The apparent exponential radiation of Phanerozoic land vertebrates is an artefact of spatial sampling biases.

There is no consensus about how terrestrial biodiversity was assembled through deep time, and in particular whether it has risen exponentially over the Phanerozoic. Using a database of 60 859 fossil occurrences, we show that the spatial extent of the worldwide terrestrial tetrapod fossil record itself expands exponentially through the Phanerozoic. Changes in spatial sampling explain up to 67% of the change in known fossil species counts, and these changes are decoupled from variation in habitable land area that existed through time. Spatial sampling therefore represents a real and profound sampling bias that cannot be explained as redundancy. To address this bias, we estimate terrestrial tetrapod diversity for palaeogeographical regions of approximately equal size. We find that regional-scale diversity was constrained over timespans of tens to hundreds of millions of years, and similar patterns are recovered for major subgroups, such as dinosaurs, mammals and squamates. Although the Cretaceous/Palaeogene mass extinction catalysed an abrupt two- to three-fold increase in regional diversity 66 million years ago, no further increases occurred, and recent levels of regional diversity do not exceed those of the Palaeogene. These results parallel those recovered in analyses of local community-level richness. Taken together, our findings strongly contradict past studies that suggested unbounded diversity increases at local and regional scales over the last 100 million years.

Hwiccewyrm trispiculum gen. et sp. nov., a new leptopleuronine procolophonid from the Late Triassic of southwest England.

The fissure fill localities of southwest England and South Wales are well-known for preserving rich assemblages of predominantly small-bodied Late Triassic to Early Jurassic tetrapods, but many aspects of these assemblages remain contentious. The age of the Late Triassic fissures is disputed, with some lines of argument suggesting a latest Triassic (Rhaetian) age, whereas other evidence suggests they may be as old as Carnian. The fissures have been hypothesized by some workers to have formed on an archipelago, with island effects invoked to explain aspects of the assemblages such as the abundance of small-bodied species. Procolophonids were a successful group of Triassic parareptiles, best known from Early to early Late Triassic assemblages, but have only recently been described from one of the fissure fill sites (Ruthin) based upon fragmentary remains. Here, we describe new procolophonid specimens from another fissure (Cromhall) that represent at least six individuals of different sizes, with much of the skeleton represented including well-preserved skull material. The Cromhall procolophonid shows strong similarities to Late Triassic procolophonids from Scotland, Brazil and North America, but both autapomorphies and a unique character combination demonstrate that it represents a new species, which we name as Hwiccewyrm trispiculum gen. et sp. nov. Phylogenetic analysis places Hwiccewyrm in a derived clade within Leptopleuroninae, together with Leptopleuron, Hypsognathus, and Soturnia. The largest specimens of Hwiccewyrm demonstrate a body size that is similar to Leptopleuron and Hypsognathus, supporting other recent work that has questioned the insular dwarfism hypothesis for the fissure fill assemblages.

Diversity dynamics of Phanerozoic terrestrial tetrapods at the local-community scale.

The fossil record provides one of the strongest tests of the hypothesis that diversity within local communities is constrained over geological timescales. Constraints to diversity are particularly controversial in modern terrestrial ecosystems, yet long-term patterns are poorly understood. Here we document patterns of local richness in Phanerozoic terrestrial tetrapods using a global data set comprising 145,332 taxon occurrences from 27,531 collections. We show that the local richness of non-flying terrestrial tetrapods has risen asymptotically since their initial colonization of land, increasing at most threefold over the last 300 million years. Statistical comparisons support phase-shift models, with most increases in local richness occurring: (1) during the colonization of land by vertebrates, concluding by the late Carboniferous; and (2) across the Cretaceous/Paleogene boundary. Individual groups, such as mammals, lepidosaurs and dinosaurs also experienced early increases followed by periods of stasis often lasting tens of millions of years. Mammal local richness abruptly tripled across the Cretaceous/Paleogene boundary, but did not increase over the next 66 million years. These patterns are consistent with the hypothesis that diversity is constrained at the local-community scale.

Lepidosaurian diversity in the Mesozoic-Palaeogene: the potential roles of sampling biases and environmental drivers.

Lepidosauria is a speciose clade with a long evolutionary history, but there have been few attempts to explore its taxon richness through time. Here we estimate patterns of terrestrial lepidosaur genus diversity for the Triassic-Palaeogene (252-23 Ma), and compare observed and sampling-corrected richness curves generated using Shareholder Quorum Subsampling and classical rarefaction. Generalized least-squares regression (GLS) is used to investigate the relationships between richness, sampling and environmental proxies. We found low levels of richness from the Triassic until the Late Cretaceous (except in the Kimmeridgian-Tithonian of Europe). High richness is recovered for the Late Cretaceous of North America, which declined across the K-Pg boundary but remained relatively high throughout the Palaeogene. Richness decreased following the Eocene-Oligocene Grande Coupure in North America and Europe, but remained high in North America and very high in Europe compared to the Late Cretaceous; elsewhere data are lacking. GLS analyses indicate that sampling biases (particularly, the number of fossil collections per interval) are the best explanation for long-term face-value genus richness trends. The lepidosaur fossil record presents many problems when attempting to reconstruct past diversity, with geographical sampling biases being of particular concern, especially in the Southern Hemisphere.