Basturs Poble bonebed

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Hadrosaur tibiae and fibulae from the Basturs Poble site Basturs poble tibiae.png
Hadrosaur tibiae and fibulae from the Basturs Poble site

The Basturs Poble bonebed is a mega-bonebed of hadrosaur dinosaur fossils, discovered in Catalonia, Spain. Hundreds of hadrosaur fossils have been found at the site, which would have been on a large island during the Late Cretaceous when the animals preserved were alive. Despite the enormous amount of specimens, taxonomically informative material has been scarce at the site, leading to extensive debate as to its nature. The number of species present, age of the individuals present in the sample, and taxonomic identity of the remains have been the primary matters of debate. Previous considered to represent Koutalisaurus , Pararhabdodon , or multiple, perhaps dwarf species, it is currently thought that a single, indeterminate species of lambeosaurine was present at the site, and that individuals of many different ages were present.

Contents

Discovery and extent

Map of the stratigraphic, geographic, and geologic context of the Basturs Poble bonebed locality Basturs poble context.png
Map of the stratigraphic, geographic, and geologic context of the Basturs Poble bonebed locality

Marc Boada discovered, in the late 1990s, a new fossil-bearing locality 300 metres (980 ft) from the village of Basturs, Catalonia. As the village already lent its name to a locality where dinosaur eggs had been found, it was named the Basturs Poble (BP) locality, "poble" derived from the word for village. The locality belongs to the Conques Formation of the Tremp Group, dated to the late-Early Maastrictian. [1] Over ten years, from 2001 through 2011, field work was done at this site, and a massive bonebed - consisting of some one thousand skeletal elements - was discovered. [1] [2] The discovery was first reported in the literature in an abstract at the "55th Symposium for Vertebrate Palaeontology and Comparative Anatomy and the 16th meeting of the Symposium for Palaeontological Preparation and Conservation" conference in 2007. The authors, then, approached the bonebed under the hypothesis of a single species being present at the site. It was reported that all ontogenetic stages were present. [3]

The stratigraphic layer where the bonebed was found is 1.5 metres (4.9 ft) in thickness. Around 95% of all remains from the site belong to dinosaurs; nearly all identified specimens from this sample belong to hadrosaurs. Of prepared material, 270 skeletal elements can confidently be identified as belonging to hadrosauroids. [1] [2] The number of hadrosaur specimens from the site in total could number over 500, and possibly as much as 900. [2] [4] In addition to the large numbers of fossils of the animals themselves, the BP locality is one of many sites from the Tremp Syncline where hadrosaurs tracks have been found. At least one track is preserved at the site, in a sandstone block from a stratigraphic level similar to that of the bonebed. [2] [5] It is the richest hadrosaur bonebed ever found in Europe, and the largest ever found in one palaeoinsular locale; as such, it is the most important hadrosauroid site of the eastern Tremp Syncline. [1] [2]

Number of species present

A selection of dentary material from the Basturs Poble bonebed Basturs poble dentaries.png
A selection of dentary material from the Basturs Poble bonebed

The primary subject of research regarding the bonebed has been the taxonomic identity of the hadrosaurs there - namely, whether only one taxon or multiple taxa are present. The question has been complicated by a rarity of cranial elements in the sample, since they are the most taxonomically informative areas of the skeleton in hadrosaurs. [1] Initially, it was assumed that only a single species was present in the sample. [3] In 2015, a study by Alejandro Blanco and colleagues performed multiple types of morphometric analysis to investigate hadrosaur diversity in from the Pyrenees using dentaries. The number of alveolar positions in the dentary was the primary metric for distinctiveness, as more basal taxa possess less, though it was noted this can also increase with age as an individual grows. Three morphotypes (groupings of specimens with a distinct anatomy from the other groupings) were found in the sample, of which two (numbers two and three) were present in the BP bonebed. Growth trajectory analysis of the different morphotypes supported their separation. Sexual dimorphism being the reason for different anatomy was considered unlikely due to different results in regression analysis. Morphotype three was interpreted as belonging to dwarf, potentially relictual hadrosauroids; and morphotype two was considered to represent larger, lambeosaurine animals. It was stressed each morphotype was not necessarily just one species, but merely representative of a distinct lineage of hadrosaurs. [4]

In 2018, a more extensive study of the material was conducted by Víctor Fondevilla and colleagues, and the question of how many taxa were present of the sample was investigated using specimens from multiple parts of the body, as opposed to just the dentaries. Overall, a large amount of variation was noted in the specimens, but as it was gradual, rather than a case of two obvious morphotypes, the authors attributed it to individual variation. Regarding the dentaries, they too were not found to obviously sort into multiple morphotypes due to a high amount of variation between all of them. One particular dentary was noted to be rather distinct from the rest, but taphonomic damage or improper restoration during preparation of the fossil were considered possible explanations as opposed to taxonomic distinctiveness. The authors favored identification of the hadrosaurs present in the bonebed as a singular taxon of lambeosaurine. [1]

Size and age of the individuals

Growth trajectories of several tibiae, taken from a 2018 study of the taxonomy and ontogeny of the specimens Basturs poble growth curve.png
Growth trajectories of several tibiae, taken from a 2018 study of the taxonomy and ontogeny of the specimens

An intertwined subject to the taxonomy which has been noted and studied has been the size and ontogeny (growth and development) of the individuals represented. Individuals of multiple sizes are represented in the sample, but overall they represent smaller animals than the hadrosaurs known from Asia and North America. Despite this, a small number of individuals falling into the adult size ranges of other hadrosaurs are indicated by isolated elements, such as femurs. [1] [2] Multiple hypotheses have been put forward to explain this: firstly, juvenile individuals may have been the dominant life stage living in the area where the bonebed was preserved, due to ecological differences over lifespan; secondly, it's possible they represent dwarfed animals dominating the environment relative to rare a larger species; lastly, it's possible the large individuals were comparative giants of a single dwarfed species, near the extreme end of individual variation in the species. [2] The 2015 analysis of hadrosaur dentaries from the Pyrenees found two morphotypes to be present in the bonebed, so they favored the second hypothesis. [4]

The more in-depth 2018 study used histological analysis in order to assess the age of different individuals, using tibiae. Of the several tibiae tested, four (between 390–450 millimetres (15–18 in) long) were identified as juveniles between the ages of one and three based on comparisons to tissue patterns in other ornithopods, and three more were found to likely be juveniles based on similarities to the former four. The eighth (550 millimetres (22 in) long) and ninth (720 millimetres (28 in) long) were found to belong to an early adult and subadult individual respectively. A 720 millimetres (28 in) femur was found to share subadult growth features with the subadult tibia. Three tibiae (550 millimetres (22 in), 600 millimetres (24 in), and around 900 millimetres (35 in) respectively) were found to belong to adult individuals. Growth trajectory was found to be consistent between the tibiae, supporting the idea that the site represents a large variety of age stages of a single species. [1]

Taxonomic identity

Holotype dentary of "Koutalisaurus kohlerorum", a species the bonebed material was once assigned to Koutalisaurus.jpg
Holotype dentary of "Koutalisaurus kohlerorum", a species the bonebed material was once assigned to

A poster presentation and abstract at a conference suggested the specimens may have belonged to Koutalisaurus kohlerorum , but they cautioned an adult dentary from the bonebed would need to be discovered to test the hypothesis. [6] This was later rejected, as Koutalisaurus was found to be an invalid nomen dubium. [7] Fondevilla et al. (2018) compared to the material from Basturs Poble to recognized species of lambeosaurine from geographically and temporally similar locations in the Pyrenees. The jugals from the sample were found to be distinct from those known from Blasisaurus and Arenysaurus (possible synonyms of Pararhabdodon ), and the frontals were also distinguishable from the latter. Identity as specimens as either of those taxa was therefore ruled out. Overlapping taxonomically informative material with Pararhabdodon was not found from the bonebed, excepting a maxilla too fragmentary for use in referral. However, both the type locality of SRA and the locality of the referred maxilla MCD 4919 are close to the area the bonebed was found. As such, it was considered most likely the Basturs Poble hadrosaurs are members of the species P. isonensis, but the poor state of preservation of the only comparable material in the bonebed prevented a secure referral. [1] Later, a 2019 study by Prieto-Márquez et al. described a new genus and species of Spanish lambeosaurine, Adynomosaurus arcanus . They compared the material of their species to that of the bonebed, and though similarities were found, their tooth crowns and ilia had different shapes, and the scapulae from the site lacked the distinct shallow nature that characterizes A. arcanus. Accordingly, they considered it unlikely the bonebed belonged to their species. Additionally, they commented on the lack of tsintaosaurin synapomorphies in the bonebed material and so kept a referral as indeterminate lambeosaurines rather than considering it to represent Pararhabdodon. [8]

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<i>Aralosaurus</i> Extinct genus of dinosaurs

Aralosaurus was a genus of hadrosaurid dinosaur that lived during the Late Cretaceous in what is now Kazakhstan. It is known only by a posterior half of a skull and some post-cranial bones found in the Bostobe Formation in rocks dated from the Upper Santonian-Lower Campanian boundary, at about 83.6 Ma. Only one species is known, Aralosaurus tuberiferus, described by Anatoly Konstantinovich Rozhdestvensky in 1968. The genus name means Aral Sea lizard, because it was found to the northeast of the Aral Sea. The specific epithet tuberiferus means bearing a tuber because the posterior part of the nasal bone rises sharply in front of the orbits like an outgrowth. Aralosaurus was originally reconstituted with a nasal arch similar to that of North American Kritosaurus. For many years, Aralosaurus was thus placed in the clade of the Hadrosaurinae. This classification was invalidated in 2004, following the re-examination of the skull of the animal which allowed to identify in Aralosaurus many typical characters of Lambeosaurinae. In particular, this study revealed that Aralosaurus had a hollow bony structure located far in front of the orbits, which communicated with the respiratory tract. This structure being broken at its base, its shape and size remains undetermined. More recently, Aralosaurus has been identified as the most basal Lambeosaurinae, and placed with its close relative Canardia from the upper Maastrichtian of France in the new clade of Aralosaurini.

<i>Eolambia</i> Extinct genus of dinosaurs

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<i>Tsintaosaurus</i> Hadrosaurid ornithopod dinosaur from Late Cretaceous period

Tsintaosaurus is a genus of hadrosaurid dinosaur from China. It was about 8.3 metres (27 ft) long and weighed 2.5 tonnes. The type species is Tsintaosaurus spinorhinus, first described by Chinese paleontologist C. C. Young in 1958.

<i>Pararhabdodon</i> Extinct genus of dinosaurs

Pararhabdodon is a genus of tsintaosaurin hadrosaurid dinosaur, from the Maastrichtian-age Upper Cretaceous Tremp Group of Spain. The first remains were discovered from the Sant Romà d’Abella fossil locality and assigned to the genus Rhabdodon, and later named as the distinct species Pararhabdodon isonensis in 1993. Known material includes assorted postcranial remains, mostly vertebrae, as well as maxillae from the skull. Specimens from other sites, including remains from France, a maxilla previously considered the distinct taxon Koutalisaurus kohlerorum, an additional maxilla from another locality, the material assigned to the genera Blasisaurus and Arenysaurus, and the extensive Basturs Poble bonebed have been considered at different times to belong to the species, but all of these assignments have more recently been questioned. It was one of the last non-avian dinosaurs known from the fossil record that went extinct during the Cretaceous-Paleogene extinction event.

<i>Koutalisaurus</i> Extinct genus of dinosaurs

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<span class="mw-page-title-main">Tremp Formation</span>

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<i>Blasisaurus</i> Extinct genus of dinosaurs

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<span class="mw-page-title-main">Tsintaosaurini</span> Extinct tribe of dinosaurs

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<span class="mw-page-title-main">Timeline of hadrosaur research</span>

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<i>Adynomosaurus</i> Extinct genus of dinosaurs

Adynomosaurus is a genus of lambeosaurine dinosaur from the Late Cretaceous of what is now Catalonia, Spain. First discovered in 2012, it was named in 2019 with the type and only species being Adynomosaurus arcanus. It is only known from scant material, but is distinguished from other hadrosaurs by its weakly developed shoulder blade which would have had underdeveloped musculature, which lends it its scientific name, partially from the Greek word for "weak". Its exact relationships with other hadrosaurs remain unresolved, with it not consistently being recovered as a relative of any other specific genera, though some studies have allied it with Tsintaosaurini or even found it outside of Hadrosauridae. It would have lived as part of a diverse coastal estuary ecosystem, made up of meandering rivers and mud flats. The discovery of Adynomosaurus adds to the very incomplete fossil record of hadrosaurid dinosaurs in the Late Cretaceous of Europe, and it fits into a picture of major ecological turnover that was occurring during the Maastrichtian stage in the region.

<span class="mw-page-title-main">Arenysaurini</span> Extinct tribe of dinosaurs

Arenysaurini is a proposed tribe of primitive lambeosaurine hadrosaurs. It is composed of genera found in Europe and North Africa during the end of the Cretaceous period, and has been suggested to unite all lambeosaurs from the former continent into a singular monophyletic group.

<i>Tamarro</i> Genus of troodontid dinosaur from the Late Cretaceous period

Tamarro is a genus of troodontid theropod from the Late Cretaceous Talarn Formation of Spain. The genus contains a single species, Tamarro insperatus, known from a partial metatarsal described in 2021.

<i>Fylax</i> Genus of hadrosauroid dinosaur from the Late Cretaceous period

Fylax is a genus of hadrosauroid ornithopod from the Late Cretaceous Figuerola Formation of Spain. The genus contains a single species, Fylax thyrakolasus, known from a nearly complete left dentary.

References

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