Revisiting Jablonski (1993): cladogenesis and range expansion explain latitudinal variation in taxonomic richness

The increase in diversity towards the equator arises from latitudinal variation in rates of cladogenesis, extinction, immigration and/or emigration of taxa. We tested the relative contribution of all four processes to the latitudinal gradient in 26 marine invertebrate orders with extensive fossil records, examined previously by David Jablonski. Coupling Jablonski's estimates of latitudinal variation in cladogenesis with new data on patterns of extinction and current distributions, we show that the present-day gradient in diversity is caused by higher rates of cladogenesis and subsequent range expansion (immigration) at lower latitudes. In contrast, extinction and emigration were not important in the creation of the latitudinal gradient in ordinal richness. This work represents one of the first simultaneous tests of the role of all four processes in the creation of the latitudinal gradient in taxonomic richness, and suggests that low tropical extinction rates are not essential to the creation of latitudinal diversity gradients.     

A temperate palaeodiversity peak in Mesozoic dinosaurs and evidence for Late Cretaceous geographical partitioning

Aim Modern biodiversity peaks in the tropics and declines poleward, a pattern that is potentially driven by climate. Although this latitudinal biodiversity gradient (LBG) also characterizes the marine invertebrate fossil record, distributions of ancient terrestrial faunas are poorly understood. This study utilizes data on the dinosaur fossil record to examine spatial patterns in terrestrial biodiversity throughout the Mesozoic. Location We compiled data on fossil occurrences across the globe. Methods We compiled a comprehensive dataset of Mesozoic dinosaur genera (738), including birds. Following the utilization of sampling standardization techniques to mediate for the uneven sampling of the fossil record, we constructed latitudinal patterns of biodiversity from this dataset. Results The dominant group of Mesozoic terrestrial vertebrates did not conform to the modern LBG. Instead, dinosaur diversity was highest at temperate palaeolatitudes throughout the 160 million year span of dinosaurian evolutionary history. Latitudinal diversity correlates strongly with the distribution of land area. Late Cretaceous sauropods and ornithischians exhibit disparate LBGs. Main conclusions The continuity of the palaeotemperate peak in dinosaur diversity indicates a diminished role for climate on the Mesozoic LBG; instead, dinosaur diversity may have been driven by the amount of land area among latitudinal belts. There is no evidence that the tropics acted as a cradle for dinosaur diversity. Geographical partitioning among major clades of herbivorous dinosaurs in the Late Cretaceous may result from the advanced stages of continental fragmentation and/or differing responses to increasing latitudinal climatic zonation. Our results suggest that the modern‐day LBG on land was only established 30 million years ago, following a significant post‐Eocene recalibration, potentially related to increased seasonality.     

Taxonomic diversity gradients through geological time

Abstract. There is evidence from the fossil record to suggest that latitudinal gradients in taxonomic diversity may be time‐invariant features, although almost certainly not on the same scale as that seen at the present day. It is now apparent that both latitudinal and longitudinal gradients increased dramatically in strength through the Cenozoic era (i.e. the last 65 my) to become more pronounced today than at any time in the geological past. Present‐day taxonomic diversity gradients, in both the marine and terrestrial realms, are underpinned by the tropical radiations of a comparatively small number of species‐rich clades. Quite why these particular taxa proliferated through the Cenozoic is uncertain, but it could be that at least part of the explanation involves the phenomenon of evolutionary escalation. This is, in essence, a theory of biological diversification through evolutionary feedback mechanisms between predators and prey; first one develops an adaptive advantage, and then the other. However, there may also have been some form of extrinsic control on the process of tropical diversification, and this was most likely centred on the phenomenon of global climate change. This is especially so over the last 15 my Various Late Cenozoic (Neogene) vicariant events effectively partitioned the tropics into a series of high diversity centres, or foci. It has been suggested that, in the largest of these in the marine realm (the Indo‐West Pacific or IWP centre), a critical patterns of islands acted as a template for rapid speciation during glacioeustatic sea level cycles. The same process occurred in the Atlantic, Caribbean and East Pacific (ACEP) centre, though on a lesser scale. Tropical terrestrial diversity may also have been promoted by rapid range expansions and contractions in concert with glacial cycles (a modified refugium hypothesis). We are beginning to appreciate that an integrated sequence of Neogene tectonic and climatic events greatly influenced the formation of contemporary taxonomic diversity patterns.     

Evolutionary dispersal drives the latitudinal diversity gradient of stony corals

The diversity of stony corals displays one of the most exemplary latitudinal gradients on the planet, yet the evolutionary dynamics that produced this pattern remains unclear. Using both paleontological and distributional data, we compare the origination, extinction and immigration levels between low and high latitudes since the earliest proliferation of the group during the mid‐Triassic. Altogether, first and last occurrence localities in the fossil record do not support a positive preference towards either latitudinal bin. Nonetheless, considering past and present scleractinian fauna, the process of extinction has been apparently more pronounced at higher latitudes based on face values and correlation coefficients. Far above these differences, immigration of extant taxa has been substantially higher towards the tropics than to temperate regions. While the net dispersal toward low latitudes persists in all temporal intervals, the gradient of diversity was largely built up during the Cenozoic Era and only becomes significantly steep from the Neogene Period onwards. This dynamic supports the ‘into the tropical museum’ model, which suggests that tropics have historically acted as a center of accumulation for marine biodiversity.     

Mitogenomic Evidence for an Indo-West Pacific Origin of the Clupeoidei (Teleostei: Clupeiformes)

The clupeoid fishes are distributed worldwide, with marine, freshwater and euryhaline species living in either tropical or temperate environments. Regional endemism is important at the species and genus levels, and the highest species diversity is found in the tropical marine Indo-West Pacific region. The clupeoid distribution follows two general pattern of species richness, the longitudinal and latitudinal gradients. To test historical hypotheses explaining the formation of these two gradients, we have examined the early biogeography of the Clupeoidei in reconstructing the evolution of their habitat preferences along with their ancestral range distributions on a time-calibrated mitogenomic phylogeny. The phylogenetic results support the distinction of nine main lineages within the Clupeoidei, five of them new. We infer several independent transitions from a marine to freshwater environment and from a tropical to temperate environment that occurred after the initial diversification period of the Clupeoidei. These results combined with our ancestral range reconstruction hypothesis suggest that the probable region of origin and diversification of the Clupeoidei during the Cretaceous period was the tropical marine precursor to the present Indo-West Pacific region. Thus, our study favors the hypotheses of “Region of origin” and “Tropical conservatism” to explain the origins of the longitudinal and latitudinal gradients of clupeoid species richness, respectively. Additional geological and paleontological evidence further define the tropical marine paleo-region of origin as the eastern Tethys Sea region. The Cretaceous fossil record of the Clupeoidei is partially incongruent with the results here as it contains taxa found outside this region. We discuss three possible causes of conflict between our biogeographical hypothesis and the distributions of the Cretaceous clupeoid fossils: regional extinction, incomplete taxonomic sampling and incorrect timescale estimation.     

Northward shift in faunal diversity is a general pattern of evolution of the phanerozoic marine biota

The analysis of two global databases on spatio-temporal distribution of fossil marine animal genera (Sepkoski's compendium and The Paleobiology Database) has revealed the presence of the latitudinal diversity gradient (LDG) in the marine realm throughout the Phanerozoic. Within each time interval, LDG is characterized by two parameters: the latitudinal position of peak diversity and the steepness of monotonous decline of diversity with increasing distance from the zone of the highest diversity. During the Phanerozoic, peak diversity has drifted gradually from the tropics and subtropics of the Southern hemisphere into northern midlatitudes. The shift in peak diversity is not likely to be an artifact of incompleteness of the fossil record or uneven sampling of different regions. The shift proceeded in a stepwise manner, with periods of relatively fast changes separated by longer periods of little or no change. The latitudinal shift in peak diversity was probably due to a combination of several causes: tectonic (northward shift in the latitudinal distribution of continental shelf area), climatic (as demonstrated by the fact that peak diversity tended to occur near equator during the cold epochs and in midlatitudes during the warm epochs), and historical ("evolutionary inertia" of local faunas).     

The early Eocene birds of the Messel fossil site: a 48 million‐year‐old bird community adds a temporal perspective to the evolution of tropical avifaunas

Birds play an important role in studies addressing the diversity and species richness of tropical ecosystems, but because of the poor avian fossil record in all extant tropical regions, a temporal perspective is mainly provided by divergence dates derived from calibrated molecular analyses. Tropical ecosystems were, however, widespread in the Northern Hemisphere during the early Cenozoic, and the early Eocene German fossil site Messel in particular has yielded a rich avian fossil record. The Messel avifauna is characterized by a considerable number of flightless birds, as well as a high diversity of aerial insectivores and the absence of large arboreal birds. With about 70 currently known species in 42 named genus‐level and at least 39 family‐level taxa, it approaches extant tropical biotas in terms of species richness and taxonomic diversity. With regard to its taxonomic composition and presumed ecological characteristics, the Messel avifauna is more similar to the Neotropics, Madagascar, and New Guinea than to tropical forests in continental Africa and Asia. Because the former regions were geographically isolated during most of the Cenozoic, their characteristics may be due to the absence of biotic factors, especially those related to the diversification of placental mammals, which impacted tropical avifaunas in Africa and Asia. The crown groups of most avian taxa that already existed in early Eocene forests are species‐poor. This does not support the hypothesis that the antiquity of tropical ecosystems is key to the diversity of tropical avifaunas, and suggests that high diversification rates may be of greater significance.     

The impact of shifts in marine biodiversity hotspots on patterns of range evolution: Evidence from the Holocentridae (squirrelfishes and soldierfishes)

One of the most striking biodiversity patterns is the uneven distribution of marine species richness, with species diversity in the Indo-Australian Archipelago (IAA) exceeding all other areas. However, the IAA formed fairly recently, and marine biodiversity hotspots have shifted across nearly half the globe since the Paleogene. Understanding how lineages have responded to shifting biodiversity hotspots represents a necessary historic perspective on the formation and maintenance of global marine biodiversity. Such evolutionary inferences are often challenged by a lack of fossil evidence that provide insights into historic patterns of abundance and diversity. The greatest diversity of squirrelfishes and soldierfishes (Holocentridae) is in the IAA, yet these fishes also represent some of the most numerous fossil taxa in deposits of the former West Tethyan biodiversity hotspot. We reconstruct the pattern of holocentrid range evolution using time-calibrated phylogenies that include most living species and several fossil lineages, demonstrating the importance of including fossil species as terminal taxa in ancestral area reconstructions. Holocentrids exhibit increased range fragmentation following the West Tethyan hotspot collapse. However, rather than originating within the emerging IAA hotspot, the IAA has acted as a reservoir for holocentrid diversity that originated in adjacent regions over deep evolutionary time scales.     

Large-scale heterogeneity of the fossil record: implications for Phanerozoic biodiversity studies

Patterns of origination, extinction and standing diversity through time have been inferred from tallies of taxa preserved in the fossil record. This approach assumes that sampling of the fossil record is effectively uniform over time. Although recent evidence suggests that our sampling of the available rock record has indeed been very thorough and effective, there is also overwhelming evidence that the rock record available for sampling is itself distorted by major systematic biases. Data on rock outcrop area compiled for post-Palaeozoic sediments from Western Europe at stage level are presented. These show a strongly cyclical pattern corresponding to first- and second-order sequence stratigraphical depositional cycles. Standing diversity increases over time and, at the coarsest scale, is decoupled from surface outcrop area. This increasing trend can therefore be considered a real pattern. Changes in standing diversity and origination rates over time-scales measured in tens of millions of years, however, are strongly correlated with surface outcrop area. Extinction peaks conform to a random-walk model, but larger peaks occur at just two positions with respect to second-order stratigraphical sequences, towards the culmination of stacked transgressive system tracts and close to system bases, precisely the positions where taxonomic last occurrences are predicted to cluster under a random distribution model. Many of the taxonomic patterns that have been described from the fossil record conform to a species-area effect. Whether this arises primarily from sampling bias, or from changing surface area of marine shelf seas through time and its effect on biodiversity, remains problematic.