Andrew Z. Krug

Phanerozoic trends in the global diversity of marine invertebrates.

It has previously been thought that there was a steep Cretaceous and Cenozoic radiation of marine invertebrates. This pattern can be replicated with a new data set of fossil occurrences representing 3.5 million specimens, but only when older analytical protocols are used. Moreover, analyses that employ sampling standardization and more robust counting methods show a modest rise in diversity with no clear trend after the mid-Cretaceous. Globally, locally, and at both high and low latitudes, diversity was less than twice as high in the Neogene as in the mid-Paleozoic. The ratio of global to local richness has changed little, and a latitudinal diversity gradient was present in the early Paleozoic.

Out of the tropics, but how? Fossils, bridge species, and thermal ranges in the dynamics of the marine latitudinal diversity gradient.

Latitudinal diversity gradients are underlain by complex combinations of origination, extinction, and shifts in geographic distribution and therefore are best analyzed by integrating paleontological and neontological data. The fossil record of marine bivalves shows, in three successive late Cenozoic time slices, that most clades (operationally here, genera) tend to originate in the tropics and then expand out of the tropics (OTT) to higher latitudes while retaining their tropical presence. This OTT pattern is robust both to assumptions on the preservation potential of taxa and to taxonomic revisions of extant and fossil species. Range expansion of clades may occur via “bridge species,” which violate climate-niche conservatism to bridge the tropical-temperate boundary in most OTT genera. Substantial time lags (∼5 Myr) between the origins of tropical clades and their entry into the temperate zone suggest that OTT events are rare on a per-clade basis. Clades with higher diversification rates within the tropics are the most likely to expand OTT and the most likely to produce multiple bridge species, suggesting that high speciation rates promote the OTT dynamic. Although expansion of thermal tolerances is key to the OTT dynamic, most latitudinally widespread species instead achieve their broad ranges by tracking widespread, spatially-uniform temperatures within the tropics (yielding, via the nonlinear relation between temperature and latitude, a pattern opposite to Rapoport’s rule). This decoupling of range size and temperature tolerance may also explain the differing roles of species and clade ranges in buffering species from background and mass extinctions.

Incumbency, diversity, and latitudinal gradients

Physical environmental factors have been seen as paramount in determining many large-scale biodistributional patterns in time and space. Although this is probably correct for many situations, this view has become so pervasive that it has led to the neglect of the role of biotic interactions in setting large-scale diversity patterns. (In this paper diversity denotes taxonomic richness.) New approaches to this perennial debate on the roles of physical and biotic forces in paleoecology and macroevolution are needed, and here we explore an argument for the role of incumbency or priority effects in the dynamics behind the most dramatic spatial pattern in biodiversity, the latitudinal diversity gradient. A global analysis of the fossil record of living marine bivalve genera and subgenera (hereafter simply genera) of the continental shelves provides perhaps the strongest evidence for the Out of the Tropics (OTT) dynamic associated with the formation of the present marine latitudinal diversity gradient (LDG) (Jablonski et al. 2006). The marine LDG appears to be driven primarily by the origin of novel lineages in the Tropics, some of which then expand their ranges into higher, extratropical latitudes (see Jablonski 1993, 2005; Clark and Crame 2003; Goldberg et al. 2005; Jablonski et al. 2006; Martin et al. 2007). Support for this pattern comes from the overwhelmingly tropical first fossil occurrences of living bivalve genera and their subsequent appearances in higher latitudes. Some genera are first found in the extratropical fossil record, but these never match, at any latitude, the number or proportion of genera that have expanded from the Tropics (the ratio is generally ∼3:1), and even this smaller number is probably an overestimate, because the extratropical post-Paleozoic fossil record is so much better sampled than that of the Tropics (Allison and Briggs 1993; Jablonski 1993; Jackson …

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.

Global environmental predictors of benthic marine biogeographic structure

Analyses of how environmental factors influence the biogeographic structure of biotas are essential for understanding the processes underlying global diversity patterns and for predicting large-scale biotic responses to global change. Here we show that the large-scale geographic structure of shallow-marine benthic faunas, defined by existing biogeographic schemes, can be predicted with 89–100% accuracy by a few readily available oceanographic variables; temperature alone can predict 53–99% of the present-day structure along coastlines. The same set of variables is also strongly correlated with spatial changes in species compositions of bivalves, a major component of the benthic marine biota, at the 1° grid-cell resolution. These analyses demonstrate the central role of coastal oceanography in structuring benthic marine biogeography and suggest that a few environmental variables may be sufficient to model the response of marine biogeographic structure to past and future changes in climate.

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/Gs falling within the null expectation, and tropical and polar S/Gs 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.

Signature of the end-Cretaceous mass extinction in the modern biota.

The long-term effects of mass extinctions on spatial and evolutionary dynamics have been poorly studied. Here we show that the evolutionary consequences of the end-Cretaceous [Cretaceous/Paleogene (K/Pg)] mass extinction persist in present-day biogeography. The geologic ages of genera of living marine bivalves show a significant break from a smooth exponential distribution, corresponding to the K/Pg boundary. The break reflects a permanent increase in origination rates, intermediate between the Mesozoic rate and the post-extinction recovery pulse. This global rate shift is most clearly seen today in tropical bioprovinces and weakens toward the poles. Coupled with the modern geographic distributions of taxa originating before and after the K/Pg boundary, this spatial pattern indicates that tropical origination rates after the K/Pg event have left a permanent mark on the taxonomic and biogeographic structure of the modern biota, despite the complex Cenozoic history of marine environments.

Geographic variation in turnover and recovery from the Late Ordovician mass extinction

Abstract Understanding what drives global diversity requires knowledge of the processes that control diversity and turnover at a variety of geographic and temporal scales. This is of particular importance in the study of mass extinctions, which have disproportionate effects on the global ecosystem and have been shown to vary geographically in extinction magnitude and rate of recovery. Here, we analyze regional diversity and turnover patterns for the paleocontinents of Laurentia, Baltica, and Avalonia spanning the Late Ordovician mass extinction and Early Silurian recovery. Using a database of genus occurrences for inarticulate and articulate brachiopods, bivalves, anthozoans, and trilobites, we show that sampling-standardized diversity trends differ for the three regions. Diversity rebounded to pre-extinction levels within 5 Myr in the paleocontinent of Laurentia, compared with 15 Myr or longer for Baltica and Avalonia. This increased rate of recovery in Laurentia was due to both lower Late Ordovician extinction rates and higher Early Silurian origination rates relative to the other continents. Using brachiopod data, we dissected the Rhuddanian recovery into genus origination and invasion. This analysis revealed that standing diversity in the Rhuddanian consisted of a higher proportion of invading taxa in Laurentia than in either Baltica or Avalonia. Removing invading genera from diversity counts caused Rhuddanian diversity to fall in Laurentia. However, Laurentian diversity still rebounded to pre-extinction levels within 10 Myr of the extinction event, indicating that genus origination rates were also higher in Laurentia than in either Baltica or Avalonia. Though brachiopod diversity in Laurentia was lower than in the higher-latitude continents prior to the extinction, increased immigration and genus origination rates made it the most diverse continent following the extinction. Higher rates of origination in Laurentia may be explained by its large size, paleogeographic location, and vast epicontinental seas. It is possible that the tropical position of Laurentia buffered it somewhat from the intense climatic fluctuations associated with the extinction event, reducing extinction intensities and allowing for a more rapid rebound in this region. Hypotheses explaining the increased levels of invasion into Laurentia remain largely untested and require further scrutiny. Nevertheless, the Late Ordovician mass extinction joins the Late Permian and end-Cretaceous as global extinction events displaying an underlying spatial complexity.

The sampling and estimation of marine paleodiversity patterns: implications of a Pliocene model

Abstract Data that accurately capture the spatial structure of biodiversity are required for many paleobiological questions, from assessments of changing provinciality and the role of geographic ranges in extinction and originations, to estimates of global taxonomic or morphological diversity through time. Studies of temporal changes in diversity and global biogeographic patterns have attempted to overcome fossil sampling biases through sampling standardization protocols, but such approaches must ultimately be limited by available literature and museum collections. One approach to evaluating such limits is to compare results from the fossil record with models of past diversity patterns informed by modern relationships between diversity and climatic factors. Here we use present-day patterns for marine bivalves, combined with data on the geologic ages and distributions of extant taxa, to develop a model for Pliocene diversity patterns, which is then compared with diversity patterns retrieved from the literature as compiled by the Paleobiology Database (PaleoDB). The published Pliocene bivalve data (PaleoDB) lack the first-order spatial structure required to generate the modern biogeography within the time available (<3 Myr). Instead, the published data (raw and standardized) show global diversity maxima in the Tropical West Atlantic, followed closely by a peak in the cool-temperate East Atlantic. Either todays tropical West Pacific diversity peak, double that of any other tropical region, is a purely Pleistocene phenomenon—highly unlikely given the geologic ages of extant genera and the topology of molecular phylogenies—or the paleontological literature is such a distorted sample of tropical Pliocene diversity that current sampling standardization methods cannot compensate for existing biases. A rigorous understanding of large-scale spatial and temporal diversity patterns will require new approaches that can compensate for such strong bias, presumably by drawing more fully on our understanding of the factors that underlie the deployment of diversity today.

Contrarian clade confirms the ubiquity of spatial origination patterns in the production of latitudinal diversity gradients

The latitudinal diversity gradient (LDG), wherein the number of species and higher taxa peaks in the tropics and decreases toward the poles, is the best-documented large-scale diversity pattern on Earth, but hypotheses explaining the standard LDG must also account for rare “contrarian” taxa that show diversity maxima outside of the tropics. For marine bivalves, one of the few groups that provide spatially explicit temporal data on a global scale, we show that a major contrarian group, the Anomalodesmata, unexpectedly exhibits the same large-scale dynamics as related clades having normal LDGs in two key respects. First, maxima in standing genus diversity and genus origination rates coincide spatially. Second, the strength of a clades present-day LDG is significantly related to the proportion of its living genera that originated in the tropics during the late Cenozoic, with the contrarian gradient strength at both species and genus level predicted quantitatively by the values for the other clades. Geologic age distributions indicate that the anomalous LDG results from origination that is damped in the tropics rather than heightened in the temperate zones. The pervasive role of spatial origination patterns in shaping LDGs, regardless of the position of their diversity maxima, corroborates hypotheses based on clades showing standard gradients and underscores the insights that contrarian groups can provide into general principles of diversity dynamics.