Philip D. Mannion

Testing the effect of the rock record on diversity: a multidisciplinary approach to elucidating the generic richness of sauropodomorph dinosaurs through time

The accurate reconstruction of palaeobiodiversity patterns is central to a detailed understanding of the macroevolutionary history of a group of organisms. However, there is increasing evidence that diversity patterns observed directly from the fossil record are strongly influenced by fluctuations in the quality of our sampling of the rock record; thus, any patterns we see may reflect sampling biases, rather than genuine biological signals. Previous dinosaur diversity studies have suggested that fluctuations in sauropodomorph palaeobiodiversity reflect genuine biological signals, in comparison to theropods and ornithischians whose diversity seems to be largely controlled by the rock record. Most previous diversity analyses that have attempted to take into account the effects of sampling biases have used only a single method or proxy: here we use a number of techniques in order to elucidate diversity. A global database of all known sauropodomorph body fossil occurrences (2024) was constructed. A taxic diversity curve for all valid sauropodomorph genera was extracted from this database and compared statistically with several sampling proxies (rock outcrop area and dinosaur‐bearing formations and collections), each of which captures a different aspect of fossil record sampling. Phylogenetic diversity estimates, residuals and sample‐based rarefaction (including the first attempt to capture ‘cryptic’ diversity in dinosaurs) were implemented to investigate further the effects of sampling. After ‘removal’ of biases, sauropodomorph diversity appears to be genuinely high in the Norian, Pliensbachian–Toarcian, Bathonian–Callovian and Kimmeridgian–Tithonian (with a small peak in the Aptian), whereas low diversity levels are recorded for the Oxfordian and Berriasian–Barremian, with the Jurassic/Cretaceous boundary seemingly representing a real diversity trough. Observed diversity in the remaining Triassic–Jurassic stages appears to be largely driven by sampling effort. Late Cretaceous diversity is difficult to elucidate and it is possible that this interval remains relatively under‐sampled. Despite its distortion by sampling biases, much of sauropodomorph palaeobiodiversity can be interpreted as a reflection of genuine biological signals, and fluctuations in sea level may account for some of these diversity patterns.

Geological and anthropogenic controls on the sampling of the terrestrial fossil record: a case study from the Dinosauria

Abstract Dinosaurs provide excellent opportunities to examine the impact of sampling biases on the palaeodiversity of terrestrial organisms. The stratigraphical and geographical ranges of 847 dinosaurian species are analysed for palaeodiversity patterns and compared to several sampling metrics. The observed diversity of dinosaurs, Theropoda, Sauropodomorpha and Ornithischia, are positively correlated with sampling at global and regional scales. Sampling metrics for the same region correlate with each other, suggesting that different metrics often capture the same signal. Regional sampling metrics perform well as explanations for regional diversity patterns, but correlations with global diversity are weaker. Residual diversity estimates indicate that sauropodomorphs diversified during the Late Triassic, but major increases in the diversity of theropods and ornithischians did not occur until the Early Jurassic. Diversity increased during the Jurassic, but many groups underwent extinction during the Late Jurassic or at the Jurassic/Cretaceous boundary. Although a recovery occurred during the Cretaceous, only sauropodomorphs display a long-term upward trend. The Campanian–Maastrichtian diversity ‘peak’ is largely a sampling artefact. There is little evidence for a gradualistic decrease in diversity prior to the end-Cretaceous mass extinction (except for ornithischians), and when such decreases do occur they are small relative to those experienced earlier in dinosaur evolution. Supplementary material: The full data set and details of analyses are available at www.geolsoc.org.uk/SUP18487 The same materials (in the form of an Excel workbook) are also available from the first author on request.

A quantitative analysis of environmental associations in sauropod dinosaurs

Abstract Both the body fossils and trackways of sauropod dinosaurs indicate that they inhabited a range of inland and coastal environments during their 160-Myr evolutionary history. Quantitative paleoecological analyses of a large data set of sauropod occurrences reveal a statistically significant positive association between non-titanosaurs and coastal environments, and between titanosaurs and inland environments. Similarly, “narrow-gauge” trackways are positively associated with coastal environments and “wide-gauge” trackways are associated with inland environments. The statistical support for these associations suggests that this is a genuine ecological signal: non-titanosaur sauropods preferred coastal environments such as carbonate platforms, whereas titanosaurs preferred inland environments such as fluvio-lacustrine systems. These results remain robust when the data set is time sliced and jackknifed in various ways. When the analyses are repeated using the more inclusive groupings of titanosauriforms and Macronaria, the signal is weakened or lost. These results reinforce the hypothesis that “wide-gauge” trackways were produced by titanosaurs. It is commonly assumed that the trackway and body fossil records will give different results, with the former providing a more reliable guide to the habitats occupied by extinct organisms because footprints are produced during life, whereas carcasses can be transported to different environments prior to burial. However, this view is challenged by our observation that separate body fossil and trackway data sets independently support the same conclusions regarding environmental preferences in sauropod dinosaurs. Similarly, analyzing localities and individuals independently results in the same environmental associations. We demonstrate that conclusions about environmental patterns among fossil taxa can be highly sensitive to an investigators choices regarding analytical protocols. In particular, decisions regarding the taxonomic groupings used for comparison, the time range represented by the data set, and the criteria used to identify the number of localities can all have a marked effect on conclusions regarding the existence and nature of putative environmental associations. We recommend that large data sets be explored for such associations at a variety of different taxonomic and temporal scales.

Completeness metrics and the quality of the sauropodomorph fossil record through geological and historical time

Abstract Despite increasing concerns about the effect of sampling biases on our reading of the fossil record, few studies have considered the completeness of the fossil remains themselves, and those that have tend to apply non-quantitative measures of preservation quality. Here we outline two new types of metric for quantifying the completeness of the fossil remains of taxa through time, using sauropodomorph dinosaurs as a case study. The “Skeletal Completeness Metric” divides the skeleton up into percentages based on the amount of bone for each region, whereas the “Character Completeness Metric” is based on the number of characters that can be scored for each skeletal element in phylogenetic analyses. For both metrics we calculated the completeness of the most complete individual and of the type specimen. We also calculated how well the taxon as a whole is known from its remains. We then plotted these results against both geological and historical time, and compared curves of the former with fluctuations in sauropodomorph diversity, sea level, and sedimentary rock outcrop area. Completeness through the Mesozoic shows a number of peaks and troughs; the Early Jurassic (Hettangian–Sinemurian) is the interval with highest completeness, whereas the mid-to-Late Cretaceous has completeness levels that are consistently lower than the rest of the Mesozoic. Completeness shows no relationship to rock outcrop area, but it is negatively correlated with sea level during the Jurassic–Early Cretaceous and correlated with diversity in the Cretaceous. Completeness of sauropodomorph type specimens has improved from 1830 to the present, supporting the conclusions of other recent studies. However, when this time interval is partitioned, we find no trend for an increase in completeness from the 1990s onward. Moreover, the 2000s represent one of the poorest decades in terms of average type specimen completeness. These results highlight the need for quantitative methods when assessing fossil record quality through geological time or when drawing conclusions about historical trends in the completeness of taxa. The new metrics may also prove useful as sampling proxies in diversity studies.

New information on the anatomy and systematic position of Dinheirosaurus lourinhanensis (Sauropoda: Diplodocoidea) from the Late Jurassic of Portugal, with a review of European diplodocoids

Although diplodocoid sauropods from Africa and the Americas are well known, their European record remains largely neglected. Here we redescribe Dinheirosaurus lourinhanensis from the Late Jurassic of Portugal. The holotype comprises two posterior cervical vertebrae, the dorsal series and a caudal centrum. Redescription demonstrates its validity on the basis of three autapomorphies: (1) posteriorly restricted ventral keel on posterior cervical vertebrae; (2) three small subcircular fossae posterior to the lateral coel on posterior cervical neural spines; (3) accessory lamina linking the hyposphene with base of the posterior centrodiapophyseal lamina in middle-posterior dorsal vertebrae. Phylogenetic analysis places Dinheirosaurus as the sister taxon to Supersaurus, and this clade forms the sister taxon to other diplodocines. However, this position should be treated with caution as Dinheirosaurus displays several plesiomorphic features absent in other diplodocids (including unbifurcated presacral neural spines, and dorsolaterally projecting diapophyses on dorsal vertebrae) and only four additional steps are required to place Dinheirosaurus outside of Flagellicaudata. We identify Amazonsaurus as the basal-most rebbachisaurid and recover Zapalasaurus outside of the South American Limaysaurinae, suggesting the biogeographic history of rebbachisaurids is more complex than previously proposed. Review of the European diplodocoid record reveals evidence for the earliest known diplodocid, as well as additional diplodocid remains from the Late Jurassic of Spain. A Portuguese specimen, previously referred to Dinheirosaurus, displays strong similarities to Apatosaurus from the contemporaneous Morrison Formation of North America, indicating the presence of a second Late Jurassic Portuguese diplodocid taxon. Along with Dinheirosaurus, these Portuguese remains provide further evidence for a Late Jurassic palaeobiogeographic connection between Europe and North America. No dicraeosaurids are currently known from Europe, but rebbachisaurids are present in the Early Cretaceous, with weak evidence for the earliest known representative from the Late Jurassic of Spain; however, more complete material is required to recognize early members of this clade.

A reassessment of Mongolosaurus haplodon Gilmore, 1933, a titanosaurian sauropod dinosaur from the Early Cretaceous of Inner Mongolia, People's Republic of China

The Early Cretaceous of China has yielded a diverse sauropod fauna, dominated by basal titanosauriforms. Known from a basicranium, teeth and the first three cervical vertebrae, Mongolosaurus haplodon is a poorly understood taxon whose evolutionary relationship to other sauropods has remained uncertain. Redescription of Mongolosaurus identifies key features that confirm its titanosaurian affinities and demonstrate its validity, based on several autapomorphies and a unique combination of characters. Diagnostic features include: a low neural spine which does not extend above the epipophyses; the dorsal edges of the posterior articular surfaces of anterior cervical vertebrae projecting more posteriorly than the ventral edges; and postaxial spinoprezygapophyseal laminae forming a dorsoventrally elongate X-shape in anterior view. Mongolosaurus is incorporated into three independent cladistic analyses; however, its precise phylogenetic position within Titanosauria remains unclear. This is partly a consequence of the limited material, but also reflects the unusual combination of basal and derived features for this clade and the need for a better understanding of titanosaur intra-relationships. Reassessment of this animal provides new information on the atlas-axis complex, which is extremely poorly known in titanosaurs, and highlights several new characters that may prove useful in future phylogenetic analyses, such as the ratio of the transverse widths of the basal tubera and occipital condyle. A number of features pertaining to both the cranial material and cervical vertebrae are similar to those seen in some diplodocoids (including an anteroventral projection on the atlantal intercentrum and distal bifurcation of anterior cervical neural spines), providing further evidence for convergence between these two major neosauropod radiations. Other remains previously attributed to Mongolosaurus from China and Russia cannot be referred to this genus. In summary, Mongolosaurus represents one of the earliest known titanosaurs from eastern Asia and adds to our understanding of this diverse clade of sauropods.