Alexander M. Dunhill

Assessing the quality of the fossil record: insights from vertebrates

Abstract Assessing the quality of the fossil record is notoriously hard, and many recent attempts have used sampling proxies that can be questioned. For example, counts of geological formations and estimated outcrop areas might not be defensible as reliable sampling proxies: geological formations are units of enormously variable dimensions that depend on rock heterogeneity and fossil content (and so are not independent of the fossil record), and outcrop areas are not always proportional to rock exposure, probably a closer indicator of rock availability. It is shown that in many cases formation counts will always correlate with fossil counts, whatever the degree of sampling. It is not clear, in any case, that these proxies provide a good estimate of what is missing in the gap between the known fossil record and reality; rather they largely explore the gap between known and potential fossil records. Further, using simple, single numerical metrics to correct global-scale raw data, or to model sampling-driven patterns may be premature. There are perhaps four approaches to exploring the incompleteness of the fossil record, (1) regional-scale studies of geological completeness; (2) regional- or clade-scale studies of sampling completeness using comprehensive measures of sampling, such as numbers of localities or specimens or fossil quality; (3) phylogenetic and gap-counting methods; and (4) model-based approaches that compare sampling as one of several explanatory variables with measures of environmental change, singly and in combination. We suggest that palaeontologists, like other scientists, should accept that their data are patchy and incomplete, and use appropriate methods to deal with this issue in each analysis. All that matters is whether the data are adequate for a designated study or not. A single answer to the question of whether the fossil record is driven by macroevolution or megabias is unlikely ever to emerge because of temporal, geographical, and taxonomic variance in the data.

Disentangling rock record bias and common-cause from redundancy in the British fossil record

The fossil record documents the history of life, but the reliability of that record has often been questioned. Spatiotemporal variability in sedimentary rock volume, sampling and research effort especially frustrates global-scale diversity reconstructions. Various proposals have been made to rectify palaeodiversity estimates using proxy measures for the availability and sampling of the rock record, but the validity of these approaches remains controversial. Targeting the rich fossil record of Great Britain as a highly detailed regional exemplar, our statistical analysis shows that marine outcrop area contains a signal useful for predicting changes in diversity, collections and formations, whereas terrestrial outcrop area contains a signal useful for predicting formations. In contrast, collection and formation counts are information redundant with fossil richness, characterized by symmetric, bidirectional information flow. If this is true, the widespread use of collection and formation counts as sampling proxies to correct the raw palaeodiversity data may be unwarranted.

Problems with using rock outcrop area as a paleontological sampling proxy: rock outcrop and exposure area compared with coastal proximity, topography, land use, and lithology

Abstract Fossil specimens can be recovered easily only from exposed localities, so rock exposure area should represent a better proxy for rock availability than the frequently used outcrop (i.e., map) area. Data collected via remote sensing and GIS show that map area does not consistently correlate with exposure area in different regions. Proportional rock exposure is not geographically consistent and is influenced by a number of variables that are independent of outcrop area, including proximity to the coast, elevation, bedrock age, land use and lithology. These variables appear to be non-independent in their influence on rock exposure, and are not consistent in their effects across continents. The inconsistency in the correlation between outcrop and exposure area, and the variability in the influence of different factors on rock exposure, suggests that using outcrop area as a sampling proxy is poorly supported. The weaknesses in using outcrop area as a sampling proxy, highlighted by the lack of correlation with exposure area, suggest that a single accurate global sampling proxy may never be attained and it is premature to assume that paleodiversity curves can be corrected using such proxies. It is therefore preferable to work on a regional scale, comparing regional fossil collection data with a number of proxies representing all aspects of sampling. The lack of correlation between outcrop and exposure area suggests that the covariance detected between outcrop area and paleodiversity might be better explained by a common-cause model, and that geological megabiases may not have had as profound an effect on paleodiversity curves as previously thought.

Using remote sensing and a geographic information system to quantify rock exposure area in England and Wales: Implications for paleodiversity studies

Rock outcrop area and number of sedimentary formations have been widely used as sampling proxies in paleodiversity studies and have often been found to correlate with apparent paleodiversity. However, rock exposure area is a better proxy for the amount of sedimentary rock available for study than the widely used measures of outcrop area (i.e., map area) or number of formations. With the use of remote sensing and a geographic information system (GIS), it is possible to quantify rock exposure area accurately on a regional scale. Rock exposure area does not correlate well with either outcrop area or number of sedimentary formations, and the proportion of rock exposed in different areas can vary considerably with proximity to the coast, bedrock age, lithology, land use, and elevation. Therefore, this suggests that the correlation of paleodiversity and rock volume estimates is best explained by a common-cause hypothesis, rather than a geologic sampling bias hypothesis. GIS methods offer an efficient and accurate method of quantifying regional sedimentary rock exposure and may provide a helpful approach to assessing rock record bias on patterns in the fossil record.

Geographic range did not confer resilience to extinction in terrestrial vertebrates at the end-Triassic crisis.

Rates of extinction vary greatly through geological time, with losses particularly concentrated in mass extinctions. Species duration at other times varies greatly, but the reasons for this are unclear. Geographical range correlates with lineage duration amongst marine invertebrates, but it is less clear how far this generality extends to other groups in other habitats. It is also unclear whether a wide geographical distribution makes groups more likely to survive mass extinctions. Here we test for extinction selectivity amongst terrestrial vertebrates across the end-Triassic event. We demonstrate that terrestrial vertebrate clades with larger geographical ranges were more resilient to extinction than those with smaller ranges throughout the Triassic and Jurassic. However, this relationship weakened with increasing proximity to the end-Triassic mass extinction, breaking down altogether across the event itself. We demonstrate that these findings are not a function of sampling biases; a perennial issue in studies of this kind.

Completeness of the fossil record and the validity of sampling proxies: a case study from the Triassic of England and Wales

Many studies have highlighted correlations between palaeodiversity and sampling proxies. These correlations have been interpreted as evidence for bias, common cause, or redundancy between signals. Here, we compare a number of sampling proxies representing sedimentary rock volume, rock accessibility and worker effort with palaeodiversity through the predominantly terrestrial Triassic System in England and Wales. We find that proxies for sedimentary rock volume and accessibility do not correlate with palaeodiversity until the removal of facies-related preservational and palaeoecological factors. This indicates that a weak sampling signal may be present, but the effects of changing palaeoenvironments are far more important at the regional scale. Significant correlations between worker effort and palaeodiversity are detected, although this is likely to be a result of the preferential sampling of formations already known to be rich in fossils. The fact that there is little evidence for sedimentary rock bias in the fossil record of the Triassic of England and Wales suggests that either (1) sampling bias is not a major source of error at the regional scale or (2) sampling proxies are inadequate representations of geological and human sampling bias. Supplementary material: Correlation test results and raw time series data from the study are available at www.geolsoc.org.uk/SUP18567.

Assessing sampling of the fossil record in a geographically and stratigraphically constrained dataset: the Chalk Group of Hampshire, southern UK

Taphonomic, geological and sampling processes have been cited as biasing richness measurements in the fossil record, and sampling proxies have been widely used to assess this. However, the link between sampling and taxonomic richness is poorly understood, and there has been much debate on the equivalence and relevance of proxies. We approach this question by combining both historical and novel data: a historical fossil occurrence dataset with uniquely high spatial resolution from the Upper Cretaceous Chalk Group of Hampshire, UK, and a newly compiled 3D geological model that maps subsurface extent. The geological model provides rock volumes, and these are compared with exposure and outcrop area, sampling proxies that have often been conflated in previous studies. The extent to which exposure area (true rock availability) has changed over research time is also tested. We find a trend of low Cenomanian to high Turonian to Campanian raw richness, which correlates with, and is possibly driven by, the number of specimens found. After sampling standardization, an unexpected mid-Turonian peak diversity is recovered, and sampling-standardized genus richness is best predicted by rock volume, suggesting a species–area (or ‘genus–area’) effect. Additionally, total exposure area has changed over time, but relative exposure remains the same. Supplementary materials: A locality list, abundance matrix and all correlation and modelling results are available at https://doi.org/10.6084/m9.figshare.c.3592208.

Decoupled taxonomic and ecological recoveries from the Permo-Triassic extinction

After the enormous Permian-Triassic mass extinction, marine animals high in the food chain recovered the most quickly. The Permian-Triassic mass extinction was the worst crisis faced by life; it killed >90% of marine species in less than 0.1 million years (Ma). However, knowledge of its macroecological impact over prolonged time scales is limited. We show that marine ecosystems dominated by non-motile animals shifted to ones dominated by nektonic groups after the extinction. In Triassic oceans, animals at high trophic levels recovered faster than those at lower levels. The top-down rebuilding of marine ecosystems was still underway in the latest Triassic, ~50 Ma after the extinction, and contrasts with the ~5-Ma recovery required for taxonomic diversity. The decoupling between taxonomic and ecological recoveries suggests that a process of vacant niche filling before reaching the maximum environmental carrying capacity is independent of ecosystem structure building.

Modelling determinants of extinction across two Mesozoic hyperthermal events

The Late Triassic and Early Toarcian extinction events are both associated with greenhouse warming events triggered by massive volcanism. These Mesozoic hyperthermals were responsible for the mass extinction of marine organisms and resulted in significant ecological upheaval. It has, however, been suggested that these events merely involved intensification of background extinction rates rather than significant shifts in the macroevolutionary regime and extinction selectivity. Here, we apply a multivariate modelling approach to a vast global database of marine organisms to test whether extinction selectivity varied through the Late Triassic and Early Jurassic. We show that these hyperthermals do represent shifts in the macroevolutionary regime and record different extinction selectivity compared to background intervals of the Late Triassic and Early Jurassic. The Late Triassic mass extinction represents a more profound change in selectivity than the Early Toarcian extinction but both events show a common pattern of selecting against pelagic predators and benthic photosymbiotic and suspension-feeding organisms, suggesting that these groups of organisms may be particularly vulnerable during episodes of global warming. In particular, the Late Triassic extinction represents a macroevolutionary regime change that is characterized by (i) the change in extinction selectivity between Triassic background intervals and the extinction event itself; and (ii) the differences in extinction selectivity between the Late Triassic and Early Jurassic as a whole.