularly promising test case, an evolutionary time series of populations displaying armor reduction in a fossil stickleback lineage could not be consistently distinguished from a null model of neutral drift, despite excellent temporal resolution and an abundance of indirect evidence implicating natural selection. Here, we revisit this case study, applying analyses that differ from standard approaches in that: (1) we do not treat genetic drift as a null model, and instead assess neutral and adaptive explanations on equal footing using the Akaike Information Criterion; and (2) rather than constant directional selection, the adaptive scenario we consider is that of a population ascending a peak on the adaptive landscape, modeled as an Orstein–Uhlenbeck process. For all three skeletal features measured in the stickleback lineage, the adaptive model decisively outperforms neutral evolution, supporting a role for natural selection in the evolution of these traits. These results demonstrate that, at least under favorable circumstances, it is possible to infer in fossil lineages the relationship between evolutionary change and features of the adaptive landscape.

Species‐Level Heritability Reaffirmed: A Comment on “On the Heritability of Geographic Range Sizes”

For many current issues in macroevolution and macroecology, it is important to know to what degree the attributes of species are shared among closely related lineages, a concept sometimes referred to as species‐level heritability. Recently, Webb and Gaston proposed a new method for analyzing the heritability of geographic range size and concluded that range size is not heritable in Cretaceous gastropods (data from Jablonski) and modern birds (their data). Here we show that Webb and Gaston’s method is flawed in that it implicitly assumes that range sizes are uniformly distributed. When range size distributions show their characteristic strong right skew, Webb and Gaston’s method spuriously tends to find that range sizes of closely related pairs of species are more dissimilar than the random expectation. A reanalysis of Jablonski’s data finds range size to be robustly and strongly heritable in Cretaceous gastropods and less strongly but still significantly heritable in present‐day birds.

The use of sighting records to infer species extinctions: an evaluation of different methods

In the absence of long-term monitoring data, inferences about extinctions of species and populations are generally based on past observations about the presence of a particular species at specified places and times (sightings). Several methods have been developed to estimate the probability and timing of extinctions from records of such sightings, but they differ in their computational complexity and assumptions about the nature of the sighting record. Here we use simulations to evaluate the performance of seven methods proposed to estimate the upper confidence limit on extinction times under different extinction and sampling scenarios. Our results show that the ability of existing methods to correctly estimate the timing of extinctions varies with the type of extinction (sudden vs. gradual) and the nature of sampling effort over time. When the probability of sampling a species declines over time, many of the methods perform poorly. On the other hand, the simulation results also suggest that as long as the choice of the method is determined by the nature of the underlying sighting data, existing methods should provide reliable inferences about the timing of past extinctions.

A macroevolutionary perspective on species range limits

Understanding the factors that determine the geographic range limits of species is important for many questions in ecology, evolution and conservation biology. These limits arise from complex interactions among ecology and dispersal ability of species and the physical environment, but many of the underlying traits can be conserved among related species and clades. Thus, the range limits of species are likely to be influenced by their macroevolutionary history. Using palaeontological and biogeographic data for marine bivalves, we find that the range limits of genera are significantly related to their constituent species richness, but the effects of age are weak and inconsistent. In addition, we find a significant phylogenetic signal in the range limits at both genus and family levels, although the strength of this effect shows interoceanic variation. This phylogenetic conservatism of range limits gives rise to an evolutionary pattern where wide-ranging lineages have clusters of species within the biogeographic provinces, with a few extending across major boundaries.

Phylogenetic Conservatism of Extinctions in Marine Bivalves

Honing Bivalve History What are the lasting effects of extinction, both persistent background extinctions and major events, on surviving lineages? Roy et al. (p. 733) examined the excellent fossil record of marine bivalves over the past 200 million years, which spans the end-Cretaceous extinction. Background extinctions tended to be higher within certain lineages and depended on the previous history of extinctions within those lineages. Cenozoic taxa are still reflecting the end-Cretaceous event. Extinction rates of fossil bivalves tended to be higher in certain lineages and were influenced by prior events. Evolutionary histories of species and lineages can influence their vulnerabilities to extinction, but the importance of this effect remains poorly explored for extinctions in the geologic past. When analyzed using a standardized taxonomy within a phylogenetic framework, extinction rates of marine bivalves estimated from the fossil record for the last ~200 million years show conservatism at multiple levels of evolutionary divergence, both within individual families and among related families. The strength of such phylogenetic clustering varies over time and is influenced by earlier extinction history, especially by the demise of volatile taxa in the end-Cretaceous mass extinction. Analyses of the evolutionary roles of ancient extinctions and predictive models of vulnerability of taxa to future natural and anthropogenic stressors should take phylogenetic relationships and extinction history into account.

Temporal latitudinal-gradient dynamics and tropical instability of deep-sea species diversity

A benthic microfaunal record from the equatorial Atlantic Ocean over the past four glacial-interglacial cycles was investigated to understand temporal dynamics of deep-sea latitudinal species diversity gradients (LSDGs). The results demonstrate unexpected instability and high amplitude fluctuations of species diversity in the tropical deep ocean that are correlated with orbital-scale oscillations in global climate: Species diversity is low during glacial and high during interglacial periods. This implies that climate severely influences deep-sea diversity, even at tropical latitudes, and that deep-sea LSDGs, while generally present for the last 36 million years, were weakened or absent during glacial periods. Temporally dynamic LSDGs and unstable tropical diversity require reconsideration of current ecological hypotheses about the generation and maintenance of biodiversity as they apply to the deep sea, and underscore the potential vulnerability and conservation importance of tropical deep-sea ecosystems.

Measuring rates of phenotypic evolution and the inseparability of tempo and mode

Abstract Rates of phenotypic evolution are central to many issues in paleontology, but traditional rate metrics such as darwins or haldanes are seldom used because of their strong dependence on interval length. In this paper, I argue that rates are usefully thought of as model parameters that relate magnitudes of evolutionary divergence to elapsed time. Starting with models of directional evolution, random walks, and stasis, I derive for each a reasonable rate metric. These metrics can be linked to existing approaches in evolutionary biology, and simulations show that they can be estimated accurately at any temporal resolution via maximum likelihood, but only when that metrics underlying model is true. The estimation of generational rates of a random walk under realistic paleontological conditions is compared with simulations to that of a prominent alternative approach, Gingerichs LRI (log-rate, log-interval) method. Generational rates are estimated poorly by LRI; they often reflect sampling error more than the actual pace of change. Further simulations show that under some realistic conditions, it is simply not possible to infer generational rates from coarsely sampled populations. These modeling results indicate a complex dependence between evolutionary mode and the measurement of evolutionary rates, and that there is unlikely to be a rate metric that works well for all traits and time scales. Compilations of paleontological and phylogenetic data indicate that all of the three rate metrics derived here show some relationship with interval length. Although there is no perfect rate metric, at present the most practical choices derive from the parameters of the stasis and random walk models. The latter, called the step variance, is particularly promising as a rate metric in paleontology and comparative biology.